Neotropical Simuliidae: Diptera, Insecta (Biodiversidad Aquatica En America Latina)

Aquatic Biodiversity in Latin America Biodiversidad Acuática en América Latina

ABLA Vol. 3

STUDENTS FROM NUMEROUS COUNTRIES HAVE COMPLAINED ABOUT THE LACK OF “TOOLS” TO IDENTIFY AQUATIC INVERTEBRATES FROM LATIN AMERICA. KEYS FOUND IN ACCEPTED TEXTBOOKS ARE MOSTLY LIMITED, SUPERFICIAL OR TOO POPULAR-SCIENCE. ON THE OTHER HAND, MORE ADVAN-

Volume 3 This book is unique in coverage, summarizing all available information concerning the American simuliid fauna south of the United States. It also includes morphological diagnoses of females, males, pupae and larvae, as well as keys (in English and Spanish), illustrations of characters, mapped distributions, and bionomics. This region appears to support 359 species grouped in two tribes, 12 genera and 18 subgenera. The description of each taxon is provided with a list of the available literature, as well as all other relevant information. The book is addressed to taxonomists, limnologists, ecologists, veterinarians and biologists in general.

Sixto Coscarón & Cecilia L. Coscarón Arias

CED KEYS IN ACADEMIC BOOKS ON AQUATIC INVERTEBRATES FAIL TO COVER NEOTROPICAL REPRESENTATIVES IN DUE DETAIL. THE FEW THAT DO, HOWEVER, ALSO SHOW TOO MANY DEFECTS IN SCOPE AND PRESENTATION TO BE CONSIDERED UP-TO-DATE. AN INTERNATIONAL TEAM OF EDITORS COMBINED THEIR EFFORTS WITH PENSOFT PUBLISHERS TO LAUNCH A NEW MAJOR SERIES ON THE AQUATIC BIODIVERSITY IN LATIN AMERICA. ABOUT 15 MONOGRAPHS, WRITTEN BY SCIENTISTS FROM VARIOUS COUNTRIES, WILL OFFER A NEW, UNRIVALLED VIEW OF THE MARVELLOUS AQUATIC WORLD OF SOUTH AMERICA! THE SERIES IS ADDRESSED TO ZOOLOGISTS, ECOLOGISTS, HYDROBIOLOGISTS, BIOGEOGRAPHERS, CONSERVATIONISTS AND STUDENTS INTERESTED IN AQUATIC BIODIVERSITY. THE SERIES WILL BE AN IMMINENT TOOL FOR ANY BIOLOGICAL LIBRARY.

Neotropical Simuliidae (Diptera: Insecta)

FOR MANY YEARS,

ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales

Aquatic Biodiversity in Latin America Biodiversidad Acuática en América Latina

Volume 3

Neotropical Simuliidae (Diptera: Insecta) Volumen 3

Simuliidae Neotropicales (Diptera: Insecta)

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Volumen 3

SIMULIIDAE NEOTROPICALES (DIPTERA: INSECTA) por

Sixto Coscarón & Cecilia L. Coscarón Arias Lector Científico a los Editores: Peter H. Adler Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2007

ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales

AQUATIC BIODIVERSITY IN LATIN AMERICA Volume 3

NEOTROPICAL SIMULIIDAE (DIPTERA: INSECTA) by

Sixto Coscarón & Cecilia L. Coscarón Arias Scientific Reader to the Editors: Peter H. Adler Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2007

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

AQUATIC BIODIVERSITY

IN

BIODIVERSIDAD ACUÁTICA

LATIN AMERICA

EN

AMÉRICA LATINA

Series Editors/Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen VOLUME 3. NEOTROPICAL SIMULIIDAE (DIPTERA: INSECTA) by Sixto Coscarón & Cecilia L. Coscarón Arias Scientific Reader to the Editors: Peter H. Adler

VOLUMEN 3. SIMULIIDAE NEOTROPICALES (DIPTERA: INSECTA) por Sixto Coscarón & Cecilia L. Coscarón Arias Lector Científico a los Editores: Peter H. Adler

Front cover: Male of the newly described Simulium nunesdemelloi HAMADA, PEPINELLI & HERNÁNDEZ, 2006 (Simuliidae) and Amazon River near Manaus, Brazil (photos: M. Cutrim, W.J. Junk; design: Zheko Aleksiev & Elke Bustorf) Aquatic Biodiversity in Latin America Vol. 3 ISSN 1312-7276 First published 2007 ISBN: 978-954-642-293-4

The book should be cited as follows: Sixto Coscarón & Cecilia L. Coscarón Arias. 2007. Neotropical Simuliidae (Diptera: Insecta). In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.): Aquatic Biodiversity in Latin America (ABLA). Vol. 3. Pensoft, Sofia-Moscow, 685 pp. © PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria [email protected] www.pensoft.net Printed in Bulgaria, March 2007

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AQUATIC BIODIVERSITY IN LATIN AMERICA BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen Biodiversity is a key word in science and global management schemes; however very few people are able to identify the species and their ecology that make up “biodiversity”. For many years, researchers and students from numerous countries complain about the lack of “tools” to identify aquatic invertebrates from Latin America. Keys found in accepted entomological textbooks are mostly highly limited, superficial and rarely cover Neotropical biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology is scattered throughout the literature in many single publications. An international team of editors have combined their efforts with Pensoft Publishers to launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Their goal was to find experts who combine the current state of knowledge in taxonomy and ecology, in order to produce a concise and affordable handbook for each group. About 15 separate monographs, written by reference scientists from various countries will offer a new, unrivalled view on the aquatic fauna of South America. Information on the ecology and status of the taxa (written in English) is combined with illustrated identification keys to families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists, hydrobiologists, biogeographers, conservationists and students interested in aquatic biodiversity. The series will be an essential tool for any biological library. Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and photos (15 plates in color); publication date: March 2006. Volume 2: Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-259-2, ISBN-13: 978-954-642259-0, 165x240 mm, 646 pp., including 234 plates of figures in line drawings and photos (16 plates in color); publication date: May 2006.

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarón & Cecilia L. Coscarón Arias. Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-293-4, 165x240 mm, 685 pp., including 135 plates of figures in line drawings and 19 maps of distribution; publication date: March 2007. Simuliid larvae and pupae inhabit freshwater streams, from sea level to about 5000 m altitude in the Neotropics. The larvae are filter feeders and, along with pupae, live attached to different underwater strata for support. These insects play an important role in the bioenergetic cycle of continental waters. Imagoes are strong flyers, and females are well known for their biting habits as well public health importance, being vectors of several pathogens. This book compiles the current known information on the American simuliid fauna south of the United States. It also includes a morphological diagnosis of females, males, pupae and larvae, along with keys, illustrations of characters, distributions with maps, and bionomics for the various taxa. In this region, there are 359 known species, grouped in two tribes, 12 genera and 18 subgenera. The description of each taxon includes a list of the available literature as well as the most relevant information. The book is addressed to systematists, limnologists, ecologists, veterinarians and biologists in general.

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ADDRESSES Authors: DR. SIXTO COSCARÓN Museo de Ciencias Naturales de La Plata Paseo del Bosque S/N 1900 La Plata, Argentina e-mail: [email protected] DR. CECILIA L. COSCARÓN ARIAS LIBIQUIMA (Laborartorio de Investigaciones Bioquímicas y de Medio Ambiente) Universidad Nacional del Comahue Buenos Aires 1400, 8300 Neuquen, Argentina e-mail: [email protected] Series Editors: PROF. DR. JOACHIM ADIS Max-Planck-Institute for Limnology Tropical Ecology Working Group Postfach 165 24302 Plön, Germany e-mail: [email protected] DR. JORGE R. ARIAS 5870 Colfax Avenue Alexandria, Virginia 22311, USA e-mail: [email protected]

M.Sc. GUILLERMO RUEDA-DELGADO Grupo de Investigación en Cuencas y Humedales Tropicales GICHT UNIMAGADALENA Universidad de Bogotá Jorge Tadeo Lozano Laboratorio de Limnología Carrera 4 No. 22-61 Bogotá, D.C. Colombia e-mail: [email protected] DR. KARL MATTHIAS WANTZEN Institute of Limnology University of Konstanz Postfach M 659 78457 Konstanz, Germany e-mail: [email protected] Scientific Reader to the Editors: PROF. DR. PETER H. ADLER Division of Entomology Clemson University P.O. Box 340315, 114 Long Hall Clemson, SC 29634-0315, USA e-mail: [email protected]

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

FOREWORD Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarón & Cecilia L. Coscarón Arias Research on the Simuliidae in the Neotropical area has been ongoing for over a century. It largely consisted of sporadic accounts of species by various authors with little coordination of effort until the 1930s. Enderlein then produced many names of new species and genera and a classification, which through various convoluted pathways helped to form the taxonomic framework in use today. The driving force behind the subsequent increase in taxonomic studies has been largely medical. Simuliid flies in Latin America are responsible for two problems. Firstly, the presence of enormous biting populations in certain areas that can preclude development of land for agriculture and deter tourism. Secondly, the incrimination of several species in transmitting two filarial worms to man, Mansonella ozzardi and Onchocerca volvulus, responsible for the diseases mansonelliasis and onchocerciasis. The major impetus to these studies was the need to incriminate the vectors of human onchocerciasis in the 1940s in Guatemala and Mexico. This led to the pioneering work on vector incrimination and simuliid taxonomy by Luis Vargas and his colleague Alfonso Díaz Nájera in Mexico. These authors provided a good taxonomic foundation for the Central American fauna. In the 1950s Rachou’s work in the Amazon basin of Brazil on mansonelliasis, a disease already suspected for some time to be associated with Simuliidae in Argentina, inspired continuing studies on Simuliidae of the region. At about this time Pedro Wygodzinsky started work on the family in Argentina, soon to be joined by Sixto Coscarón. Their classic work on the family, with methodical collection of species and further development of the taxonomic framework culminated in Coscarón’s overview of the Neotropical Simuliidae in 1987. This publication, in conjunction with Crosskey’s world inventory of Simuliidae produced at about the same time, has had monumental significance in Latin America. For the first time simuliid researchers in Latin America were able to access information on valid names for simuliid species, their classification and references that supply data on simuliid biology and medical importance. This latest Neotropical overview of Coscarón & Coscarón-Arias brings us up to date with the considerable advances emanating from the discovery of human onchocerciasis in Amazonia in the 1970s. Its publication in English now provides ready access for simuliid specialists at an international level. This new book has brought together the relevant voluminous literature in Portuguese, Spanish, French and English on simuliids from a vast area of the planet. It provides a systematic analysis that largely corresponds to the scheme in Crosskey’s world annotated catalogue. Main characters of species are covered, with ample references to descriptions by these and other authors and very comprehensive illustrations and species distribution maps. Methods used in collecting and preparing specimens for

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subsequent examination and the terminology used in their description are described. Synopses of the biology, medical importance and control of simuliids in Latin America are also provided. The authors regard this work as a “snapshot view” of the complicated Neotropical simuliid fauna for which much taxonomic study is still necessary. It provides a sound basis for future in depth morphological studies of certain species, where warranted, on economic or medical grounds, and for studies integrating morphological, cytological and molecular methods for phylogenetic purposes. TONY SHELLEY The Natural History Museum London, U.K.

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

PREFACE Simuliidae are present in most freshwater streams where the larval stages develop and the adults are well known to the nearby human population by the nuisance of the female’s bite. The immature stages are present in currents of Neotropical creeks, streams and rivers, in slow to torrential waters. During the larval and pupal stages, they live attached to various substrates in waters of variable temperatures, salinity and turbidity, showing great adaptability to different habitats. Larvae feed on algae, bacteria, organic particles of dead organisms and inorganic material, showing a great capacity to metabolize different materials. This aspect of their life history, added to the fact that they serve as food for fishes, birds and other arthropods, gives an idea of the important role they play in the energy cycle of continental freshwater ecosystems. Adults are strong fliers with high dispersal qualities, and they have variable feeding habits, ranging from plant nectar to vertebrate blood. This great adaptability is represented by about 350 known species in the Neotropical region. Certainly this number will increase since many areas are unstudied or have been scarcely studied. This volume is a compilation of the known Neotropical black flies found from south of the United States to the south of Tierra del Fuego, Argentina. The biogeography does not coincide with political limits, but in this case it approximates the extension of this region. The aim of this volume is to make a compilation of the recognizable species, arranged in supraspecific taxa and arranged in accordance with the relationships obtained by cladistic studies. Here we consider l2 genera and 284 species with keys, diagnoses, illustrations of morphology and distribution maps. We do not include 65 species of Gigantodax and Pedrowygomyia, because they were treated in a previous review (WYGODZINSKY & COSCARÓN, l989). An introductory section on general morphology is included to understand the anatomical names used in the descriptions. Also, we add information on biotic conditions related to the species, as well as their public health importance and control. The present volume was planned to help less experienced people that desire to investigate this interesting family of Diptera. In order to address the Latin American audience, keys and illustration legends are also offered in Spanish. Carmencita (Carmen ENRIQUETA ANNUASI) is heartly thanked for her great patience and continuous support. We are very grateful to Nélida CALIGARIS for her help in typing and arranging most of the illustrations, which are based on her illustrations made during many years of our publications. Also, we are indebted to ALICIA GARCIA for her help in typing part of this manuscript. We are indebted to all the Neotropical simuliidologists that have provided information from their publications used here or have provided study material, namely Victor PY-DANIEL, Anthony SHELLLEY, Sergio IBÁÑEZ-BERNAL, Marilza MAIA-HERZOG, Milton STRIEDER, Neusa HAMADA, Gilson MOREIRA, Paulino LUNA-DIAS, Peter ADLER, Leonardo Gil AZEVEDO, Marcia

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COSTA, Jaime RAMÍREZ PÉREZ, Paulina MUÑOZ DE HOYOS, and specially to Rafael MIRANDA ESQUIVEL for the cladistic studies and his preoccupation to solve the phylogeny of the Simuliidae. We are also grateful to several institutions that have provided study material, including the American Museum of Natural History (New York), the United States National Museum, (Smithsonian Institution, Washington, D.C.), The Natural History Museum ( London), the Museum für Naturkunde (Berlin), INDRE, (Mexico) and the Museu de Zoologia (São Paulo). In the first institution we were also allowed to use information from articles published in the Bulletin and Novitates. Our special acknowledgment goes to Zootaxa for permission to use information published in volume 396 (2004), and to the Memoirs on Entomology International for the information extracted from volume 14 (1999). The authors are much indebted to CONICET for financial support. We thank the editors and the scientific reader of the book for their valuable help received while preparing the manuscript for publication. We extend special acknowledgment to our late master, Pedro WYGODZISNKY, who gave all his knowledge and enthusiasm to improve the study of black flies with modern methods in morphology and phylogenetic analysis, and especially for encouraging us to explore the interesting Simuliidae world. THE AUTHORS

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Table of Contents 1I

NTRODUCTION

Bionomics Behavior Public health and socioeconomic importance Control

2M 3C 4S 5 S Keys to the genera of Neotropical Simuliidae

ORPHOLOGY AND CHARACTERS OF TAXONOMIC VALUE

15 15 16 17 18 22

OLLECTION AND PREPARATION OF MATERIAL

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TUDY OF MATERIAL

36

YSTEMATICS

Claves de géneros de simúlidos de la región Neotropical Mayacnephia WYGODZINSKY & COSCARÓN Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA Araucnephia WYGODZINSKY & COSCARÓN Araucnephiodes WYGODZINSKY & COSCARÓN Cnesia ENDERLEIN Gigantodax ENDERLEIN Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL Cnesiamima WYGODZINSKY & COSCARÓN Paraustrosimulium WYGODZINSKY & COSCARÓN Lutzsimulium D’ANDRETTA & D’ANDRETTA Kempfsimulium PY-DANIEL & NUNES DE MELLO

38 39 45 52 63 66 76 81 90 91 93 99 102 110

ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales

Simulium LATREILLE Keys to Simulium subgenera Claves de subgéneros de Simulium Simulium (Eusimulium) ROUBAUD Simulium (Nevermannia) ENDERLEIN Simulium (Byssodon) ENDERLEIN Simulium (Psilozia) ENDERLEIN Simulium (Pternaspatha) ENDERLEIN NIGRISTRIGATUM species group NEMORALE species group Simulium (Notolepria) ENDERLEIN Simulium (Chirostilbia) ENDERLEIN PERTINAX species group SUBPALLIDUM species group Simulium (Inaequalium) COSCARÓN & WYGODZINSKY INAEQUALE species group BOTULIBRANCHIUM species group Simulium (Psaroniocompsa) ENDERLEIN INCRUSTATUM species group SIOLII species group Simulium (Cerqueirellum) PY-DANIEL Simulium (Coscaroniellum) PY-DANIEL QUADRIFIDUM species group QUADRIVITTATUM species group Simulium (Ectemnaspis) ENDERLEIN BICOLORATUM species group ROMANAI species group PERFLAVUM species group DINELLII species group Simulium (Psilopelmia) ENDERLEIN ESCOMELI species group BLANCASI species group Simulium (Aspathia) ENDERLEIN

114 115 125 137 137 138 138 139 159 188 224 238 247 259 266 273 289 295 302 319 325 349 354 364 366 385 403 409 414 419 437 463 471

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Simulium (Hemicnetha) ENDERLEIN PAYNEI species group BRACHYCLADUM species group MEXICANUM species group OVIEDOI species group Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA JUAREZI species group CAROLINAE species group Simulium (Trichodagmia) ENDERLEIN Simulium (Thyrsopelma) ENDERLEIN

6M 7R 8S 9A

APS OF

DISTRIBUTION

EFERENCES

517 531 540 547 556 561 571 586 597 612 627 648

UBJECT INDEX

671

DDENDUM

682

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1 INTRODUCTION Bionomics Black flies can be found in highly variable environments from very cold water (about 0 °C) to warm water exceeding 25 °C, from sea level to an altitude of 5000 m, from freshwater to water with a certain degree of salinity, from clear to turbid water, and from oxygen-saturated to oxygen-poor water. The only distinctive condition that is necessary for their development is continuously running water. With this extensive range of possibilities, they can be found practically everywhere flowing water exists, from tiny streams to large rivers (COSCARÓN ARIAS, 2001). Larval black flies inhabit aquatic environments, constituting an important link in the trophic chain of lotic biotopes. Generally, they choose places with continuous and fastflowing water. They are found near the surface where higher oxygen tension exists and leaves, branches, stones or plastic substrates, to which they attach, are present and are free of algae and fungus. Physical and chemical factors regulate the presence, richness and abundance in the micro- and macrodistribution of the species (COSCARÓN ARIAS, 2001). The more important factors are the following: • Turbidity: Larvae prefer clear water but they sometimes tolerate water with some degree of turbidity during certain periods or permanently. Also, they require water that receives direct light. Watercourses that run long distances through poorly lighted areas usually have no larvae or pupae. • Salinity: They live in oligohaline waters, which have a solid residue lower than 5 g/l. Most of them live in waters with less than 1 g/l, but others, such as Simulium wolffhuegelii, can breed with a concentration close to 4 g/l. • pH: The habitat with a value near neutral is typical; however, some species can live in water with pH values above or below neutral, ranging from 6.56 to 8.90. • Oxygen: The high content of dissolved oxygen in micro-bubbles that elevate the O2 tension to saturation in torrential waters is a factor that benefits most black flies. Most of the dissolved oxygen comes from the photosynthetic process that results in sunny places. • Temperature: The larvae are eurythermic and they can live in water with temperatures close to 0 °C, such as the species found in the Puna (Argentina), high mountain ranges or in Tierra del Fuego (Argentina) where they are found in streams covered by snow. They can also be found in watercourses with temperatures of 25 °C such as those in

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

tropical areas. The great majority of species prefer water temperatures ranging from 7 °C to 15 °C that are more frequently found in springs. Altitude: They can be found from sea level up to 4700 m. The altitude is correlated with temperature. For example, in Argentina, the highest percentage of species collected (41%) is found between 500 and 1000 m. This is followed by those species that are found between sea level and 500 m. Support strata: The larvae use a large variety of substrates to adhere to, but they always require a solid support. They avoid places with many unicellular algae. The most frequent substrates used are stones, leaves, sticks, roots, branches or tree trunks; even though they rarely adhere to dead vegetation, 53% of the Argentinean species adhere to vegetation. Some species have a predilection for a particular type of substrate but there are others that are indifferent and use any kind of support, including pieces of iron, plastic, and wood. Depth: The majority of larvae and pupae live on trailing vegetation in water that is not deeper than 50 cm. Species are only observed in deeper water when there is no turbidity. Current speed: Simuliid larvae and pupae are positively reophilic. Since they filter the water to obtain food and dissolved oxygen, the greater the volume of water they filter, the more beneficial it is to them. Current speed is an important factor to define their habitat because they need to maintain attachment to the substrate, keep their cephalic fans erect and allow their muscles to contract when folding the fans to take the food to the mouth. The species that live in the plains will survive in water with minimum speeds of around 2 cm/s; however, there are species that can live in water with a lesser speed. Most of the species prefer water speeds between 10 and 50 cm/s, although there are some species that only live in torrential courses of 100 cm/s.

More information about the environmental factors in relation to the different species of simuliids of the Neotropics can be found in COSCARÓN et al. (1996b; 2000), COSCARÓN & COSCARÓN ARIAS (2000 b), GRILLET & BARRERA (1997), HAMADA & MCCREADI (1999), MOREIRA et al. (1994). Behavior Simuliids are primarily diurnal insects. Both sexes feed on nectar from plants or flowers; only the females of some species are haematophagous. Although some species can be autogenous (i.e., they don’t suck blood), others require blood of vertebrates to lay viable eggs. Depending on their haematophagic preferences, they can be recognized as zoophilic (feeding on animals), ornithophilic (on birds), mammalophilic (on mammals) and anthropophilic (on humans). Immature states (eggs, larvae and pupae) are aquatic. Copulation occurs at the site where the adults emerge from their pupal casings. The females produce 200 to 500 eggs in a single gonotrophic cycle and some are able to distribute their eggs freely on the surface of the water (during flight) or in mass ovipositing on leaves or branches near the water. Some females will crawl under the water to deposit their eggs (PETERSON, 1981). The incubation period of the eggs varies from 4 to

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30 days, depending on the species and the water temperature, being much longer in those species in which the eggs undergo diapause (COLBO, pers. commun.). The eggs develop under humid conditions or below water. Hatching occurs a few days after oviposition and the larvae subsequently adhere to a substrate by the disk at the posterior end of their body (abdominal proleg). Sometimes they will also use the thoracic proleg, as well as a fine silk thread that is secreted through the mouth from their salivary glands. The larvae are nonselective filter feeders, trapping small organisms (zooplankton and phytoplankton) and organic particles of other insects circulating in the water column. The larvae occasionally drift downstream, being carried by the water currents. The final larval instars build a cocoon of silk secreted from the salivary glands. This serves as a refuge for pupation and transformation to the adult stage. The pupae have a slipper or shoe shape, with a series of filaments, which are the branches of gills (breathing organs), anteriorly. Simuliids remain in the pupal stage from 7 to 20 days, depending on the water temperature. During this period, they undergo a complicated metamorphosis to the adult stage, from which the adult fly emerges wrapped in a bubble of air that pops to the surface, where the fly becomes airborne to restart the cycle. At the beginning of the adult stage, the female flies look for their host. With few exceptions, the females show no host specificity. After taking a bloodmeal, the females look for a place to rest and digest the meal for egg maturation. Once the eggs have matured, the females select the appropriate site for oviposition. Simuliids generally feed during daylight hours, although some species prefer less sun and will feed either at dawn or during twilight hours, as in other haematophagous Diptera. Some species exhibit bimodal patterns of haematophagous activity while other species remain active throughout the day. It is believed that solar light stimulates haematophagous activity, but this is probably not the only factor, as feeding activity has been observed in dark places or on cloudy days. On the other hand, wind decreases feeding behavior, reducing the incidence of bites as wind velocity increases. The adults generally remain close to the streams in which they breed, generally in nearby humid microclimates, such as trees. The largest densities are observed within 100 m of the breeding sites, in shaded areas. Simuliids can disperse by intrinsic and extrinsic flight mechanisms to distances as great as 15 km or even as far as 40 km from their breeding sites, with the aid of wind (COSCARÓN, 1981b) and in Africa up to 400 km (CROSSKEY, 1990). Simuliid longevity is not well understood under field conditions but it is believed to vary around two months in temperate areas. The distribution of black flies is a consequence of their adaptive capacity to the environment and ancestral origin; a significant correlation is evident between distribution and endemism in the Neotropical realm (COSCARÓN & COSCARÓN ARIAS, 1995), as also evidenced in intercontinental relationships (MIRANDA ESQUIVEL & COSCARÓN, 2004). Public health and socioeconomic importance Black flies are not only annoying bloodsucking flies, but they are also vectors of important infectious diseases. Knowledge of this group will allow the reader to appreciate their

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importance as entomo-epidemiological agents. Black flies are known vectors of pathogens, such as filarial worms, viruses (myxomatosis and encephalitis) and protozoans (e.g., Leucocytozoon in birds). Of the parasites transmitted by this family, the most dangerous one to humans is Onchocerca volvulus, the etiological agent of onchocerciasis, where in extreme case it can produce blindness. Other species of Onchocerca parasitize cattle and wild animals (e.g., primates, birds) (Onchocerca lienalis, O. reticulata, O. gutturosa and O. cervicalis). Another filarial worm that can be transmitted to humans by simuliids is Mansonella ozzardi, although its resultant pathogenic action is slight. Leucocytozoon smithi (a protozoan parasite) has caused serious problems and numerous deaths on turkey farms, and L. simondi does the same in ducks in the USA. Leucocytozoon spp. also parasitize wild birds, particularly raptors, but they also have been found in doves (L. bonasae), chickens (L. andrewai), and even in penguins in New Zealand (L. tawaki). To date, no studies dealing with this parasite exist in Latin America. Simuliids are also vectors of arboviuses. They have been identified as being responsible for the transmission of Venezuelan equine encephalitis (VEE) and eastern equine encephalitis (EEE). In Australia, they are incriminated as vectors of the myxoma virus among others. As well as the annoyance that they produce, the saliva that is excreted during biting causes severe allergic reactions and may be responsible for secondary infections in people when they scratch. Black flies also mechanically transmit tularemia and other diseases (causing local reactions with itch and edema). Simuliids also cause economic losses at a regional level by affecting agriculture. The flies inhibit people from working normally outdoors, thereby reducing agricultural production. They also impact the cattle industry because the irritation caused by the simuliids to the cattle inhibit them from feeding properly, reducing the production of meat and milk. Losses to tourism must also be added to the list of socioeconomic problems; the annoyance caused by these insects can be so intense as to suspend recreational activities. Control There are different methods of black fly control: chemical, mechanical and biological; a general review of this topic for the Neotropical realm can be found in CAMPOS & ANDRADE (2001). Types of Control Chemicals are the best-known methods for insect control; synthetic insecticides have been especially effective. However, their use, particularly the wide-spectrum, residual insecticides, have altered ecosystems. Pesticides have been used against both the adult flies that attack us and the larvae, interrupting their life cycle; yet the methods to control the adult stage are not recommended because the insecticides that have to be applied to large areas result in adverse effects on non-target organisms, which include beneficial insects such as pollinators, predators, fish, birds, and amphibians. Chemicals used in the control of adult flies have included products such as DDT, methoxyclor, malathion, naled (commercially

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sold as Dibron) (ROMAÑA & AVALOS, 1948; ANDRADE, 1989) or diverse pyrethroids (SMITH, 1973; ELLIOT & POTTER, 1978); extreme caution should be taken so that these products do not contaminate rivers, streams or ponds. Adulticides should be used only in the case of an epidemic, to stop nuisance fly populations in restricted areas and only for short periods, as in the case of social events. For these reasons, the larvae are targeted in control strategies because they are limited to more defined areas, i.e. watercourses. There are extensive lists of products that have been used efficiently in the control of black flies, but for diverse reasons, such as high residual action, adverse effects on non-target organisms and resistance, many products have been replaced by newer less persistent insecticides. One of the latest insecticides being used has been temephos, mainly due to its low toxicity to vertebrates. However, resistance and toxicity to other insects has led to its replacement, especially by biological larvicides that have a more selective action. As yet there is no commercial chemical product available that is specific against simuliids that can be applied on a large scale that will not be harmful to the other organisms. The most advisable technique for black fly control is the use of biological control agents, including the use of live organisms or products that are derived from the black flies themselves and that are aimed at the reduction or elimination of the damages caused by species that are harmful to humans, domestic animals or agriculture. There are several biological control methods, but those with better results are the ones derived from bacteria, fungi and viruses. Natural enemies of simuliids in watercourses include organisms such as predators (e.g., fish, coleopteran larvae), parasites (Mermithidae), protozoa (microsporidia), fungi (especially Coelomycidium), viruses and bacteria. In regions where natural enemies exist and do not reach sufficient levels to be effective as a control measures, it is necessary to establish artificial rearing facilities to produce significant densities of the pathogens and parasites. This would not be easy to implement because they usually have complex cycles with diverse intermediary hosts and would be difficult to produce in sufficient quantities to achieve a viable level of control. Furthermore, there is still insufficient knowledge about the biological cycle of microsporidia and fungi to say whether they could offer better options. The search for a practical larvicide has led to the entomopathogen Bacillus thuringiensis var. israelensis (Bti), which was found to be the ideal solution. Bacillus thuringiensis is a bacterium that forms spores and produces protoxin crystals (delta-endotoxin) activated inside the alkaline intestine of the insects, and which become toxic, causing the degeneration of the epithelial lining of the intestine, paralysis and rapid death. Another control method is the mechanical cleaning and elimination of substrates, especially vegetation, to which the simuliid larvae and pupae can adhere. These methods could be used in conjunction with the application of the B.t.i. as a form of integrated control. Abundant subaquatic vegetation helps the development of black fly larvae. It has also been shown that this vegetation notably reduces the flow and speed of the water current that negatively impacts the effect of the biocide (LACEY & UNDEEN, 1988). The reduced flow not only limits the spread of the biocide, but also facilitates its adsorption by the vegetation and the periphyton. Consequently, it requires a higher dosage and increased frequency of application if vegetation is present. It was estimated that it would be

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necessary to apply the product at shorter intervals, requiring a 15- to 20-fold higher rate of application in certain areas. In large-scale control programs, the impact on the environment is of great importance, particularly when the control measures are used in recreational areas or in areas that are a source of potable water. For this reason, regulations exist in numerous countries concerning the use of chemical or biological products in watercourses in order to prevent the contamination or adverse effect on the quality of the water. Bacillus thuringiensis var. israelensis has been registered as a larvicide that does not produce adverse effects among the 10 most important groups of invertebrate aquatic organisms associated with black flies (W.H.O., 1982). To obtain the parameters to define a control strategy using Bti in the field, bioassays have to be carried out under laboratory conditions, keeping in mind the susceptibility of local species to different dosages, times of exposure, distances and the possible timing of application. Laboratory trials The system and procedure for testing B.t.i. are as follows. A rearing tank consists of a system of six flasks (5 for trials and 1 for control), with capacity of 2000 ml (approximately) each filled with 1000 ml of water collected from the same stream where the larvae were collected (COSCARÓN ARIAS, 1998a). The use of water originating from the appropriate streams assures that there is sufficient food for the larvae during the experiments. A system of aquarium air pumps oxygenates the water with diffusion stones to create a turbulence that maintains a constant agitation in the water. One hundred larvae of the different instars, preferably not of the last instar, are placed in each flask, using flexible forceps to avoid damaging the larvae. The suggested dosages of B.t.i. are 5.0, 7.5, 10.0, 15.0 and 25.0 ppm, with exposure times of 1, 5, 10, 30 and 60 min, respectively. Each of the five experimental flasks, along with the control flask, has to be labeled according to the time the larvae are to be exposed. The appropriate quantity of B.t.i. suspension is introduced into the flasks by means of a pipette, while maintaining uniform agitation in the flask. Once the pipette has been completely emptied, the time is recorded. After the specified exposure time period has elapsed, the water with the B.t.i. is filtered carefully to avoid the loss of larvae; next, the water is replaced with 1000 ml of water from the collection site. After 24 hours, the larvae, along with the possible pupae that formed in each flask, are counted. Finally, the percentage relationship of larvae that survived is calculated. The experiments are repeated three times with the same five dosages. Mortality data should be discarded for the trials in which the larvae from the control flasks exceed 10%. Results of the tests that have a mortality rate between 5 and 10% can be corrected by means of Abbott’s formula (COSCARÓN ARIAS, 1998a). Field trials The laboratory trials provide a basis for determining the concentration of B.t.i. and exposure time necessary during the field studies. The effect that the B.t.i. will have on the larvae will depend on the environmental conditions that determine the maximum dispersal distance and settling rates along the watercourses.

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Techniques, such as the one used by GRAY et al. (1999), are recommended. These techniques entail the application of the B.t.i. in concentrations according to the laboratory results. Prior to the application of the B.t.i., the velocity and flow of the water and the temperature must be recorded. To determine the distance that the B.t.i. carries downstream and the effect on the larval population, samples should be collected between three and five hours after the application at different distances downstream from the site of application (to be determined according to the characteristics of the watercourse). The samples will be taken 500 m beyond where the last larvae are observed to have been affected by the B.t.i. At each observation point, a sample of the substrate with larvae stuck to it will be placed in a container with one liter of water and placed in a cooler and refrigerated for transport to the laboratory. Once in the laboratory, aerators are placed in each container, and after 24 hrs the larvae are recounted and the percentage mortality is calculated. Adults are collected in the field at the time of application, as well as at various intervals after treatment, using the same technique, in order to compare the effects on the population after the application of B.t.i. The dates and times are recorded so that the trials can be repeated in the same area in the future. Based on the results obtained from the trials and the observations made, the trials must be repeated to determine the time taken for repopulation between the controls events.

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2 MORPHOLOGY AND CHARACTERS OF TAXONOMIC VALUE The description of the various characters will be presented in the order used in the species diagnoses, beginning with the female and followed by the male, pupa and larva. Adults: The body size is relatively small (2.2-6.0 mm long), but the wing length generally is used and does not change when the specimen dries out. The color is generally blackish gray to yellowish brown; commonly the female shows lighter coloration than the male. The head is well developed with compound eyes (Fig. 1 B: o) occupying a large portion of the cephalic capsule; they are composed of numerous ommatidia. Males have proportionally larger eyes (Fig. 1F), with upper macrofacets (mc) and lower microfacets (mi). Female eyes are separated by the frons and basally by the clypeus (Fig. 1B: c). At the base of the frons are the insertions of the antennae, where the eyes have an area without ommatidia, the fronto-ocular triangle (fo) (nudi-ocular area) of different depths. The male is holoptic, with the frons highly reduced. Some males are dichoptic, such as Gigantodax femineum, having an obvious frons. Males with different frons widths generally are intersexes (= specimens with male and female characters), produced by parasites, especially mermithids. Intersexes can be observed in different species, the genitalia showing parts of both sexes. Antennae (Figs. 1 A: a, B) are relatively short, composed of 11 segments (exceptionally 10, as in Paraustrosimulium anthracinum) of similar shape, although the two basal segments, scape (s) and pedicel (p), can be more elongate and in some species living at high altitudes (over 3000 m asl), they show long hairs (Figs. 38 P, Q). The apex of the head is the proboscis (Fig. 1 B) with relatively short mouthparts. Mandibles (md) and laciniae (lc) have serrations on the margin of one (Fig. 26 D) or both sides (Fig. 13 D) in females when they are functional. These serrations are absent or vestigial in males and non-haematophagous females. The lacinia is connected with the maxillary palp (mp) (Fig. 1 B: mp), which has five segments or palpomeres; the third palpomere has the sensory vesicle (sv), an organ with tubercles and which opens to the outside. The sensory vesicle is bigger in females and is of a different size in each species. Mandibles and laciniae are covered dorsally by a subtriangular plate, the labrum (lr). The clypeus is dorsal and the labium is basal and lateral, presenting the flexible labella (la) apically. Internally are the cibarium and hypopharynx. The basal portion of the cibarium (Fig. 1C) has two well-sclerotized branches laterally, the cornuae (co), and a median portion that can be smooth (Fig. 15 F) or have denticles in different positions and with different shapes, showing great variation in Neotropical species (Figs. 76 X, N; 80 D, P, Y; 84 I-W).

ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales M1 Rs

ast

ma

Cu1

R+M M sb Sc C Bas

st

F M2 mc

R1

A

23

mi

Cu2 An1 An2 abdt

pr

G

a an

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m ps pl

tr cx h

f

t

E ba

tm cl

ca

pe fl

o

p s

D

fo ca

c mp sv md

lr lc

co

la

B

C

Figure 1. A: Schematic black fly female in lateral view (a = antenna, abd t = abdominal tergites, an = anepisternum, ba = basitarsus, cl = claw, cx = coxa, f = femur, h = halter, k = katepisternum, m = mesoepimeron, mp = maxillary palp, pl = pleural sulcus, pr = pronotum, ps = posterior spiracle, st = scutum, t = tibia, tm = tarsomeres 1, 2, 3, 4, tr = trochanter; wing veins: An1 = anal 1, An2 = anal 2, Bas = basal cell, C = costa, Cu1 = cubitus 1, Cu2 = cubitus 2, M = media, M1 = media 1, M2 = media 2, mc = medial-cubital fold, R1 = radius 1, Rs = secondary radius (radius 2), R+M = radial-medial, sb = basal sector of radius, Sc = subcosta). B: Schematic female head in frontal view (c = clypeus, fl = flagellum, fo = fronto-ocular triangle, la = labellum, lc = lacinia, lr = labrum, md = mandible, mp = maxillary palp, o = compound eyes, p = pedicel, s = scape, sv = sensory vesicle). C: Basal portion of cibarium with denticles, S. (In.) subnigrum (co = cornuae). D: Basitarsus apex and tarsomere 1 (ca = calcipala, pe = pedisulcus). E: claw with small subbasal tooth. F-G. Schematic male head; F: lateral view (ma = macrofacets, mi = microfacets). G: frontal view.

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The thorax is well developed with great variation in prescutum (pr) and scutum (st) coloration as well as hair microstructure and arrangement (Figs. 5 A, B). The scutellum and metanotum (postnotum) have uniform morphology but variable coloration. The pleurae are generally without hair, except in Tlalocomyia, which presents some on the upper third of the pleural membrane (Fig. 10 B). The anepisternum (an) and katepisternum (k) can show some variation in sheen. The pleurae have anterior stigmata (= spiracles) (ast) and posterior stigmata (ps) on the upper area. The wings (Fig. 1 A) are wide, with a hyaline membrane covered by microtrichia, and along the anterior margin they have well developed veins with hairs and setae. The costa (c), subcosta (Sc) and radius (R), all end before the wing apex. The radius is bifurcated; the anterior branch is called R1 and the posterior is the radial sector (Rs ); the portion before the bifurcation is the basal sector of the radius (sb). The following veins are not readily evident. The media is bifurcated (M1 and M2). The median and cubital fold (mc) represent the false vein. The cubital vein is also bifurcated (Cu1 and Cu2) and two anal veins (An1 and An2) are present. Cu2 is generally curved but in Gigantodax and Pedrowygomyia it is straight. A basal cell (Bas) at the base of M can be present or not. Halteres (h) do not usually show variation in shape or coloration and are not used to differentiate species. Legs are relatively short and robust, with variable pigmentation and thickness, which is more accentuated in males, especially on the tibia and basitarsus. The legs have the following segments: coxa (cx), trochanter (tr), femur (f), tibia (t), basitarsus (ba), four tarsomeres (tm) and claws (cl). The legs are covered with hair with variable morphology, from filiform to flattened (Figs. 5 C, D). The hind basitarsus can have a flattened expansion on the apex, the calcipala (Fig. 1 D: ca), and tarsomere I can have a dorsal incision, the pedisulcus (Fig. 1 D: pe). The male claws have different morphology than those in the females and are always very uniform. Female claws vary greatly in shape among different taxa, and may or may not have a basal or subbasal tooth of various sizes (Figs. 1 E; 5 E, F). The abdomen is robust and can have different colors, similar to the thorax, particularly on the tergal area that also is well sclerotized; the pleurae have extensive membranous areas. The sternites are less developed than the tergites. The abdomen has only nine evident segments; the other two are transformed into genitalic organs. The first abdominal segment is short, scale-like and with long hairs. The female terminalia (Figs. 2 A, B) are composed of the eighth sternite (VIII st), a wide sclerite with abundant hairs laterally, and which project medially as a pair of foliaceous gonapophyses (go); the cerci (c); and paraprocts (p) (= anal lobes), all with various morphologies. Internally are the genital fork (g) and the spermathecal duct (sd). The genital fork has three branches and can have anterior apodemes on lateral branches, as in Cnesia, Pedrowygomyia and Gigantodax (Figs. 17 H, Q; 20 I, J). The spermatheca (Fig. 2 A: sp) is well sclerotized and subovoidal, with a neck at the union with the spermathecal duct that can be sclerotized or not. The male terminalia are composed of the gonopods and the aedeagus with associated parts (Fig. 2 C). The gonopods are comprised of two pieces; the basal one is the gonocoxite (gc) (= basimere or basistyle) and the distal one is the gonostylus (gs) (= distimere or dististyle). These show distinct interspecific morphological variations. The gonostylus generally has one apical or subapical spur (= spine) or several (2-5 or more). The aedeagus

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A

sd

c go g

sp

p

VIII st

B

gc

vp

C e gs

ms a

Figure 2. A: Schematic female terminalia, lateral view, Simulium sp. (c = cercus, g = genital fork, go = gonapophysis, p = paraproct, sd = spermathecal duct, sp = spermatheca, VIII st = eighth sternite). B: Female genitalia, ventral view (left cercus and paraproct) mounted on slide position, S. (E.) romanai. C: Male genitalia, ventral view, S. (E) mayuchuspi (a = aedeagal membrane, gc = gonocoxite, gs = gonostylus, e = endoparamere, ms = median sclerite, vp = ventral plate).

can be associated with several sclerotized structures such as the ventral plate (vp) transversely positioned and with variable morphology; the median sclerite (ms) on the middle of the genitalia, showing a distal, deep incision; and the endoparameres (e) (= parameres), composed of 1+1 well sclerotized branches, frequently with robust teeth (= hooks or spines) on the distal portion. Bordering the muscular aedeagus is the aedeagal membrane (a), with abundant minute spiculae.

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Pupa: Size variable (2.0-8.0 mm); the measures that are generally used are the cocoon length dorsally and basally and the gill length. The pupal body and frequently the gills are protected by a cocoon, which is secreted by the larva. The cocoon weave can be compact, loose, or spongy,, sometimes not showing the threads. The common shape is that of a slipper or shoe, with great variations such as ventral or dorsal projections, dorsal prominences (e.g., a carina), or reduction to only a posterior covering of the pupal abdomen. The pupal body shows great uniformity. It has a resistant integument, generally well sclerotized, with yellowish to dark grayish brown tones. It has tubercles, trichomes, and spines, with great variety of shape, size, number and position. The trichomes can be single or multibranched, generally filiform (Figs. 3 B, C), but sometimes flattened (Figs. 33 S, T) or star shaped (Figs.

g

A c

fr

f

B

C

rf g

D

Figure 3. Black fly pupa, S. (Pt.) limay. A: lateral view (c = cocoon, g = gills). B: frontoclypeus with tubercles and trichomes (f = facial trichome, fr = frontal trichomes). C: thoracic plate with gill (g), respiratory filaments (rf) and trichomes. D: abdominal chaetotaxy.

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53 S, T); tubercles can be flattened or elevated with smooth or wrinkled surfaces or apically acuminate; the spines can be isolated or grouped like a comb, with one or several branches, straight or curved like a hook (Fig. 3 D). The frontoclypeus, the antennae and eye sheaths are evident on the cephalic plate. The frontoclypeus can show great variation according to the number, type and disposition of tubercles. The trichomes are generally the frontal (fr) and facial (f) trichomes but can also include epicranial trichomes. For the thorax, special attention is given to the dorsal exposed portion, with the presence or absence of tubercles and different types of trichomes. The abdomen has nine evident segments, ending with a pair of spines (Figs. 6 E, F). The tergal plates and sternal, pleural or intersegmental membranes have different integumentary processes such as trichomes, spines and hooks that are of taxonomic value (Fig. 3 D). On the anterior portion of the pupal body, there are two prothoracic expansions (with a respiratory function), the gills (g) (Figs. 3 A, C). The gills have great morphological variations and are very important for taxonomy. Generally, the gills have a basal trunk from which emerge several filamentous branches with arborescent disposition (Fig. 6 A). These filaments are sclerotized with varied porosity and are hollow internally; externally they are smooth (Figs. 6 B, C) or with furrows (Fig. 6 D); the gill apex can be acute (Fig. 6 B) or blunt (Fig. 6 C). The gill also can show thick and asymmetrical branches with varied morphology (Figs. 66 L, M; 70 A-C, G-I, P-R; 122-124). The number of filaments varies from one to more than two hundred; the numbering of branches is from the dorsal to ventral position, following a clockwise direction. The branches emerging from the primary branches are the secondary branches, and those branching from the secondary branches are the tertiary branches, and so on. Larva: Maximum length in the last larval stage varies from 4.0 to 12.0 mm. The larvae have a subcylindrical shape with a well differentiated head; the posterior third of the body is more expanded (Fig. 4 A). Coloration is brown, gray, yellow to greenish with distinct intensity and tones from light to dark, especially on the dorsum and head. The lateral thoracic area can have spots from light to dark brown corresponding to the appendage histoblasts and especially to the gill histoblasts. On the ventral part of the body, it is often possible to see the ventral nerve cord and the prothoracic ganglion at the base of the head. The head has a hard consistency due to the sclerotized cephalic capsule. Generally it has a constant coloration, although it is lighter when ecdysis has been recent. Unique coloration is shown dorsally on the cephalic apotome (= frontoclypeal apotome) (Fig. 4 B: ca), a median plate delimited by the ecdysial line (el). Generally the cephalic apotome has several spots arranged in groups that can be positive or negative. According to their position, they are referred to as anteromedial (as), posteromedial (pm), anterolateral (al) and posterolateral spots (pl). Lateral to the ecdysial line on each side is the postgena (pg), with ocelli (o) in the mid area. Anteroventrally, there is a projection, the hypostomium (Fig. 4 C: h) with wellsclerotized teeth on the anterior margin (Fig. 4 D). These teeth are the median (md) (one tooth), intermediate (i) (= sublateral) (generally 3 teeth on each side), lateral (l) (one tooth on each side), and paralateral (pl) (several teeth) followed sometimes by serrations. Anterolaterally, the hypostomium has a row of variably numbered setae in a diagonal arrangement. Separating the hypostomium from the postgena is the hypostomial groove and the space between this sulcus and postgenal cleft (pc) is the postgenal bridge. The

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pg ca

pl po

H

F

ce r

l m

me

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E

pb

d

p

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cf

Figure 4. Black fly larva. A: schematic general view in lateral position (a = anal sclerite, ap = abdominal proleg, gh = pupa gill histoblast, rp = rectal papillae, tp = thoracic proleg, vt = ventral tubercles). B: schematic head, dorsal view (a = antenna, al = anterolateral spots, as = anteromedial spot, ca = cephalic apotome, ce = cervical sclerite, cs = cephalic fan stem, el = ecdysis line, m = mandible, o = ocellus, pg = postgena, pl = posterolateral spots, pm = posteromedial spots, po = postocciput). C: schematic head, ventral view (cf = primary cephalic fan, h = hypostomium, lh = labiohypopharynx, m = mandible, mp = maxillary palp, mx = maxilla, pb = postgenal bridge, pc = postgenal cleft, r = cephalic fan rays , sf = secondary cephalic fan, t = tentorial pit). D: hypostomium, anterior border (i = intermediate teeth, l = lateral tooth, md = median tooth, pl = paralateral teeth, s = lateral serrations). E: antenna, S. (Ch.) subpallidum (d = distal, me = medial, p = proximal). F: maxillary palp. G: mandible apex, S. (Ec.) albanense (in = inner teeth, lm = latero-mandibular process, ms = mandibular serrulations, o = outer teeth, ap= apical tooth, pa = preapical teeth). H: anal sclerite with abundant hair, S. (Ec.) albanense.

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A

B

C

D

E

F

Figure 5. Female. A-B, D-E: S. (N.) exiguum; A: scutum hair disposed in groups; B: scutum scalelike seta; D: filiform and scale-like setae of leg; E: claw without subbasal tooth. C: S. (Pt.) stelliferum, hind leg with filiform setae; F: Gigantodax clandestinus, claw with subbasal tooth.

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A

B

C

D

E

F

Figure 6. Pupa. A-B: Gigantodax igniculus; A: gill; B: apex of gill branch. C: G. marginalis, apex of gill branch. D: S. (Tr.) lahillei, apex of gill branch. E-F: G. igniculus, abdomen apical portion in lateral and dorsal view showing terminal spur, pleural membrane, tubercles, spine combs and hooks.

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postgenal cleft shows variation in shape and depth and has taxonomic importance. At the postgenal base, ventrally on each side, is the posterior tentorial pit (p), and all around the posterior margin is the well sclerotized postocciput (po). The postocciput, over the median cephalic portion of the apotome, is thinner, and has the cervical sclerites (ce), which meet the postocciput. Below the epidermis and near the postgenal cleft is the subesophageal ganglion, which is sometimes darkly pigmented. Dorsolaterally on the anterior portion of the head are the prominent cephalic fans (= labral fans) (Fig. 4 C: cf), with a robust cephalic fan stalk (Fig. 4 B: cs) and variable number of rays (r). Cephalic fan rays have thinner teeth arranged like a comb, showing variable length and position (Fig. 7 B). The largest fan rays are the primary rays which number from 20 to 80; there are other smaller ones know as secondary rays that are fewer in number and located more basally (Fig. 4 C: sf); a third type consists of short, flat, straight blades, the median rays, that form the scale fan (sensu CROSSKEY, 1990: 146). Behind the cephalic fan are the antennae (Fig. 4 B: a), composed of three segments (Fig. 4 E): proximal (p), medial (me) and distal (d) ending in a short and acute sensillum. Relative proportions of segment lengths are useful in taxonomy. Ventrally there are several mouthparts. Apicoventrally, the mandibles (Fig. 4 C: m) have several brushes and teeth (Figs. 4 G; 7 E). The teeth are more important taxonomically, showing variation in number and size. The outer teeth (o) are the first 2 or 3 large teeth. Following these is the largest one, the apical tooth (ap) and it is followed by the preapical teeth (pa) and the inner teeth (in) that can be present in several rows, and lastly the marginal serrations (ms) which are also variable in number. Some species can have a latero-mandibular process (lm) that can be single or double and cylindrical or a little flattened. In some supraspecific taxa, a row of supramarginal setae (Fig. 79 B) can be present. The larvae have maxillae with abundant long hairs (Fig. 4 C: mx); the maxillary palp (mp) is cylindrical (Fig. 4 F), with apical sensillae of variable morphology (Fig. 7 A). The labrohypopharynx is setose and has a uniform appearance (Fig. 4 C: lh). The labrum is in front of the head and mostly covered by setae having little taxonomic importance. The body is slightly curved ventrally but it can be strongly curved while moving from one place to another. It shows little segmentation particularly on the thorax. The body integument is generally covered by microscopic trichomes but can show different epithelial processes of varied morphology, some of them being of larger size (Figs. 4 D; 135 G, L, S). Trichomes and scales are frequently found around the anal sclerite (Fig. 4 H). On the thorax are whitish histoblasts that correspond to wings, legs and pupal gills (Fig. 4 A: gh). Ventrally there is an elongated cylindrical appendage, the thoracic proleg (= prothoracic proleg) (Fig. 4 A: tp). On its apex, it has a row of hooks and subapically 1+1 sclerotized, subtriangular to subtrapezoidal lateral sclerites, with a comb-like structure of elongated teeth frequently arranged in groups (Figs. 7 F; 8 X; 12 M). The larval abdomen has external pseudoannulations, sometimes evident as thin, pale rings. Laterally, there are minute rounded dark spots corresponding to nonfunctional spiracles. Posteriorly, on the ninth segment there are 1+1 ventral tubercles (= ventral papillae) (Fig. 4 A: vt) with variable presentation in different taxa. On the posterior apex is the abdominal proleg (ap) (= posterior proleg), which looks like a sucker, bordered by hooks arranged in rows and disposed around the proleg as the anal ring (= posterior

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A

B

C

D

E

F

Figure 7. Larva. A: Gigantodax herreri, apical sensillae of maxillary palp. B: Gigantodax sp., microtrichia on cephalic fan of primary ray. C: S. (Ps.) auripellitum, terminal portion of abdomen showing rectal papillae, abdominal proleg, ring of hooks and cuticular hair. D: Simulium sp., cuticular scale-like hair. E-F: G. igniculus. E: mandible portion showing teeth. F: apex of thoracic proleg showing lateral sclerite.

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circlet) (Fig. 7 C). The number of rows of hooks is variable (40 to more than 300) and is very useful in taxonomy. Anterior to the abdominal proleg is a well-sclerotized structure, the anal sclerite (a) (Fig. 4 A: a). In Neotropical species, it has an X shape with ventral branches longer than the dorsal ones (Fig. 4 H). In some taxa, such as Gigantodax, Pedrowygomyia and partially in Hearlea, the ventral branches are very elongated and joined ventrally to form a circle (Fig. 20 S) around the anal ring. In some species of Hearlea, there also are well-sclerotized accessory plates with different shapes near the anal sclerite (Figs. 127 B-J). Dorsal to the abdominal proleg is the anus. From the distal portion of the rectum, larvae can extrude a membranous, elongate organ, the rectal papillae (Fig. 4 A: rp). Basically, the rectal papillae is composed of three cylindrical lobes commonly present in the primitive simuliid groups; as an apparent apomorphic variation each lobe can have one or more diverticula (= lobules) on each lobe (Fig. 7 C). There can be over 30 diverticula on each lobe. The rectal lobes frequently remain inside the larva after fixation and it is not possible (without dissection) to see this character, which has some taxonomic value. Eggs: The eggs are ovoid to subtriangular and uniform in color, which varies from light yellowish gray to dark gray, depending on the time of ovoposition. There are no published studies on Neotropical simuliid eggs for species identification. It is possible to find eggs when they are deposited in large masses where several females gather to oviposit. The curved larva and its dark head with the ruptor ovi (= egg buster) dorsally positioned in the center of the head can be seen in the eggs before eclosion.

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3 COLLECTION AND PREPARATION OF MATERIAL Adults: The easier of the two sexes to obtain is the female because of the haematophagous habits, which attract them to humans or domestic animals. Females can be collected by nets or traps such as Malaise, Shannon or Manitoba traps, which are commonly used to capture other blood feeding insects. When using a trap, it is useful to have an attractant such as a person, an animal or CO2. To capture biting females, it is best to put a small vial over the fly when it is feeding and not use forceps so as to avoid damaging the specimen. For taxonomic studies, it is preferable to kill the flies and maintain part of the material dry to be pinned afterwards in the lab. To kill adults, ethyl acetate, ether, chloroform, cyanide, etc. can be used, but not applied over the specimen. It is desirable to place pinned material in a freezer for at least five weeks to avoid having the abdomen retract. The remaining material should be fixed in 80% ethyl alcohol and maintained as long as it is necessary for further studies. It is better to see coloration in dry material; however, if fixed material is to be used for the observation of color, it should be placed in ethyl acetate for at least ten minutes and dried on filter paper to evaporate all the liquid. After this, the hairs will become more erect and better show the characteristics. To collect males, it is necessary to use a hand net and try to find a swarm or to catch them as they fly above the aforementioned traps. Larval stages: Larvae and pupae are found together in different types of freshwater streams and on various substrates. Generally, they are found near the surface, rarely below one meter in depth. The best place to find them is where the current is fast. The material can be collected with forceps and maintained alive or can be fixed directly in alcohol or other fixative. It is convenient to keep material alive in a vial placed in a cooler while transporting it to the lab where the flies can be reared or fixed for future studies. For taxonomic studies, it is best to keep them in 80% ethanol. Carnoy’s fixative (100% ethanol: glacial acetic acid = 3:1) should be used for cytological studies or 100% alcohol for molecular studies. For the last two studies, it is convenient to change the fixative after 3060 minutes and then keep the vials in a refrigerator. Color information should be obtained from fresh material (before fixation). Pupae can be fixed and stored in 80% alcohol. It is possible to rear black flies under laboratory conditions to obtain different stages. Larvae can be reared if water current is assured. There are different methods, some of which are more sophisticated (EDMAN & SIMMONS 1985 a, b), and others less so. One of the easier ways to obtain pupae is to maintain the larvae in a vial with an air flux at the

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base and with a magnetic stirrer. Pupae can also be reared to obtain imagoes. They can be maintained in a vial on humid paper with a cotton stopper, avoiding water condensation on the walls. It is convenient to rear pupae individually in order to identify the adults and have associated pupal exuviae.

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4 STUDY OF MATERIAL Taxonomic studies start with the observation of external characters such as size, coloration and morphology. It is necessary to use a stereomicroscope that can also be used for dissections and specimen assembly on slides. Anatomical parts used for the identification of specimens are chitinous. To clarify them adequately, it is useful to digest the accompanying tissue with NaOH or KOH at about 5% to 10%. This can be done by boiling the material for 30-60 seconds, but since this procedure can destroy delicate structures, it is better to leave the material in the solution for 12-18 hours at room temperature instead of boiling it. In adults, the wings should be removed first, to avoid damage. In larvae, this treatment needs to be done in less time, and some delicate parts, such as the rectal gill and antennae, are better seen without the use of these treatments. After this procedure, it is necessary to neutralize the material with acetic acid or phenol and dissection. The parts can be stored in glycerin or mounted on glass slides in Canada balsam or Euparal. Before mounting, it is desirable to study the different pieces with a compound microscope to see the different structures in diverse positions and to gain better knowledge of the morphology. There are different methods used to clear the pieces before they are slide mounted. A fast method is to transfer from NaOH or KOH directly to phenol (= phenic acid) for 10-15 minutes for clarifying. It is also possible to dissect the material at this time, with histological needles. Materials treated with phenol can be observed with a compound microscope in temporary preparations, or can be assembled on a slide for a permanent preparation. Phenol can crystallize at low temperatures (in air conditioning). In this case, it must be warmed and several drops of 100% alcohol added to the remaining liquid. When the material needs to be cleared further after passing through phenol, it is convenient to leave it for an hour in creosote. It is also possible to leave the material for more than one day in the phenol or creosote without causing any damage. If the material is in phenol for several hours, exposed to ambient conditions, it will be necessary to replace it with new phenol, because it can hydrolyze, and the preparation may be not clear enough. When the structures are too clear, they can be dyed with a coloring solution. Leaving the material for 30 minutes or more in “lignin pink”, dissolved in creosote gives good coloration. Gill histoblasts can be seen best if they are transferred to acetic acid diluted at 1020%. This helps to uncurl the filaments and, therefore, makes it is easier to count them. Phase-contrast microscopes and electron microscopes are useful for seeing some structures. For electron microscopy, it is best not to use material that has been preserved

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for a long time. To clean the adhered particles, ultrasonic equipment can be used with some detergent. It is important to control time in order not to lose some parts such as hairs or to break others. It is important to check the material after 10-15 seconds and repeat the procedure if necessary. After using the detergent, the structures must be rinsed with water, alcohol and acetone. Before affixing the specimen to the support for metal coating, it is necessary to dry the material carefully with filter paper. During the analysis of the material it is useful to make drawings for future comparisons.

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5 SYSTEMATICS Neotropical Simuliidae are all included in the subfamily Simuliinae, according to the general classification used by ADLER et al. (2004). In the present volume, it comprises 12 genera and 359 species, following the arrangement given by WYGODZINSKY & COSCARÓN (1973a). The first ten genera were previously considered in the tribe Prosimuliini, as described by CROSSKEY & HOWARD (1997, 2004). The validity of Prosimuliini was questioned by PY-DANIEL (l994). Presently, the most representative of these ten genera are included in the tribe Simuliini (ADLER et al., 2004). The genus Simulium is the most abundant with l8 subgenera, as described in COSCARÓN (1987). The CROSSKEY & HOWARD (2004) taxonomic arrangement is very close, yet with some small differences in Kempfsimulium, Psaroniocompsa, Ectemnaspis and Trichodagmia. These Simulium subgenera are considered genera by PY-DANIEL & MOREIRA SAMPAIO (1994, 1995). Here, we will not deal with this supraspecific category, considering that a study of the world black flies would be the best forum to address this issue. In this compilation, the species of each taxon are arranged based on the arrangement given in previous cladistic studies, particularly those in COSCARÓN (1987); COSCARÓN & COSCARÓN ARIAS (1996b, 1997); COSCARÓN et al. (1996, 1999, 2004); MIRANDA-ESQUIVEL & COSCARÓN (2001, 2004); and STRIEDER & PY-DANIEL (2002). Generic and specific diagnoses are restricted to those morphological characters that permit taxon identification, and are accompanied by figures facilitating their identification. Diagnosis is accompanied by geographic distribution, including states or provinces of each country and their corresponding maps. Succinct bionomic information is included with each taxon. We have attempted to compile most of the available information, but recognizing the vast extent of this realm, with areas yet to be explored or collected, we consider that the information presented corresponds only to a snapshot view of the Neotropical black flies. With this volume we intend to aid researchers that are immersed in the complex world of limnology and who are trying to understand the interrelationships of these systems while trying to protect them from the damage “progress” has caused. Also, we aim to help those that try to control nuisance or disease-bearing black flies. If rational control is planned, it is necessary to know which species are pests and which are innocuous species in order to save the latter which play an important role in competing with the more dangerous species.

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KEYS TO THE GENERA

OF

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NEOTROPICAL SIMULIIDAE

Adults 1 – 2(1) – 3(2)

–

4(2) – 5(4)

–

6(5) – 7(6)

–

Pedisulcus well developed (Fig. 1 D: pe); basal cell invariably absent; length of basal section of R (as measured from humeral cross-vein) less than 1/3 distance from base of Rs to wing tip ................................................................................ Simulium Pedisulcus absent (Figs. 8 G, N); basal cell absent or present; length of basal section of R about 1/3 distance mentioned or more ................................................ 2 Cu2 and A1 straight (Fig. 20 C); basal cell absent ........................................................ 3 Cu2 and A1 curved (Fig. 8 E); basal cell absent (Fig. 26 F) or present (Figs. 8 E; 15 G) ................................................................................................................................................ 4 First flagellomere as long as wide; hair of basal flagellomeres and palpomeres shorter than segment; clypeus longer than wide (Fig. 20 B); female mouthparts normal, functional; calcipala present (Fig. 20 E); subbasal tooth on claw well developed (Fig. 20 G) ....................................................................................... Gigantodax First flagellomere as long as wide; hairs of basal flagellomeres and palpomeres longer than segment; clypeus as long as wide (Fig. 20 A); female mouthparts reduced, non functional; calcipala absent (Fig. 20 D), subbasal tooth on claw obsolescent (Fig. 20 F) .................................................................................Pedrowygomyia Upper portion of pleural membrane hairy (Fig. 10 B); apical segment of maxillary palp more than twice as long as penultimate segment .............................. Tlalocomyia Pleural membrane entirely bare; apical segment of maxillary palp generally less than twice as long as penultimate segment (Figs. 13 C; 15 C; 21 B) ...................... 5 Antennae with 10 segments each; arms of furcasternum with conspicuous projections (Fig. 23 A); male genitalia with median sclerite geniculate, composed of slender basal Y-shaped portion and 2 subparallel elongate apical arms (Fig. 23 H); apical portion of endoparameres with numerous elongate denticles; genital fork of female almost entirely unpigmented, its stem very stout, not longer than arms (Fig. 23 E) .......................................................................................Paraustrosimulium Antennae with 11 segments each (Fig. 19 A); arms of furcasternum lacking conspicuous projections (Figs. 8 D; 13 F) [except Lutzsimulium (Fig. 24 L)]; median sclerite of male not as above, or apical portion of endoparameres obsolescent, lacking denticles (Figs. 8 Q; 24 I); genital fork distinctly pigmented, with stem not as above (Figs. 8 L; 10 E; 11F; 13 M; 17 H, Q) .......................................................... 6 R1 only with hair-like setae, spiniform setae not developed (Fig. 21 F); basal tooth of claw of female large (Fig. 21 J) ................................................................ Cnesiamima R1 with hair-like and spiniform setae (Figs. 13 H, I); if spiniform setae not very distinct, then basal tooth of claw of female small (Fig. 24 D) ................................ 7 Basal cell absent (Fig. 26 F); mandible of female toothed only on internal margin (similar to Fig. 26 D); claws of female with small subbasal tooth (Figs. 24 D; 26 K); male genitalia with teeth of endoparameral organ not perceptible (Figs. 24 I; 27 A) ..................................................................................................................................... 8 Basal cell present (Fig. 8 E); mandible of female as above or with well-developed teeth on internal and external margins (Fig. 13 D); claws of female with larger

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8(7) – 9(7)

– 10(9) – 11(10) – 12(11)

–

13(10) – 14(13) –

basal tooth (Figs. 13 G; 15 I); male genitalia with teeth of endoparameral organ distinct (Figs.11K; 13T; 17K: ed) .................................................................................... 9 Frons longer than wide with median sulcus relatively short and without basal bifurcated branches (Fig. 24 A); Sc with about 50 setae (Fig. 24 H); furcasternal branches with large projections (Fig. 24 L) .............................................. Lutzsimulium Frons about as long as wide with median sulcus elongated and bifurcated, with long basal branches (Fig. 26 A); Sc without setae (Fig. 26 F); furcasternal branches with very small projections (Fig. 26 L) .................................................... Kempfsimulium Wings with slight curvature of Cu2; A1 almost attaining wing margin (Fig. 17 B); male genitalia with basal portion of endoparameres obsolescent, denticles of distal portion well developed but few in number, forming tight group of characteristic arrangement (Fig. 17 K); female with arms of genital fork bearing 2 forwardly directed apodemes (Figs. 17 H, Q) ......................................................Cnesia Wings with pronounced curvature of Cu2; A1 ending well before wing margin (Fig. 8 E); endoparameres of male not as above; genital fork of female without apodemes (Figs. 8 L; 11 F) ............................................................................................. 10 Males ................................................................................................................................... 11 Females ............................................................................................................................... 13 Median sclerite with basal transverse satellite plate continuous distally with deep cleft and bifurcated apically (Fig. 8 R); endoparameres lacking teeth (similar to Fig. 8 Q) .................................................................................................................... Mayacnephia Median sclerite of complex geniculate shape, with distal half divided into 2 wide but short and divergent arms (Figs. 11 J, K; 13 R, S); endoparameres with numerous teeth (Fig. 11 K) ............................................................................................ 12 Genitalia with ventral plate subtriangular and basal arms narrow, straight (Figs. 11 I; 13 P); arms of median sclerite parallel (Fig. 13 S); endoparameres with basal portion large and denticles of apical portion well developed (Figs. 11 K; 13 T) .. .............................................................................................................................. Araucnephia Genitalia with ventral plate subrectangular and basal arms stout, rounded at apex and curved (Figs. 15 P, Q); arms of median sclerite sharply diverging apically (Fig. 15 U); endoparameres with basal portion small and denticles of apical portion obsolescent .................................................................................................. Araucnephioides Basal tooth of claws narrow, hook like (Fig. 15 I) .............................. Araucnephioides Basal tooth of claws larger, subtriangular or spatulate (Figs. 8 H, J; 13 G) ........ 14 Frons very narrow (Fig. 8 A), frontal angle approximately 50°; inner surface of spermatheca without spiculae (Mesoamerica) .......................................... Mayacnephia Frons wider (Fig. 15 A), frontal angle approximately 85°; inner surface of spermatheca with scattered spiculae (southern South America) ........... Araucnephia

Females, based on genitalia 1 –

Arms of genital fork with long cephalad-directed apodemes (Figs. 17 H, Q; 20 I, J) ................................................................................................................................................ 2 Arms of genital fork lacking long apodemes .............................................................. 3

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–

7(5) – 8(4) – 9(8) –

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Anterior branch of genital fork of uniform width (Figs. 17 H, Q; 20 J) ................ .................................................................................................................. Cnesia, Gigantodax Anterior branch of genital fork expanded on basal 1/3 (Fig. 20 I ) ....... Pedrowygomyia Genital fork almost entirely unpigmented, its stem wide and very short, not longer than arms (Fig. 23 E) .............................................................................Paraustrosimulium Genital fork not as above ................................................................................................. 4 Sclerotization of spermatheca continuing onto base of spermathecal duct (Figs. 26 P, Q) ................................................................................................................................ 5 Spermathecal duct inserted on small membranous area of spermatheca (Figs. 8 J; 21L) ....................................................................................................................................... 8 Area of insertion of spermathecal duct on spermatheca with network-like structure extending to base of duct (Fig. 24 G) ......................................................... 6 Not as above ....................................................................................................................... 7 Sclerotized area of spermathecal duct also covering spermathecal neck and portion of spermathecal duct with meshwork (Fig. 24 G); spermatheca internally with spiculae; eighth sternite with sclerotized area reduced to medial area (Fig. 24 E) .... ............................................................................................................................ Lutzsimulium Sclerotized area of spermathecal duct reduced to neck, without meshwork on spermatheca union (Figs. 26 P, Q); spermatheca internally without spiculae; eighth sternite with sclerotized area covering all area from one side to other (Fig. 26 M) .......................................................................................................................... Kempfsimulium Paraproct very small, triangular; surface of spermatheca conspicuously reticulate, its inner surface without spiculae; sclerotization of spermathecal duct ending abruptly (Fig. 10 D) .......................................................................................... Tlalocomyia Paraproct of normal size, rounded (Fig. 21 M); surface of spermatheca smooth, its inner surface with few spiculae; sclerotized portion of spermathecal duct gradually fading into unpigmented portion (Fig. 21 L) ............................ Cnesiamima Inner surface of spermatheca without spiculae ....................................... Mayacnephia Inner surface of spermatheca with scattered spiculae ............................................... 9 Paraproct about as long as wide (Fig. 13 L); inner surface of spermatheca not rugose ................................................................................................................. Araucnephia Paraproct about one half shorter than wide (Fig. 15 L); inner surface of spermatheca rugose ................................................................................... Araucnephioides

Males, based on genitalia 1 – 2(1)

Endoparameral teeth not discernible; median sclerite comparatively simple, wide basally, elongate lamella, apically divided into 2 short arms connected by membrane (Figs. 24 I; 27 A) ............................................................................................ 2 At least some portion of endoparameral organ discernible; median sclerite different, generally of complex structure ..................................................................... 3 Gonocoxite about 2 times longer than gonostylus; gonostylus gradually acuminate distally, apex relatively blunt, with 3 (2) terminal teeth (Figs. 24 I, M); ventral plate distally with soft rounded curvature; median sclerite very elongated (Fig. 24 J); pedisulcus absent ............................................................................................ Lutzsimulium

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3(1) – 4(3) – 5(4) – 6(3) – 7(6) – 8(7) – 9(7) –

Gonocoxite about 3 times longer than gonostylus; gonostylus strongly acuminate distally with 2 terminal teeth (Figs. 27 A-C); ventral plate showing irregular border distally (Figs. 27 A, D); median sclerite wider, not elongated and distally gradually acuminate; pedisulcus absent to well developed ................................... Kempfsimulium Basal portion of endoparameral organ not discernible or only very weakly developed; denticulate portion composed of 1 or 2 large and several very closely associated, often oppositely directed smaller denticles (Fig. 17 K) ........................ 4 Basal portion of endoparameral organ distinct, plate like; denticles of distal portion, varying from absent to well developed, but not as described above ...... 6 Ventral plate subpentagonal, about as long as wide, not conspicuously emarginated apically (Fig. 17 N) ......................................................................................................Cnesia Ventral plate in most cases elongate-rhomboid, with distinct apical emargination ................................................................................................................................................ 5 Ventral plate short, with lateral margins at midlength concave and apical margin bilobed (Fig. 20 L); endoparameral teeth elongated .................................. Gigantodax Ventral plate long, about 2 times longer than wide, with lateral margins at midlength straight and apical margin straight or convex (Fig. 20 K); endoparameral teeth short ..........................................................................Pedrowygomyia Median sclerite consisting of transverse basal flange and deeply incised median portion perpendicular to flange (Figs. 8 R; 10 G); endoparameres reduced to basal plate-like portion, and without denticles (Fig. 8 Q) ........... Mayacnephia, Tlalocomyia Median sclerite different; endoparameres with or without denticles ...................... 7 Median sclerite strongly geniculate in side view (Fig. 21 U), roughly M-shaped when seen in full view (Figs. 21 V; 23 H), with inner arms of M partly fused and connected to basal flange ................................................................................................. 8 Median sclerite not as above, its shape complex and occupying 3 different planes ................................................................................................................................................ 9 Endoparameral organ with free plate-like basal portion and numerous elongate subparallel apical denticles not connected to basal portion (Fig. 23 H) .................. .....................................................................................................................Paraustrosimulium Endoparameral organ reduced to subquadrate plate-like structure; denticles not developed (Fig. 21 U) ....................................................................................... Cnesiamima Ventral plate subpentagonal or subtriangular (Fig. 11 I), its basal arms in shape of elongate slender rods (Fig. 13 P); branches of median sclerite parallel (Fig. 13 S); endoparameres with conspicuous denticles distally (Fig. 11K) .............. Araucnephia Ventral plate subrectangular, its basal arms stout, apically knobbed (Fig. 15 P); branches of median sclerite diverging apically (Fig. 15 U); endoparameres with denticles of distal portion obsolescent (Fig. 15 S) ............................. Araucnephioides

Pupae 1

Abdominal sterna VI and VII divided longitudinally along middle by membranous, striate area (Figs. 14 D; 18 D); hooks of terga VI-VIII invariably simple; abdominal segments VIII and IX in many cases with strongly curved, twisted, looped or grapnel-like strong setae (Figs. 18 E; 23 J; 25 E) .......................................................... 2

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– 2(1) – 3(2)

–

4(2) – 5(4) – 6(5) – 7(5) – 8(7) – 9(8) – 10(9) –

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Abdominal sterna VI and VII entire; in many cases, some hooks on terga VI-VIII bifid or trifid; abdominal terga VIII and IX without above-mentioned setae ........ ................................................................................................................................... Simulium Terminal processes of abdomen short and pointed (Fig. 23 J), or absent (Fig. 10 L) ................................................................................................................................................ 3 Terminal processes of abdomen elongate (Figs. 12 E; 18 E; 25 E) ....................... 4 Cocoon reduced to small pad on which terminal abdominal segments are inserted (Fig. 10 H); gills shaped like thick stem with few thread-like filaments (Fig. 10 I); abdomen strongly sclerotized, terga and sterna with large number of supernumerary spines or hooks (Figs. 10 J, K); apex of abdomen blunt; terminal processes absent (Fig. 10 L) ............................................................................ Tlalocomyia Cocoon well developed, of definite shape, covering whole body of pupa except gills, latter in shape of twisted pseudo-segmented lamella (Figs. 22C; 23 I); abdomen less sclerotized than thorax; abdomen without supernumerary hooks, and with short terminal processes (Fig. 23 J) ........................................... Cnesiamima, Paraustrosimulium Abdominal terga with spine-combs (Fig. 14 D); tergum VIII without large hooks ................................................................................................................................................ 5 Abdominal terga without spine-combs; tergum VIII with 4 large hooks ..... Cnesia Facial trichomes (Figs. 12 B; 27 E), thoracic trichomes (Figs. 12 D; 27 F, G), and setae of eighth and ninth abdominal segments tightly looped apically (Fig. 25 E) ................................................................................................................................................ 6 At least facial and thoracic trichomes not looped apically........................................ 7 Gill with 12-22 branches; frontoclypeus and thorax with tubercles (Fig. 25 D) .... ............................................................................................................................ Lutzsimulium Gill with 12 branches; frontoclypeus and thorax without tubercles (Fig. 27 E) .... .......................................................................................................................... Kempfsimulium Head sclerite with frontal, facial, epicranial, lateral, and genal hair-like trichomes (Figs. 12 B; 14 B) .............................................................................................. Araucnephia Head sclerite not with all trichomes mentioned; facial trichomes hair-like or spinelike ......................................................................................................................................... 8 Thorax with tubercles arranged in circles (Fig. 16 E); basal portion of gills with minute plate-like cuticular structures; thoracic trichomes hair-like ....................... ........................................................................................................................ Araucnephioides Thorax with tubercles not arranged in circles; basal portion of gills without aforementioned cuticular structures; thoracic trichomes hair-like or spine-like ................................................................................................................................................ 9 Cephalic sclerite with 2+2 or 3+3 frontal trichomes; branches of gills more or less tubular (Figs. 9 A-J) ........................................................................................ Mayacnephia Cephalic sclerite with 1+1 or without frontal trichomes; gill branches varied in shape ................................................................................................................................... 10 Frontal and labial-frontoclypeal trichomes absent (Fig. 20 N); clypeus comparatively narrow (Fig. 20 B) .................................................................. Gigantodax Frontal and labial-frontoclypeal trichomes present (Fig. 20 M); clypeus comparatively wide (Fig. 20 A) ...................................................................Pedrowygomyia

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Larvae 1 – 2(1) – 3(1) – 4(3) – 5(4)

–

6(5) – 7(3) – 8(7) –

Anal sclerite with accessory sclerite forming complete ring around posterior end of body (Fig. 20 S); mandibles each with 3 outer teeth (Fig. 20 P) ........................ 2 Ring-shaped accessory sclerite not developed (Fig. 12 N); mandibles each with 2 (Fig. 12 J) or 4 (Fig. 18 J) outer teeth ............................................................................ 3 Hypostomium with lateral tooth, projecting beyond level of first intermediate tooth (Fig. 20 R); cephalic apotome without basal spots ......................... Gigantodax Hypostomium with first intermediate tooth generally projecting beyond level of lateral tooth (Fig. 20 Q); cephalic larval apotome with basal spots (Fig. 20 O) ..... ...........................................................................................................................Pedrowygomyia Teeth of hypostomium arranged in 3 conspicuous groups (Fig. 8 W); mandible with marginal serrations numerous, more than 5 (Fig. 12 J) .................................... 4 Teeth of hypostomium more evenly distributed, not arranged in conspicuous groups (Figs. 18 H; 23 M; 25 I); mandible with marginal serrations less numerous, not more than 5, generally only 2 (Fig. 25 K) ............................................................. 7 Antennae approximately as long as stem of cephalic fan (Fig. 16 I); hypostomium with either 13 (Fig. 8 W) or 17 teeth (Fig. 16 N) ........................................................ 5 Antennae much shorter than stem of cephalic fan (Fig. 14 H); hypostomium with 15 teeth (Fig. 14 K) .......................................................................................... Araucnephia Cervical sclerite very small, isolated in membrane (Fig. 16 H); mandibles with accessory teeth at level of inner teeth, with second preapical tooth minute (Fig. 16 L) and with short basal setae slender, only faintly dentate; hypostomium with 17 teeth (Fig. 16 N) ......................................................................................... Araucnephioides Cervical sclerite small or transverse and isolated in membrane (Fig. 8 S), or transverse and fused with upper ends of postocciput (Fig. 10 O); mandibles without accessory serrations at level of inner teeth, with second preapical tooth as long as first (Fig. 10 M) or only slightly shorter (Fig. 8 U) and with short basal setae bearing conspicuous elongated denticles (Fig. 8 V); hypostomium with 13 teeth (Fig. 8 W) ........................ 6 Proximal antennal segment more than half as long as medial segment (Fig. 8 T) ............................................................................................................................. Mayacnephia Proximal antennal segment at most half as long as medial segment .......... Tlalocomyia Antenna much longer than stem of cephalic fan; distal antennal segment distinctly longer than proximal and medial segments combined (Figs. 22 I; 23 L); backwardly directed struts underlying main body of anal sclerite (Fig. 23 N) ................ Cnesiamima, Paraustrosimulium Antenna not longer than stem of cephalic fan (Fig. 27 H); distal segment not or only slightly longer than proximal and medial segments combined (Figs. 18 G; 25 J); anal sclerite without struts (Figs. 18 M; 25 L) ........................................................ 8 Postgenal cleft very shallow (Fig. 19 N); hypostomium with 17 teeth, the 4+4 intermediate ones similar to remaining teeth (Fig. 19 M); mandibles with 4 outer teeth (Fig. 18 J) ...........................................................................................................Cnesia Postgenal cleft well developed (Fig. 25 H); hypostomium with at most 13 teeth (Figs. 25 I; 27 J), the 1+1 lateral ones, when perceptible, somewhat similar to lateral serrations; mandible with 2 outer teeth (Figs. 25 K; 27 K) .......................... 9

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9(8) – 10(9)

–

CLAVES

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Preapical teeth of mandible subequal in size (Fig. 33 X), or decreasing in size from first to third (Fig. 38 G) ....................................................................................... Simulium Preapical teeth of mandible with second smaller than either first or third (Figs. 25 K; 27 K) ............................................................................................................................. 10 Body integument with (Fig. 25 N) or without setae; without cylindrical projections; rays of cephalic fan with longer setae alternately disposed among smaller setae; antenna with basal segments not overly wide (Fig. 25 J); antenna slightly shorter than or as long as stem of cephalic fan (Fig. 25 G); ninth abdominal segment with 1+1 ventral tubercles (Fig. 25 F); anal ring with about 75 rows and 13 hooks each ............................................................................... Lutzsimulium Body integument without setae; cylindrical projections present (Fig. 27 L); rays of cephalic fan with setae arranged homogeneously, without bigger setae; antenna with basal segments very wide (Fig. 27 I); antenna shorter than stem of cephalic fan (Fig. 27 H); ninth abdominal segment without ventral tubercles; anal ring with 110-130 rows and about 22 hooks each. ................................................. Kempfsimulium DE GÉNEROS DE SIMÚLIDOS DE LA REGIÓN

NEOTROPICAL

Adultos 1 – 2(1) – 3(2)

–

4(2) – 5(4)

Pedisulco bien desarrollado (Fig. 1 D: pe); célula basal siempre ausente; largo del sector basal de R (medido desde la nervadura transversal humeral) menos de 1/3 de la distancia desde la base de Rs al extremo del ala .................................. Simulium Pedisulco ausente (Figs. 8 G, N); célula basal ausente o presente; largo del sector basal de R aproximadamente 1/3 de la distancia mencionada o mayor ................ 2 Cu2 y A1 rectas (Fig. 20 C); célula basal ausente ......................................................... 3 Cu2 y A1 curvadas (Fig. 8 E); célula basal ausente (Fig. 26 F) o presente (Figs. 8 E; 15 G) ..................................................................................................................................... 4 Primer flagelómero tan largo como ancho; pelos del flagelómero basal y palpómeros más cortos que el artejo; clípeo más largo que ancho (Fig. 20 B); piezas bucales de la hembra normales; calcipala presente (Fig. 20 E); diente subbasal de la uña bien desarrollado (20 G) ............................................... Gigantodax Primer flagelómero tan largo como ancho; pelos del flagelómero basal y palpómeros más largos que el artejo; clípeo tan largo como ancho (Fig. 20 A); piezas bucales de la hembra, reducidas, no funcionales; calcipala ausente (Fig. 20 D); diente subbasal de la uña reducido (Fig. 20 F) ................................ Pedrowygomyia Porción superior de membrana pleural con pelos (Fig. 10 B); segmento apical del palpo maxilar más del doble de largo que el penúltimo ........................... Tlalocomyia Membrana pleural totalmente desnuda; segmento apical del palpo maxilar generalmente menos del doble de largo que el penúltimo (Figs. 13 C; 15 C; 21 B) ................................................................................................................................................ 5 Antenas con 10 artejos; ramas del furcasternum con conspicuas proyecciones (Fig. 23 A); genitalia del macho con esclerito geniculado, compuesto de una porción basal en forma de Y y 2 ramas subparalelas alargadas apicalmente (Fig. 23 H); porción apical de endoparámeros con numerosos dentículos alargados;

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horquilla genital casi enteramente sin pigmento, su pedúnculo muy robusto, no más largo que los brazos (Fig. 23 E) ..................................................Paraustrosimulium – Antenas con 11 artejos (Fig. 19 A); ramas del furcasternum sin conspicuas proyecciones (Figs. 8 D; 13 F) [excepto Lutzsimulium (Fig. 24 L)]; esclerito mediano del macho no como arriba, o porción apical del los endoparámeros reducida, faltando dentículos (Figs. 8 Q; 24 I); horquilla genital bien pigmentada, con pedúnculo no como arriba (Figs. 8 L; 10 E; 11 F; 13 M; 17 H, Q) ................ 6 6(5) R1 solamente con pelos como cerdas, sin cerdas con forma de espina (Fig. 21 F); diente basal de la uña de la hembra, grande (Fig. 21 J) ............................ Cnesiamima – R1 con pelos como cerdas y en forma de espinas (Figs. 13 H, I); si las cerdas en forma de espina son poco evidentes, el diente basal de la uña de la hembra es pequeña (Fig. 24 D) ........................................................................................................... 7 7(6) Célula basal ausente (Fig. 26 F); mandíbula de hembra dentada solo en el margen interno (similar a figura 26 D); uñas de la hembra con un pequeño diente subbasal (Figs. 24 D; 26 K); genitalia del macho con dientes no perceptibles en endoparámeros (Figs. 24 I; 27 A) ................................................................................... 8 – Célula basal presente (Fig. 8 E); mandíbula de la hembra como arriba o con dientes bien desarrollados en margen interno y externo (Fig. 13 D); uñas de la hembra con diente basal grande (Figs. 13 G; 15 I); genitalia del macho con dientes perceptibles en endoparámeros (Figs. 11 K; 13 T; 17 K: ed) ................................... 9 8(7) Frente más larga que ancha con surco mediano relativamente corto y sin ramas basales bifurcadas (Fig. 24 A); Sc con alrededor de 50 tricomas (Fig. 24 H); ramas del furcasternum con grandes proyecciones (Fig. 24 L) ..................... Lutzsimuliumm – Frente aproximadamente tan larga como ancha con surco mediano alargado y bifurcado, con largas ramas (Fig. 26 A); Sc sin tricomas (Fig. 26 F); ramas del furcasternum con muy pequeñas proyecciones (Fig. 26 L) ................. Kempfsimulium 9(7) Alas con leve curvatura de Cu2; A1 casi llegando al borde del ala (Fig. 17 B); genitalia del macho con porción basal del endoparámero poco desarrollada, y con dentículos de la porción distal bien desarrollados, pero pocos en número, formando un apretado grupo de disposición característica (Fig. 17 K); hembra con ramas de la horquilla genital con 2 apodemas dirigidos hacia delante (Figs. 17 H, Q) ............................................................................................................................. Cnesia – Alas con curvatura de Cu2 pronunciada; A1 terminando antes del margen (Figs. 8E) ; endoparámeros de machos diferentes a dilema anterior; horquilla genital sin apodemas (Figs. 8 L; 11 F) ............................................................................................. 10 10(9) Machos ............................................................................................................................... 11 – Hembras ............................................................................................................................. 13 11(10) Esclerito mediano con una placa saliente basal transversa continuada distalmente con una profunda incisión y bifurcada apicalmente (Fig. 8 R); endoparámeros sin dientes (similar a Fig. 8 Q) ............................................................................ Mayacnephia – Esclerito mediano de forma compleja curvada con la mitad distal dividida en 2 ramas anchas y cortas (Figs. 11 J, K; 13 R, S); endoparámeros con numerosos dientes (Fig. 11 K) ............................................................................................................ 12

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12(11) Genitalia con placa ventral subtriangular; ramas basales angostas y rectas (Figs. 11 I; 13 P); ramas del esclerito mediano paralelos (Fig. 13 S); endoparámeros con porción basal grande y dientes de la porción distal bien desarrollados (Figs. 11 K; 13 T) .................................................................................................................... Araucnephia – Genitalia con placa ventral subrectangular, sus ramas basales robustas, redondeadas en el ápice y curvadas (Figs. 15 P, Q); ramas del esclerito mediano notoriamente divergentes apicalmente (Fig. 15 U); endoparámeros con porción basal pequeña y dentículos de la porción apical reducidos ............... Araucnephioides 13(10) Diente basal de la uña en forma de gancho angosto (Fig. 15 I) ...... Araucnephioides – Diente basal de la uña más grande, subtriangular ó espatulado (Figs. 8 H, J; 13 G) .............................................................................................................................................. 14 14(13) Frente muy angosta (Fig. 8 A), ángulo frontal de 50° aproximadamente; superficie interna de la espermateca sin espículas (Meso América) ....................... Mayacnephia – Frente más ancha (Fig. 15 A); ángulo frontal de 85° aproximadamente; superficie interna de la espermateca con espículas dispersas (Sur de Suramérica) .................. .............................................................................................................................. Araucnephia Hembras, basadas en la genitalia 1 – 2(1) – 3(1) – 4(3) – 5(4) – 6(5)

–

Ramas de la horquilla genital con largos apodemas dirigidos hacia delante (Figs. 17 H, Q; 20 I, J) ................................................................................................................. 2 Ramas de la horquilla genital sin largos apodemas ..................................................... 3 Rama anterior de horquilla genital de ancho uniforme (Figs. 17 H, Q; 20 J.) ........ .................................................................................................................. Cnesia, Gigantodax Rama anterior de horquilla genital expandida en 1/3 basal (Fig. 20 I) .................... ...........................................................................................................................Pedrowygomyia Horquilla genital casi totalmente sin pigmento, su tallo ancho y muy corto, no más largo que las ramas (Fig. 23 E) ............................................................. Paraustrosimulium Horquilla genital no como arriba ................................................................................... 4 Esclerotización de la espermateca continuando en la base del ducto espermático (Figs. 26 P, Q) ..................................................................................................................... 5 Ducto espermático inserto en una pequeña área membranosa de la espermateca (Figs. 8 J; 21 L) ................................................................................................................... 8 Área de inserción del ducto espermático con una peculiar estructura como una red, extendiéndose a la base del ducto (Fig. 24 G) ..................................................... 6 No como arriba .................................................................................................................. 7 Área esclerotizada del ducto espermático también cubriendo el cuello de la espermateca y porción del ducto espermático con aspecto de malla (Fig. 24 G); espermateca internamente con espículas; octavo esternito con porción esclerotizada reducida al área mediana (Fig. 24 E) ................................. Lutzsimulium Área esclerotizada del ducto espermático reducida al cuello, sin aspecto de malla en la unión con espermateca (Figs. 26 P, Q); espermateca internamente sin espículas; octavo esternito con porción esclerotizada cubriendo todo el área desde un costado al otro (Fig. 26 M) ................................................................... Kempfsimulium

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ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Neotropical Simuliidae

Paraprocto muy pequeño, triangular; superficie de la espermateca conspicuamente reticulada; su superficie interna sin espículas; esclerotización del ducto espermático terminando abruptamente (Fig. 10 D) .................................. Tlalocomyia Paraprocto de tamaño normal, redondeado (Fig. 21 M); superficie de la espermateca lisa, su superficie interna con pocas espículas; porción esclerotizada del ducto espermático uniéndose gradualmente con la porción no pigmentada (Fig. 21 L) ........................................................................................................... Cnesiamima Superficie interna de la espermateca sin espículas ................................... Mayacnephia Superficie interna de la espermateca con espículas dispersas .................................. 9 Paraprocto aproximadamente tan largo como ancho (Fig. 13 L); superficie interna de la espermateca no rugosa ......................................................................... Araucnephia Paraprocto aproximadamente la mitad más corto que ancho (Fig.. 15 L); superficie interna de la espermateca rugosa ........................................................... Araucnephioides

Machos, basados en la genitalia 1 – 2(1)

–

3(1)

– 4(3) – 5(4)

Dientes de endoparámeros no perceptibles; esclerito mediano comparativamente simple, ensanchado basalmente, lamela alargada, apicalmente dividida en 2 cortas ramas conectadas por una membrana (Figs. 24 I; 27 A) ........................................... 2 Por lo menos una porción del endoparámero distinguible; esclerito mediano diferente, generalmente de estructura compleja .......................................................... 3 Gonocoxito alrededor de 2 veces más largo que el gonostilo; gonostilo suavemente aguzado distalmente; ápice relativamente romo con 3 (2) dientes terminales (Figs. 24 I, M); placa ventral distalmente con suave curvatura redondeada; esclerito mediano muy alargado (Fig. 24 J); pedisulco ausente .......... ............................................................................................................................ Lutzsimulium Gonocoxito alrededor de 3 veces más largo que el gonostilo; gonostilo fuertemente aguzado distalmente, con 2 dientes terminales (Figs. 27 A-C); placa ventral presentando distalmente un borde irregular (Figs. 27 A, D); esclerito mediano ensanchado, no alargardo, distal mente aguzado; pedisulco desde ausente a bien insinuado ............................................................................................ Kempfsimulium Porción basal de endoparámeros no visible o apenas muy poco desarrollado; porción denticulada compuesta de 1 o 2 grandes dientes y de varios pequeños dentículos muy estrechamente asociados, a veces dirigidos en opuesta dirección (Fig. 17K) ............................................................................................................................. 4 Porción basal de endoparámeros distinta, en forma de placa; dientes de la porción distal variando desde ausentes a bien desarrollados, pero no ordenados como fue descrito arriba ..................................................................................................................... 6 Placa ventral subpentagonal, aproximadamente tan larga como ancha, no con conspicuo reborde apical (Fig. 17 N) ....................................................................Cnesia Placa ventral en la mayoría de los casos en forma romboide alargado, con notorio reborde apical ...................................................................................................................... 5 Placa ventral corta, con márgenes laterales en la mitad de su largo cóncavos y margen apical bilobulado (Fig. 20 L); dientes de endoparámeros alargados .......... ............................................................................................................................... Gigantodax

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– 6(3)

– 7(6) – 8(7) – 9(7)

–

49

Placa ventral larga alrededor de 2 veces más larga que ancha, con márgenes laterales en la mitad de su largo rectas y margen apical recto y convexo (Fig. 20 K); dientes de endoparámeros cortos ..............................................................Pedrowygomyia Esclerito mediano compuesto de una placa basal saliente transversal continuada distalmente por una profunda incisión bifurcada apicalmente (Figs. 8 R; 10 G); endoparámeros reducidos a una porción basal en forma de placa y sin dentículos (Fig. 8Q) ....................................................................................... Mayacnephia, Tlalocomyia Esclerito mediano diferente; endoparámeros con o sin dientes .............................. 7 Esclerito mediano fuertemente curvo en vista lateral (Fig. 21 U), en vista de conjunto con forma aproximada de M (Figs. 21 V; 23 H), con ramas internas de la M parcialmente fusionadas y conectadas al reborde basal ........................................ 8 Esclerito mediano no como arriba, en forma de una estructura compleja ocupando 3 diferentes planos .......................................................................................... 9 Endoparámeros compuestos de una porción basal en forma de placa libre y apicalmente numerosos dentículos alargados subparalelos no conectados a la porción basal (Fig. 23 H) .......................................................................Paraustrosimulium Endoparámeros reducidos a una estructura subcuadrada en forma de placa; dentículos no desarrollados (Fig. 21 U) ....................................................... Cnesiamima Placa ventral subpentagonal o subtriangular (Fig. 11 I), con ramas basales en forma de delgadas varillas (Fig. 13 P); ramas de esclerito mediano paralelas (Fig. 13 S); endoparámeros con conspicuos dentículos en la porción distal (Fig. 11 K) .............................................................................................................................. Araucnephia Placa ventral subrectangular con ramas basales robustas apicalmente, redondeadas (Fig. 15 P); ramas del esclerito mediano divergiendo apicalmente (Fig. 15 U); endoparámeros con dentículos de la porción distal no desarrollados (Fig. 15 S) ..................................................................................................... Araucnephioides

Pupas 1

– 2(1) – 3(2)

Esternitos abdominales VI y VII divididos longitudinalmente a lo largo del medio por un área estriada, membranosa (Figs. 14 D; 18 D); ganchos de los tergitos VIVIII invariablemente simple; segmentos abdominales VIII y IX en muchos casos con cerdas robustas curvas, enruladas bífidas o en forma de gancho (Figs. 18 E; 23 J; 25 E) ............................................................................................................................ 2 Esternitos VI y VII del abdomen enteros; en muchos casos con cerdas bífidas o trífidas sobre los tergitos VI y VIII; tergitos VIII y IX sin las setas mencionadas ................................................................................................................................... Simulium Proceso terminal del abdomen corto y puntiagudo (Fig. 23 J), o ausente (Fig. 10 L) ................................................................................................................................................ 3 Proceso terminal del abdomen alargado (Figs. 12 E; 18 E; 25 E) .......................... 4 Capullo reducido a una pequeña capa ubicada sobre los segmentos terminales del abdomen (Fig. 10 H); branquias en forma de un grueso tallo con unos pocos filamentos en forma de hebras (Fig. 10 I ); abdomen fuertemente esclerotizado, tergitos y esternitos con un número elevado, supernumerario de espinas o ganchos (Figs. 10 J, K); ápice del abdomen romo; procesos terminales ausentes (Fig. 10 L) .............................. Tlalocomyia

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4(2) – 5(4) – 6(5) – 7(5) – 8(7) – 9(8) – 10(9) –

Capullo bien desarrollado, de forma definida, cubriendo todo el cuerpo de la pupa excepto branquias, las que son dos, en forma de lámina y pseudosegmentadas (Figs. 22 C; 23 I); abdomen menos esclerotizado que el tórax; abdomen sin espinas supernumerarias y con cortos procesos terminales (Fig. 23 J) ..................................... ............................................................................................... Cnesiamima, Paraustrosimulium Tergitos abdominales con peines de espinas (Fig. 14 D); tergito VIII sin grandes ganchos ................................................................................................................................. 5 Tergitos abdominales sin peines de espinas; tergito VIII con 4 grandes ganchos ........................................................................................................................................ Cnesia Tricomas faciales (Figs. 12 B; 27 E), tricomas torácicos (Figs. 12 D; 27 F, G) y cerdas del octavo y noveno segmento abdominal notoriamente curvado apicalmente (Fig. 25 E) ..................................................................................................... 6 Al menos tricomas faciales y torácicos no curvado apicalmente ............................ 7 Branquias con 12-22 ramas; frontoclípeo y tórax con tubérculos (Fig. 25 D) ....... ............................................................................................................................ Lutzsimulium Branquias con 12 ramas; frontoclípeo y tórax sin tubérculos (Fig. 27 E) ............... .......................................................................................................................... Kempfsimulium Esclerito cefálico con tricomas frontal, facial, epicranial, lateral y genal en forma de pelo (Figs. 12 B; 14 B) ............................................................................... Araucnephia Esclerito cefálico no con todos los tricomas mencionados; tricomas faciales con forma de pelo o de espina ................................................................................................ 8 Tórax con tubérculos ordenados en círculos (Fig. 16 E); porción basal de las branquias con pequeñas estructuras cuticulares en forma de placa; tricomas torácicos en forma de pelo ...................................................................... Araucnephioides Tórax con tubérculos no ordenados en círculos; porción basal de las branquias sin las estructuras mencionadas; tricomas torácicos en forma de pelo o espina ........... 9 Esclerito cefálico con 2+2 o 3+3 tricomas frontales; ramas de las branquias más o menos tubulares (Figs. 9 A-J) ....................................................................... Mayacnephia Esclerito cefálico 1+1 o sin tricomas faciales; ramas de las branquias de forma variada ................................................................................................................................. 10 Frontoclípeo sin tricomas frontal y labial (Fig. 20 N); clípeo comparativamente angosto (Fig. 20 B) ............................................................................................ Gigantodax Frontoclípeo con tricomas frontal y facial (Fig. 20 M); clípeo comparativamente ancho (Fig. 20 A) ...........................................................................................Pedrowygomyia

Larvas 1 – 2(1)

Esclerito anal con esclerito accesorio formando un anillo completo alrededor de la porción posterior del cuerpo (Fig. 20 S); mandíbulas con 3 dientes externos (Fig. 20 P) ............................................................................................................................ 2 Esclerito accesorio en forma de anillo no desarrollado (Fig. 12 N); mandíbulas con 2 (Fig. 12 J) o 4 dientes externos (Fig. 18 J) ........................................................ 3 Hipostomio con diente lateral proyectándose sobre el nivel del primer diente intermedio (Fig. 20 R); apotoma cefálico sin manchas basales ............... Gigantodax

ABLA Vol. 3 – Sixto Coscarón & Cecilia L. Coscarón Arias: Simuliidae Neotropicales

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6(5) – 7(3)

– 8(7) –

9(8) –

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Hipostomio con primer diente intermedio generalmente proyectado por sobre el nivel del diente lateral (Fig. 20 Q); apotoma cefálico con manchas basales (Fig. 20 O) ......................................................................................................................Pedrowygomyia Dientes del hipostomio ordenados en 3 grupos conspicuos (Fig. 8 W); serrulaciones del margen de la mandíbula numerosas más de 5 (Fig. 12 J) .......... 4 Dientes del hipostomio uniformemente distribuidos, no ordenados en grupos conspicuos (Figs. 18 H; 23 M; 25 I); serrulaciones del margen de las mandíbulas menos numerosas, no mayor de 5, generalmente solo 2 (Fig. 25 K) ...................... 7 Antena aproximadamente tan larga como el tallo del abanico cefálico (Fig. 16 I); hipostomio con 13 (Fig. 8 W) o 17 dientes (Fig. 16 N) ............................................. 5 Antena más corta que el tallo del abanico cefálico (Fig. 14 H); hipostomio con 15 dientes (Fig. 14 K) ............................................................................................ Araucnephia Escleritos cervicales muy pequeños, aislados en la membrana del cuello (Fig.16 H); mandíbulas con dientes accesorios al nivel de los dientes internos, con el segundo diente preapical pequeño (Fig. 16 L) y con cerdas basales cortas y finas; dentición del hipostomio corta, con 17 dientes (Fig. 16 N) ............. Araucnephioides Escleritos cervicales pequeños o transversales y aislados en la membrana del cuello (Fig. 8 S), o transversales y fusionados con los bordes superiores del occiput (Fig. 10 O); mandíbulas sin dientes accesorios al nivel de los dientes internos, con segundo diente preapical tan largo como el primero (Fig. 10 M) o ligeramente más corto (Fig. 8 U ) y con cortas cerdas basales llevando conspicuos dentículos alargados (Fig. 8 V); hipostomio con 13 dientes (Fig. 8 W) ..................................... 6 Artejo proximal de la antena más de la mitad del largo del mediano (Fig. 8 T) .... ............................................................................................................................. Mayacnephia Artejo proximal de la antena por lo menos tan largo como el mediano ....... Tlalocomyia Antena más larga de que el tallo del abanico cefálico; artejo distal de la antena notoriamente más largo que el proximal y mediano juntos (Figs. 22 I; 23 L); sostenes dirigidos hacia atrás a los costados del cuerpo principal del esclerito anal (Fig. 23 N) ............................................................................................... Cnesiamima, Paraustrosimulium Antena no más larga que el tallo del abanico cefálico (Fig. 27 H); artejo distal no o solamente ligeramente más largo que el proximal y mediano juntos (Figs. 18 G; 25 J); esclerito anal sin los mencionados sostenes (Figs. 18 M; 25 L) .......................... 8 Hendidura postgenal muy superficial (Fig. 19 N); hipostomio con 17 dientes, los 4+4 intermedios similares a los restantes dientes (Fig. 19 M); mandíbula con 4 dientes externos (Fig. 18 J) ......................................................................................Cnesia Hendidura postgenal bien desarrollada (Fig. 25 H); hipostomio cuando más 13 dientes (Figs. 25 I; 27 J), los 1+1 laterales, cuando son perceptibles, se parecen a las serrulaciones laterales; mandíbula con 2 dientes externos (Figs. 25 K; 27 K) ................................................................................................................................................ 9 Dientes preapicales de mandíbula subiguales en tamaño (Fig. 33 X), o decreciendo en tamaño del primero al tercero (Fig. 38 g) .................................................. Simulium Dientes preapicales de la mandíbula con el segundo diente más pequeño que el primero o el tercero (Figs. 25 K; 27 K) ...................................................................... 10

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Tegumento del cuerpo con (Fig. 25 N ) o sin cerdas; sin proyecciones epidérmicas cilíndricas; rayos de los abanicos cefálicos con dientes más largos alternativamente dispuestos entre otros menores; antena con artejo basales no notoriamente gruesos (Fig. 25 J); antena ligeramente más corta que el largo del abanico cefálico (Fig. 25 G); noveno segmento abdominal con 1+1 tubérculo ventral (Fig. 25 F); anillo anal con alrededor de 75 hileras con 13 ganchos cada una ............................. Lutzsimulium Tegumento del cuerpo sin cerdas; presencia de proyecciones epidérmicas cilíndricas (Fig. 27 L); rayos del abanico cefálico con dientes ordenados homogéneamente sin dientes más largos; antena con artejos basales notoriamente más gruesos (Fig. 27 I); antena más corta que el tallo del abanico cefálico (Fig. 27 H); noveno segmento abdominal sin tubérculos ventrales; anillo anal con 110-130 hileras de alrededor de 22 ganchos cada una ......................................... Kempfsimulium

Mayacnephia WYGODZINSKY & COSCARÓN (Figs. 8, 136.) Mayacnephia WYGODZINSKY & COSCARÓN, 1973a: 144-148. Type-species: Simulium pachecolunai DE LEON, 1945.

Adults: Antennae with 11 segments; male holoptic. Female with frons narrow (Fig. 8 A), frontal angle 50 degrees; mandibles and maxillae of female normally developed, mandibles with teeth on both margins; apical segment of maxillary palp in both sexes less than twice as long as penultimate (Fig. 8 B); sensory vesicle well developed, with abundant tubercles (Fig. 8 C); mesepisternal sulcus wide, shallow, evanescent anteriorly; katepisternum glabrous, in profile about as long as high; pleural membrane glabrous; branches of furcasternum lacking projections (Fig. 8 D); metanotum glabrous; wings with basal cell present (Fig. 8 E); R1 with hair and spine like setae (Fig. 8 F); basal section of R hairy; Rs hairy below and hairy above on its apical half, beginning at level of junction of R1 with C; Rs not forked; M appearing duplicated; Cu2 conspicuously curved; A2 ending well before wing margin; calcipala present, bluntly rounded apically (Fig. 8 G); hind basitarsus of male strongly widened (Fig. 8 N); claws of female with large, bluntly subtriangular basal tooth (Figs. 8 H, I). Female with spermatheca normally sclerotized, with small depression on one side, reticulate and without spicules internally, with small circular membranous area at insertion of spermathecal duct, the latter not sclerotized (Fig. 8 J); gonapophysis large, subtriangular, apices rounded, medial borders approximated, surface with numerous microtrichia and a few setae (Fig. 8 K); lateral arms of genital fork with small anterior processes and expanded distally into subrectangular plate (Fig. 8 L); stem slender, at least as long as arms; paraproct short with rounded rim (Fig. 8 M); male with gonostylus about as long as gonocoxite, conical, not flattened with two apical spines (Fig. 8 O); ventral plate somewhat wider than long, subsemicircular, setose (Fig. 8 P). Endoparameres reduced to plate like sclerite; denticles absent or only weak (Fig. 8 Q); median sclerite with transverse basal flange, deeply narrow incision and bifurcated apically (Fig. 8 R). Pupa: Cocoon irregularly shaped, covering part or all of body; body of pupa sclerotized throughout; head, and thorax with pterothecae, and abdominal segments tuberculate; gills with 2-15 thickened, tubular, occasionally divided (or basally fused) branches, arising from short common base, all with superficial but mostly conspicuous

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bc

E Sc

C

A

B

C R1

Rs

F H

G D

N

I M K L T J

O

P

R U

V

Q W

X

S

Figure 8. A-Q, S-X: Mayacnephia aguirrei; A-M: Female; A: frons; B: maxillary palp; C: sensory vesicle of maxillary palp with high magnification; D: furcasternum; E: wing, general view (bc = basal cell); F: portion of anterior wing veins, showing pilosity (C = costa, R1= radius 1, Rs = secondary radius (radius 2), Sc = subcosta); G: calcipala and tarsomere I; H-I: claw with large basal tooth; J: spermatheca; K: eighth sternite and gonapophyses; L: genital fork; M: cercus and paraproct; N-Q: Male; N: calcipala and tarsomere I; O: gonocoxite and gonostylus; P: ventral plate, with portion of median sclerite; Q: endoparameres with portion of median sclerite and aedeagus membrane; R: median sclerite positioned on ventral plate; S-X: Larva; S: cervical sclerite; T: antenna; U: teeth of mandible; V: short basal setae of mandible; W: teeth of hypostomium; X: lateral sclerite of proleg. R: M. roblesi; ventral plate and median sclerite. (A-X from WYGODZINSKY & COSCARÓN, 1973a).

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transverse annulations with spiculae; head with 2+2 or 3+3 frontal and 1+1 facial trichomes; all hair like. Thorax with approximately five simple trichomes; abdomen pointed posteriorly. Distinct pleural plate absent; abdominal segments without perceptible transversal ridges; terga III and IV with 4+4 or 5+5 hooks posteriorly, in some cases 5+5 hooks also on tergum II, and 1+1 hooks on V. Terga V-IX, or VI-IX, with continuous spine-combs anteriorly; faint spine-combs also perceptible on III and IV; sternum IV with 1+1 strong setae or hooks; sternum V with from 2+2 to 4+4 hooks; sternal plates VI and VII either each with 1+1 hooks accompanied by no or 1+1 hooks in pleural membrane, or sternal plates VI with 3+3 and sternal plates VII with 2+2 hooks, accompanied by 1+1, 2+2 or 3+3 hooks in pleural membrane; all hooks simple. Terminal hooks well developed, relatively stout, not sinuous; posterior segments with few irregularly formed hook- or grapnel-shaped setae. Larva: Head spots positive; cephalic apotome broadest well before hind margin; cervical sclerites transverse, small or large, either free or fused to upper ends of postocciput (Fig. 8 S); postgenal cleft very shallow; antennae as long as stern of cephalic fans; third segment slightly darker than remaining. First antennal segment over half as long as second (Fig. 8 T); cephalic fans normal; mandibles with two outer and one apical tooth; third preapical tooth largest, but not so large as apical tooth; inner teeth arranged in single row; mandibular serrations very numerous (15-22) (Fig. 8 U); short basal setae stout, with conspicuous denticles along almost their entire length (Fig. 8 V); hypostomium with 13 teeth arranged in three conspicuous groups, with median tooth and lateral teeth by far largest; 2+2 lateral teeth situated only slightly below level of remaining teeth (Fig. 8 W); lateral sclerite of thoracic proleg well developed, about as long as wide, subrectangular, lower border subparallel to longitudinal axis of proleg; setae of lateral sclerite less than half as long as sclerite, arranged in irregular double row, this row extending slightly beyond sclerite below (Fig. 8 X); larval cuticle bare, except simple setae at base of anal sclerite, and few perianal scales; anal sclerite normal, X-shaped. Terminal ventral tubercles absent or present. Distribution: Canada: Alberta, British Columbia, Yukon; Highlands of Mexico to Panama; USA: Arizona, California, Idaho, Oregon, Washington (Fig. 136). Discussion: This genus comprises 15 species, 11 of which are found in the Neotropical Region south of the United States. Mayacnephia is most closely related to Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA; the two genera form a monophyletic group, Tlalocomyia being the more apomorphic component. ADLER et al. (2004) synonymized Mayacnephia with Tlalocomyia. Here they are maintained separate. One Neotropical species, Mayacnephia alticola (DYAR & SHANON), is recorded from Mexico, but we could not obtain more information and do not consider it to be a recognizable species. KEY

TO

MAYACNEPHIA

SPECIES

We include 10 species from Mexico to Panama considered as Mayacnephia by the authors. We present only a key to pupae, which have good characters, especially in the shape of the respiratory organs that help to identify species; our key is in part based on that of RAMÍREZ PÉREZ et al. (1988), and includes the known Mayacnephia species at this time. We don’t include keys to females, males or larvae because of the lack of enough specimens; based on the available descriptions, these have very similar characters. Females

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are unknown in Mayacnephia atzopensis and M. mixensis, and males, in addition to these two species, also are unknown in Mayacnephia muzquicensis and M. tadai. DALMAT (1955) gave keys to larva, pupae, males and females of three species: Mayacnephia aguirrei, M. pachecolunai and M. roblesi. DÍAZ NÁJERA (1962) made a larval key to five species, including the three species presented by DALMAT, as well as Mayacnephia atzopensis and M. mixensis. The characters used by DÍAZ NÁJERA show great variation and do not offer security in identification. Only distinctive characters are considered in the species descriptions. Those characters not mentioned are in accordance with the generic description. Key to Mayacnephia pupae 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(4) – 7(6) – 8(2) – 9(8) –

Gill with 2 tubular and relatively swollen and elongated branches (Fig. 9 A) ........ ................................................................................................................ Mayacnephia aguirrei Gill with variable shape, 3-15 branches ......................................................................... 2 Gill with 3-7 branches ....................................................................................................... 3 Gill with 11-15 branches .................................................................................................. 8 Gill with 3 branches (Fig. 9 E), (based on Fig. 20 of DÍAZ NÁJERA, 1962) ............. ...............................................................................................................Mayacnephia mixensis Gill with 4-7 branches ....................................................................................................... 4 Gill with 4 branches .......................................................................................................... 5 Gill with 6-7 branches ....................................................................................................... 6 Gill tubular, subannulated, apically acuminate (Fig. 9 C) ........... Mayacnephia grenieri Gill swollen, smooth, apically rounded (Fig. 9 I) (based on Fig. 2 of RAMÍREZ PÉREZ et al., 1988) ................................................................................. Mayacnephia salasi Gill with 6 smooth tubular branches with short stalks, subequal in length and thickness (Fig. 9 J), (based on Fig. 6 of OCHOA & TAKAOKA, 1993) ......................... .................................................................................................................... Mayacnephia tadai Gill with 7 terminal branches .......................................................................................... 7 Gill with terminal filamentous branches originating each in number of 1 or 2 on 4 thick tubular subannulated stalks (Fig. 9 H) ................................... Mayacnephia roblesi Gill with terminal clavate branches originating on 4 relatively thin tubular and smooth stalks (sometimes with thumb-like process on anterior internal branch) (Fig. 9 D), (based on Fig. 8 of PETERSEN, 1985) ..................... Mayacnephia fortunensis Gill with 11 terminal filaments of different lengths, with thick base and gradually becoming thin to apex, (Fig. 9 B) (based on Fig. 9 of DÍAZ NÁJERA, 1962) ........... ............................................................................................................ Mayacnephia atzopensis Gill with 14 to 15 filaments ............................................................................................. 9 Gill with 12 or 13 elongated filamentous branches and 1 medial internal short, horizontally disposed (Fig. 9 G) ............................................... Mayacnephia pachecolunai Gill with 14 or 15 arborescent filamentous branches emerging from 4 thick stalks (Fig. 9 F), (based on Fig. 18 of DÍAZ NÁJERA, 1971 ........... Mayacnephia muzquicensis

Clave de especies de Mayacnephia basada en pupas 1

Branquias tubulares y con 2 ramas relativamente infladas y alargadas (Fig. 9 A) ................................................................................................................ Mayacnephia aguirrei

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– 2(1) – 3(2) – 4(3) – 5(4) – 6(4) – 7(6) – 8(2) – 9(8) –

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Branquias con 3-15 ramas ................................................................................................ 2 Branquias con 3-7 ramas .................................................................................................. 3 Branquias con 11-15 ramas .............................................................................................. 8 Branquias con 3 ramas (Fig. 9 E) (basada en figura 20 de DÍAZ NÁJERA, 1962) .... ...............................................................................................................Mayacnephia mixensis Branquias con 4-7 ramas .................................................................................................. 4 Branquia con 4 ramas ........................................................................................................ 5 Branquia con 6-7 ramas .................................................................................................... 6 Branquias tubulares, subanilladas, apicalmente aguzadas (Fig. 9 C) .......................... ................................................................................................................. Mayacnephia grenieri Branquias infladas, lisas, apicalmente redondeadas (Fig. 9 I) (basada en figura 2 de RAMÍREZ PÉREZ et al., 1988) ................................................................. Mayacnephia salasi Branquias con 6 ramas tubulares con corto tallo, casi iguales en longitud y grosor, y con superficie lisa (Fig. 9 J) (basado en OCHOA & TAKAOKA, 1993) ...................... .................................................................................................................... Mayacnephia tadai Branquias con 7 ramas terminales .................................................................................. 7 Branquias con ramas terminales filamentosas originadas cada una en número de 1 o 2 sobre 4 tallos tubulares gruesos subanillados (Fig. 9 H) ...... Mayacnephia roblesi Branquias con ramas terminales en forma de clava, originadas sobre 4 tallos relativamente finos, tubulares y lisos (Fig. 9 D) (basado en figura 8 de PETERSEN, 1985) ................................................................................................. Mayacnephia fortunensis Branquias con 11 filamentos terminales de diferente longitud con gruesa base y afinándose suavemente hacia el ápice (Fig. 9 B) (basado en figura 9 de DÍAZ NÁJERA, 1962) .................................................................................. Mayacnephia atzopensis Branquia con 14-15 filamentos terminales ................................................................... 9 Branquia con 12-13 ramas filamentosas, alargadas y una mediana interna corta, dispuesta horizontalmente (Fig. 9 G) ..................................... Mayacnephia pachecolunai Branquia con 14-15 ramas filamentosas, arborescentes, emergiendo de 4 tallos gruesos (Fig. 9 F) (basado en DÍAZ NÁJERA, 1971) ............. Mayacnephia muzquicensis

Mayacnephia aguirrei (DALMAT) (Figs. 8 A-Q, S-W; 9 A.) Simulium aguirrei DALMAT, 1949: 544. Cnephia aguirrei DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1952: 53; DALMAT, 1955: 89; VARGAS & DÍAZ NÁJERA, 1957b: 268; DÍAZ NÁJERA, 1962: 273. Mayacnephia aguirrei WYGODZINSKY & COSCARÓN, 1973a: 149-150; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on DALMAT, 1955: 89). Female: Wing length 3.4 mm; frons, clypeus and palpi blackish; scutum velvety black with gray pruinosity, with appressed silvery and few yellow hairs, metanotum velvety dark brown; legs mostly dark brown; abdominal tergites dark brown, with silvery white hair, sternites yellowish brown; frons narrow (Fig. 8 A); maxillary palp as in figure 8 B, with sensory vesicle (Fig. 8 C) about 1/3 of basal segment length; furcasternum as in figure 8 D; wing as in figure 8 E, with basal cell well developed; Sc completely pilose; hair and spinelike setae of C, R1 and Rs as in figure 8 F; calcipala as in figure 8 G; claw subbasal

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tooth elongate (Fig. 8 H-I); spermatheca as in figure 8 J, spermathecal duct ending in a membranous area; eighth sternite with about 19 trichomes (Fig. 8 K); genital fork as in figure 8 L; paraproct subovoidal distally (Fig. 8 M). Male: Wing length 3.0 mm; clypeus and palpi blackish; scutum velvety black with short black pilosity; metanotum velvety black; legs dark brown; abdominal segments I-II blackish, others velvety dark brown to black, with posterior margin brown to white pruinose; Sc pilose on basal third; length/width ratio of hind basitarsus III = 3.1; gonocoxite and gonostylus as in figure 8 O; ventral plate, endoparameres and median sclerite as in figures 8 P, Q, R. Pupa: Cocoon length at base 2.0 mm covering one half of abdomen, weave composed of loose threads; thorax and abdomen with abundant granulosity; gills composed of 2 long, tubular, subannulated branches tapered at apex (Fig. 9 A); maximum length 2.5 mm. Larva: Total length 6.3 mm; general color grayish, with very long dark patches on segments 6 and 7; cephalic fan with 30-35 rays; cervical sclerite as in figure 8 S; antenna as in figure 8 T; ratio of proximal, medial and distal segments of antenna = 1:1.2:1.7; mandible with 23 marginal teeth (Fig. 8 U), basal setae of mandible as in figure 8 V; hypostomium with 13 apical teeth (Fig. 8 W), 3 hairs on each side and laterally with 6 serrations; anal ring with 72-75 rows of hooks; area between anal sclerite arms with few bifid and trifid scales; eighth sternite with 2 tubercles; thoracic proleg sclerite with about 28 setae (Fig. 8 X). Distribution: Highlands of Guatemala; Mexico: Chiapas. Bionomics: DALMAT (1955) reported that this species was found between 900 and 1825 m, most frequently above 1525 m. The aquatic instars occurred in streams with a temperatures ranging between 14 °C and 23 °C, with most specimens at 19 °C and below. None of the streams where this species breeds was wider than 1.5 m, and almost all were no deeper than 30 cm. The preferred current was between 2.5 and 25 cm per second, and never over 50 cm per second. Pupae preferred stones or rocks as a substrate. Less than 10 percent were found on leaves and twigs. Mayacnephia atzopensis (DÍAZ NÁJERA) (Fig. 9 B.) Cnephia atzopensis DÍAZ NÁJERA, 1962: 273. Mayacnephia atzopensis WYGODZINSKY & COSCARÓN, 1973a: 150.

(Based on original description). Female, Male and Pupa: Unknown. Larva: Body length 9.0 mm; general coloration yellowish; cephalic fan with 25 rays; mandible with marginal serrations of about 21 teeth; hypostomium with 13 teeth, 6 or 7 lateral setae on each side; ratio of proximal, medial and distal segments of antenna = 1:1.3:2.1; pupal gill histoblast with 11 tubular branches of irregular length, with two pairs on each side with well-developed stalk (Fig. 9 B); ninth abdominal segment without ventral tubercles; anal ring with 78-80 rows of hooks; area of anal sclerite with scarce thin spines. Distribution: Mexico: Morelos. Bionomics: Larvae live in small creeks, in shaded and cold water. Discussion: The peculiar shape of the gill histoblast allows this species to be differentiated from other species.

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C A

B

D

E

F

H

G

I

J

Figure 9. Mayacnephia. A-J: Gills; A: M. aguirrei; B: M. atzopensis; C: M. grenieri; D: M. fortunensis E: M. mixensis; F: M. muzquicensis; G: M. pachecolunai; H: M. roblesi; I: M. salasi; J: M. tadai.

Mayacnephia fortunensis PETERSEN (Fig. 9 D.) Mayacnephia fortunensis PETERSEN, 1985: 80; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on original description). Female: Wing length 3.2 mm; antennae, frons and clypeus brown, palp dark brown, scutum dark brown densely covered with yellowish hair, legs brown, with tarsi light brown; Sc sparsely pilose. Male: Wing length 2.5 mm; general color dark brown; clypeus brown with silver pruinosity and pale yellowish to brown hair; antenna light brown with yellowish base; palp dark brown; legs as in female; abdominal tergites dark brown with distal border light brown.

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Pupa: Cocoon roughly pocket-shaped, loosely woven; gill with 7 thick tubular branches emerging from 4 primary branches (Fig. 9 D), sometimes with small internal protuberance like a short process on frontal internal branch; head and thorax with small tubercles in irregular distribution. Larva: Body length about 7.5 mm; general color dull cream yellow mottled from gray green to dark brown; hypostomium with 13 teeth; mandible with marginal serrations of about 22 teeth. Distribution: Panama: Chiriqui. Bionomics: Larvae live in small streams less than 1 m wide, on roots, leaves or rocks, in shaded areas at water temperatures of 14-20 °C. Mayacnephia grenieri (VARGAS & DÍAZ NÁJERA) (Fig. 9 C.) Cnephia grenieri VARGAS & DÍAZ NÁJERA, 1948b: 325; 1957b: 270; DÍAZ NÁJERA, 1962: 273. Mayacnephia grenieri WYGODZINSKY & COSCARÓN, 1973a: 149; CROSSKEY & HOWARD, 1997: 16.

(Based on original description). Female: Wing length 3.5 mm; general coloration blackish; frons black with golden pilosity, clypeus and palp black; scutum black with light brown pilosity, pleura black, legs blackish with black hair and tarsi whitish; abdomen black, tergites laterally with silvery pilosity; Sc with hair on entire length. Male: Wing length 3.0 mm; general coloration as in female; scutum uniformly black; abdomen whitish with silver pilosity; tergites blackish elongated; sternites also with blackish coloration; Sc without hair. Pupa: Cocoon length 3.0 mm, conical in shape, covering only abdomen; body integument granulose; gill with four tubular elongated branches disposed in two pairs with primary branches very short and emerging from small stalk (Fig. 9 C). Larva: Total length 8.0-9.0 mm; general coloration light yellowish and darkened distally on ventral side; antenna as long as stem of cephalic fan; hypostomium shape typical for genus but with 9 teeth; 2 or 3 lateral setae on each side, and marginal serrations with 5 small teeth; anal ring with 76 rows of hooks. Distribution: Mexico: Veracruz. Bionomics: The aquatic instars of this species were found at an altitude of 1500 m, in a small clear stream 60 cm below its origin from a spring. Mayacnephia mixensis (DÍAZ NÁJERA) (Fig. 9 E.) Cnephia mixensis DÍAZ NÁJERA, 1962: 284. Mayacnephia mixensis WYGODZINSKY & COSCARÓN, 1973a: 150; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on original description). Female, Male and Pupa: Unknown. Larva: Length 8.0 mm; general color yellowish; cephalic fan with 28 rays; mandible with marginal serrations of about 16 teeth; hypostomium with 15 teeth, 3 or 4 lateral setae on each side; ratio of proximal, medial and distal segments of antenna = 1:1.5:2.2; pupal gill histoblast with 3 thick tubular branches emerging from same place on very short stalk

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(Fig. 9 E); ninth sternite without ventral tubercles; anal ring with 94 rows of hooks; area of anal sclerite without scales or spines. Distribution: Mexico: Oaxaca. Bionomics: Larval habitats unknown. Discussion: The peculiar pupal gill shape with 3 branches allows us to separate Mayacnephia mixensis from all the known Mayacnephia species. Mayacnephia muzquicensis (DÍAZ NÁJERA) (Fig. 9 F.) Cnephia (Cnephia) muzquicensis DÍAZ NÁJERA, 1971: 239. Mayacnephia muzquicensis PY-DANIEL, 1982: 311; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on DÍAZ NÁJERA, 1971). Female: Wing length 4.1 mm; general coloration blackish; clypeus yellowish; palp blackish; scutum black with coppery shine and golden pilosity; metanotum reddish; pleura blackish; abdomen blackish; Sc mostly haired. Male: Unknown. Pupa: Cocoon length 4.0 mm, weave soft, without definite shape; thorax with scarce granulosity; gill arborescent with 14 or 15 filamentous branches emerging from 7 bifid primary branches from thick stalk (Fig. 9 F). Larva: Body length 8.5 mm; color darkened dorsally and whitish ventrally; head black; cephalic fan with 27 rays; mandible with about 14 marginal teeth; ratio of proximal, medial and distal segments of antennae = 1:2.5:2.4; hypostomium with 13 teeth; 5 or 6 lateral setae on each side; without scales; ninth segment without ventral tubercles. Distribution: Mexico: Coahuila. Bionomics: Larvae live on stones in small creeks of clear and cold water, without vegetation. Feeding habits of the females are unknown. Mayacnephia pachecolunai (DE LEÓN) (Fig. 9 G.) Simulium pachecolunai DE LEÓN, 1945: 67. Cnephia pachecolunai DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1954: 67; DALMAT, 1955: 43; DÍAZ NÁJERA, 1962: 273. Mayacnephia pachecolunai WYGODZINSKY & COSCARÓN, 1973a: 148; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on DALMAT, 1955). Female: Wing length 4.1 mm; frons, clypeus, palpi and scutum blackish with gray pruinosity; pleura dark brown, legs brown and abdomen blackish with posterior margin brown; Sc pilose along basal three fourths. Male: Wing length 3.6 mm; clypeus dark brown, palpi black, scutum velvety black with long yellow and black hair, pleura dark brown to black, legs dark brown to black on tarsi; abdomen velvety brown to black with posterior margin white-pruinose with black hair; length/width ratio of hind basitarsus = 3.1-3.4. Pupa: Cocoon length at base 4.0 mm, weave very soft without regular shape and appearing to cling to pupa, covering abdomen and one fourth of thorax; thorax and abdomen

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covered by tubercles; gills with 12-13 filamentous branches with median interior shorter; branches flattened in one plane arising a little behind anterior margin of thorax (Fig. 9 G). Larva: Length 5.9 mm; general color yellowish brown; cephalic fan with 25 branches; mandible with 25 marginal teeth; hypostomium with 13 teeth and 3 or 4 setae, lateral margins with 8 serrations; anal ring with 80-81 rows of hooks; ninth abdominal segment with 2 ventral tubercles; area between anal sclerite arms without scales. Distribution: Guatemala: Solola and Totonicapan; Mexico: Chiapas. Bionomics: The following data are based on DALMAT (1955). One adult was found to bite humans, but the real hosts of this species are not known. Normally, Mayacnephia pachecolunai occurs only above 2100 m and breeds in cold streams of not more than 17 °C, although the concentration of specimens is greatest at water temperatures below 11 °C. The breeding grounds were “almost entirely minute trickles of water that pass over a swampy area supporting large quantities of vegetation; such rivulets usually dried up during the months of November through March.” These rivulets were most frequently less than 15 cm wide and 13 cm or less deep; the current was generally very slow, less than 38 cm per second. About half of the pupae collected were attached to rocks, and the remainder to leaves or some few to branches or twigs. Mayacnephia roblesi (DE LEÓN) (Figs. 8 R, 9 H.) Simulium roblesi DE LEÓN, 1943: 97; 1945: 67. Cnephia roblesi DALMAT, 1951: 48; VARGAS & DÍAZ NÁJERA, 1954: 64; DALMAT, 1955: 97. Mayacnephia roblesi WYGODZINSKY & COSCARÓN, 1973a: 149; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

Female: Wing length 4.3 mm; frons and clypeus dark brown with gray pruinosity, scutum black with golden colored hair, with 3 longitudinal stripes of gray pruinosity; pleura brown with gray pruinosity, legs dark brown with femur a little lighter; abdomen dark brown with gray pruinosity, tergites with black or brown hair; Sc completely pilose. Male: Wing length 4.0 mm; palpi and clypeus black; scutum velvety black with appressed golden hair; pleura brown with gray pruinosity; legs mostly dark brown; abdominal tergites velvety dark brown to black; sternites yellowish brown. Ventral plate and median sclerite as figure 8 R. Pupa: Cocoon length of base 5.3 mm; loosely woven, cocoon covering abdomen and one fourth of thorax; thorax and abdomen without granulosity; gill with seven filamentous terminal branches emerging from 4 inflated saclike, subannulated basal branches, 3 with 2 terminal branches and the frontal external with only one (Fig. 9 H). Larva: Length 8.3 mm; general color yellowish brown; cephalic fan with 24 rays; mandible with 29-32 marginal teeth; ratio of proximal, medial and distal segments of antennae = 1:1.1:1.5; hypostomium with 13 apical teeth and 15 lateral serrations; anal ring with 95-100 rows of hooks; area between anal sclerite arms with scarce bifid and trifid scales; ninth sternite with 2 ventral tubercles. Distribution: Highlands of Guatemala. Bionomics: The feeding habits of the adults are not known. This is a highland species breeding at altitudes above 2100 m. The aquatic instars were found in streams with a water temperature below 13 °C, not infrequently between 10 °C and 11 °C. Mayacnephia

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roblesi breeds in the same types of small rivulets in swampy areas as M. pachecolunai (see above), and is restricted to rivulets of less than 15 cm wide, with depths of generally less than 3 cm; the current is slow, with speeds of 25 cm per second, or less. Contrary to the situation for M. pachecolunai, the pupae of M. roblesi are never found on rocks or stones; they are in most cases attached to leaves, with only a few found on branches or twigs. Mayacnephia salasi RAMÍREZ PÉREZ, PETERSON & VARGAS (Fig. 9 I.) Mayacnephia salasi RAMÍREZ PÉREZ et al., 1988: 66; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on original description). Female: Wing length 3.4 mm; antenna and palp dark brown; scutum dark brown to blackish with lateral margins paler with grayish polinosity, covered by yellowish pilosity; pleura dark brown mottled with some darker areas; legs dark brown; abdomen yellowish brown. Male: Wing length 3.1-3.4 mm; general color velvety dark brown. Pupa: Cocoon loosely woven; gill with four cylindrical, globose branches originating from a common short base (Fig. 9 I); head and thorax without granulosity. Larva: Length 6.5-8.5 mm; color pale creamy brown; ratio of proximal, medial and distal segments of antenna = 1:7.7:2.6; labral fan with 25-33 rays; hypostoma with 13 teeth; 3-6 lateral serrations; mandible with 16-25 marginal serrations; anal ring with 62-65 rows and 8-10 hooks each; ninth segment with 2 ventral tubercles; area of anal sclerite with 8-10 thin setae. Distribution: Costa Rica: San José. Bionomics: Mayacnephia salasi lives in small temporary streams at an elevation of 3150 m in an area heavily shaded by forest, and with temperatures of 10-14 °C and a pH of 7.1-7.8. Mayacnephia tadai OCHOA & TAKAOKA (Fig. 9 J.) Mayacnephia tadai OCHOA & TAKAOKA, 1993: 5; CROSSKEY & HOWARD, 1997: 16; 2004: 7.

(Based on original description). Female: Frons, clypeus and scutum dark brown with whitish pollinosity; wing and legs lost; abdomen dorsally dark brown. Male: Unknown. Pupa: Cocoon small, short, slipper shaped, loosely woven, covering only abdomen; body length 4.0 mm; integument moderately covered by minute tubercles; gill with 6 tubular filaments with tapered apex arranged in three pairs with a short stalk (Fig. 9 J). Larva: Length 6.5 mm; color pale yellow; antenna as long as stem of cephalic fan; ratio of proximal, medial and distal segments = 1:0.85:1.1; cephalic fan with 36 rays; mandible with about 22 marginal serrations; hypostomium with 13 teeth, and 4 lateral setae; ninth segment with 2 ventral tubercles; anal ring with 76 rows of 18 hooks each. Distribution: Guatemala: Esquintla. Bionomics: Larvae live on twigs in small temporary streams, at 1000 m of altitude, together with M. aguirrei. Feeding habits of the female are unknown.

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Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA (Figs. 10, 136.) Tlalocomyia WYGODZINSKY & DÍAZ NÁJERA, 1970: 83-108. Type-species: Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA, 1970: 88. Mayacnephia WYGODZINSKY & COSCARÓN, 1973a: 144-150; ADLER et al., 2004: 290 (synonym).

Adults: Frons very narrow; frontal angle 30 degrees; apical segment of maxillary palp more than twice as long as penultimate; pleural membrane hairy on upper third (Fig. B); claw with subbasal tooth subovoidal spermatheca reticulate on outer surface, not spiculate on inner surface; membranous area at insertion of spermathecal duct with evident sclerotization, the duct sclerotized at base and this sclerotization terminating abruptly (Fig. D); genital fork with median and lateral branches well sclerotized (Fig. E); paraproct relatively short, subtriangular; ventral plate subovoidal distally; endoparameres with weak basal plates, no spines; median sclerite with posterior portion divided in two short wide and divergent branches (Fig. G). Pupa: Cocoon obsolescent, composed of closely woven basal cushion and irregular threads loosely covering, at most, posterior half of abdomen of pupa (Fig. H); body of pupa very heavily sclerotized throughout; gills composed of elongate, basally thickened trunk from which arise sequentially a few threadlike filaments (Fig. I); filaments smooth; facial trichomes spinelike; frontoclypeal tubercles large, acute and few in number; abdomen rounded-truncate posteriorly; spine-combs of abdominal terga absent (Fig. J); abdominal terga and sterna each with median transverse ridge; ridges on terga II-VIII and sterna III-V each with one row of very numerous simple spines or hooks; sternal plates of segments VI-VII with several irregularly arranged hooks (Fig. K); terminal hooks not developed; pleural membranes of posterior segments with 4-5 additional spines or hooks per segment (Fig. L); all hooks simple. Larva: Head with normal shape; cephalic apotome well pigmented before posterior margin; cervical sclerites large, fused to terminal portion of postocciput (Fig. O); antenna as long as stem of cephalic fan, third preapical teeth longer than others but shorter than apical; mandible with abundant marginal teeth (about 20) (Fig. M); hypostomium with 13 teeth arranged in three groups (Fig. N); postgenal cleft not deep; lateral sclerite of proleg well developed, subtriangular; body cuticle bare except scarce perianal scales; anal sclerite X shaped; rectal papillae trilobate without additional diverticula. Distribution: Mexico (Fig. 136). Discussion: Tlalocomyia is very close to Mayacnephia. They share a combination of characters such as the loosely woven cocoon covering only a part of the body (generally abdomen), reticulated and one flattened side of the spermatheca, deeply incised median sclerite of the male genitalia and arrangement of the hypostomial teeth, confirming a monophyletic group. ADLER et al. (2004) synonymized Mayacnephia with Tlalocomyia and subdivided Tlalocomyia into three species groups: PACHECOLUNAI which comprises all Neotropical species including Tlalocomyia revelata and T. andersoni, T. osborni, T. ramifera and T. stewarti that are from North America. The following characters of Tlalocomyia help separate it from Mayacnephia: accentuated differences between sexes, viz. sensory vesicle of the maxillary palp which is very big in females but small in males; scutum bright orange-yellow in the female, black in the male;

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maxillary palp of last segment twice as long as the second; anepisternal membrane with hairs on the upper area; spermatheca strongly reticulate; spermathecal duct sclerotized at base, terminating abruptly; median sclerite with wide incision and thick branches; subovoidal tooth of basal claw; pupal gill composed of a thick tube from which several thick short filaments emerge without spiculae on the annulations; facial spine-like B A

C

E G

D

F

I

H

M

J

K

N

L

O

Figure 10. Tlalocomyia revelata. A-E: Female; A: sensory organ of maxillary palp; B: thorax, lateral view; C: claw; D: spermatheca. E: genital fork; F-G: Male; F: hind basitarsus and tarsomere I; G: median sclerite. H-L: Pupa; H: general view; I: gill; J: chaetotaxy of abdomen, dorsal view; K: idem, ventral view; L: idem of apex, lateral view. M-O: Larva; M: teeth of mandible; N: teeth of hypostomium; O: cervical sclerites (A-O from WYGODZINSKY & COSCARÓN, 1973a).

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trichomes; abdominal segments II-VIII and ventral segments III-VIII with abundant hooks or robust spines (described in detail by PY-DANIEL & MOREIRA SAMPAIO, 1994: 139); no terminal hooks; larva with proximal antennal segment not over half as long as medial, and scale fan not evident. Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA (Fig. 10.) Tlalocomyia revelata WYGODZINSKY & DÍAZ NÁJERA, 1970: 88; WYGODZINSKY & COSCARÓN, 1973a: 151; CROSSKEY & HOWARD, 1997: 24; 2004: 7.

Female: Wing length 4.5-5.0 mm; general color yellowish-orange; frons and clypeus brownish to orange; palpi brownish; scutum, scutellum, metanotum, pleurae and legs yellowish orange; scutum with golden pilosity; fronto-ocular triangle longer than wide; cibarium smooth; lacinia with 42-46 teeth; last palpal segment longer than the two before; palpal sensory organ as long as the diameter of the segment with abundant sensory vesicles (Fig. A); wings with well-developed basal cell; Sc with basal sector of R hairy; length/width ratio of hind basitarsus about 7; claw with wide subovoidal subbasal tooth (Fig. C). Male: General coloration blackish; scutum black with golden appressed hair, pleura black; abdomen with tergites blackish and posterior border lighter; hind basitarsus wide (Fig. F); length/width ratio about 3; median sclerite deeply bifurcated with wide branches (Fig. G). Pupa: Body length 4.2-4.5 mm; general aspect as in figure H; tubercles rounded on head, acuminate on thorax; maximum gill length 4.0 mm; gill with 5 branches (Fig. I); trichomes simple, with 3+3 frontal and 1+1 facial, thicker than others; thoracic trichomes simple; number of spines on abdominal tergites as follows: II: 16-17; III: 17-18; IV: 18; V: 14; VI: 12; VII: 10-11 and VIII: 4. Larva: Maximum length 9.0-9.5 mm; general color whitish; head dark brown; ratio of proximal, medial and distal antennal segments = 1:1.9-2.3:1.8-2.1; cephalic fan with 3034 rays; mandible with 20-24 teeth; hypostomium with 13 teeth (Fig. N); 8-11 lateral setae, irregularly disposed; anal sclerite with perianal scales; anal ring with 72-74 rows with 14-17 hooks each. Distribution: Mexico: Distrito Federal. Bionomics: Larvae live in small temporary creeks, attached to stones and vegetation. SPECIES INCERTAE SEDIS ?Tlalocomyia alticola (DYAR & SHANNON), 1927 Eusimulium alticolum DYAR & SHANNON, 1927: 24; VARGAS, 1945b: 110; VARGAS & DÍAZ NÁJERA, 1957b: 230. Mayacnephia alticola CROSSKEY & HOWARD, 1997: 16; 2004: 7. Tlalocomyia alticola ADLER et al., 2004: 290.

According to the original female description, based especially on the genital fork and the elongated basal tooth of the claw, T. alticola is closer to Mayacnephia and Tlalocomyia than to the other Neotropical genera; it doesn’t have enough characters to include it in the keys and to differentiate it from other species. More material would be necessary to adequately differentiate this species.

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Araucnephia WYGODZINSKY & COSCARÓN (Figs. 11-14, 136.) Araucnephia WYGODZINSKY & COSCARÓN, 1973a: 165. Type-species: Simulium montanum PHILIPPI.

Adults: Antennae with 11 segments; male holoptic; female with frons comparatively wide (Fig. 13 A); frontal angle 85 to 100 degrees; mandibles denticulate on both margins (Fig. 13 D); apical segment of maxillary palp of both sexes less than twice as long as penultimate (Fig. 13 C); mesepisternal sulcus shallow, evanescent anteriorly; katepisternum glabrous, in profile higher than wide; pleural membrane glabrous; metanotum glabrous; arms of furcasternum without projections (Fig. 13 F); wings with basal cell present; R1 with spine-like setae, more delicate than those of C (Figs. 13 H-I); Sc setose; basal section of R with hair arranged in several irregular rows; Rs entirely hairy below, hairy above only on apical half; Rs generally simple, very rarely faintly forked apically; M appearing duplicated for most of its length; Cu2 conspicuously curved in both sexes (Fig. 13 E); A2 ending well before wing margin; calcipala present although comparatively small, bluntly rounded; claws of female with medium-sized pointed subtriangular tooth (Figs. 11 D; 13 G); male with sclerotized areas of abdominal terga increasing in size toward posterior segments; sclerotized areas of abdominal sterna small, transverse; spermatheca of normal size, well sclerotized, with circular membranous area at insertion of spermathecal duct, the latter not sclerotized at base; surface of spermatheca not sculptured; internal surface smooth, with scattered spicules; gonapophyses short, subtriangular, apex rounded, their median borders narrowly pigmented, approximated; surface of gonapophyses with microtrichia and few setae (Figs. 11 E; 13 K); paraprocts subovoidal distally (Fig. 13 L); genital fork without forwardly directed apodemes; arms narrow (Figs. 11 F; 13 M); male with gonostylus slightly shorter than gonocoxite, broadly conical, with two apical spines; median surface of gonostylus smooth, glabrous, delimited by two carinae (Figs. 11 G-H; 13 N, O); ventral plate transverse to subtriangular, its basal arms in shape of elongate parallel-sided sclerotized rods (Figs. 11 I; 13 P, Q); median sclerite in shape of complex geniculate plate; basal half entire, fitting into concavity of ventral plate; distal half arising at sharp angle with basal half, divided into two straight parallel arms narrowed toward their apex (Figs. 13 R-S); endoparameres with large plate-like basal portion connected to numerous denticles forming apical portion (Figs. 11 K; 13 T); median sclerite expanded basal and distally (Figs. 11 J-K). Pupa: Cocoon large, shapeless, covering entire pupa except respiratory organs (Figs. 12 A; 14 A); body of pupa well sclerotized throughout; head and thorax with or without tubercles; gills arborescent, divided very close to base into primary and secondary branches; general structure of branches filamentous (Fig. 12 C); head sclerite with 1+1 frontal and facial, and 2+2 epicranial, labral and genal trichomes, all hair like (Figs. 12 B; 14 B); thorax with 6-8+6-8 simple hair like trichomes (Figs. 12 D; 14 C); abdomen pointed posteriorly; distinct pleural areas not developed; onchotaxy of abdomen as follows: Terga III and IV with 4+4 hooks posteriorly; terga VI-IX with continuous spine-combs anteriorly; sternum V, or IV and V, with 2+2 hooks posteriorly; sternal plates VI and VII with 1+1 hooks, 1+1 hooks each also in pleural membranes of segments VI and VII (Fig. 14 D); terminal spines well developed, from stout to slightly sinuous (Fig. 12 E); posterior segments with several strong S-shaped or apically curved or looped setae, only very rarely branched.

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Larva: Head spots positive; cephalic apotome broadest well before hind margin (Figs. 12 G; 14 F); cervical sclerites wide to faintly fused with upper ends of postocciput (Fig. 14 J); postgenal cleft shallow but distinct, inverted U-shaped (Fig. 12 L); length of hypostomium approximately equal to length of postgenal bridge; antennae much shorter than stern of cephalic fan (Fig. 14 H); proximal segment of antennae very short, less than half as long as medial; distal segment distinctly darker than remainder (Figs. 12 H; 14 G); cephalic fans with rays unusually stout and heavily sclerotized (Fig. 12 I); mandibles with two outer teeth and one apical tooth; third preapical tooth larger than remaining teeth, but shorter than apical tooth; first and second preapical teeth subequal in size (Figs. 12 J; 14 I); inner teeth arranged in several rows; one large mandibular serration followed by numerous (10-15) very unequal smaller ones; short basal setae of mandibles very stout, with conspicuous barbules; maxillary palp relatively elongated (Fig. 12 K); hypostomium with 15 teeth arranged in three conspicuous groups, with median tooth and lateral teeth by far the largest; 3+3 lateral teeth descending below level of remaining teeth (Fig. 14 K); lateral sclerite of proleg about as long as wide, subtriangular, narrow at base, strongly widened apically (Figs. 12 M; 14 L); setae of lateral sclerite less than half as long as sclerite, inserted on distinct individual tubercles, arranged in irregular double row, this row extending slightly beyond sclerite below; larval cuticle bare, except for perianal scales; anal sclerite normal, X-shaped (Fig. 12 N); terminal ventral tubercles absent. Distribution: Northeast of Argentina; south of Brazil and Mediterranean climate area of central Chile (Fig. 136). Discussion: Araucnephia differs from the Mexican Mayacnephia and Tlalocomyia by the specialized median sclerite of the male genitalia and also by the presence of distinct endoparameral spines; by the spiculae of the spermatheca; the single pair of frontal trichomes and by the rarity of branched setae at the end of the abdomen of the pupa. In the larva the first segment of the antenna is not more than half as long as the second, as it is in Mayacnephia and Tlalocomyia. There are only two South American genera with comparable hypostomial characters, Gigantodax and Araucnephioides. Gigantodax is easily distinguished from Araucnephia by the absence of a basal cell and its specialized wing venation and the different genitalia in both sexes, and the accessory sclerite forming a complete ring at the posterior end of the body of the larvae. The pupae are not so easy to distinguish, but those of Gigantodax do not have the full complement of cephalic trichomes, as described for Araucnephia. Araucnephia is possibly closest to Araucnephioides; this is discussed under the heading of the latter genus. Cnesiamima differs from Araucnephia in the absence of spinelike setae on R1 and specialized genitalic features, such as the structure of the median sclerite in the male and that of the spermatheca in the female. KEYS

TO SPECIES OF

ARAUCNEPHIA

Females –

Wing length 2.7-2.8 mm; mandible with about 40 (14+26) teeth; length/width ratio of hind basitarsus = 8.5; spermatheca subovoidal with some superficial tubercles .............................................................................................Araucnephia iberaensis

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–

Wing length 3.3-4.3 mm; mandible with about 55 (21+34) teeth; length/width ratio of hind basitarsus = 6.5; spermatheca spheroidal without superficial tubercles .............................................................................................. Araucnephia montana

Males – –

Wing length 2.6-2.7 mm; scutum brown, lighter on posterolateral border; abdomen brownish; gonostylus with small depression before subapical spurs (Figs. 11 G, H) ..................................................................................Araucnephia iberaensis Wing length 3.5-3.6 mm; scutum velvety black; abdomen blackish; gonostylus without depression before subapical spurs (Figs. 13 O, N) ..... Araucnephia montana

Pupae – –

Cocoon length 3.6-4.2 mm; gill with 9 terminal branches; facial and thoracic trichomes curved distally (Figs. 12 B, D); cephalothorax with abundant tubercles ..............................................................................................................Araucnephia iberaensis Cocoon length 7.0-8.3 mm; gill with about 14 terminal branches; facial and thoracic trichomes not curved distally (Figs. 14 B, C); cephalothorax without abundant tubercles ............................................................................ Araucnephia montana

Larvae –

–

CLAVES

Maximum length 6.6-7.6 mm; ratio of proximal, medial and distal antennal segments = 1:1.2-1.5:2.4-2.8; cephalic fan with 48-53 rays; comb of thoracic proleg sclerite with about 15-17 teeth (Fig. 12 M); anal ring with 64-66 rows with 14-16 hooks each. ............................................................................Araucnephia iberaensis Maximum length 11.0 mm; ratio of proximal, medial and distal antennal segments = 1:2.7-3.0:2.7-3.2; cephalic fan with 23-26 rays; comb of thoracic proleg sclerite with about 61 teeth (Fig. 14 L); anal ring with 80-100 rows with 18-22 hooks each. ............................................................................................................... Araucnephia montana DE ESPECIES DE

ARAUCNEPHIA

Hembras – –

Longitud ala 2,7-2,8 mm; mandíbula con alrededor de 40 (14+26) dientes; relación largo/ancho de basitarso posterior = 8,5; espermateca subovoidal con algunos tubérculos superficiales ....................................................Araucnephia iberaensis Longitud ala 3,3- 4,3 mm; mandíbula con alrededor de 55 (21+34) dientes; relación largo/ancho de basitarso posterior = 6,5; espermateca esferoidal sin tubérculos superficiales .................................................................... Araucnephia montana

Machos –

Longitud ala 2,6-2,7 mm; escudo castaño, más claro en el borde latero posterior; abdomen castaño; gonostilo con una pequeña depresión delante de los espolones subapicales (Figs. 11 G, H) ............................................................Araucnephia iberaensis

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–

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Longitud ala 3,5-3,6 mm; escudo negro aterciopelado; abdomen negruzco; gonostilo sin depresión delante de los espolones subapicales (Figs. 13 O, N) ....... ............................................................................................................... Araucnephia montana

Pupas – –

Longitud del capullo 3,6-4,2 mm; branquia con 9 ramas terminales; tricomas faciales y torácicos curvados distalmente (Figs. 12 B, D); cefalotórax con abundantes tubérculos ...........................................................................................Araucnephia iberaensis Longitud del capullo 7,0-8,3 mm; branquia con alrededor de 14 ramas terminales; tricomas faciales y torácicos no curvados distalmente (Figs. 14 B, C); cefalotórax sin abundantes tubérculos ............................................................... Araucnephia montana

Larvas –

–

Longitud máxima 6,6-7,6 mm; relación de los artejos antenales proximal mediano y distal = 1:1,2-1,5:2,4-2,8; abanico cefálico con 48-53 rayos; peine del esclerito de la propata torácica con alrededor de 15-17 dientes (Fig. 12 M); anillo anal con 6466 hileras con 14-16 ganchos cada una .......................................Araucnephia iberaensis Longitud máxima 11,0 mm; relación de los artejos antenales proximal mediano y distal = 1:2,7-3,0:2,7-3,2; abanico cefálico con 23-26 rayos; peine del esclerito de la propata torácica con alrededor de 61 dientes (Fig. 14 L); anillo anal con 80-100 hileras con 18-22 ganchos cada una .............................................. Araucnephia montana

Araucnephia iberaensis COSCARÓN & COSCARÓN ARIAS (Figs. 11 A-K; 12 A-N.) Araucnephia iberaensis COSCARÓN & COSCARÓN ARIAS, 2002: 81-87.

This species is very close to Araucnephia montana and here we include only the differences from A. montana. Female: Wing length 2.7-2.8 mm; frons, clypeus, antenna, palp and occiput black with grayish pollinosity; scutum and metanotum blackish, ornamented as in figure 11 A; legs brownish darkened on tarsi; abdomen blackish with gray pollinosity; frontal angle = 100°; fronto-ocular triangle as in figure 11 B; last segment of maxillary palp about twice length of penultimate segment; sensory vesicle small about 1/5 of third palpomere length (Fig. 11 C); mandible denticulate on both sides with 14 (15)-26 (27) teeth; lacinia with about 25 teeth; length/width ratio of hind basitarsus = 8.5; calcipala small subovoidal; eighth sternite mostly black; gonapophyses with 6-7 hairs (Fig. 11 E); spermatheca subovoidal, showing some superficial tubercles; genital fork with lateral branches distally expanded (Fig. 11 F). Male: Wing length 2.6-2.7 mm Scutum brown, lighter on posterolateral border; abdomen brownish, a longitudinal blackish stripe composed of subtrapezoidal sclerotized tergal plates on segments II-V and 1+1 sublateral spots on segments VI-VIII; ventrally with subquadrate spots blackish on segments III-VIII; length/width ratio of hind basitarsus = 4.4; gonostylus distally with depression before two apical spurs (Figs. 11 GH); ventral plate subtrapezoidal (Fig. 11 I); median sclerite and aedeagus as in figures 11 JK, endoparameres with some well-developed denticles (Fig. 11 K).

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F B

A C

D

E

I

G H J K Figure 11. Araucnephia iberaensis. A-F: Female; A: head and thorax, dorsal view; B: fronto-ocular triangle; C: sensory vesicle of maxillary palp; D: claw; E: eighth sternite and gonapophyses; F: Genital fork. G-K: Male; G: general aspect of genitalia dorsal view, showing gonocoxite, gonostylus, ventral plate, median sclerite and endoparameres; H: gonostylus; I: ventral plate; J-K: different view of aedeagus membrane, base and scarcely evident hooks of endoparameres and median sclerite.

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B A

C

E D F

o.t.

G I

J

a.t. p.t.

H

i.t. m.s.

M

L

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N

Figure 12. Araucnephia iberaensis. A-E: Pupa; A: general aspect; B: frontoclypeus, with trichomes and tubercles magnified; C gill; D: thoracic trichomes; E: abdominal terminal area, showing curled trichomes. F-N: Larva; F: head, lateral view; G: cephalic apotome; H: antenna; I: teeth of cephalic fan rays; J: apical portion of mandible with teeth (a.t. = apical teeth, i.t. = inner teeth, m.s. = marginal serrations, o.t. = outer teeth, p.t. = preapical teeth); K: maxillary palp; L: postgenal cleft and hypostomium; M: lateral sclerite of thoracic proleg; N: anal sclerite with scales and hairs.

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Pupa: Cocoon total length on dorsum 3.6-4.2 mm; total pupa length 6.0-6.3 mm; cocoon not compacted, generally covering pupa up to base of gill (Fig. 12 A). frontoclypeus and thorax with abundant tubercles mostly verrucose (Fig. 12 B); abdomen with abundant tubercles; gill with 9 terminal branches with 3 short primary branches (Fig. 12 C); trichomes of head and thorax single and mostly curved distally (Fig. 12 D) as for terminal area of the abdomen (Fig. 12 E). Larva: Maximum length 6.6-7.6 mm; head light brown, darkened around eyes (Fig. 12 F); cephalic apotome with positive spots blackish brown and very evident (Fig. 12 G); antenna about same length as stem of cephalic fan; ratio of proximal, medial and distal antennal segments = 1:1.2-1.5:2.4-2.8 (Fig. 12 H); cephalic fan with 48-54 rays; disposition of ray setae as in figure 12 I; mandible with marginal serrations of 8-12 teeth (Fig. 12 J); maxillary palp as in figure 12 K; hypostomium and postgenal cleft as in figure 12 L; comb of thoracic proleg sclerite with 15-17 teeth (Fig. 12 M); anal ring with 64-66 rows with 1416 hooks each. Distribution: Argentina: Corrientes; Brazil: Paraná (unpubl. information). Bionomics: The species breeds in small creeks 0.5-3.0 m wide and 0.1-0.5 m deep in crystalline water and strong but not torrential currents; larvae and pupae are found on vegetation. Females are haematophagus and bite humans, producing a hard spot followed by an allergic reaction. Discussion: The morphological characters allow it to be differentiated from Araucnephia montana, as seen in the key. Araucnephia montana (PHILIPPI) (Figs. 13 A-T; 14 A-L.) Simulium montanum PHILIPPI, 1865: 633. Simulium (Cnephia) montanum: EDWARDS, 1931: 149. Cnephia montanum SMART, 1945: 498. Araucnephia montana WYGODZINSKY & COSCARÓN, 1973a: 168-179; COSCARÓN, 1991: 14-15; CROSSKEY & HOWARD, 1997: 11; 2004: 2.

Female: Length of wing, 3.3-4.3 mm, generally almost 4.0 mm (type 3.9 mm); head black; occiput, frons, and clypeus gray pruinose; antennae, palpi, and labrum brownish black; pilosity of antenna very short, that of palpi, clypeus, frons, and occiput somewhat longer, its color from silvery gray to black; thorax dull black, with silvery gray pollinosity; metapleura and wing base brownish; hair of thorax silvery gray; wings hyaline; veins light brown, hair and spines blackish; tufts at wing base from silvery to brownish yellow; halteres light brown; legs from light brown with darkened apices of femora, tibiae and tarsi, to entirely dark brown or black; most specimens with legs dark, concolorous; hair of legs from silvery gray to black; abdomen from dark grayish brown to dull black; dorsal surface always darkest; hair of abdomen from silvery gray to black; frons as shown in figure 13 A; frontal angle 85-90 degrees; fronto-ocular triangle (Fig. 13 B) approximately as long as wide; last segment of maxillary palp (Fig. 13 C) slightly less than twice as long as penultimate; diameter of sensory vesicle smaller than half diameter of third palpomere; lacinia with 30-35 teeth; mandible with about 75 teeth (Fig. 13 D); furcasternum as in figure 13 F; wings as in generic description and figure 13 E; spine-like setae of R1 often

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A C B E G D C

H

I

R1 Rs

F L K P

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Q

N

O

R

T S

Figure 13. Araucnephia montana. A-L: Female; A: frons; B: fronto-ocular triangle; C: maxillary palp; D: mandible; E: wing, general view; F: furcasternum; G: claw; H-I: portion of wings showing part of anterior veins with chaetotaxy in different specimens (c = costa, R1= radius 1, Rs= secondary radius (radius 2) ); J: calcipala and tarsomere I; K: eighth sternite and gonapophyses; L: cercus and paraproct; M: genital fork. N-T: Male; N: gonocoxite and gonostylus, dorsolateral view; O: gonostylus, showing membranous inner surface; P: ventral plate, anteroventral view; Q: ventral plate and median sclerite in lateral views; R-S: median sclerite in different position; T: endoparameres. (A-T from WYGODZINSKY & COSCARÓN, 1973a).

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A D

C

B

E

F

K H

G J

I

Figure 14. Araucnephia montana. A-D: Pupa; A: general view; B: cephalic sclerite with trichomes; C: portion of thorax with trichomes; D: abdominal chaetotaxy. E-L: Larva; E: general aspect in lateral view; F: cephalic apotome; G: antenna; H: portion of head dorsal view showing short antenna; H: portion of mandible with teeth; J: posterior region of head, with cervical sclerites; K: anterior portion of hypostomium; L: sclerite of thoracic proleg. (A-L from WYGODZINSKY & COSCARÓN, 1973a).

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distinctly more slender than those of C (Fig. 13 H-I); hind basitarsus 6.1-6.9 times as long as wide; claws as in figure 13 G; second hind tarsomere with strong rugosity (Fig. 13 J); eighth abdominal sternum as shown in figure 13 K; pigmentation variable, from only central area darkened to entire sternum dark; gonapophysis with abundant microtrichia and 10-14 short setae; cerci and paraprocts as illustrated in figure 13 L; cerci much wider than high; paraprocts short, their free extremity somewhat truncate; genital fork with stem and apical portion of arms well pigmented (Fig. 13 M). Male: Length of wing, 3.5-3.6 mm; head black; clypeus dark gray pollinose; antennae, palpi, and proboscis brownish black; antennae with short pubescence; setae of basal segments longer than that of remaining segments; occiput, line between eyes, clypeus and basal segments of maxillary palps with several long hairs; thorax velvety black; pleura with silvery gray pollinosity; scutum with decumbent hair from light silvery to light brassy; distal portion of scutum as well as scutellum and metanotum with black, erect setae; color of wings, halteres and legs as in female; abdomen blackish, dark dorsally, grayish laterally and below; hair of abdomen colored as on thorax; chaetotaxy of wings as in female, but fewer setae on Sc; spiniform setae of R1 distinctly smaller than those of C; in some specimens, Rs with short bifurcation apically; hind basitarsus 3.9-4.6 times as long as wide; calcipala narrowly rounded apically; gonostylus without apical depression (Figs. 13 N, O); ventral plate with arms and basal margin distinctly sclerotized, disc uniformly pigmented (Figs. 13 P-Q); median sclerite and endoparameres as in figures 13 R, S and T. Pupa: Cocoon covering pupa completely or almost so (Fig. 14 A), in some cases also including part or all of gills; cocoon closely woven, threads not perceptible under medium magnification; color light grayish brown; length along dorsum, 7.0-8.3 mm; total length of pupa 4.0-4.5 mm; cephalopterothecal length, 3.0 mm; gill composed of six short primary branches arising from extremely short common trunk; the first two of three dorsal primary branches generally several times as long as wide, the third much shorter than others; all giving rise to two filaments; two median branches giving origin to three filaments each, filaments arising at variable relative positions; ventral primary branch very short, with two filaments delicately pitted; head and thorax of pupa light brown; trichomes of head as in figure 14 B; epicranial, lateral and genal trichomes in pairs, others single; all trichomes simple; head sclerite lacking tubercles; thorax without tubercles and with not more than eight trichomes on each side on disc (Fig. 14 C); abdominal segments distinctly sclerotized although somewhat less so than thorax, intensity of sclerotization diminishing toward posterior segments; onchotaxy and accessory hair and setae of abdomen as in generic description and figure 14 D; tubercles on abdomen absent or few in number. Larva: Maximum length, 11.0 mm, maximum width of head capsule, 1.2 mm. General shape as shown in figure 14 E; color of larva light grayish brown, head often very dark; cephalic apotome (Fig. 14 F) yellowish brown, darker along middle, along hind border, and with one or two sublateral spots on each side; antennae as in figure 14 G; ratio of lengths of proximal, medial and distal segments = 1:2.75-3.0:2.75-3.2; cephalic fan with 23-26 rays; mandibles as in figure 14 I, with 13-20 inner teeth and 7-15 marginal serrations irregularly decreasing in size; cervical sclerite as in figure 14 J; hypostomium as shown in figure 14 K, its anterior border strongly pigmented; lateral border of hypostomium with 3-8 distinct serrations; hypostomial setae arranged in one, rarely in two

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irregular series, with 5-7 setae in each group; disc of hypostomium with few scattered short hairs; postgenal cleft about half as long as postgenal bridge; lateral sclerite of thoracic proleg as in figure 14 L; anal sclerite with numerous perianal scales, two to fivepronged; crochet ring with 80-100 rows with 18-22 hooks each. Distribution: Chile: Aconcagua, Coquimbo, Curico, Santiago, Valparaiso. Bionomics: We have collected the aquatic stages of Araucnephia montana from the surface of stones and rocks exclusively in small, clear, temporary streams, toward the end of the wet season. This is apparently a species that completes its cycle in late winter or very early spring (September or very early October). Discussion: The closest relative of Araucnephia may be Araucnephioides; both genera are sympatric, and individuals are often difficult to distinguish. We, therefore, present a detailed enumeration of differential characters to help to identify specimens. Differences between Araucnephia and Araucnephioides Araucnephia Female: Basal tooth of claws subtriangular (Figs. 11 D; 13 G); calcipala broadly rounded (Fig. 13 J}; inner surface of spermatheca not rugose. Male: Genitalia with subtriangular ventral plate, its basal arms narrow (Figs. 11 I; 13 P), straight; arms of median sclerite parallel (Fig. 13 S); endoparameral organ with basal portion well developed, plate like, and teeth of distal portion conspicuous (Figs. 11 K; 13 T). Pupa: Gill with 14-17 filaments; head sclerite with full complement of trichomes, i.e., frontal, facial, epicranial, ocular, and genal (Figs. 12 B; 14 B). Larva: Antenna much shorter than stem of cephalic fan (Figs. 12 F; 14 H); rays of cephalic fan very heavily sclerotized (Fig. 12 I). Araucnephioides Female: Basal tooth of claws narrowly hook like (Fig. 15 I); calcipala subtriangular (Fig. 15 H). Male: Genitalia with ventral plate subrectangular, its basal arms stout, rounded at apex and recurved (Fig. 15 P); arms of median sclerite sharply diverging apically (Fig. 15 U); endoparameral organ with basal portion small and denticles of apical portion faint. Pupa: Gill with 10-13 filaments; head sclerite without or with poorly developed frontal trichomes (Fig. 16 D). Larva: Antenna almost as long as stem of cephalic fan (Fig. 16 I); sclerotization of rays of cephalic fan normal. Araucnephioides WYGODZINSKY & COSCARÓN (Figs. 15, 16, 136.) Araucnephioides WYGODZINSKY & COSCARÓN, 1973a: 179. Type-species: Araucnephioides schlingeri WYGODZINSKY & COSCARÓN, 1973a.

Adults: Female with frons medium wide (Fig. 15 A); frontal angle 60-70 degrees; antenna with 11 segments (Fig. 15 B); male holoptic; mandibles of female denticulate on both margins; apical segment of maxillary palp of both sexes less than twice as long

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as penultimate segment (Fig. 15 C); cibarium with basal portion smooth and with strong concavity (Fig. 15 F); mesepisternal sulcus shallow, evanescent anteriorly; katepisternum glabrous, in profile higher than wide; pleural membrane bare; metanotum glabrous; arms of furcasternum without projections (Fig. 17 J); wings with basal cell present (Fig. C

B

A I

G

E H

L D

F

K M

N S O P

Q

U

T J

R

Figure 15. Araucnephioides schlingeri. A-M: Female; A: frons; B: Antenna; C: maxillary palp; D: sensory vesicle of maxillary palp; E: fronto-ocular triangle; F: basal portion of cibarium; G: wing, general view; H: calcipala and tarsomere I; I: claw; J: furcasternum; K: genital fork; L: cercus and paraproct; M: eighth sternite and gonapophyses. N-U: Male; N: gonocoxite and gonostylus; O: gonostylus, showing membranous inner area; P: ventral plate with median sclerite visible through it; Q: idem side view; R: idem full view; S: idem, with median sclerite and endoparameres; T-U: median sclerite in different positions. (A-U from WYGODZINSKY & COSCARÓN, 1973a).

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15 G); R1 with spinelike setae more delicate than those of C; Sc setose; basal section of R with hair arranged in several irregular rows; Rs entirely hairy below, hairy above only on apical half; Rs not forked apically; M appearing duplicated for most of its length; Cu2 conspicuously curved in both sexes; A2 ending well before wing margin; calcipala small, narrowly subtriangular (Fig. 15 H); claws of female with medium-sized, pointed, and curved hook-like basal tooth (Fig. 15 I); male with sclerotized areas of abdominal terga of subequal size on all segments; sclerotized areas of abdominal sterna subquadrate; spermatheca globular, of normal size, well sclerotized, with circular membranous area at insertion of spermathecal duct, the latter not sclerotized at base; surface of spermatheca not sculptured; internal surface irregularly rugose and with scattered spicules; gonapophyses short, subtriangular, apically rounded, their median borders pigmented, approximated; surface of gonapophysis with microtrichia and setae (Fig. 15 M); genital fork without forwardly directed apodemes; posterior arms wide (Fig. 15 K); male with gonostylus shorter than gonocoxite, broadly conical, with two or three apical spines; median surface of gonostylus smooth, glabrous, delimited by two carinae (Figs. 15 N, O); ventral plate subquadrate, its anterior border truncate or slightly emarginated; its basal arms with terminal knobbed sclerotization (Figs. 15 P-S); median sclerite in shape of complex geniculate plate; basal half entire, transverse, fitting into concavity of ventral plate; distal half arising at sharp angle with basal half, divided into two apically, outwardly bent strong arms (Figs. 15 T-U); endoparameral organ with narrow plate-like basal portion tapering into very narrow obsolescent apical portion; denticles weakly developed. Pupa: Cocoon large but shapeless, covering entire pupa except gills (Fig. 16 A); head and thorax with numerous tubercles (Figs. 16 D-E); body of pupa well sclerotized throughout; gills arborescent, consisting of several primary branches and not very numerous (approximately 10) narrow filaments arranged in two more or less distinct groups (Fig. 16 B); head sclerite without or with 1-2+1-2 very short frontal and with 1+1 hair-like well-developed facial trichomes; thorax with 6-8+6-8 simple hair-like trichomes; abdomen pointed posteriorly; its structure and onchotaxy as in Araucnephia. Larva: Body slightly wider posteriorly on preapical portion (Figs. 16 F); head spots positive, distinct (Fig. 16 G); cervical sclerites small, isolated in membrane (Figs. 16 HI); antenna almost as long as stern of cephalic fan (Fig. 16 I); proximal antennal segment very short, less than half as long as medial segment (Fig. 16 J); distal segment only very slightly darker than remaining; cephalic fans with rays normal, slender, not heavily sclerotized; setae of all rays distinctly of two sizes; maxillary palp normal (Fig. 16 K); mandibles with two external teeth and one apical tooth; third preapical tooth larger than the two others, but shorter than apical tooth; second preapical tooth minute; inner teeth arranged in several rows; one large basal mandibular serration followed by numerous smaller ones; accessory mandibular serrations situated on border of mandible at level of internal teeth (Fig. 16 L); short setae of mandible slender, only very faintly dentate or not at all; postgenal cleft shallow but distinct, biarcate (Fig. 16 M); hypostomium shorter than length of postgenal bridge; hypostomium with 17 teeth arranged in three conspicuous groups, with median tooth and lateral teeth by far largest; 4+4 lateral teeth descending below level of remaining teeth (Fig. 16 N); lateral sclerite of thoracic proleg

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as in Araucnephia; larval cuticle bare except for perianal scales; anal sclerite normal, Xshaped; ventral tubercles absent. Distribution: Mediterranean climate area of central Chile.

A E F D

B

I

C J

M N

K L

G P

H O

Figure 16. Araucnephioides schlingeri. A-E: Pupa; A: general view; B: gill; C: basal portion of gill; D: frontoclypeus; E: arrangement of tubercles of exposed portion of thorax; F-P: Larva; F: dorsal and lateral views; G: cephalic apotome; H: posterior region of head with reduced cervical sclerites; I: portion of cephalic capsule showing antenna and stem of cephalic fan; J: antenna; K: maxillary palp; L: portion apical mandible and teeth; M: postgenal cleft and hypostomium; N: anterior border of hypostomium; O: anal sclerite; P: rectal papillae. (A-P from WYGODZINSKY & COSCARÓN, 1973a).

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Araucnephioides schlingeri WYGODZINSKY & COSCARÓN (Figs. 15 A-U; 16 A-P.) Araucnephioides schlingeri WYGODZINSKY & COSCARÓN, 1973a: 181-184; COSCARÓN, 1991: 17; CROSSKEY & HOWARD, 1997: 11.

Female: Length of wing, 3.2-4.0 mm, in most specimens more than 3.5 mm; head blackish, eyes dark piceous; occiput, frons and clypeus grayish brown pollinose; antennae and palpi dark brown, labrum light grayish brown; pilosity of antennae very short, except on two basal segments, that of palpi, clypeus, frons, and occiput somewhat longer; hair from brassy to black; scutum dull black, with gray pollinosity and decumbent hair of same color as that of occiput; pleura brownish, with silvery gray pollinosity; base of wings light brown; scutellum piceous, hair semi-erect, from light gray to dark brown; metanotum and katepisternum brown to black, with silvery pollinosity; wings and veins hyaline, veins light brown; halteres piceous; legs in most specimens dark brown, with apices of femora, tibiae, and tarsi piceous; in some specimens legs lighter, with dark apical regions of leg segments more conspicuous; hair from silvery gray to blackish; color of abdomen dark grayish brown to blackish; frons as shown in figure 15 A; frontal angle 60-70 degrees; frontoocular triangle slightly wider than high (Fig. 15 E); shape and proportions of antennal segments as shown in figure 15 B; mandible with 36-40 teeth; last segment of maxillary palp (Fig. 15 C) distinctly less than twice as long as penultimate segment; diameter of sensory vesicle (Fig. 15 D) less than half the diameter of third segment; maxillae with about 30 teeth; wing as in figure 15 G, calcipala about as long as wide at base (Fig. 15 H); claws as in figure 15 I; hind basitarsus 6.3-7.6 times as long as wide; eighth abdominal sternum with abundant setae; gonapophyses each with 10-14 medium-sized hairs and with numerous microtrichia (Fig. 15 M); paraprocts short (Fig. 15 L); genital fork with wide lateral branches (Fig. 15 K). Male: Length of wing, 2.9-3.3 mm; color of head, body and appendages brown dark to blackish; last segment of maxillary palp less than twice as long as penultimate segment; diameter of sensory vesicle smaller than half diameter of third segment; hind basitarsus 3.7-4.6 times as long as wide; genitalia as in generic description (Figs. 15 N-U). Pupa: Cocoon (Fig. 16 A) covering entire pupa, in some cases including gills and often also containing larval exuviae; color of cocoon very light brown, loosely woven with body of pupa discernible; threads visible under medium magnification; length of cocoon along dorsum, 4.0-6.0 mm; total length of body of pupa 3.5-4.3 mm, cephalopterothecal length 2.05-2.10 mm; gills (Figs. 16 B-C) about half as long as body of pupa, or slightly longer, with 10-12, rarely 13, filaments; two primary branches arising from very short common trunk; dorsal primary branch with very short secondary branch immediately beyond base, divided into two branches; secondary branch in some cases arising directly from base, thus constituting third primary branch; dorsal primary branch with four or five, rarely six, additional filaments; ventral primary branch generally giving origin to four, rarely five filaments, two originating very close to base, the others slightly above; surface of filaments delicately pitted, and with irregularly shaped and distributed minute plate-like sclerotized structures basally; head and thorax of pupa light brown; frontoclypeus with numerous tubercles (Fig. 16 D); thorax with numerous tubercles, arranged in roughly circular groups, composed of five to eight tubercles each (Fig.16 E); abdomen somewhat

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lighter than thorax; onchotaxy much as in Araucnephia montana, but spine-combs on tergum V often absent, and composed of fewer spines (about 15) on tergum VI; all terga with numerous tubercles in addition to usual hair and spines; specialized setae of eighth and ninth segments with strong curvature distally. Larva: Maximum length, 8.0 mm; maximum width of head capsule, 0.75 mm; body shape as shown in figure 16 F; color of larva light grayish brown; cephalic apotome yellowish brown, darker along middle, and with several spots as shown in figure 16 G; antennae as in figure 16 J; ratio of length of proximal, medial and distal segments = 1:2.2:2.9-3; cephalic fan with 37-42 rays; mandible as in figure 16 L, with 15-17 inner teeth and 10-15 marginal serrations; hypostomial teeth as shown in figure 16 N; anterior area of hypostomium strongly pigmented; lateral border of hypostomium with 6-9 serrations; hypostomial setae arranged on each side in one irregular row composed of about four setae; disc of hypostomium with few scattered, short setae; postgenal cleft short and subquadrate (Fig. 16 M); anal sclerite with perianal scales as shown in figure 16 O; crochet ring with 60-67 rows composed of 18-20 hooks each. Distribution: Same as for genus. Cnesia ENDERLEIN (Figs. 17-19, 136.) Cnesia ENDERLEIN, 1934b: 273; STONE, 1963: 4; WYGODZINSKY & COSCARÓN, 1973a: 151; COSCARÓN, 1991: 18; CROSSKEY & HOWARD, 1997: 11. Type-Species: Simulium (Cnephia) dissimile EDWARDS, 1931 (as Simulium (Cnephia) gynandrum EDWARDS, 1931).

Adults: Antennae with 11 segments; male holoptic; female with frons medium wide; frontal angle 50-67 degrees; mandibles with well-developed teeth only on internal margin, external border weakly serrated (Fig. 17 A); apical segment of maxillary palp in both sexes less than twice as long as penultimate segment; mesepisternal sulcus shallow, evanescent anteriorly; katepisternum glabrous, in profile slightly longer than high; pleural membrane glabrous; metanotum with hair in some but not all species; arms of furcasternum lacking conspicuous projections (Fig. 17 E); wings with basal cell present (Fig. 17 B); R1 with hair and spine-like setae, the latter not more delicate on R1 than on C (Fig. 17 I); Sc and basal section of R with hair arranged in several irregular series; Rs entirely hairy below, hairy above only on apical half; Rs not forked apically; M not appearing double; Cu2 only weakly curved in both sexes; A2 almost attaining wing margin; calcipala present, large, bluntly rounded apically (Fig. 17 C); claws of female with large subtriangular basal tooth (Fig. 17 D); spermatheca of normal size, normally sclerotized, oval; internal surface minutely wrinkled and with small irregularly scattered spicules; area of insertion of spermathecal duct membranous, distinctly larger than diameter of duct; duct not sclerotized (Fig. 17 F); gonapophyses subtriangular, transverse, their medial borders narrowly pigmented, approximated, their surface with microtrichia and few setae; paraproct short (Fig. 17 G); posterior arms of genital fork wide, with small but distinct forwardly directed apodemes (Figs. 17 H-Q); male with gonostylus distinctly shorter than gonocoxite, narrowly conical, with two apical spines; medial surface of gonostylus smooth, delimited by two carinae (Figs. 17 K-L); ventral plate subpentagonal, about as long as wide, setose, with very low

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median longitudinal keel (Figs. 17 M-N; 19 C-E); median sclerite with flaring base, then narrowed, apically widened again and with two wing-like pointed subapical flanges (Figs. 17 O-P; 19 F-G); endoparameral organs with basal portion strongly reduced, apical portion composed of a group of one or two large and several oppositely directed medium-sized denticles. Pupa: Cocoon large, shapeless, covering pupa to base of respiratory organs (Fig. 18 A); body of pupa well sclerotized throughout; head and thorax with or without tubercles; gills bushy, much shorter than pupa proper, composed of 35-40 slender filaments arising from about 12 primary branches very close to their base (Figs. 18 B; 19 H); head with 1+1 prefrontal, 1+1 or 2+2 frontal, 1+1 facial, and occasionally 1+1 labial trichomes; prefrontal and single or anterior frontal trichomes spinelike; posterior frontal, facial and labial trichomes hair like (Figs. 18 C; 19 I); disc of thorax with approximately 5+5 trichomes, two or three dorsalmost ones in shape of strong spines, others hair like; abdomen pointed posteriorly; distinct pleural areas not developed; terga with numerous small tubercles, those near anterior margins of sclerite arranged in very conspicuous transverse ridges; onchotaxy of abdomen as follows: terga III and IV with 4+4 short hooks posteriorly; tergum VIII with 2+2 very elongate spines posteriorly; terga without spines-comb; sterna IV and V with 3+3 hooks posteriorly, those of IV arranged in one median and 1+1 submedian groups of two spines each, those on V arranged in 1+1 closely approximated groups of three spines each; sternal plates VI with 2+2 hooks, VII with 1+1 hooks; 1+1 hooks each also in pleural membrane of segments VI and VII (Fig. 18 D); terminal spines well developed, stout, not sinuous; posterior segments with several strong S-shaped setae, often apically curved, looped or spiraled, occasionally branched near base or apex (Fig. 18 E). Larva: Body shape with wider preapical region (Fig. 19 K); head spots positive; cephalic apotome broadest well before hind margin (Fig. 18 F); cervical sclerites very small, isolated in membrane; postgenal cleft very shallow, broadly U-shaped (Figs. 18 I; 19 N); antennae about as long as stem of cephalic fans; medial antennal segment about two times as long as proximal, distal segment darker than remaining segments (Figs. 18 G; 19 L); mandibles with four outer teeth in addition to apical tooth; third preapical tooth longer and more strongly sclerotized than other preapical teeth, as long as apical tooth; first inner tooth as long and stout as third preapical tooth; inner teeth arranged in three or more series; two large mandibular serrations followed by two or three small ones (Fig. 18 J); hypostomium slightly bi-emarginate anteriorly, with 17 teeth, not arranged in conspicuous groups; median tooth and lateral teeth somewhat larger than remaining teeth; 4+4 lateral teeth inserted almost at same level as other teeth (Figs. 18 H; 19 M); lateral sclerite of thoracic proleg well developed, rather narrow for most of length but abruptly flaring below and above at anterior margin (Fig. 18 K); setae conspicuously decreasing in size from dorsal to ventral portion of anterior border of sclerite; lowermost setae barbed, some others once or twice branched on their apical half; setae arranged in one or two irregular rows, not arranged in groups and not inserted upon distinct tubercles; larval cuticle bare, except perianal scales; anal sclerite normal, X-shaped (Fig. 18 M); terminal ventral tubercles absent. Distribution: Valdivia and Magellanic areas of southern Chile and western Patagonia, with slight intrusion into Mediterranean climate area of central Chile (Fig. 136).

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Discussion: Cnesia differs from Cnephia by the much smaller spermatheca that is unusually large in Cnephia, from Stegopterna by the simplified hypostomial teeth of the larva, and from both genera by the large basal tooth on the claw of the female. Among the Neotropical Prosimuliini there are two others in which the teeth of the hypostomium of the larva are not arranged in three prominent groups: Lutzsimulium and Paraustrosimulium. Cnesia shares with Gigantodax the apomorphic structure of the genitalia in both sexes, and on this basis alone the two genera are indistinguishable. Cnesia and Gigantodax differ considerably in other ways (wing venation, onchotaxy of the abdomen of the pupa, several characters of the larvae), but their synapomorphic genitalia suggest common ancestry not shared by any other known genus. KEYS

TO SPECIES OF

CNESIA

Females 1 – 2(1) –

Metanotum glabrous ...................................................................................... Cnesia ornata Metanotum with short hair posteriorly ......................................................................... 2 Scutum pale reddish brown, dark brown on disc in some specimens; wing length more than 3.0 mm ..................................................................................... Cnesia dissimilis Scutum black, wing length 3.0 mm or less ............................................... Cnesia pusilla

Males 1 – 2(1) –

Scutum bright reddish brown; metanotum glabrous .............................. Cnesia ornata Scutum varying from dark brown to black; metanotum with short hair posteriorly ................................................................................................................................................ 2 Wing length more than 3.0 mm; scutum varying from dark reddish brown to black ........................................................................................................................ Cnesia dissimilis Wing length 3.0 mm or less; scutum black ............................................... Cnesia pusilla

Pupae 1 – 2(1) –

Head and thorax with numerous tubercles; those on thorax generally arranged in circles (Fig. 19 J) ............................................................................................. Cnesia ornata Head and thorax without tubercles ................................................................................ 2 Cephalopterothecal length approximately 2.5 mm ............................. Cnesia dissimilis Cephalopterothecal length approximately 2.0 mm ................................. Cnesia pusilla

The larvae of the species of Cnesia are so similar that we have been unable to find any differential characters, but larvae of Cnesia pusilla from Argentina can be distinguished from those of the two other species by their smaller head width. CLAVES

DE ESPECIES DE

CNESIA

Hembras 1 –

Metanoto glabro ............................................................................................. Cnesia ornata Metanoto posteriormente con cortos pelos ................................................................. 2

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2(1) –

Escudo castaño rojizo claro, castaño oscuro sobre el disco en algunos especimenes; longitud del ala mayor de 3,0 mm ................................. Cnesia dissimilis Escudo negro, longitud del ala 3,0 mm o menos .................................... Cnesia pusilla

Machos 1 – 2(1) –

Escudo castaño rojizo brillante; metanoto glabro .................................. Cnesia ornata Escudo variando desde castaño oscuro a negro; metanoto posteriormente con cortos pelos ......................................................................................................................... 2 Longitud ala mayor de 3,0 mm; escudo variando desde castaño rojizo oscuro a negro ............................................................................................................. Cnesia dissimilis Longitud ala 3,0 mm; escudo negro........................................................... Cnesia pusilla

Pupas 1 – 2(1) –

Cabeza y tórax con numerosos tubérculos, estando los del tórax generalmente ordenados en círculos (Fig. 19 J) ................................................................ Cnesia ornata Cabeza y tórax sin tubérculos .......................................................................................... 2 Longitud de cefalopterotecas aproximadamente de 2,5 mm ............ Cnesia dissimilis Longitud de cefalopterotecas aproximadamente 2,0 mm ..................... Cnesia pusilla

Las larvas de las especies de Cnesia son tan similares que no hemos podido encontrar caracteres diferenciables; pero la larva de Cnesia pusilla de Argentina puede ser separada de las otras2especies por su cabeza que es menos ancha que los otros. Cnesia dissimilis (EDWARDS) (Figs. 17 A-P; 18 A-M.) Simulium (Cnesia) dissimilis EDWARDS, 1931: 145. Cnephia dissimilis SMART, 1945: 498. Cnesia dissimilis VARGAS & DÍAZ NÁJERA, 1951: 166; COSCARÓN & WYGODZINSKY, 1962: 310; COSCARÓN, 1968: 66; 1991: 20-21; WYGODZINSKY & COSCARÓN, 1973a: 155-159; COSCARÓN, 1991: 20; CROSSKEY & HOWARD, 1997: 12; 2004: 3. Stegopterna dissimilis ENDERLEIN, 1934b: 274. Simulium (Cnephia) gynandrum EDWARDS, 1931: 147. Cnephia gynandrum SMART, 1945: 498. Cnesia gynandra ENDERLEIN, 1934b: 278. Cnesia gynandrum VARGAS & DÍAZ NÁJERA, 1951: 166. Cnephia (Stegopterna) gynandrum: RUBTZOV, 1940: 132. Simulium gynandrum VARGAS, 1945b: 142.

Female: Wing length 4.2-5.3 mm; head brown; scutum light reddish brown to dark brown in some specimens; scutellum light yellowish gray; metanotum brown; pleura brown violaceous; hyaline, wing with veins light brown with hair and spiniform setae black; legs light brown; abdomen dark grayish brown; mandible as in figure 17 A; calcipala and claw in accordance with Figs. 17 C, D; furcasternum as in figure 17 E; eighth sternite with abundant setae (Fig. 17 F); paraproct short (Fig. 17 G); genital fork with wide branches (Fig. 17 H).

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Figure 17. Cnesia dissimilis. A-H : Female; A: mandible; B: wing, general view; C: calcipala and tarsomere I; D: claw; E: furcasternum; F: eighth sternite and gonapophyses, genital fork visible through them and spermatheca; G: cercus and paraproct; H: genital fork. I-P: Male; I: anterior portion of wing, showing chaetotaxy of costa, radial 1 and radial secondary veins; J: hind basitarsus; K: gonocoxite, gonostylus ventral plate (v.p.) and endoparameres (e.d.), ventral view; L: gonocoxite and gonostylus, dorsal view; M-N: ventral plate in different position; O-P: median sclerite; O: sublateral aspect; P: dorsal view. Q-R: C. pusilla; Q: Female, genital fork; R: Male, hind basitarsus. (A-P from WYGODZINSKY & COSCARÓN, 1973a).

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Figure 18. Cnesia dissimilis. A-E: Pupa; A: general view; B: gill; C: frontoclypeus of female; D: abdomen chaetotaxy; ventral view; E: terminal portion of abdomen, ventral view. F-M: Larva; F: cephalic apotome; G: antenna; H: hypostomium; I: cephalic capsule, ventral view; J: apex of mandible and teeth; K: sclerite of thoracic proleg; L: gill histoblast; M: anal sclerite. (A-L from WYGODZINSKY & COSCARÓN, 1973a).

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Male: Wing length 3.4-4.5 mm; color as in female but a little darker; hind basitarsus wide (Fig. 17 J); ratio of length/width = 3.2-3.4; genitalia as in figures 17 K-P. Pupa: Cocoon light brown, loosely woven; covering all of body (Fig. 18 A); length of body including gills about 5.0 mm; gill length 1.7 mm; head and thorax without granulosity; gills with about 38 branches emerging from numerous primary branches implanted on short basal trunk (Fig. 18 B); frontoclypeus with prefrontal and anterior frontal trichomes spinelike (Fig. 18 C); ventral abdominal chaetotaxy as in figures 18 D, E. Larva: Length 7.5-8.0 mm; head dark brown and body yellowish brown to dark greenish; cephalic apotome and cervical sclerite as in figure 18 F; postgenal cleft not deep (Fig. 18 I); antenna as in figure 18 G; ratio of proximal, medial and distal segments = 1:1.2:1.5; cephalic fan with about 40 rays; hypostomial teeth, mandibular border, postgenal cleft, thoracic proleg sclerite, gill histoblast and anal sclerite in accordance with figures 18 H-M; thoracic proleg sclerite with about 44 teeth on comb; anal ring with 115-120 rows with 25-28 hooks each. Distribution: Cnesia dissimilis extends through all the Valdivian and Magellanic area of southern Chile and penetrates into the Mediterranean climate area of central Chile north to the province of Valparaiso. It is also very common east of the Andes in Argentina, at least from Chubut to southern Neuquen. Bionomics: The aquatic stages of this species are restricted to clear, large streams and rivers with considerable current; they are never found in trickles, small streams, irrigation ditches or any slow-flowing watercourse. The preferred water temperature varies from 10 to 12 °C. Larvae are generally found attached to branches, stems, and roots of terrestrial plants trailing in the streams; the plant involved is frequently a bamboo, Chusquea culeou DESVAUX. Pupae are found where larvae occur, but occasionally also on the undersurface of large stones or rocks resting on the river bottom. Larvae and pupae generally occur in large aggregations, with cocoons often forming irregular, multilayered masses, as in certain Prosimulium in the Northern Hemisphere. Cnesia dissimilis is a spring species, with abundant larvae and pupae found as early as October. In January, larvae become very rare, and most pupae encountered are empty. Dissection of freshly emerged females shows the species to be anautogenous. The feeding habits are unknown, but to judge from the structure of the claws of the female with their large basal process, Cnesia dissimilis is probably ornithophilic. Discussion: Although this widespread species is surprisingly uniform, some variation was observed. The wing length fluctuates between 3.2 and 4.3 mm; most specimens have wings close to 4.0 mm. Females with wing length close to 3.0 mm were observed occasionally in Chilean material. None of the Argentinean females examined had wings shorter than 3.5 mm; these variations are paralleled by variations in the measurements of pupae and larvae. The cephalopterothecal length of Argentinean pupae remains very close to 2.5 mm, but fluctuates in a Chilean population from Cautín between 2.25 and 2.50 mm. Likewise, the head width of last-instar larvae of Argentinean material varies from 0.70 to 0.75 mm, with 0.75 mm being the most frequent, but in the Chilean population mentioned it extends from 0.65 to 0.75 mm, with most specimens ranging between 0.65 and 0.70 mm. Some variation was also observed in the color of the thorax of the adults. In males, the scutum varies from dark brown to black. The scutum of the

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females, which is normally dull orange brown, may become dark brown in some specimens, especially at the center of the disc, but it is never black. Cnesia pusilla WYGODZINSKY & COSCARÓN (Figs. 17 Q, R.) Cnesia pusilla WYGODZINSKY & COSCARÓN, 1973a: 159-161; COSCARÓN, 1991: 21-22; CROSSKEY & HOWARD, 1997: 11.

Female: Wing length 2.9 mm; head blackish; scutum dull black with yellowish pronotal area; metanotum dark grayish brown; frontal angle narrow, 50 degrees; hind basitarsus 7 times as long as wide; eighth sternite, gonapophysis, paraprocts and cerci as in Cnesia dissimilis; genital fork as in figure 17 Q; terminal plates with conspicuous areas of pigment differences. Male: Length of wing 3.0 mm; overall color of head and its appendages as in female; lateral portion of thorax colored as in female; scutum velvety black; shoulders silvery gray; scutellum, metanotum, wings, and legs as in female; abdomen gray; tergal plates velvety black; basal fringe silvery gray; hind basitarsus wide (Fig. 17 R) 2.4 times as long as wide; genitalia as in Cnesia dissimilis. Pupa: Cephalopterothecal length 1.8-2.1 mm, in most specimens between 1.9 and 2.0 mm; head and thorax without tubercles; structure and onchotaxy as in Cnesia dissimilis. Larva: General structure and antenna as in Cnesia dissimilis, but specimens smaller; head width only 0.65-0.70 mm. Distribution: Argentina: Neuquen, Rio Negro. Bionomics: Larvae adhere to stones in creeks of moderate volume, about 0.5 m deep. Discussion: Cnesia pusilla seems to be very close to C. dissimilis, and may not always be easy to recognize. It differs from C. dissimilis mainly in being smaller in size in all stages, the black scutum of the female, and morphologically by the comparatively small membranous area at the insertion of the spermathecal duct on the spermatheca. Cnesia pusilla can be distinguished from C. ornata, by its smaller overall size, the presence of small hairs on the metanotum of the adults, the black scutum of the male, and the absence of tubercles on the pupa. Cnesia ornata WYGODZINSKY & COSCARÓN (Figs. 19 A-N.) Cnesia ornata WYGODZINSKY & COSCARÓN, 1973a: 161; COSCARÓN, 1991: 22-23; CROSSKEY & HOWARD, 1997: 11; 2004: 11.

Female: (Pharate) Color unknown; characters of head, eyes, antennae, mouthparts, and legs with claws as in Cnesia dissimilis, as far as discernible; thorax and abdomen not examined. Male: Length of wing 4.0 mm; scutum bright reddish brown with golden decumbent hair; scutellum pale yellow brown with long golden hair; metanotum light reddish brown, with golden pruinescence; pleura light brown; katepisternum slightly darker; wings with veins pale and hair golden brown, including basal tuft; legs light yellowish brown, their hair from golden to brown; coxae, bases and apices of femora and tibiae, and all tarsi, darkened; abdomen grayish brown, except posterior border of terga and pleural regions, which are grayish; hair of abdomen, including that of basal fringe, brass-colored.

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Figure 19. Cnesia ornata. A-G: Male; A: antenna; B: calcipala and tarsomere I; C-E: ventral plate in different position; F-G: median sclerite in different position. H-J: pupa; H: gill; I: frontoclypeus and trichomes; J: trichomes and arrangement of tubercles on thorax. K-N: Larva; K: general aspect, dorsal view; L: antenna; M: anterior border of hypostomium; N: postgenal cleft and hypostomium. (A-N from WYGODZINSKY & COSCARÓN, 1973a).

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Antenna as in figure 19 A; last segment of maxillary palp less than twice as long as penultimate segment; sensory vesicle slightly smaller than one-third diameter of third palpomere; hind basitarsus 3.8 times as long as wide; calcipala as in figure 19 B; genitalia much as in Cnesia dissimilis; ventral plate and median sclerite as in figures 19 C-G. Pupa: Cocoon light brown, covering pupa almost completely, exposing only gills and part of dorsal surface of thorax; cocoon with filaments of varying diameter, easily perceptible; cocoon often covered with foreign organic matter; length of cocoon dorsally along middle, 3.5 mm; length along base, 4.5 mm; length of body of pupa, 3.5-4.0 mm; cephalopterothecal length 2.4-2.5 mm; maximum length of gill, 1.5 mm; head and thorax of pupa light brown; gills (Fig. 19 H) very similar to those of Cnesia dissimilis; number of filaments about 35, arising from about 12 primary branches inserted on very short common trunk; some primary branches divided twice, thus giving rise to three filaments; surface of filaments minutely granulose; their apex rounded; chaetotaxy of head much as in C. dissimilis, but prefrontal and frontal anterior trichomes capitate; frontoclypeus with tubercles arranged as in figure 19 I; chaetotaxy of thorax as in C. dissimilis, but strong trichomes capitate (Fig. 19 J); exposed dorsal surface of thorax with numerous smooth tubercles arranged in circles; abdomen with general onchotaxy as in C. dissimilis; stout setae of tergum II blunt apically; tubercles of terga more numerous than in C. dissimilis. Larva: Maximum length 6.5-7.2 mm; color of body light yellowish gray-brown; head dark brown; body shape as in figure 19 K, with wider subapical area; cephalic apotome brownish with negative spots; hypostomium with about 15-20 teeth (Fig. 19 M); sublateral setae in number of 5-7 on each side; lateral serrations absent; cervical sclerites reduced and isolated from postocciput sides; antennal shape as in figure 19 L; ratio of proximal, medial and distal antennal segments = 1:1.0-1.7:1.2-1.6; postgenal cleft with small rounded concavity (Fig. 19 N); head fan with 38 rays; mandible and proleg sclerite very similar to those of Cnesia dissimilis; ninth segment without ventral tubercles; anal ring with 144-152 rows of 25-30 hooks each; anal integument without hair. Distribution: Argentina: Rio Negro; Chile: Chillán, Curacautín, Talca. Bionomics: Larvae live in torrential creeks, over “caña colihue” (Chusquea culeou). Discussion: Cnesia ornata differs from the other two species of the genus mainly by its bright reddish brown scutum in the male, the absence of setae on the metanotum, and the presence of tubercles and capitate trichomes on the head and thorax of the pupa. Gigantodax ENDERLEIN (Figs. 20 B, C, E, G, H, J, L, N, P, R, S.) Gigantodax ENDERLEIN, 1925: 205. Type-species: Gigantodax bolivianus ENDERLEIN, 1925: 205-206. Archicnesia ENDERLEIN, 1934b: 273; Type-species Simulium (Gigantodax) femineum EDWARDS, 1931: 135.

Diagnosis: Antennae with 11 segments; scutum with recumbent hair; mesepistenal sulcus shallow, evanescent anteriorly; katepisternum and pleural membrane glabrous; arms of furcasternum without projections (Fig. 20 H); wing with basal cell not developed; basal portion of R with hair arranged in several irregular series, R1 dorsally with or rarely without spinelike setae and fine hair, Rs not forked apically, Cu-A2 straight (Fig. 20 C); calcipala well developed (Fig. 20 E); female claws with large subbasal tooth of varied shapes (Fig. 20 G);

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genital fork distinctly pigmented, lateral arms with 1+1 forwardly directed well-developed sclerotized apodemes (Fig. 20 J); male gonostylus more than half as short as gonocoxite with 2-3 apical spines, ventral plate elongated with long basal arms connected by wide membrane, endoparemeres scarcely developed basally and teeth reduced to a group of 1-4 mediumsized denticles isolated on aedeagal membrane (Fig. 20 L). Pupa: Cocoon generally large; shapeless and covering most of pupal body, which is well sclerotized; frontal trichomes absent (Fig. 20 N); cephalic and thoracic trichomes spinelike; gills of varied structure, from filamentous to arborescent, tubular or globular; terminal spines thin and stout. Larva: Cephalic apotome with positive spots; cervical sclerite generally transverse and connected to upper ends of postocciput; postgenal cleft not developed; mandible with three external teeth, preapical teeth small, marginal serrations with one or two large teeth followed by several minute serrations (Fig. 20 P); hypostomial intermediate teeth shorter than lateral tooth (Fig. 20 R); larval cuticle generally bare; anal sclerite with posterior arms prolonged forming a complete ring and 1+1 backwardly directed struts underlying the X-shaped portion (Fig. 20 S); rectal papillae with 3 single lobes; abdominal segment IX with pair of ventral tubercles. Distribution: Cool and temperate western South America, high lands of Mesoamerica and one species from southwestern North America. Discussion: Gigantodax and Pedrowygomyia are differentiated from all the known Neotropical black flies by the combination of the straight Cu, An2 and structure of the larval mandible, and also by the presence in larvae of a complete anal ring and backwardly directed struts, although this anal ring is partially present in some Simulium (Hearlea) species. Gigantodax is very close to Cnesia, as was shown by COSCARÓN & MIRANDA ESQUIVEL (1998a) and reinforced by chromosomal similarity (COSCARÓN ARIAS, 1998); an interesting cladistic revision of the Gigantodax species group, using morphological and chromosomal information is presented by PINTO-SANCHEZ et al.2005. Gigantodax contains 65 species; 60 were treated in the revision of WYGODZINSKY & COSCARÓN (1989); the other species of Gigantodax are G. adleri MOULTON, 1996; G. bierigi VARGAS & RAMÍREZ PÉREZ, 1988; G. conviti RAMÍREZ PÉREZ, 1980a; G. lazoi TAKAOKA et al., 1988 and G. willei VARGAS & RAMÍREZ PÉREZ, 1988. Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL (Figs. 20 A, D, F, I, K, M, O, Q.) Pedrowygomyia COSCARÓN & MIRANDA ESQUIVEL, 1998a: 164. Type-species: Gigantodax cortesi WYGODZINSKY & COSCARÓN, 1989: 22-27.

Diagnosis: Pedrowygomyia has a close relationship with Gigantodax and the species were included by WYGODZINSKY & COSCARÓN (1989) in that genus as Gigantodax CORTESI species group; COSCARÓN & MIRANDA ESQUIVEL (1998) through cladistic studies showed the monophyly of Gigantodax species treated by the former authors, but considered that the Gigantodax CORTESI species group has enough characters to designate it as an independent genus. Here we consider only the characters that differ from Gigantodax. Adults: Frons and clypeus relatively wide, frontal median sulcus scarcely evident (Fig. 20 A); basal flagellomeres and palpomeres with long hair; scape and pedicel globose

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Figure 20. Gigantodax and Pedrowygomyia. A-B: head, frontal view; A: Pedrowygomyia cortesi, B: Gigantodax brophyi. C: wing, G. dryadicaudicis; D-E: apex of basitarsus and tarsomere 1; D: P. punapi, E: G. igniculus. FG: claws; F: P. cortesi, G: G. igniculus. H: furcasternum, G. antarctiucs. I-J: genital fork; I: P. cortesi; J: G. rufidulus. K-L: male genitalia, ventral view, K: P. jatunchuspi, showing gonocoxite, gonostylus, ventral plate and median sclerite; L: G. igniculus, showing gonocoxite, gonostylus, ventral plate, median sclerite, aedeagus membrane and teeth of endoparameres. M-N: Pupa, frontoclypeus; M: P. jatunchuspi; N: G. marginalis. O-S: Larva; O: P. jatunchuspi, head dorsal view; P: G. igniculus, mandible teeth; Q-R: hypostomial teeth, anterior margin; Q: P. cortesi, R: G. minor; S: anal sclerite, with scales and lateral struts, G. igniculus.

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and first flagellomere elongate, female mandible and maxilla obsolescent; scutum with erect and recumbent hair; R1 without spines but with numerous placoid sensilla; calcipala absent (Fig. 20 D) and claws without transverse sulcus and very short subbasal tooth (Fig. 20 F); genital fork with anterior branch wide basally, becoming thin toward apex (Fig. 20 I); male with hair on interocular space, scutum relatively flattened, ventral plate comparatively elongated (Fig. 20 K) and lateral margins at midlength convex, endoparameral teeth very short. Pupa: Frontoclypeus with frontal and facial trichomes and clypeus comparatively thinner (Fig. 20 M). Larva: Large body size, up to 9.6 mm; cephalic apotome widest near base and with basal spots (Fig. 20 O); hypostomium with first intermediate tooth longer than lateral tooth; body cuticle with small single setae implanted in dark spot. Distribution: Pedrowygomyia is restricted to the high plateau (Puna) of the Andes from northern Chile to northwestern Argentina and central Peru. This genus contains four species: Pedrowygomyia chacabamba, P. cortesi, P. jatunchuspi and P. punapi. Detailed descriptions of these species can be found in WYGODZINSKY & COSCARÓN (1989). Cnesiamima WYGODZINSKY & COSCARÓN (Figs. 21, 136.) Cnesiamima WYGODZINSKY & COSCARÓN, 1973a: 184. Type-Species: Simulium (Cnephia) atroparvum EDWARDS, 1931: 148.

Adults: Antennae with 11 segments; male holoptic; female with frons relatively wide (Fig. 21 A); mandibles of female with well-developed denticles on both margins (Fig. 21 C); apical segment of maxillary palp of both sexes less than twice as long as penultimate segment (Fig. 21 B); mesepisternal sulcus shallow, although deeper than in Cnesia, somewhat evanescent anteriorly; katepisternum glabrous, in profile slightly wider than high; pleural membrane bare; metanotum glabrous; arms of furcasternum without projections (Fig. 21 G); wings with basal cell distinct (Fig. 21 E); Sc with numerous hairs arranged in single row; basal section of R with hair arranged in several rows (Fig. 21 F); R1 with hair only, spiniform setae absent, Rs entirely hairy below, hairy above on its apical half; Rs not forked apically; M appearing thickened on most of its length; Cu2 rather weakly curved in female, more strongly so in male; A2 almost attaining wing margin; calcipala present, large, bluntly rounded apically (Fig. 21 O); claws of female with large basal tooth (Fig. 21 J); spermatheca of normal size, about as large as gonapophyses; globular, its internal surface faintly wrinkled and with few scattered spicules; whole spermatheca and base of spermathecal duct sclerotized; sclerotized portion gradually fading into unpigmented portion; gonapophyses widely subtriangular, somewhat transverse, their medial borders slightly pigmented, approximated; their surface with microtrichia and few setae (Fig. 21 L); genital fork distinctly pigmented, with stem slender, longer than arms, latter wide, lacking forwardly directed apodemes; male with gonostylus slightly shorter than gonocoxite (Figs. 21 P, Q); gonocoxite with semicircular membranous area opposite inner basal angle of gonostylus, the latter with corresponding bulge at base; gonostylus only slightly narrowed toward apex, with two distal spines (Figs. 21 Q, R); medial surface of gonostylus smooth, delimited by two carinae; ventral plate subtriangular, about as log as wide, setose, with low median longitudinal keel

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(Figs. 21 S, T); median sclerite geniculate in side view (Fig. 21 U), roughly M-shaped in full view, with the inner arms of the M partly fused and connected to basal flange (Fig. 21 V); endoparameres reduced to subquadrate plates connected to arms of ventral plate (Fig. 21 U); denticles not developed. Pupa: Cocoon well defined, covering pupa except anterior extremity and gills; shape broadly oval, flattened, shield like; borders of anterior opening slightly but distinctly reinforced, especially at center, somewhat salient at middle (Fig. 22 A-B); cocoon without ventral layer; gills cigar-shaped, inserted perpendicularly on extremely short trunk at posterior third of body of gill, the gill projected forward and downward and somewhat laterally; gills about as long as maximum width of body of pupa, with several irregularly transverse septa; surface of gills rugose-reticulate; gills bearing delicate scattered respiratory filaments extending perpendicularly to longitudinal axis of gill, length of filaments over twice that of diameter of gill, their surface minutely pitted, their apex slightly clavate (Fig. 22 C); head only with 1+1 hair-like facial trichomes (Fig. 22 D); disc of thorax with 5-6+5-6 trichomes, the three dorsal ones in shape of apically bent strong spine-like setae, the others hair like; abdomen pointed posteriorly; distinct pleural areas not developed; onchotaxy of abdomen as follows: tergum I glabrous; tergum II with several fine hairs, 1+1 submedian ones slender, spinelike; terga III and IV with 3+3 short hooks posteriorly; terga V-VIII with spine-combs; terminal spines short, sharply pointed; sternum V with 2+2 elongate spine-like hooks; sternal plates VI and VII with 1+1 similar hooks; 1+1 hooks also in pleural membrane of segments VI and VII; pleural area of terminal segments with several extremely complex grapnel-shaped setae (Fig. 22 E). Larva: Abdomen relatively elongated, with 1+1 ventral tubercles (Fig. 22 F); cephalic apotome widened before posterior border; cervical sclerites of reduced size and isolated from postocciput margin (Fig. 22 G); postgenal cleft obsolescent (Fig. 22 H); antenna surpassing base of cephalic fan stem; distal antennal segment very thin and more than twice length of basal segments and medial segment very short (Figs. 22 I, J); palp elongated; rays of cephalic fan very thin showing larger setae alternately; mandible with scarce marginal teeth (Figs. 22 K, L); hypostomium with anterior border surpassing the teeth except the angle tooth (Fig. 22 M); thoracic proleg subquadrate; anal sclerite Xshaped, without or only faint rods (Fig. 22 N); body cuticle nude. Distribution: Valdivian and Magellanic areas of southern Chile, and western Patagonia. Discussion: The adults of Cnesiamima are superficially very similar to those of Paraustrosimulium, as already stated by EDWARDS (1931); this similarity extends to the general shape of the median sclerite of the male genitalia, but the structure of the endoparameres and of the female genitalia as found in the two genera is quite different, and does not support a hypothesis of close relationship. The following enumeration summarizes the significant differences between the adults of Cnesiamima atroparva and Paraustrosimulium anthracinum. In regard to the remaining South American Prosimuliini, the adults of Cnesiamima can be distinguished from the Chilean-Patagonian Cnesia, Araucnephia, and Araucnephioides by, among other characters, the absence of spine-like setae on R1, the sclerotized neck of the spermatheca, and certain features of the male genitalia, such as the bulge at the base of the distimere, the absence of denticles on the endoparameres, and the M-shaped median sclerite. Cnesiamima can be further distin-

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Figure 21. Cnesiamima atroparva. A-M: Female; A: frons; B: maxillary palp; C: mandible; D: frontoocular triangle; E: wing, general view; F: portion of wing with C and R1; G: furcasternum; H: sensory vesicle of maxillary palp; I: hind leg; J: claw; K: genital fork; L: eighth sternite and gonapophyses, genital fork and spermatheca; M: cercus and paraproct: right, ventral view, left mounted on slide. NV: Male; N: hind leg; O: calcipala and tarsomere I; P: gonocoxite, gonostylus, ventral plate and median sclerite in ventral view; Q: gonocoxite and gonostylus, dorsal view; R: gonostylus, showing inner membranous surface; S-T: ventral plate in different position; U: ventral plate, median sclerite and plate of endoparameral organ; V: median sclerite. (A-V from WYGODZINSKY & COSCARÓN, 1973a).

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guished from Araucnephia and Araucnephioides by A2 which almost attains the wing margin, and from the latter genus also by the larger basal tooth of the claws of the female. The differences between Cnesiamima and Gigantodax are obvious in wing venation and the anal sclerite of the larva. Lutzsimulium differs by the absence of spiniform setae on R1, the large tooth of the claw of the female, and by the peculiarities of the male and female genitalia. Cnesiamima differs from Mayacnephia by the absence of spiniform setae on R1, the katepisternum which is wider than high, the sclerotized neck of the spermatheca, and by the quite different structure of the median sclerite of the male. Cnesiamima shares with Tlalocomyia a spermatheca with a sclerotized neck, although there are differences in detail, but the apical segment of the maxillary palp of Tlalocomyia is very long in both sexes, the pleural membrane is hairy on its upper portion, spiniform setae are on R1, and the median sclerite of the male is quite different. Cnesiamima atroparva (EDWARDS) (Figs. 21 A-V; 22 A-O.) Simulium (Cnephia) atroparvum EDWARDS, 1931: 148. Cnesiamima atroparva WYGODZINSKY & COSCARÓN, 1973a: 187-189; 1973b: 5; COSCARÓN, 1985: 67; 1991: 74-75; CROSSKEY & HOWARD, 1997: 12; 2004: 3. Cnephia atroparvum SMART, 1945: 498. Cnesia atroparvum VARGAS & DÍAZ NÁJERA, 1951: 165. Simulium atroparvum VARGAS, 1945a: 116. Simulium (Astega) atroparvum RUBTSOV, 1940: 532.

Female: Wing length 3.1-3.5 mm; color of head and its appendages black; eyes, antenna, palp, proboscis, and occiput dark grayish brown; frons and clypeus gray pollinose; pilosity of antenna very short, that of palp, clypeus, frons, and occiput somewhat longer; hair from brass-colored to black; scape and pedicel slightly lighter than flagellum, and with longer hair; scutum dark gray, with silvery to nacreous pruinosity; shoulders reddish; appressed setae of scutum brass-colored; scutellum reddish brown, its hair from brass-colored to dark brown; metanotum black, silvery pollinose; wings hyaline, veins medium brown; spines and hair from dark brown to black; hair on wing base from yellowish to piceous; halteres from yellowish to pale brown; pleural tuft brass-colored; pleura dark grayish brown, silvery pollinose; legs rather uniformly light brown to piceous, their hair from brass-colored to black; abdomen brownish or grayish, tergal plates darker; membranous areas light grayish brown; hair of abdomen, including basal fringe, brasscolored; frons (Fig. 21 A) medium wide, frontal angle 75 degrees; fronto-ocular triangle distinctly deeper than wide (Fig. 21 D); maxillary palp as shown in figure 21 B; third segment somewhat swollen; structure of sensory vesicle as shown in figure 21 H, its diameter about half that of third palpomere; lacinia with 25-28 denticles, mandible with 3741 denticles (Fig. 21 C); structure of wing as in generic description; shape and proportion of segments of hind leg as shown in figure 21 I; calcipala shorter than width at base; claws as shown in figure 21 J; tergal plates of abdomen wider than long, except on terga II-IV where they are as wide as long; ventral plates not developed; eighth sternum with 1+1 groups of 16-20 long setae; disc of eighth sternite uniformly pigmented except darker subsemicircular band anteriorly (Fig. 21 L); paraprocts short, rounded distally (Fig. 21 M).

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Male: Wing length 2.9-3.2 mm; head black; eyes red in live specimens, mostly black in dried material; antennae, palp, and clypeus black; occiput dark gray pollinose; proboscis dark grayish brown; antennae with short silvery to black pubescence; scape and pedicel, palpi, clypeus and occiput with long brown or black hair; scutum velvety black, scutellum dark brown, with brown to black hair; metanotum black, with silvery pruinosity; color of wings, legs, pleura, and pleural tuft as in female; abdomen velvety black dorsally, grayish at sides and below; hair of abdomen from brass-colored to black; third palpomere more slender than in female; sensory vesicle only slightly smaller than in female; wings as in female, but Cu2 with stronger curvature; shape and proportions of segments of hind leg as shown in figure 21 N; hind basitarsus 3.5-3.9 times as long as wide; calcipala slightly shorter than width at base (Fig. 21 O); gonocoxite subtrapezoidal (Figs. 21 P-Q), longer than wide, with membranous area apically on inner surface projecting in some cases; gonostylus elongate, subcylindrical (Figs. 21 P-R); ventral plate as shown in figures 21 S, T, U; lateral projections and basal margin conspicuously sclerotized; median sclerite as shown in figures 21 V-U Endoparameral organ as in generic description and partially in figure 21 U. Pupa: Cocoon grayish brown, with distinct threads; shape as in figures 22 A-B; length of cocoon 3.3-4.0 mm, its maximum width 2.7 mm; total length of pupa (gills not included) 2.7-2.9 mm; tubercles weakly developed, perceptible only on thorax, even there ill defined; gill length 1.2-1.4, its maximum width 0.25-0.30 mm; gills each with 13-17 septa, and with about 35 rather regularly spaced respiratory filaments; filaments inserted on spaces between septa (Fig. 22 C); onchotaxy of head, thorax, and abdomen as in generic description and figures 22 D-E; spine-combs of posterior terga with 12-22 widely spaced teeth. Larva: Maximum length 6.0-6.5 mm; cephalic capsule width, 0.7 mm; general color light grayish green, head light yellowish brown; shape as in figure 22 F; cephalic apotome with positive dark brown spots, and darkened on posterior margin (Fig. 22 G); cephalic fan with unknown number of rays; ratio of proximal, medial and distal antennal segments = 1: 0.16-0.27:2.5-3.4; shape as in figures 22 I, J; mandible with 10-12 internal teeth and 35 marginal denticles (Figs. 22 K, L); hypostomium with median tooth lower than lateral teeth (Fig. 22 M); 3-5 lateral setae on each side (Fig. 22 H); thoracic proleg lateral sclerite with 14-17 teeth; anal sclerite with hair, not scales (Fig. 22 N); anal ring with 75-80 rows of 13-15 hooks each; pupa gill histoblast as in figure 22 O. Distribution: Argentina: Neuquén, Río Negro, Tierra del Fuego; Chile: Curicó, Llanquihue, Magallanes, Talca, Valdivia. Bionomics: The pupae were collected in crystal-clear, small mountain streams at water temperatures of 0-6 °C. The larvae were found in Tierra del Fuego in a creek covered by ice and snow; pupae were found singly, on the upper surface of stones or on leaves of shrubs trailing in the water. Discussion: Cnesiamima has a great similarity with Paraustrosimulium in characters of male genitalia, pupa and larva. They are easy to separate by the pupal gills that are quite different superficially, those of Paraustrosimulium being lamellate and glabrous and those of Cnesiamima cigar-shaped and beset with numerous respiratory filaments.

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Figure 22. Cnesiamima atroparva. A-E: Pupa; A-B: general aspect in dorsal and lateral views. C: gill. D: frontoclypeus and trichome. E: apical portion of abdomen, ventral view, with trichomes and terminal spur. F-O: Larva; F: general aspect, lateral view; G: cephalic apotome; H: postgenal cleft and hypostomium; I: antenna, general view; J: portion of antenna showing medial and part of proximal and distal articles; K: apical portion and teeth of mandible; L: marginal serrations of mandible; M: anterior portion of hypostomium; N: anal sclerite and hair; O: gill histoblast. (A-E from WYGODZINSKY & COSCARÓN, 1973b).

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Paraustrosimulium WYGODZINSKY & COSCARÓN (Figs. 23, 136.) Type-species: Simulium anthracinum BIGOT, 1888. Austrosimulium (Paraustrosimulium) WYGODZINSKY & COSCARÓN, 1962: 244; STONE, 1963: 14. Paraustrosimulium CROSSKEY, 1969: 17.

Adults: Antenna with 10 segments; male holoptic; female with frons relatively narrow (frontal angle about 65 degrees); mandibles of female with well-developed denticles on both sides; apical segment of maxillary palp of both sexes less than twice as long as penultimate segment; mesepisternal sulcus deep, well defined anteriorly; katepisternum glabrous, in profile much wider than high; metanotum glabrous; arms of furcasternum with conspicuous projections (Fig. 23 A); pleural membrane bare; wings with basal cell distinct; Sc with numerous hairs arranged in single row; basal section of R with hair arranged in several rows; R1 with hair and very slender spine-like setae arranged in one or two irregular rows; Rs entirely hairy below, hair arranged in single row at base and apex of vein, in several rows on central portion of vein; apical portion of Rs dorsally on apical half with one series of hairs; Rs not forked apically; M appearing thickened on most of its length; curvature of Cu2 normal, somewhat more pronounced in male than in female; A2 falling distinctly short of wing margin; calcipala present, large, rounded apically; claws of female with medium-sized subtriangular basal tooth (Fig. 23 B); spermatheca larger than usual, larger than gonapophysis; its internal surface smooth, with very small number of scattered spiculae; spermatheca fully sclerotized, except very small subcircular membranous area at insertion of spermathecal duct; paraproct short with truncate apical border (Fig. 23 C); gonapophyses widely subtriangular, somewhat transverse, and closely approximated, their surface with microtrichia and few very short setae, medial border slightly pigmented (Fig. 23 D); genital fork lacking conspicuous pigment; stem very short and wide, not longer than arms, the latter also very wide; forwardly directed apodemes absent (Fig. 23 E); male with gonostylus slightly shorter than gonocoxite, subtriangular, with two to four spines; medial surface of gonostylus smooth, delimited by two carinae (Fig. 23 F); ventral plate subpentagonal, with distinct median carina (Fig. 23 G); median sclerite distinct, geniculate in lateral view; basal portion narrow proximally, somewhat widened and deeply incised distally; apical portion composed of two simple divergent arms (Fig. 23 H); endoparameres composed of distinct free plate-like basal portion and group of elongate parallel denticles not connected by sclerotized area to plate-like portion. Pupa: Cocoon well defined, wall-pocket shaped, viz., without anteroventral collar; undersurface of cocoon reduced to few irregular strands near base of cocoon; body of pupa distinctly sclerotized throughout, although not strongly so; respiratory organs in shape of twisted pseudo-segmented lamella (Fig. 23 I); head sclerite with frontal and facial trichomes, all hair like; disc of thorax with approximately 5+5 hair like trichomes, furthermore with 3+3 spine-like trichomes dorsally near midline; abdomen pointed posteriorly; distinct pleural areas not developed; terga III and IV with 4+4 hooks posteriorly; terga V to IX with continuous spinecombs anteriorly; sternum V with 4+4 hooks posteriorly; sternal plates VI and VII with 1+1 hooks, and 1+1 hooks each also in pleural membrane of segments VI and VII; terminal spines in shape of short, pointed tubercle; posterior segments with several grapnel-shaped setae (Fig. 23 J). Larva: Head spots positive; cephalic apotome widest slightly before hind margin; cervical sclerites composed of small subcircular sclerotized area and transverse less strongly sclerotized area, the latter not fused to upper ends of postocciput (Fig. 23 K); postgenal cleft not developed;

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N Figure 23. Paraustrosimulium anthracinum. A-E: Female; A: furcasternum; B: claw; C: cercus and paraproct; D: eighth sternite and gonapophyses; E: genital fork. F-H: Male; F: gonocoxite and gonostylus, dorsal view; G: ventral plate; H: aedeagus with median sclerite, endoparameres and ventral plate visible through it. I-J: Pupa; I: general aspect dorsal view; J: apex of abdomen with trichomes, spines and terminal spurs. K-N: Larva; K: cephalic apotome; L: antenna; M: hypostomium; N: anal sclerite, with lateral rods, hair and spines. (A-N from WYGODZINSKY & COSCARÓN, 1973a).

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hypostomium much shorter than postgenal bridge; antennae slightly longer than stem of cephalic fans; their two basal segments very dark, the apical two very light-colored (Fig. 23 L); cephalic fans with rays normal; mandibles with two external and one apical tooth; first and second preapical teeth of identical size, rather wide; third preapical tooth larger than others, but much smaller than apical tooth, and indistinguishable from internal teeth; the latter arranged in single row; one large mandibular serration followed by from two to four smaller ones; short basal setae of mandible very slender, glabrous; hypostomium with 13 teeth, not arranged in conspicuous groups, median tooth and lateral teeth slightly larger than others; the 2+2 lateral teeth inserted almost at same level as remaining teeth (Fig. 23 M); teeth of hypostomium underlain by lamellar extension of hypostomium; lateral sclerite of thoracic proleg subtriangular, slightly wider than long rounded at base, strongly widened apically; setae of lateral sclerite shorter than sclerite, arranged in two or three irregular rows, inserted individually on barely perceptible tubercles; abdominal cuticle bare, except for conspicuous perianal scales; anal sclerite X-shaped; 1+1 backwardly directed struts under lying main body of sclerite (Fig. 23 N). Distribution: Valdivian and Magellanic areas of southern Chile, and western Patagonia (Fig. 136). Bionomics: SILVA FIGUEROA (1917) mentioned that P. anthracinum (here Simulium moorei) bites humans on the Taitao Peninsula of Chile, and can become very bothersome. EDWARDS (1931) examined a specimen that “bit, causing painful swelling of hand.” We have not recorded anthropophilic behaviour in this species. The aquatic instars are found in clear, cold, small and large streams. Pupae and larvae were found attached to rocks, leaves, pieces of wood, and dead branches trailing in the stream, invariably close to the surface. Discussion: Paraustrosimulium presents an array of autapomorphic characters that distinguish it from all other Western Hemisphere Simuliini genera, i.e., the 10-segmented antennae, the peculiar endoparameral organ and the almost unpigmented genital fork with its short and wide stem; the well-formed cocoon and the unique gills of the pupa. The extremely elongate third antennal segment in the larva are shared with Cnesiamima; the backwardly directed struts underlying the anal sclerite are also found in Gigantodax. Paraustrosimulium shares with Lutzsimulium and Simulium the presence of conspicuous projections on the arms of the furcasternum, an apomorphic character within the framework of the family. Paraustrosimulium anthracinum (BIGOT) (Fig. 23.) Simulium anthracinum BIGOT, 1888: 15. Simulium (Austrosimulium) anthracinum EDWARDS, 1931: 143. Austrosimulium anthracinum SMART, 1945: 499; WYGODZINSKY, 1953a: 293; DUMBLETON, 1960: 543. Austrosimulium (Paraustrosimulium) anthracinum WYGODZINSKY & COSCARÓN, 1962: 242; COSCARÓN, 1968: 66. Paraustrosimulium anthracinum CROSSKEY, 1969: 73; WYGODZINSKY & COSCARÓN, 1973a: 191-192; COSCARÓN 1991: 76-78. Simulium moorei SILVA FIGUEROA, 1917: 30. Simulium (Austrosimulium) moorei EDWARDS, 1931: 144. Austrosimulium moorei SMART, 1945: 499.

Female: Wing length 2.8-3.3 mm; general coloration blackish; scutum black; scutellum grayish-brown, metanotum black with silvery pollinosity; hyaline wing with brown veins,

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with hair and setae black; legs dark brown, abdomen grayish brown; palpal sensory organ about 1/3 segment length; length/width ratio of hind basitarsus = 6.1; calcipala about 1/3 tarsomere I length; furcasternum (Fig. A), claw (Fig. B) and genitalia (Figs. C-E) as in generic description. Male: Wing length 3.0 mm; coloration as for female; palpal sensory organ about half of segment length; length/width ratio of hind basitarsus = 4.5-4.8; calcipala about ¼ tarsomere length; abdominal tergal plates large, sternal plates small; genitalia (Figs. F-H) as in generic description. Pupa: Cocoon color light yellowish brown, weave compact; reinforced on anterior margin, covering most of body but leaving gill free (Fig. I); length on dorsum 3.2-3.5 mm, on base 3.5-4.4 mm; gill length 2.0-2.2 mm; gill foliaceus, annulated with 2 branches basally, internal larger than external; cephalothorax with abundant small tubercles; chaetotaxy as in generic description; thorax trichomes very strong and terminal spines very short (Fig. J). Larva: Maximum length 6.5 mm; general coloration light grayish yellow; cephalic apotome darkened basally and medially, with positive spots (Fig. K); antennal shape as in figure L; ratio of proximal, medial and distal antennal segments = 1:0.8-0.9:2.1-2.2; hypostomial teeth shorter than frontal margin of hypostomium; with 5 or 6 lateral setae on each side (Fig. M); sclerite comb of pseudopod with about 33 teeth; rectal papillae with 70 rows of 15-18 hooks each; anal sclerite with abundant small scales (Fig. N); rectal papillae with three single lobes. Distribution: Neantarctic biogeographic region (Central Chile plus subAntarctic realms); Argentina: Chubut, Neuquén, Tierra del Fuego; Chile: Aysen, Chiloe, Llanquihue, Magallanes, Malleco, Osorno, Valdivia (Fig. 136). Bionomics: It lives on vegetation in creeks and small rivers, not too torrential. Lutzsimulium D’ANDRETTA & D’ANDRETTA (Figs. 24, 25, 136.) Lutzsimulium D’ANDRETTA & D’ANDRETTA, 1947: 402. Type-Species: Simulium hirticosta LUTZ, 1909: 135 (as Lutzsimulium cruzi D’ANDRETTA & D’ANDRETTA).

Adults: Antennae 11-segmented; male holoptic; female with frons comparatively wide (Fig. 24 A); frontal angle about 90 degrees; mandibles of female toothed only on inner margin; apical segment of maxillary palp of both sexes only very slightly longer than penultimate segment; mesepisternal sulcus rather deep, not evanescent anteriorly; katepisternum glabrous, in profile about as high as wide; arms of furcasternum with conspicuous projections (Fig. 24 L); metanotum glabrous; wings without basal cell; R1 with spine-like setae (Fig. 24 H); Sc and basal section of R setose; Rs entirely hairy below, setose above on distal half; Rs not forked apically; M appearing duplicated for most of its length; Cu2 conspicuously curved, in some specimens with rudimentary branch submedially; A2 ending well before wing margin; calcipala large, widely rounded apically (Fig. 24 C); hind basitarsus of male not strongly widened; claws of female with one very small, pointed, subbasal tooth (Fig. 24 D); gonapophyses large, transversely subtriangular, their apex bluntly rounded; medial borders approximated, slightly pigmented (Fig. 24 E); surface of gonapophysis with microtrichia and few short setae; genital fork distinctly pigmented; stem slender, only slightly longer than arms, the latter wide, without apodemes (Fig. 24 F); spermatheca of normal size,

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fully sclerotized, not reticulate on either surface; internal surface with scattered spiculae; subcircular area at insertion of spermathecal duct with peculiar network-like structure extending to base of duct, there gradually fading (Fig. 24 G); male with gonostylus slightly shorter than gonocoxite, subconical, with from two to four apical spines (Figs. 24 I, M); ventral plate subquadrate or subrectangular, with conspicuous keel (Fig. 24 J); median sclerite somewhat S-shaped in lateral view; in frontal view with proximal portion entire, wide basally, narrowed distally (Fig. 24 J); apical portion in shape of two short, diverging arms connected by membrane; short ends of arms free, backwardly curved; endoparameres scarcely perceptible, reduced to basal plates (Fig. 24 I). Pupa: Cocoon without definite shape, covering abdomen and only part of thorax of pupa; body of pupa well sclerotized throughout (Fig. 25 A); head and thorax with or without tubercles; many trichomes of head and thorax and setae of terminal abdominal segments tightly looped apically (Fig. 25 E); gills arborescent, divided into primary and secondary branches close to base; general structure of branches filamentous (Figs. 25 B, C); head sclerite with 1+1 facial trichomes; frontal trichomes present or absent; thorax with 4+4 or 5+5 trichomes; abdomen pointed posteriorly; distinct pleural areas not developed; onchotaxy: terga III and IV with 4+4 hooks posteriorly; terga VI-IX with continuous spine-combs anteriorly; sternum IV without or with 1+1, V with 2+2 hooks posteriorly; sternal plates VI and VII with 1+1 hooks; 1+1 hooks each also in pleural membranes of segments VI and VII; terminal spines well developed, long and slender, strongly curved at base (Fig. 25 E); setae of segments VIII and IX strong, looped apically, never branched. Larva: Head spots positive; cephalic apotome widest well before hind margin (Fig. 25 G); cervical sclerites either wide, fused to upper end of postocciput, or small and free; postgenal cleft large (Fig. 25 H); antennae as long or slightly shorter than stem of cephalic fan (Fig. 25 G); third segment distinctly darker than preceding; cephalic fans normal with large setae alternating among shorter setae; mandibles with two outer teeth in addition to apical tooth; second preapical tooth much smaller than either first or third; third smaller or larger than first, and distinctly larger than first inner tooth; inner teeth arranged in three series; mandibular serrations numbering two, first much larger than second; short basal setae slender, glabrous (Fig. 25 K); hypostomium with anterior border slightly emarginated; teeth numbering 13, subequal in size, not arranged in conspicuous groups; 2+2 lateral teeth inserted much below level of lateral teeth (Fig. 25 I); lateral sclerite of proleg well developed, subtriangular, about as long as wide (Fig. 25 M); setae of lateral sclerite less than half as long as sclerite, arranged in irregular row extending slightly beyond main body of sclerite below; larval cuticle bare or conspicuously hairy (Fig. 25 N); perianal scales well developed; ninth segment with ventral tubercles; anal sclerite normal, Xshaped (Fig. 25 L). Distribution: This genus has been found in Misiones (Argentina) and in the coastal ranges of southern Brazil, from the State of Rio de Janeiro south to Paraná (Fig. 136). Bionomics: The aquatic instars have been found in fast- and slow-flowing mountain streams, generally at elevations of 1000 m or above, although a few sites of lower elevation are known. LUTZ (1910) collected females of L. pernigrum on horses. Discussion: In many respects, Lutzsimulium fits the features normally associated with the Prosimuliini, as understood by CROSSKEY (1969): the short apical segment of the maxillary palp

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of the adults, the absence of a pedisulcus, multiple apical spines on the gonostylus, the structure of the median sclerite of the genitalia of the male not unlike that found in Prosimulium or Stegopterna, the shapeless loosely woven cocoon, the comparatively heavily sclerotized pupal abdomen, the membranous area dividing sterna VI and VII of the pupa along their middle, the long terminal spines of the pupal abdomen and the modified setae on the terminal abdominal segments, and the dark third segment of the antenna of the larva. On the other hand, Lutzsimulium shows an array of apomorphic characters generally associated with the tribe Simuliini sensu CROSSKEY: the rather deep mesepisternal sulcus, the absence of a basal cell in the wing, the conspicuous apodemes of the arms of the furcasternum, the Simulium-like hypostomium of the larva, the small number of mandibular serrations, and the extremely well-developed postgenal cleft. It is not possible to place Lutzsimulium, using conventional criteria, unequivocally in one or the other tribe accepted by CROSSKEY (loc. cit.). The combination of the peculiar autapomorphic characters of Lutzsimulium, such as the meshlike surface structure of part of the spermatheca, the reduced endoparameral organ in the genitalia of the male, the apically looped trichomes of the thorax of the pupa, and the size ratios of the preapical teeth of the mandible of the larvae are sufficient to distinguish Lutzsimulium from any described genus-group in the Simuliidae. KEYS

TO

LUTZSIMULIUM

SPECIES

Adults Lutzsimulium flavopubescens adults are not included due to lack of sufficient information. –

–

Female and male scutum reddish brown with coppery pilosity; female sensory organ of maxillary palp big (about 0.5 of palpomere length, Fig. 24 B); third segment of palp relatively thick; male median sclerite without strong constriction on median area (Fig. 24 J) ............................................................ Lutzsimulium hirticosta Female and male scutum dark velvety brown with brown to reddish pilosity; female sensory organ of maxillary palp small (about 0.3 of palpomere length, Fig. 24 K); third segment of palp relatively thin; male median sclerite with strong constriction on median area .......................................................Lutzsimulium pernigrum

Pupae 1 – 2(1) –

Gill branches 20-22; frontoclypeus and thorax with abundant tubercles (Fig. 25 D) ............................................................................................................ Lutzsimulium hirticosta Gill branches 12-14; frontoclypeus and thorax with or without tubercles ............ 2 Gill branches 14; frontoclypeus and thorax without tubercles .................................. ...........................................................................................................Lutzsimulium pernigrum Gill branches 12; frontoclypeus and thorax with tubercles ........................................ .................................................................................................... Lutzsimulium flavopubescens

Larvae –

Postgenal cleft deep, with postgenal bridge half length of hypostomium; cervical sclerites isolated from postocciput border; anal ring with 72-74 rows of 12-13 hooks each ....................................................................................... Lutzsimulium hirticosta

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Postgenal cleft not deep, with postgenal bridge as long as or longer than hypostomium; cervical sclerites wide at postocciput border; anal ring with 96-110 rows of 18-25 hooks each .......... Lutzsimulium pernigrum, Lutzsimulium flavopubescens DE ESPECIES DE

LUTZSIMULIUM

Adultos No se incluye Lutzsimulium flavopubescens por ser escasa la información. –

–

Escudo de hembra y macho castaño rojizo con pilosidad cobriza; vesícula sensorial del palpo maxilar de hembra grande (aproximadamente 0,5 de la longitud del palpómero, Fig. 24 B); tercer artejo del palpo relativamente grueso; esclerito mediano del macho sin fuerte constricción en área mediana (Fig. 24 J) .................... ............................................................................................................ Lutzsimulium hirticosta Escudo de hembra y macho castaño oscuro aterciopelado, con pilosidad castaño rojiza; vesícula sensorial del palpo maxilar pequeña (aproximadamente 0,3 del palpómero, Fig. 24 K); tercer artejo del palpo relativamente fino; esclerito mediano del macho con fuerte constricción sobre el área mediana ......................... ...........................................................................................................Lutzsimulium pernigrum

Pupas 1 – 2(2) –

Ramas de las branquias en número de 20-22; frontoclípeo y tórax con abundantes tubérculos (Fig. 25 D) ................................................................... Lutzsimulium hirticosta Ramas de las branquias en número de 12-14; frontoclípeo y tórax con o sin tubérculos ............................................................................................................................ 2 Ramas de las branquias en número de 14; frontoclípeo y tórax sin tubérculos ..... ...........................................................................................................Lutzsimulium pernigrum Ramas de las branquias en número de 12; frontoclípeo y tórax sin tubérculos ..... .................................................................................................... Lutzsimulium flavopubescens

Larvas ––

Hendidura postgenal profunda, con puente postgenal la mitad del largo del hipostomio; escleritos cervicales aislados del borde del postocciput; anillo anal con 72-74 hileras de 12-13 ganchos cada una ......................... Lutzsimulium hirticosta Hendidura postgenal no profunda, con puente postgenal tan largo o más que el hipostomio; escleritos cervicales anchos, llegando al borde del postocciput; anillo anal con 96-110 hileras de 18-25 ganchos cada una ..................................................... .......................................................... Lutzsimulium pernigrum, Lutzsimulium flavopubescens

Lutzsimulium hirticosta (LUTZ) (Figs. 24 A-J; 25 A-N.) Simulium hirticosta LUTZ, 1909: 135; 1910: 226; PINTO, 1931: 756. Lutzsimulium hirticosta VULCANO, 1967: 4; WYGODZINSKY & COSCARÓN, 1973a: 197; PY-DANIEL, 1982: 305; COSCARÓN, 1991: 80-81. Lutzsimulium cruzi D’ANDRETTA & D’ANDRETTA, 1947: 401; 1948: 639.

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M Figure 24. A-J: Lutzsimulium hirticosta. A-G: Female; A: frons; B: third palpomere and sensory vesicle; C: calcipala and palpomere I; D: claw; E: portion of eighth sternite and gonapophyses; F: genital fork; G: portion of spermatheca with insertion base of spermathecal duct. H-J: Male; H: chaetotaxy of wing anterior veins; I: genitalia, ventral view showing gonocoxite and gonostylus; J: ventral plate and portion of median sclerite. K-N: L. pernigrum. K-L: Female; K: third palpomere with sensory vesicle; L: furcasternum. M-N: Male; M: gonocoxite and gonostylus; N: Pupa, portion of thorax with gill and trichomes.

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A

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L N Figure 25. Lutzsimulium hirticosta. A-E: Pupa; A: general view; B: gill; C: basal portion of gill, magnified; D: frontoclypeus; E: abdomen distal portion, ventral view, showing curled trichomes and terminal spur. F-N: Larva; F: general aspect, lateral view; G: head, dorsal view; H: postgenal cleft and hypostomium; I: hypostomium, anterior border; J: antenna; K: mandible apex showing teeth; L: anal sclerite, with scales and hairs; M: lateral sclerite of thoracic proleg; N: body integument hair.

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Female: Wing length 2.4-2.7 mm; head blackish; antenna, palp and proboscis dark brown to blackish; frons light grayish brown, scutum reddish brown with coppery pilosity and abdomen dark grayish brown; legs brownish with darker areas blackish, hair and spines dark brown to black; frons with faint median sulcus and abundant pilosity (Fig. 24 A); sensory organ of maxillary palp about 2.5 times length of third palpomere (Fig. 24 B); Sc with about 50 hairs; calcipala as figure 24 C; eighth sternite with about 27 hairs; gonapophyses with truncate border (Fig. 24 E); genital fork with median branch well sclerotized (Fig. 24 F); spermatheca with insertion base of spermathecal duct not totally sclerotized, mesh like(Fig. 24 G). Male: Wing length 2.4-2.6 mm; coloration as in female but more darkened and with more abundant blackish pilosity; distribution of wing vein hair and setae as in figure 24 H; length/width ratio of hind basitarsus = 6; gonocoxite, gonostylus, endoparameres and ventral plate as shown in figure 24 I; median sclerite expanded distally and without strong constriction medially (Fig. 24 J). Pupa: Cocoon leaving thorax and gills free (Fig. 25 A); length at base 3.0-3.5 mm, dorsally 2.5-3.1 mm; gill length 2.0-2.5 mm; frontoclypeus and thorax dorsally with abundant tubercles (Fig. 25 D); frontal trichomes only present in male; gill arborescent with 20-22 branches arriving from short basal trunk with primary and secondary branches relatively short (Figs. 25 B-C); abdomen with tubercles; distal portion of abdomen with trichomes apically curved and terminal spines strong (Fig. 25 E). Larva: Maximum length 5.0-5.5 mm; general aspect as figure 25 F; color grayish yellow; cephalic apotome with abundant hair and positive spots well developed; cervical sclerites small and isolated in postocciput (Fig. 25 G); postgenal bridge nearly half hypostomium height and 1/4 of postgenal cleft (Fig. 25 H); ratio of length of proximal, medial and distal antennal segments = 1:0.8:2.0; hypostomium with median tooth at same height as lateral teeth (Fig. 25 I) with intermediate teeth well developed; antenna as figure 25 J; mandibular dentition as in figure 25 K; thoracic proleg sclerite with about 30 teeth (Fig. 25 M); anal sclerite area with abundant hair and minute single scales (Fig. 25 L); anal ring with 72-74 rows with 12-13 hooks each; larval cuticle with hair more abundant on distal portion of body (Fig. 25 N). Distribution: Argentina: Misiones; Brazil: Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Lutzsimulium hirticosta breeds on vegetation in creeks with a small volume of crystalline water. Discussion: The closest species is L .pernigrum; differences are given in the keys. Lutzsimulium pernigrum (LUTZ) (Figs. 24 K-N.) Simulium pernigrum LUTZ, 1910: 263; PINTO, 1931: 708-756. Simulium (Eusimulium) montanum LUTZ, 1909: 128 (nec PHILIPPI). Simulium (Friesia) pernigrum RUBTZOV, 1940: 124. Lutzsimulium pernigrum D’ANDRETTA & D’ANDRETTA, 1952: 312-324; WYGODZINSKY & COSCARÓN, 1973a: 197; PY-DANIEL, 1982: 304-305; COSCARÓN, 1991: 81-82.

Female: Wing length 2.5-3.1 mm; head blackish with dark brown appendages; scutum dark brown with brown to reddish hair; legs dark brown; sensory organ of

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maxillary palp small, about 1/5 of third palpomere length (Fig. 24 K); furcasternum as in figure 24 L; genitalia similar to those of Lutzsimulium hirticosta. Male: Wing length 2.8-3.0 mm; coloration velvety dark brown, homogeneous, without special ornamentation and abundant coppery pilosity; length/width ratio of hind basitarsus about 6.3; gonocoxite and gonostylus as in figure 24 M; median area of median sclerite with strong constriction. Pupa: Cocoon with irregular weave, leaving thorax free dorsally; body length 3.8-4.2 mm; gill length 2.3-2.8 mm; frontoclypeus and thorax dorsally without tubercles; gill arborescent with about 14 branches, with primary branches shorter than in Lutzsimulium hirticosta (Fig. 24 N); frontal trichomes in 2 pairs on male; abdominal trichomes abundant, curved distally. Larva: Body length 7.5-8.0 mm; general color grayish brown; head yellowish brown; cephalic apotome with positive spots not well pigmented; postgenal bridge as long as hypostomium height and postgenal cleft smaller than either one; cervical sclerites elongated and reaching occiput; cephalic fan with 35-41 rays; Ratio of proximal, medial and distal antennal segments = 1:1.1-1.6:2.1-2.6; mandible with third preapical tooth larger than first; thoracic proleg sclerite with about 45 teeth; anal ring with 96-102 rows with 20-25 hooks each; larval cuticle without trichomes. Distribution: Argentina: Corrientes; Brazil: Rio de Janeiro, Rio Grande do Sul, São Paulo. Bionomics: Lutzsimulium pernigrum is found on vegetation in small creeks with clear water. Lutzsimulium flavopubescens (LUTZ) Simulim flavopubescens LUTZ, 1910: 248-249, 263, 265; PINTO, 1931: 702. Eusimulium flavopubescens LANE & PORTO, 1939: 169; LANE & VULCANO, 1943: 431, 434-436. Lutzsimulim flavopubescens PY-DANIEL, 1982: 308; COSCARÓN 1991: 82.

Female (based on lectotype): Frontal sulcus very evident and frons wider than in Lutzsimulium hirticosta; fronto-ocular triangle about as long as wide; claws, calcipala and genitalia similar to those of Lutzsimulium hirticosta; sensory organ of palp about half of basal segment length. Male: Unknown. Pupa: Cocoon with soft weave; length dorsally 2.9 mm, ventrally 3.6 mm; gill length 1.8-2.6 mm; gill arborescent with 12 branches arising from short primary branches and these from very short trunk; head, thorax and abdomen with tubercles; head with 2+2 frontal and 1+1 facial single trichomes; thorax dorsally with 5+5 trichomes, 3 of them curved apically; trichomes on distal portion of abdomen from gradually curved to strongly curved at apex. Larva: Ratio of proximal, medial and distal antennal segments = 1:1.5-1.6:2.3-2.4; postgenal cleft not deep, postgenal bridge two times longer than hypostomium length; cervical sclerites wide, reaching postocciput; anal ring with 108-110 rows with 18-22 hooks each. Distribution: Brazil: Rio de Janeiro. Bionomics: Unknown. Discussion: There are few specimens of Lutzsimulium flavopubescens; most of them are mounted on slides, and it is difficult to see some characters of taxonomic value. The bionomic information is dubious and very poor. The best information comes from pupae. Available information, based especially on pupal and larval morphology, suggests that Lutzsimulium flavopubescens is very similar to L. pernigrum.

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Kempfsimulium PY-DANIEL & NUNES DE MELLO (Figs. 26, 27, 136.) Kempfsimulium PY-DANIEL, 1982: 294. Type-species: Simulium simplicicolor LUTZ, 1910: 251.

Adults: Antennae each with 11 segments; frons of female as long as wide with strong median sulcus bifurcated basally (Fig. 26 A); mandible with only internal teeth; apical segment of maxillary palp about as long as subapical segment; mesepisternal sulcus well developed; katepisternum wider than high; furcasternum with small lateral projections (Fig. 26 L); wings (Fig. 26 F) with Sc without hair-like setae; R1 with hair and setae, Cu2 well curved distally; calcipala well developed and rounded apically; pedisulcus from absent to faint in some males (Fig. 26 J); female claw with small subtriangular subbasal tooth (Fig. 26 K); female eighth sternite with sclerotized area extending transversely from one side to other; gonapophyses subtriangular with rounded apical border and internal margin more sclerotized (Fig. 26 M); paraproct relatively short with small distal curvature (Fig. 26 N); genital fork with median branch longer than lateral branches and well sclerotized (Fig. 26 O); spermatheca oval, with sclerotized area reduced to neck of spermathecal duct and without spiculae internally (Fig. 26 P-Q). Male: Gonostylus about 1/3 length of gonocoxite, subconical and acuminate with two apical teeth distally (Figs. 27 A-C); ventral plate subtrapezoidal, elongated and narrowed basally (Figs. 27 A, D); endoparameres with basal plates and denticles inserted on membranous area that cover also aedeagus and median sclerite (Fig. 27 A); median sclerite subovoidal. Pupa: Cocoon with loose weave and evident threads; leaving gill and portion of thorax free; gill with filamentous branches emerging close to base (Fig. 27 F); body well sclerotized; frontoclypeus and thorax dorsally without tubercles; cephalothorax trichomes single with 1+1 frontal and 2+2 facials small (Fig. 27 E); and 5-6 trichomes mostly curved distally on each side of thorax disc (Fig. 27 F); abdominal chaetotaxy very similar to that of Lutzsimulium, but showing two transverse carinae on tergites. Larva: Cephalic apotome wider before posterior border, with median and lateral punctiform spots scarcely evident (Fig. 27 H); cervical sclerites small and free from postocciput; postgenal cleft deep with postgenal bridge shorter than hypostomium height; antenna shorter than stem of cephalic fan length, with wide basal segments (Fig. 27 I); combs of cephalic fan with similar setae of similar length; mandible with 3 preapical teeth, median tooth reduced, 11 or 12 internal, and 2 marginal teeth (Fig. 27 K); hypostomium teeth not arranged in groups (Fig. 27 J); lateral sclerite of thoracic proleg subtriangular; larval cuticle without setae but with short cylindrical projections on cuticle (Fig. 27 L); anal sclerite X-shaped, with abundant perianal scales; anal gill with 3 single lobes; ninth abdominal segment without ventral tubercles. CROSSKEY & HOWARD 1997, 2004 maintain Kempfsimulium in Lutzsimulium as a subgenus. Differences between Lutzsimulium and Kempfsimulium females: –

General coloration brown-blackish, with coppery brown to blackish pilosity; Sc with about 50 trichomes (Fig. 24 H); furcasternum branches with large projections (Fig. 24 L); frons longer than wide with median sulcus relatively short and without

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basal bifurcation (Fig. 24 A); spermatheca internally with spiculae; mesh-like sclerotized area of spermathecal ducts also covering spermatheca neck and part of spermathecal duct (Fig. 24 G); sclerotization of eighth sternite reduced to medial area (Fig. 24 E). ................................................................................... Lutzsimulim General coloration brown-yellow greenish to reddish brown with light yellowish brown pilosity; Sc without trichomes (Fig. 26 F); furcasternum branches with small projections (Fig. 26 L); frons about as long as wide, with median sulcus long with elongated basal bifurcate branches (Fig. 26 A); spermatheca internally without spiculae; sclerotized area of spermathecal duct reduced to neck, without mesh-like sclerotization at spermatheca union (Fig. 26 P); sclerotized area of eighth sternite covering area from one side to other (Fig. 26 M). .... Kempfsimulium

Differences in the male, pupa and larva are given in the generic keys Diferencias entre las hembras de Lutzsimulium y Kempfsimulium –

–

Coloración castaño negruzco, con pilosidad cobre-castaño; Sc con aproximadamente 50 tubérculos (Fig. 24 H); ramas del furcaesterno con grandes proyecciones (Fig. 24 L); frente mas largo que ancho, con sulco mediano relativamente corto y sin bifurcación basal (Fig. 24 A); espermateca con tubérculos internos; área esclerotizada de los ductos de la spermatheca también cubriendo el cuello de la espermateca y parte de los ductos, en aspecto de malla (Fig. 24 G); área esclerotizada del octavo esternito reducido a las áreas medianas. (Fig. 24 E). ....... .............................................................................................................................. Lutzsimulim Coloración amarillo castaño verdoso a castaño rojizo con pilosidad castaño amarillento; Sc sin tubérculos (Fig. 26 F); ramas del furcasterno con pequeñas proyecciones (Fig. 26 L); frente tan largo como ancho, sulco mediano largo con alongadas ramas bifurcadas basales (Fig. 26 A); espermateca sin tubérculos internos; área esclerotizada de la espermateca reducido al cuello, sin aspecto de malla en la unión de la espermateca (Fig. 26 P); área esclerotizada del octavo esternito cubriendo toda el área transversalmente de un lado al otro (Fig. 26 M). ....................................... Kempfsimulium

Las diferencias en el macho, larva y pupa están el las claves a género. Kempfsimulium simplicicolor (LUTZ) (Figs. 26 A-Q; 27 A-L.) Simulium simplicicolor LUTZ, 1910: 251; 1922: 40; KNAB, 1911: 179; NEIVA & PENNA, 1916: 93; LUTZ, 1922: 90; PINTO, 1931: 712; FAIRCHILD, 1940: 706; VARGAS, 1945a: 197; VULCANO, 1967: 20; LACEY & CHARLWOOD, 1980: 495, 497, 500-506; LACEY, 1981: 330. Simulium hebeticolor LUTZ, 1910: 267; KNAB, 1915b: 179; SMART, 1945: 513; VULCANO, 1967: 20. Eusimulium simplicicolor SURCOUF & GONZÁLEZ RINCONES, 1911: 286. Lutzsimulium simplicicolor PY-DANIEL, 1981: 173; CROSSKEY & HOWARD, 1997: 16; 2007: 44. Kempfsimulium simplicicolor PY-DANIEL, 1982: 300-308. Lutzsimulium (Kempfsimulium) simplicicolor CROSSKEY & HOWARD, 1997: 16; 2004: 7.

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F B

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Figure 26. Kempfsimulium simplicicolor. A-Q: Female; A: frons; B: fronto-ocular triangle; C: third palpomere with sensory vesicle; D: apex of mandible; E: basal portion of cibarium; F: wing, general view; G: anterior portion of wings showing setae, hair and spiculae (C = costa, R1= radius 1, Rs= secondary radius (radius 2) ; H-J: fore, mid and hind leg, respectively; K: claw; L: furcasternum; M: eighth sternite and gonapophyses; N: cercus and paraproct; O: genital fork; P: spermatheca; Q: portion of spermatheca with insertion base of spermathecal duct.

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B A

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J L Figure 27. Kempfsimulium simplicicolor. A-D: Male; A: genitalia, ventral view showing gonocoxite, gonostylus, ventral plate, endoparameres base (end.) and aedeagal membrane (ae.); B: gonocoxite and gonostylus, dorsal view; C: gonostylus, dorsal view; D: ventral plate (v.pl.) and median sclerite (m.s.). E-G: Pupa; E: frontoclypeus; F: portion of thorax with gill, and trichomes; G: thoracic trichomes magnified. H-L: Larva; H: head, dorsal view; I: antenna; J: hypostomium, anterior border; K: apex of mandible with teeth; L: short cylindrical projections of body integument.

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Female: Wing length 2.3-2.7 mm; general color brownish; head brown, frons and clypeus brown with grayish pollinosity; scutum from brown-yellow greenish to reddish brown with decumbent hair orange to yellowish brown; scutellum and metanotum brown reddish; pleurae grayish brown; wing veins light brown; legs light grayish brown, darkened on tarsi; abdomen grayish brown with light brown hair; frons with deep frontal suture (Fig. 26 A); fronto-ocular triangle about as high as wide (Fig. 26 B); sensory organ of maxillary palp about 1/3 of basal segment (Fig. 26 C); cibarium smooth (Fig. 26 E); wing chaetotaxy with thin hair brown and robust spine-like setae dark brown on C and R1 (Fig. 26 G); leg shape and coloration as in figure 26 H-J; claw subbasal tooth well developed (Fig. 26 K); gonapophysis with abundant long hair and eighth sternite with about 21 setae on each side (Fig. 26 M); genital fork with lateral branches relatively wide (Fig. 26 O); discontinuous sclerotization of spermathecal neck reduced only to spermathecal duct (Fig. 26 P-Q). Male: Wing length 1.9-2.0 mm. general coloration as in female; gonocoxite and gonostylus laterally flattened; gonostylus acuminate with 2 strong terminal teeth (Figs. 27 A-C); ventral plate basally with strong concavity and well-sclerotized margin (Figs. 27 A, D); median sclerite and membrane of aedeagus with abundant microtrichia (Fig. 27 A). Pupa: Cocoon with threads evident; gill length 0.9-1.3 mm; frontoclypeus as in figure 27 E; gill light brown with 12 branches, arborescent, becoming thinner from base to apex (Fig. 27 F); thoracic trichomes variable from straight to well curved (Fig. 27 G); abdomen without tubercles but with robust spines and hooks. Larva: Maximum length 5.5-7.0 mm; color grayish brown-green; cephalic apotome yellowish, scarcely ornamented (Fig. 27 H), without hair; ratio of proximal, medial and distal antennal segments = 1:2.0-2.5:3.0-3.6; distal segment more darkened (Fig. 27 I); hypostomium with median teeth similar in height to intermediate and lateral teeth (Fig. 27 J); cephalic fan with 35-43 rays; anal ring with 110-130 rows with about 22 hooks in each. Distribution: Brazil: Mato Grosso, Pará, Rondônia (Fig. 136). Bionomics: The species breeds on Podostemaceae in rivers with strong current. Females are haematophagous and were collected on humans, cows, horses and monkeys. Simulium LATREILLE Simulium LATREILLE, 1802: 428. Type-species: Oestrus columbacensis SCOPOLI, 1780: 133 (monotype).

Adults: Color in general yellowish to black, scutum ornamentation homogeneous to variable with filiform to flattened hair uniformly distributed or grouped and simulating scales; wing without basal cell; basal sector of R with or without hair, shorter than distance between Rs base and wing apex; radial sector unbranched; C and distal 2/3 of R1 with spiniform setae; hind basitarsus generally with calcipala; tarsomere I with deep pedisulcus; legs with hair filiform or with petaloid shapes intermixed; female claws with or without subbasal tooth; cibarium basally from light and smooth to well sclerotized and with teeth of variable disposition; gonapophysis, paraproct, gonocoxite, gonostylus and ventral plate of variable size and shape; spermatheca without sclerotized union with spermathecal duct; endoparameres generally with abundant teeth.

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Pupa: Cocoon well developed, generally compact, with slipper to shoe shape; frontoclypeus and thorax with trichomes single to multibranched, sometimes in elevated number (nearly one hundred); abdominal segments with weak sclerotization; distal tergites without special setae; terminal spines scarcely developed. Larva: Cephalic apotome with variable ornamentation; cervical sclerites isolated; mandible generally with 1 or 2 marginal teeth; distal antennal segment no longer than two basal segments together; postgenal cleft varied from very deep to not evident; hypostomial teeth very similar and median tooth about same level as lateral teeth; rectal papillae single to having more than 10 diverticula on each lobe. Distribution: This genus is distributed all over the world, and has the greatest number of species, with great morphological variation (COSCARÓN, 1987; CROSSKEY & HOWARD, 1997, 2004). For this reason, it is subdivided into a large number of subgenera; some authors elevate these subgenera to the generic level (PY-DANIEL & MOREIRA SAMPAIO, 1994, 1995). KEYS

TO

SIMULIUM

SUBGENERA

Females 1 – 2(1) – 3(2) – 4(3)

– 5(4) – 6(5) –

General coloration black or blackish brown ................................................................ 2 General coloration brown (reddish brown to yellowish) ......................................... 22 Body without special ornamentation; scutum hairs homogeneously distributed; claws with basal tooth large, subovoid, elongated; basal portion of cibarium smooth, without reinforcement at sides ............................... Eusimulium, Nevermannia Body ornamented with light vittae or spots; scutal hairs distributed homogeneously or in groups; claws with or without subbasal tooth, but if present, it is proportionally smaller and subtriangular (except Byssodon with basal tooth) ............ 3 With hair on basal sector of R ........................................................................................ 4 Without hair on basal sector of R .................................................................................. 9 Grayish pollinose coloration; scutum with light area limited to 1 median and 1+1 sublateral stripes (Fig. 29 A); abdomen blackish or grayish with 1+1 silver transverse bands (Fig. 28 A), or with bands on posterior border of tergites III-IV (Fig. 51 A) ......................................................................................... Pternaspatha (in part) Blackish coloration; scutum with or without light submedian stripes or anterior spots; abdomen blackish without silvery transverse bands on tergites III-IV ......... 5 Paraproct subtriangular, about twice as long as wide at base (Figs. 57 D, N); basal portion of cibarium smooth ........................................................................... Chirostilbia Paraproct not subtriangular, curved distally, slightly longer than wide at base (Figs. 64 G, L, P); cibarium smooth or with teeth ................................................................. 6 Basal portion of cibarium with teeth arranged in elevated median subconical group (Fig. 64 M) ..............................................................................................Inaequalium Cibarium generally smooth or with small teeth: if teeth present, these arranged in median elevated group, scutum with 1+1 light anterior spots or 3 black longitudinal stripes separated by nacreous stripes that change tone with different light positions ...................................................................................................................... 7

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7(6) – 8(7) – 9(3) – 10(9) – 11(10) – 12(11)

–

13(11) – 14(13)

–

Cibarium with very small teeth or without teeth (Figs. 102 C, F, G, L); paraprocts relatively short not lobulate, scarcely acuminate apically and directed downwards (Figs. 103 B, D, G, J) ............................................................................. Aspathia (in part) Cibarium with 1 row of teeth (Figs. 128 D, H) or without teeth but with reinforced border (Figs. 132 B, L); paraproct relatively long, lobulate and scarcely sclerotized, not acuminate (Figs. 128 C, E, J; 132 D, G, N) ..................................... 8 Scutum with hair grouped, simulating scales (Fig. 132 I), without anterior grayish subquadrate spots; cibarium smooth (Figs. 132 B, L) .............. Thyrsopelma (in part) Scutum with hair homogeneously distributed, with anterior grayish subquadrate light spots (Fig. 128 A); cibarium with teeth arranged in 1 row (Figs. 128 D, H) ............................................................................................................. Trichodagmia (in part) Cibarium without teeth ................................................................................................... 10 Cibarium with teeth ......................................................................................................... 16 Scutum with hair grouped like scales (Fig. 5 A); claw without subbasal tooth or much reduced (Figs. 54 D, E); fronto-ocular triangle absent; paraprocts very thin distally (Figs. 54 I, Q, X) ................................................................................... Notolepria Scutum with hair homogeneously distributed; claw with or without subbasal tooth, fronto-ocular triangle well developed; paraprocts not thinner distally .................... 11 Species grayish pollinose; abdomen grayish with tergites III-V blackish or velvety brown, generally with anteromedian or 1+1 silvery lateral spots; calcipala frequently reduced or absent ......................................................................................... 12 Species blackish; abdomen blackish without silver spots on tergites III-V; calcipala well developed .................................................................................................................. 13 Claws with subbasal tooth (Fig. 51 F); scutum with blackish areas delimited by grayish median and sublateral stripes; abdominal tergites III-V blackish with 1+1 sublateral posterior whitish spots (Fig. 51 A); paraprocts subrectangular, shorter than wide (Fig. 51 H) ..................................................................................... Pternaspatha Claws without subbasal tooth (Fig. 99 C); scutum grayish uniform except 1+1 light median anterior spots; tergites III-V blackish or velvety brown, with or without spots anteromedially (Figs. 98 A, P; 99 A); paraprocts subtriangular (Fig. 99 G), as long as wide .......................................... Psilopelmia (BLANCASI sp. group) Paraproct subtriangular, with distal border arched (Figs. 118 A, B); scutum with 1+1 anterior light submedian spots continued posteriorly by thin stripe joining with grayish posterior area ..................................... Hemicnetha (OVIEDOI sp. group) Paraproct not subtriangular, with distal border truncate or acuminate posteriorly; scutum ornamentation variable ..................................................................................... 14 Paraprocts with distal edge truncate and slightly acute posteriorly (Figs. 103 B, K); scutum with median and 1+1 submedian black stripes delimited by nacreous vittae that invert coloration with change of illumination position (Figs. 101 A, B, F, G) ......................................................................................................................... Aspathia Paraprocts with distal edge truncate or rounded but not acuminate posteriorly; scutum without contrasting silvery vittae (Figs. 113 A, D, G, H; 120 A, B, K, L) .............................................................................................................................................. 15

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15(14) Paraproct as long as wide at base, with external surface mostly nude and shiny, with depressions and distal border truncated (Fig. 120 I); gonapophysis as long as wide .............................................................................................................................Hearlea – Paraproct longer than wide at base, external surface mostly covered by microtrichia, without depressions and distal border curved (Figs. 113 C, F, L, M, R, U), gonapophysis longer than wide (Figs. 113 C, K, Q, T, V) .............................. ..................................................... Hemicnetha (in part BRACHYCLADUM sp. group) 16(9) Scutum with abundant gray pollinosity, showing, with some illumination, 3-5 longitudinal light stripes; cibarium with weak concavity covered by small similar acute teeth; paraprocts subtriangular, about as long as wide; claws without teeth ...................................................................................................................................... Psilozia – Scutum with sparse pollinosity, with or without 1+1 silvery longitudinal stripes; cibarium with or without teeth, generally with median depression; paraprocts not subtriangular, with curved distal edge, generally longer than wide and sometimes thinner distally; claws with or without teeth .............................................................. 17 17(16) Claws with large subovoidal basal tooth; posterior edge of tergites III-V frequently with 1+1 transverse silvery bands .................................................. Byssodon – Claws with small subtriangular subbasal tooth or without tooth; tergites III-V without silvery transverse bands ................................................................................... 18 18(17) Cibarium generally without teeth, but if teeth present, not in median depression and at sides (Figs. 102 F, G, K, O); scutum with 1+1 submedian longitudinal silvery stripes generally reaching posterior grayish area (Figs. 101 A, F) or darker with change of light direction (Figs. 101 B, G); fronto-ocular triangle well developed ................................................................................................................ Aspathia – Cibarium with median depression sclerotized or not and with small teeth at sides (Figs. 71 P; 76 N; 80 D; 84 M; 91 Q); scutum with or without silvery longitudinal stripes that can reach to posterior silvery area; fronto-ocular triangle generally absent or reduced ............................................................................................................. 19 19(18) Cibarium with central area well sclerotized and with small rounded teeth (Figs. 80 D, P, T); infra-frontal sutures well developed (Figs. 80 A, V) .................................... ....................................................................Coscaroniellum (QUADRIFIDUM sp. group) – Cibarium with central area not sclerotized, teeth acute (Figs. 71 P; 76 N, X); infrafrontal sutures not developed ........................................................................................ 20 20(19) Scutum black, with or without silvery submedian stripe mostly frequent on anterior area and wedge shaped (Figs. 71 L, O, Q, U); when reaching posterior silvery area, without gray shadow spots on silvery stripe ................... Psaroniocompsa – Scutum grayish to black with silvery stripe reaching posterior grayish area, generally with gray shadow spots on silvery stripes (Figs. 76 A-F, H, I, L, M, Q, R) .............................................................................................................................................. 21 21(20) Cibarium with small teeth about same size and absent on median depression (Figs. 76 N, X); fronto-ocular triangle very short (Figs. 76 G, W); abdomen without silvery posterior transverse spots on tergites III-V or very short (Fig. 76 K) ........ ............................................................................................................................ Cerqueirellum

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–

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Cibarium with submedian teeth larger than other teeth and with small teeth on median area (Fig. 80 Y); fronto-ocular triangle relatively deep; abdomen with silvery posterior transverse spots on tergites III-V about 1/3 of tergite height (Fig. 80 X); scutum with submedian silvery vittae narrow (Fig. 80 W) .................... .......................................................... Coscaroniellum (QUADRIVITTATUM sp. group) Species yellowish-reddish to greenish gray; if scutum black, margins are yellowish (Figs. 84 B-D); paraprocts subtriangular (Figs. 85 D-P) longer than wide at base and distally relatively thin (Figs. 92 A-N) ................................................................... 23 Species reddish brown to dark grayish; scutum generally with lyre-shaped pattern (Fig. 128 A); paraprocts subrectangular and distally not acuminate (Figs. 128 C, E, J) .............................................................................................................................................. 28 Scutum gray-greenish to brown-yellowish; cibarium smooth not sclerotized (Fig. 60 D) .............................................................. Chirostilbia (SUBPALLIDUM sp. group) Scutum orange to reddish or black bordered by yellow and frequently with submedian silvery stripes; cibarium with sclerotized margin or with small teeth .. .............................................................................................................................................. 24 Scutum with 1+1 or more nacreous or silvery longitudinal stripes (Figs. 91 B-E); abdomen generally with 3 longitudinal rows of subquadrate blackish spots (Figs. 91 G-I); cibarium with median depression well sclerotized, bordered with weak lobules and at sides with teeth on weakly elevated areas (Figs. 91 J-Q); paraprocts thinner distally (Figs. 92 A-N) ............................ Psilopelmia (ESCOMELI sp. group) Scutum generally without 1+1 or more nacreous or silvery longitudinal stripes (Figs. 84 A-E); abdomen without subquadrate blackish spots; cibarium without teeth or with teeth generally larger and with different disposition from above (Figs. 84 I-W); paraprocts not thin distally (Figs. 85 D-K) ........... Ectemnaspis......25 Scutum yellow or black, in the latter case, totally with yellow margins (Figs. 84 AE); cibarium without or with teeth, in the latter case, with plates or small lobules at median area (Figs. 84 I-P) .............................................................................................. 26 Scutum yellow or reddish, cibarium with teeth but median area without plates .............................................................................................................................................. 27 Cibarium without teeth but with well sclerotized edge (Figs. 84 I-J) ....................... ............................................................................................... BICOLORATUM sp. group Cibarium with teeth and median area with plates or lobules (Figs. 84 K-P) .......... .......................................................................................................... ROMANAI sp. group Cibarium with well-developed teeth grouped on 1+1 weak elevations (Figs. 84 Q-S) ................................................................................................... PERFLAVUM sp. group Cibarium with very small teeth and with 1+1 submedian processes generally with teeth on apex (Figs. 84 T-W) ...................................................... DINELLII sp. group Gonapophysis subovoidal shorter than wide (Fig. 132 M); paraproct slightly acuminate distally (Fig. 132 N); scutum with hairs grouped (Fig. 132 I) and arranged in longitudinal rows ....................................................... Trichodagmia (in part) Gonapophysis subtriangular or subovoidal, longer than wide or about as long as wide; paraprocts not acuminate distally; scutum with hairs homogenously distributed or in groups, but not arranged in longitudinal rows ........................... 29

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29(28) Cibarium scarcely sclerotized on median area and without teeth or spiculae (Fig. 113 B); paraprocts slightly longer than wide, subrectangular and with abundant microtrichia (Fig. 113 C, F) ...................................... Hemicnetha (PAYNEI sp. group) – Cibarium well sclerotized on median area, with thin teeth acuminate and scarcely sclerotized (Figs. 113 J; 128 D, H); paraprocts about 2 times longer than wide, and with sparse microtrichia (Figs. 113 L, M; 128 C, E, J) ............................................. 30 30(29) Gonapophysis more than 2 times longer than wide, acute apically (Fig. 113 K); cibarium with small teeth disordered (Fig. 113 J) ......................................................... ................................................................... Hemicnetha (BRACHYCLADUM sp. group) – Gonapophysis less than 2 times longer than wide, and blunt apically (Figs. 128 B, I); cibarium with conspicuous teeth arranged along border to base of cornuae (Figs. 128 D, H) ............................................................................................... Trichodagmia Males 1 – 2(1) – 3(2) – 4(3) –

5(3) – 6(5) – 7(5) – 8(7) –

General coloration black, sometimes abdomen with greenish tones; with or without hair on basal sector of R ................................................................................... 2 General coloration variable form dark brown to yellowish, generally with hair on basal sector of R .............................................................................................................. 23 With hair on basal sector of R ........................................................................................ 3 Without hair on basal sector of R ................................................................................ 11 Gonostylus longer than gonocoxite (Figs. 133 B, C, H, J); ventral plate relatively short and with or without median process and carina (Figs. 133 D, I, K) ............ 4 Gonostylus shorter than gonocoxite; ventral plate with variable shape ................ 5 Scutum with hairs homogeneously arranged and without silvery spots; ventral plate distally with small, median but not lateral processes (Fig. 129 E) ................... ............................................................................................................................. Trichodagmia Scutum with hairs disposed in groups like scales and with 1+1 silvery anterolateral spots sometimes continued submedially reaching to posterior grayish area (Figs. 133 A, G); ventral plate distally with large median and lateral processes (Figs. 133 D, I) ..................................................................................................................... Thyrsopelma Gonostylus distal spur much reduced or absent ................................ Chirostilbia......6 Gonostylus with distal spur ............................................................................................. 7 Gonostylus without lateral crest (Figs. 58 B, F, G, I, L, O) ........................................ ......................................................................................................... PERTINAX sp. group Gonostylus with lateral crest (Figs. 61 D, F, H) ............................................................ ................................................................................ SUBPALLIDUM sp. group (in part) Scutum without ornamentation; gonostylus flattened and curved, with subtriangular internal lobe ....................................................... Eusimulium, Nevermannia Scutum, with anterior lighting, generally ornamented with spots; gonostylus with variable shape ...................................................................................................................... 8 Gonostylus (microscope slide mounted) subquadrate with distal concavity and subapical spur (Figs. 28 N; 30 R; 41 I) ............................................... Pternaspahta......9 Gonostylus (microscope slide mounted) subtriangular to subtrapezoidal or subcylindrical and with apical spur .............................................................................. 10

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Gonostylus with tubercles (Figs. 28 N; 30 R; 31 S) ..................................................... ......................................................................................... NIGRISTRIGATUM sp. group Gonostylus without tubercles (Figs. 41 I; 47 K; 52 K) ... NEMORALE sp. group Gonostylus subtriangular or subtrapezoidal, with 1 apical spur (Figs. 65 D, F-H, J-N, P); ventral plate without lateral constriction (Figs. 65 C, E, I, O) ............. Inaequalium Gonostylus subcylindrical with weak distal curvature and 1 apical spur (Figs. 86 H-N) (except wolffhuegeli, in which it is relatively short and has several apical spurs); ventral plate with weak lateral constriction (Figs. 87 A-H) ......................................... ............................. Ectemnaspis (BICOLORATUM sp. group; ROMANAI sp. group) Gonostylus much shorter than gonocoxite ................................................................ 12 Gonostylus as long as or longer than gonocoxite ..................................................... 19 Gonostylus without apical spur .................................................................................... 13 Gonostylus with apical spur .......................................................................................... 14 Scutum without ornamentation; gonostylus subovoidal with reinforced edge (Figs. 54 N, S, T) ................................................................................................. Notolepria Scutum with 1+1 submedian anterior silvery spots (Figs. 61 A, B); gonostylus subtrapezoidal with sublateral crest (Figs. 61 D, F, H, I) ............................................ ......................................................................... Chirostilbia (SUBPALLIDUM sp. group) Scutum with 1+1 submedian silvery vittae, reaching or not to grayish posterior area; gonostylus subtriangular or subtrapezoidal ...................................................... 15 Scutum without 1+1 submedian silvery vittae, but with anterior subquadrate or subtriangular silvery spots (Figs. 86 A-D; 91 A-D); gonostylus subrectanglular or subcylindrical ..................................................................................................................... 17 Gonostylus about 2 times longer than wide at base; and apical spur spatuliform (Figs. 81 C, G, H) ......................................................................................... Coscaroniellum Gonostylus only slightly longer than wide at base; terminal spur subconical .... 16 Gonostylus subtriangular with apical or subapical spur (Figs. 77 J, O; 81 L) ......... ................ Byssodon, Cerqueirellum, Coscaroniellum (QUADRIVITTATUM sp. group), Psaraniocompsa (SIOLLII sp. group) Gonostylus subquadrate or subtrapezoidal with submedian spur (Figs. 72 F, H, I) .................................................................. Psaraniocompsa (INCRUSTATUM sp. group) Gonostylus subcylindrical, generally with weak dorsal curvature and 1 apical spur (Figs. 86 H-N; 93 K; 118 F, H) ............... Ectemnaspis (BICOLORATUM sp. group; ROMANAI sp. group); Psilopelmia and Hemicnetha (OVIEDOI sp. group) Gonostylus subtrapezoidal with 2-15 marginal spurs .............................................. 18 Ventral plate more than 2 times wider than long (Fig. 87 I); gonostylus with 7-15 marginal spurs (Fig. 86 O) ................................. Ectemnaspis ( ROMANAI sp. group) Ventral plate as wide as long; gonostylus with 2-4 subapical spurs .............. Psilozia Gonostylus subcylindrical and thin basally, generally narrower than half of gonocoxite, distal edge wide and generally with basal process .............................. 20 Gonostylus basally wider than half of gonocoxite, distal edge wide and without basal process ...................................................................................................................... 21 Ventral plate generally becoming narrower apicomedially (Figs. 106 M-P); gonostylus with basal process present (Figs. 105 B, C, E-H, J-L) .............. Aspathia

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Ventral plate not becoming narrower apicomedially (Fig. 121 D, G ); gonostylus without basal process but with small subbasal carina (Figs. 121 C, F) ......... Hearlea Gonostylus flattened and sinuous, scarcely narrowed distally (Figs. 114 A, E, G, I, M, N); endoparameres with large teeth and base well sclerotized (Fig. 114 K) ..... ..................................................................................................................... Hemicnetha......22 Gonostylus subtriangular, narrowed and well curved apically (Figs. 133 B, C, H, J); endoparameres without teeth and base not well sclerotized (Fig. 133 F) ................ .............................................................................................................................. Thyrsopelma Ventral plate wider than long, with large median process (Figs. 114 B, C, F, H, L) .................................................... PAYNEI sp. group, BRACHYCLADUM sp. group Ventral plate about as long as wide, without median process but with median carina (Figs. 114 J, O) ........................................................... MEXICANUM sp. group Gonostylus almost 2 times longer than gonocoxite ................................................. 28 Gonostylus shorter or slightly longer than gonocoxite ........................................... 24 Gonostylus with 3-6 spurs or more (Fig. 129 L); endoparameres without teeth (Fig. 129 C) ....................................................................................... Trichodagmia (in part) Gonostylus with 1 or no spurs; endoparameres generally with large teeth (Figs. 61 I; 99 I) ................................................................................................................................. 25 Basal sector of R without hair; scutum dull grayish; gonostylus with internal border straight (Figs. 98 V; 99 H) ...................... Psilopelmia (BLANCASI sp. group) Basal sector of R with hair (Fig. 60 C); scutum not grayish; gonostylus with internal border curved .................................................................................................... 26 Gonostylus without apical spur, with sublateral crest and distally not curved (Figs. 61 D, F, H, I) ................................................ Chirostilbia (SUBPALLIDUM sp. group) Gonostylus with apical spur, without sublateral crest and distally slightly curved .............................................................................................................................................. 27 Ventral plate generally longer than wide at base (Figs. 87 J-L); gonostylus without distal curvature (Figs. 86 P, Q, R) ............... Ectemnaspis (PERFLAVUM sp. group) Ventral plate generally shorter than wide at base (Figs. 87 M-O; 94 A-L); gonostylus generally with distal curvature (Figs. 86 S; 93 H-R) ................................ ............................................................... Psilopelmia, Ectemnaspis (DINELLII sp. group) Scutum dark brown with hairs grouped like scales; gonostylus distally relatively thick (Fig. 133 J) ............................................................................................... Thyrsopelma Scutum brown-reddish with hairs not grouped like scales; gonostylus distally relatively thin (Figs. 129 B, D) ...................................................................... Trichodagmia

Pupae 1 – 2(1) –

Gill arborescent with relatively thick branches, with acute apices well sclerotized and spine shaped, with 12-50 branches (Figs. 134 B, E, G, K), cocoon compact and with ventral side projected anteriorly .................................. Thyrsopelma (in part) Gills with variable shape but without acute, well-sclerotized, spine-shaped apices; cocoon of variable shape ................................................................................................. 2 Gills thick, asymmetrical, generally with curvature .................................................... 3 Gills generally filiform, symmetrical, without lateral curvature ............................... 6

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Frontoclypeus and exposed portion of thorax with single trichomes (Fig. 121 H); gills with striations (Figs. 122 A-K; 123 A-O; 124 A-K); cocoon without dorsal projection ...................................................................................................................Hearlea – Frontoclypeus and exposed portion of thorax with multibranched trichomes with 2-8 branches; gill without striations; cocoon generally with dorsal projection ..... 4 4(3) Cocoon generally without dorsal projection, anterior edge straight (Figs. 66 A, B, D, L; 70 A, B, P, Q) ...........................................................................................Inaequalium – Cocoon projected dorsally (Figs. 73 H-K; 88 C, D, F; 107 I) .................................. 5 5(4) Frontoclypeus and thorax with trichomes bifid (Figs. 73 O-Q) ....... Psaroniocompsa – Frontoclypeus and thorax with trichomes of 3-8 branches ....................................... ........................................................................................................... .Ectmemnaspis, Aspatha 6(2) Cocoon with compact weave and anterior portion elevated, generally protecting gill base (Figs. 115 A-D; 59 L, O, S); gill filaments relatively thick ......................... 7 – Cocoon without compact weave and generally with anterior portion not elevated or protecting gill; gill filaments relatively thin ........................................................... 13 7(6) Cocoon anterior aperture with festoons protecting gill base (Figs. 115 A-D) ..... 8 – Cocoon anterior aperture without festoons protecting gill base ............................. 9 8(7) Gill of 8 branches with apices blunt and resistant (Figs. 116 A, B); frontoclypeus generally reinforced at base and enlarged at facial area, with female and male having similar facial area (Figs. 115 O-T) ............. Hemicnetha (PAYNEI sp. group) – Gill of10 branches frequently with membranous apices blunt, not resistant (Fig. 59 N); frontoclypeus not reinforced at base and not enlarged at facial area, with male frons basally narrower than in female ................................ Chirostilbia (in part) 9(7) Gill with 6 filaments ........................................................................................................ 10 – Gill with more than 6 filaments .................................................................................... 11 10(9) Frontoclypeus and thorax with numerous tubercles, some of them acute like short spines (Figs. 119 B, D, E); cocoon scarcely elevated anteriorly (Fig. 119 A) .......... ...................................................................................... Hemicnetha (OVIEDOI sp. group) – Frontoclypeus and thorax without tubercles (Fig. 115 R); cocoon frequently very elevated anteriorly (Fig. 115 E) .......... Hemicnetha (BRACHYCLADUM sp. group) 11(9) Gill with 18-20 branches arranged in bunch (Fig. 130 L) ...... Trichodagmia (in part) – Gill with 8-16 branches ................................................................................................... 12 12(11) Gill with 12 branches .......................................................................................................... .................................... Hemicnetha (MEXICANUM sp. group), Trichodagmia (in part) – Gill with 10 branches ............. Hemicnetha (BRACHYCLADUM sp. group in part) 13(6) Gill with 4 filamentous branches .................................................................................. 14 – Gill with more than 4 filamentous branches .............................................................. 17 14(13) Frontoclypeus and thorax with tubercles acuminate (Fig. 82 D) .............................. .......................................................................................................... Coscaroniellum (in part) – Frontoclypeus and thorax with tubercles not acuminate ........................................ 15 15(14) Cocoon reduced ventrally (Figs. 98 X; 99 K); cephalic and thoracic trichomes single (Fig. 98 a-b) ................................................. Psilopelmia (BLANCASI sp. group) – Cocoon not reduced ventrally; cephalic and thoracic trichomes with 1-8 branches .............................................................................................................................................. 16

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16(15) Cephalic and thoracic trichomes generally bifid (Figs. 73 O, P, Q) .......................... ................................................................................................. Psaroniocompsa, Cerqueirellum – Cephalic and thoracic trichomes with 1-8 branches ..................................................... .................................................................. Eusimulium, Byssodon, Nevermannia, Psilopelmia 17(13) Gill with 6 filamentous branches .................................................................................. 18 – Gill with more than 6 filamentous branches .............................................................. 20 18(17) Thoracic trichomes numbering more than 10 on each side and frequently very modified (Figs. 40 U-V; 44 R; 53 S, T; 108 G, H; 109 A, G) ..................................... ............................................................................................ Pternaspatha, Aspathia (in part) – Thoracic trichomes numbering about 5 on each side .............................................. 19 19(18) Cephalic and thoracic trichomes bifid (rarely single or trifid) ................................... ................................................................ Psaroniocompsa, Cerqueirellum, Aspathia (in part) – Cephalic and thoracic trichomes generally with more than 3 branches, not bifid ..................................................................... Byssodon, Inaequalium, Notolepria, Ectemnaspis (BICOLORATUM and ROMANAI sp. group), Psilopelmia 20(17) Gill with 8-10 branches ................................................................................................... 21 – Gill with more than 10 branches ...................................................................................... ...................... Psilozia, Chirostilbia, Aspathia, Ectemnaspis (PERFLAVUM sp. group) 21(20) Thoracic trichomes numbering more than 10 and frequently much modified (Figs. 29 N, P; 31 Y, Z); gill with 8 branches ....................................................... Pternaspatha – Thoracic trichomes numbering 5 on each side; gill with 8-10 branches .............. 22 22(21) Cocoon weave spongy with elevated anterior margin and dorsal carina (Figs. 88 B, H, I, J) ................. Ectemnaspis (BICOLORATUM and ROMANAI sp. group in part) – Cocoon weave compact, without elevated anterior margin or dorsal carina (Figs. 62 C, D, G; 78 A, G, J) ............................. Chirostilbia (SUBPALLIDUM sp. group), Notolepria, Cerqueirellum, Coscaroniellum, Psilozia, Byssodon, Aspathia, Ectemnaspis (PERFLAVUM and DINELLII sp. groups), Psilopelmia Larvae 1 – 2(2)

–

3(2)

Anal ring with more than 150 rows of hooks ............................................................. 2 Anal ring with less than 120 rows of hooks ................................................................ 6 Body cuticle with lanceolate trichomes (Figs. 135 G, L, S); hypostomium with median tooth and intermediate teeth not well differentiated (Figs. 135 D, I, N); abdomen not dorsally flattened on distal portion (Fig. 135 A); cervical sclerites enlarged, each more than 2 times wider than long (Fig. 135 Q) ........... Thyrsopelma Body cuticle without lanceolate trichomes; hypostomium with median tooth and generally intermediate teeth well differentiated (Figs. 117 N, Q; 126 J; 131 G); abdomen dorsally flattened distally (Figs. 131 A; 127 G); cervical sclerites not enlarged, each less than 2 times wider than long (Fig. 131 I) .................................. 3 Anal sclerite frequently with sclerotized reinforcement, in some cases with ventral branches enlarged and completing a ring (Figs. 127 B-J); anterior margin of hypostomium generally arc shaped with lateral and intermediate teeth reduced (Figs. 126 A, B, I, H, J) ...........................................................................................Hearlea

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Anal sclerite without sclerotized reinforcement nor with ventral branches completing a ring; anterior margin of hypostomium with variable shape ............. 4 4(3) Cephalic apotome darkened on distal half in form of triangle (Fig. 63 A) ............ ............................................................................................................................... Chirostilbia – Cephalic apotome with different ornamentation ........................................................ 5 5(4) Hypostomium with lateral edges descending abruptly from lateral teeth, giving plateau-like appearance (Figs. 117 P, Q; 131 G); mandible with preapical tooth shorter than apical tooth (Figs. 117 L; 131 E, K) ......................................................... ........................................... Hemicnetha (BRACHYCLADUM sp. group), Trichodagmia – Hypostomium with lateral borders descending gradually from lateral teeth in curved manner (Figs. 117 N, O, R); mandible with preapical tooth as long as or longer than apical tooth (Figs. 117 K, M) ...................................................................... ...................................... Hemicnetha (PAYNEI sp. group; MEXICANUM sp. group) 6(1) Anal ring with 100-120 rows of hooks; cephalic fan rays with setate of similar size (Fig. 119 J); mandible with pair of large marginal teeth (Fig. 119 K) ...................... ...................................................................................... Hemicnetha (OVIEDOI sp. group) – Anal ring with less than 100 rows of hooks; cephalic fan rays alternating between groups of small setae, and some larger ones; mandible with only 1 marginal tooth, if there are 2 the second is very small (Figs. 29 S; 79 B; 83 C, O, R; 90 Q) ........ 7 7(6) Cephalic apotome without ornamentation or with small spot on median basal area; with small numerous and short hairs on cephalic capsule (Fig. 79 C); generally with dark band around first abdominal segment; antenna thin and surpassing apex of cephalic fan stem (Fig. 79 H) ....................................................... 8 – Cephalic apotome with evident ornamentation; without or with very scarce hair on cephalic capsule; antenna generally thick and surpassing, or not, length of cephalic fan stem ................................................................................................................ 9 8(7) Mandibles with row of inferior supramarginal setae (Fig. 79 B); medial antennal segment as long as or longer than proximal segment (Figs. 79 D, E, I) .................. ............................................................................................................................ Cerqueirellum – Mandible without row of inferior supramarginal setae; medial antennal segment about 1/2 to 4/5 of proximal segment (Figs. 83 B, N, Q) ................. Coscaroniellum 9(7) Postgenal cleft scarcely developed, with postgenal bridge longer than hypostomium length (Figs. 112 A, B, D, G); rectal papillae without diverticula on lobes .................................................................................................................................... 10 – Postgenal cleft well developed, with postgenal bridge as long as or shorter than hypostomium length; rectal papillae generally with diverticula on lobes ............. 13 10(9) Antennae with thin transverse striations and surpassing apex of cephalic fan stem (Figs. 111 A, B, H, L); medial antennal segment longer than proximal segment (Figs. 111 B, H, L, M) ........................................................................................... Aspathia – Antennae without striations and not surpassing apex of cephalic fan; medial antennal segment shorter than proximal segment .................................................... 11 11(10) Ninth sternite without 1+1 tubercles ............... Psilopelmia (BLANCASI sp. group) – Ninth sternite with 1+1 conspicuous tubercles ........................................................ 12 12(11) Antennae surpassing apex of cephalic fan stem ................ Eusimulium, Nevermannia

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Antennae about same length as apex of cephalic fan stem ............................Psilozia Rectal papillae without diverticula on lobes ....... Psilopelmia (BLANCASI sp. group) Rectal papillae with diverticula on lobes ..................................................................... 14 Cephalic apotome with isolated spots: anteromedian, posteromedian, anterolateral and posterolateral spots generally positive (Figs. 30 Y; 55 C; 56 K; 75 L) .............................................................................................................................................. 15 Cephalic apotome with variable ornamentation, different from above ............... 19 Anal sclerite without scales (Fig. 63 G) ....................................................................... 16 Anal sclerite with scales (Fig. 63 P) .............................................................................. 17 Antenna surpassing apex of cephalic fan stem (Figs. 75 D, L); medial antennal segment generally shorter than distal segment (Figs. 75 A, E, M) ........................... Psaroniocompsa Antenna not surpassing apex of cephalic fan stem; medial antennal segment generally longer than distal segment (Figs. 90 M, N) ................................................... ...................... Ectemnaspis (BICOLORATUM sp. group; PERFLAVUM sp. group) Body with maximum length less than 4.5 mm ............................................. Notolepria Body with maximum length over 5.0 mm .................................................................. 18 Medial antennal segment shorter than apical segment (Fig. 63 N) ........................... ......................................................................... Chirostilbia (SUBPALLIDUM sp. group) Medial antennal segment longer than apical segment (Fig. 29 S) .............................. ............................................................................................. Byssodon, Pternaspatha (in part) Cephalic apotome darkened on basal half, leaving light window on median subbasal portion (Figs. 90 E, F; 97 A-G); anal sclerite without scales ..................... ............................. Ectemnaspis (BICOLORATUM sp. group; ROMANAI sp. group; DINELLII sp. group), Psilopelmia Cephalic apotome with variable ornamentation, different from above; anal sclerite with or without scales ..................................................................................................... 20 Anal sclerite with scales; medial antennal segment longer than distal segment (Figs. 29 S; 31 b; 35 X) .................................................................................. Pternaspatha Anal sclerite without scales (Fig. 68 C); medial antennal segment shorter than distal segment (Figs. 68 D, L) ........................................................................................ 21 Cephalic apotome with dark median basal diffuse spot that becomes thinner anteriorly, delimiting 2 light elongated areas (Figs. 68 B, M; 70 M) .......Inaequalium Cephalic apotome with strong basal pigmentation generally shaped like triangle (Fig. 63 A) .............................................................. Chirostilbia (PERTINAX sp. group) DE SUBGÉNEROS DE

SIMULIUM

Hembras 1 – 2(1)

Coloración general negro o castaño negruzco ............................................................ 2 Coloración general castaño (castaño rojizo hasta amarillento) .............................. 22 Cuerpo sin especial ornamentación; pelos del escudo homogéneamente distribuidos; uñas con diente basal grande de forma subovoidal alargada; porción basal del cibario liso, sin refuerzos a los costados ............. Eusimulium, Nevermannia

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Cuerpo ornamentado con franjas o manchas claras; escudo con pelos distribuidos homogéneamente o en grupos; uñas con o sin diente subbasal, si existe es proporcionalmente más pequeño y de forma subtriangular (excepto Byssodon con diente basal) ......................................................................................................................... 3 3(2) Con pelos en sector basal de R ....................................................................................... 4 – Sin pelos en sector basal de R ......................................................................................... 9 4(3) Coloración gris polinosa; área clara del escudo delimitada a 1 banda mediana y 1+1 banda sublateral (Fig. 29 A); abdomen negruzco o grisáceo con 1+1 bandas plateadas transversales (Fig. 28 A ), o sobre el borde posterior de los tergitos IIIIV (Fig. 51 A) ................................................................................ Pternaspatha (en parte) – Coloración negruzca; escudo con o sin bandas submedianas o manchas en área anterior claras; abdomen negruzco sin bandas plateadas transversales sobre borde posterior de tergitos III-IV .............................................................................................. 5 5(4) Paraprocto de forma subtriangular, alrededor de 2 veces más largo que ancho en la base (Figs. 57 D, N); porción basal del cibario liso ............................... Chirostilbia – Paraprocto de forma no subtriangular, curvado distalmente, un poco más largo que ancho en la base (Figs. 64 G, L, P); cibario liso o con dientes ......................... 6 6(5) Porción basal del cibario con dientes dispuestos en un grupo mediano elevado subcónico (Fig. 64 M) ......................................................................................Inaequalium – Porción basal del cibario generalmente liso o con dentículos pequeños: si presentes, están dispuestos en una elevación mediana, el escudo tiene 1+1 manchas anteriores claras o 3 bandas longitudinales separadas por bandas nacaradas que cambian de tonalidad con diferente posición de la luz ................... 7 7(6) Porción basal del cibario con dientes muy pequeños o sin ellos (Figs. 102 C, F, G, L); paraproctos relativamente cortos no lobulados, escasamente aguzados apicalmente y dirigidos hacia abajo (Figs. 103 B, D, G, J) ......... Aspathia (en parte) – Porción basal del cibario con una hilera de dientes (Figs. 128 D, H), o sin dientes pero con bordes reforzados (Figs. 132 B, L); paraprocto relativamente largo, lobulado y escasamente esclerotizado, no aguzado (Figs. 128 C, E, J; 132 D, G, N) ................................................................................................................................................ 8 8(7) Pelos del escudo agrupados simulando escamas (Fig. 132 I), sin manchas anteriores grisáceas subcuadradas; cibario liso (Figs. 132 B, L) ................ Thyrsopelma (en parte) – Pelos del escudo homogéneamente distribuidos; con manchas anteriores grisáceas claras subcuadradas (Fig. 128 A); cibario con dientes dispuestos en una hilera (Figs. 128 D, H) ............................................................................ Trichodagmia (en parte) 9(3) Porción basal del cibario sin dientes ............................................................................ 10 – Porción basal del cibario con dientes .......................................................................... 16 10(9) Pelos del escudo agrupados simulando escamas (Fig. 5 A); uña sin diente subbasal o muy reducido (Figs. 54 D, E); triángulo fronto-ocular ausente; paraproctos muy finos distalmente (Figs. 54 I, Q, X) ................................................................. Notolepria – Pelos del escudo homogéneamente distribuidos; uña con o sin diente subbasal triángulo fronto-ocular bien desarrollado; paraproctos no finos distalmente .... 11 11(10) Especies gris polinoso; abdomen grisáceo con tergitos III-V negruzcos o castaño aterciopelado, generalmente con una mancha plateada mediana anterior

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o con 1+1 manchas plateadas laterales; calcipala frecuentemente reducida o ausente ................................................................................................................................ 12 Especies negruzcas, abdomen negruzco sin manchas plateadas en tergitos III-V; calcipala bien desarrollada .............................................................................................. 13 Uñas con diente subbasal (Fig. 51 F); escudo con áreas negruzcas limitadas a bandas grisáceas mediana y sublaterales; tergitos abdominales III-V negruzcos con 1+1 manchas blanquecinas sublaterales posteriores (Fig. 51 A); paraproctos de forma subrectangular, más cortos que anchos (Fig. 51 H) ............... Pternaspatha Uñas sin diente subbasal (Fig. 99 C); escudo grisáceo uniforme, excepto 1+1 manchas anteriores medianas; tergitos III-V negruzcos o castaño aterciopelado, con o sin manchas en área mediana anterior (Figs. 98 A, P; 99 A); paraproctos de forma subtriangular (Fig. 99 G) tan largos como anchos ........................................... ................................................................................... Psilopelmia (BLANCASI sp. grupo) Paraprocto subtriangular con borde distal arqueado (Figs. 118 A, B, ); escudo con 1+1 manchas anteriores submedianas continuadas posteriormente por una fina banda, uniéndose con área posterior grisácea .................... Hemicnetha (OVIEDOI sp. grupo) Paraproctos de forma no subtriangular, con borde distal truncado o aguzado posteriormente; escudo de ornamentación variable ................................................. 14 Paraprocto con borde distal truncado y suavemente aguzado posteriormente (Figs. 103 B, K); escudo con una banda negra mediana y 1+1 submediana longitudinales limitadas por bandas nacaradas que invierten la coloración con el cambio de posición de la luz (Figs. 101 A, B, F, G) ...................................... Aspathia Paraprocto con borde distal truncado o redondeado pero no aguzado posteriormente; escudo sin bandas plateadas contrastantes (Figs. 113 A, D, G, H; 120 A, B, K, L) ................................................................................................................. 15 Paraprocto tan largo como ancho en la base, con superficie externa mayormente desnuda, brillante y con depresiones, y borde distal truncado (Fig. 120 I); gonapófisis tan largas como anchas .....................................................................Hearlea Paraprocto más largo que ancho en la base, superficie externa mayormente cubierta por microtriquias, sin depresiones y con borde distal curvado (Figs. 113 C, F, L, M, R, U); gonapófisis más larga que ancha (Figs. 113 C, K, Q, T, V) ....... .................................................. Hemicnetha (en parte BRACHYCLADUM sp. grupo) Escudo con abundante polinosidad gris, mostrando, con cierta iluminación, 3-5 bandas claras; cibario con suave concavidad cubierta por dientes pequeños agudos similares; paraprocto subtriangular con borde distal curvado, generalmente más largo que ancho y a veces más fino distalmente; uñas sin dientes ................ Psilozia Escudo con escasa polinosidad, con o sin 1+1 banda plateada longitudinal; cibario con o sin dentículos, generalmente con una depresión mediana; paraprocto no subtriangular, generalmente más largo que ancho y a veces más fino distalmente; uñas con o sin diente ....................................................................................................... 17 Uñas con gran diente basal subovoidal; borde posterior de los tergitos III-V frecuentemente con 1+1 banda plateada transversa ...................................... Byssodon Uña con pequeño diente subtriangular subbasal o ausente; tergitos III-V sin banda plateada transversal .......................................................................................................... 18

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18(17) Cibario generalmente sin dentículos, si los hay no están presentes en la depresión mediana, y con dentículos a los costados (Figs. 102 F, G, K, O ); escudo con 1+1 bandas plateadas submedianas, longitudinales, alcanzando el área grisácea posterior (Figs. 101 A, F) u obscuras con cambio de dirección de luz (Figs. 101 B, G); triángulo frontoocular bien desarrollado .................................................. Aspathia – Cibario con depresión mediana esclerotizada o no, con dientes pequeños a los costados (Figs. 71 P; 76 N; 80 D; 84 M; 91 Q); escudo con o sin banda plateada longitudinal que pueden alcanzar al área posterior plateada; triángulo frontoocular generalmente ausente o reducido ................................................................................. 19 19(18) Cibario con área central bien esclerotizada y con pequeños dientes redondeados (Figs. 80 D, P, T); suturas infrafrontales bien desarrolladas (Figs. 80 A, V) ........... ................................................................... Coscaroniellum (QUADRIFIDUM sp. grupo) – Cibario con área central no esclerotizada y con agudos dientes (Figs. 71 P; 76 N, X); suturas infrafrontales no desarrolladas ................................................................. 20 20(19) Escudo negro con o sin bandas submedianas, más frecuente en borde anterior y márgenes laterales (Figs. 71 L, O, Q, U), cuando llegan al borde posterior no presentan manchas sombreadas sobre las bandas plateadas .............. Psaroniocompsa – Escudo grisáceo a negro, con banda plateada alcanzando al área grisácea posterior, presentando manchas sombreadas sobre las bandas plateadas (Figs. 76 A-F, H, I, L, M, Q, R) ..................................................................................................... 21 21(20) Cibario con dientes de aproximadamente del mismo tamaño y ausentes de la depresión mediana (Figs. 76 N, X); triángulo frontoocular muy corto (Figs. 76 G, W); abdomen sin manchas plateadas transversas sobre tergitos III-V, o muy cortas (Fig. 76 K) ........................................................................................................ Cerqueirellum – Cibario con dientes submedianos notoriamente más grandes que los restantes, y con pequeños dientes sobre área mediana (Fig. 80 Y); triángulo frontoocular relativamente profundo; abdomen con manchas plateadas transversas sobre tergitos III-V de aproximadamente 1/3 del largo del tergito (Fig. 80 X); escudo con bandas plateadas submedianas angostas (Fig. 80 W) ............................................ .......................................................... Coscaroniellum (QUADRIVITTATUM sp. grupo) 22(1) Especies amarillo rojizo a gris verdoso; si el escudo es negro, los márgenes son amarillentos (Figs. 84 B-D); paraprocto subtriangular (Figs. 85 D-P), más largo que ancho en la base y distalmente relativamente fino (Figs. 92 A-N) ................ 23 – Especies castaño rojizo a grisáceo oscuro; escudo generalmente ornamentado en forma de lira (Fig. 128 A); paraprocto subrectangullar y no aguzado distalmente (Figs. 128 C, E, J) ............................................................................................................. 28 23(22) Escudo gris verdoso a castaño amarillento; cibario liso, no esclerotizado (Fig. 60 D) .................................................................... Chirostilbia (SUBPALLIDUM sp. grupo) – Escudo anaranjado o rojizo, o negro bordeado de amarillo y frecuentemente con bandas plateadas submedianas; cibario con bordes esclerotizados o con pequeños dientes ................................................................................................................................. 24 24(23) Escudo con 1+1 o más bandas longitudinales nacaradas o plateadas (Figs. 91 BE); abdomen generalmente con 3 hileras longitudinales de manchas negruzcas subcuadradas (Figs. 91 G-I ); cibario con depresión mediana bien esclerotizada

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bordeada de suaves lobulaciones y a los costados con dientes sobre áreas suavemente elevadas (Figs. 91 J-Q); paraproctos más finos distalmente (Figs. 92 A-N) .........................................................................Psilopelmia (ESCOMELI sp. grupo) Escudo generalmente sin 1+1 o más bandas longitudinales nacaradas o plateadas (Figs. 84 A-E); abdomen sin hileras de manchas negruscas subcuadradas; cibario sin o con dientes generalmente más grandes y con diferente disposición que en el dilema opuesto (Figs. 84 I-W); paraproctos no finos distalmente (Figs. 85 D-K) . ...................................................................................................................Ectemnaspis ......25 Escudo amarillo a negro, en el último caso totalmente bordeado de amarillo (Figs. 84 A-E); cibario sin o con dientes, en el último caso con placas o pequeñas lobulaciones en el área mediana (Figs. 84 I-P) .......................................................... 26 Escudo amarillo o rojizo; cibario con dientes pero área mediana sin placas ......... 27 Cibario sin dientes pero con borde frontal bien esclerotizado (Figs. 84 I-J) .......... ............................................................................................... BICOLORATUM sp. grupo Cibario con dientes y área mediana con placas o lobulaciones (Figs. 84 K-P) ...... .......................................................................................................... ROMANAI sp. grupo Cibario con dientes bien desarrollados y agrupados mostrando 1+1 elevaciones suaves sobre el área mediana (Figs. 80 Q-S) ................... PERFLAVUM sp. grupo Cibario con dientes muy pequeños y con 1+1 elevaciones grandes, generalmente con dientes pequeños en el ápice (Figs. 84 T-W) ................... DINELLII sp. grupo Gonapófisis subovoidales más cortas que anchas (Fig. 132 M ); paraprocto suavemente aguzado distalmente (Fig. 132 N); pelos del escudo agrupados (Fig. 132 I) y ordenados en hileras longitudinales .......................... Trichodagmia (en parte) Gonapófisis subtriangulares o subovoidales, más largas que anchas, o aproximadamente tan largas como anchas; paraproctos no aguzados distalmente; pelos del escudo homogéneamente distribuidos o en grupos, pero no dispuestos en hileras longitudinales .................................................................................................. 29 Cibario escasamente esclerotizado sobre el área mediana y sin dientes ni espículas (Fig. 113 B); paraproctos un poco más largos que anchos, subrectangulares y con abundantes microtriquias (Figs. 113 C, F) ............ Hemicnetha (PAYNEI sp. grupo) Cibario bien esclerotizado sobre el área mediana, con finos dientes aguzados y escasamente esclerotizados (Figs. 113 J; 128 D, H); paraprocto aproximadamente 2 veces más largo que ancho y con escasas microtriquias (Figs. 113 L, M; 128 C, E, J) .............................................................................................................................................. 30 Gonapófisis más de 2 veces más larga que ancha, aguzadas apicalmente (Fig. 113 K); cibario con pequeños dientes y dispuestos desordenadamente (Fig. 113 J) ..... ...................................................................Hemicnetha (BRACHYCLADUM sp. grupo) Gonapófisis menos de 2 veces más larga que ancha, y con ápice romo (Figs. 128 B, I); cibario con dientes bien evidentes, dispuestos a lo largo del borde frontal extendidos en la base de los brazos laterales (Figs. 128 D, H) .............. Trichodagmia

Machos 1

Coloración general negra, a veces el abdomen con tonalidad verdosa; con o sin pelos en sector basal de R ................................................................................................ 2

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Coloración general variable, desde castaño oscuro a amarillento; generalmente con pelos en sector basal de R ...................................................................................... 23 2(1) Con pelos en sector basal de R ....................................................................................... 3 – Sin pelos en sector basal de R ....................................................................................... 11 3(2) Gonostilo más largo que el gonocoxito (Figs. 133 B, C, H, J); placa ventral relativamente corta y con o sin una prolongación mediana y carena (Figs. 133 D, I, K) ................................................................................................................................................ 4 – Gonostilo más corto que el gonocoxito; placa ventral de forma variable ............. 5 4(3) Escudo sin pelos homogéneamente dispuestos y sin manchas plateadas; placa ventral distalmente con pequeña proyección mediana y sin proyecciones laterales (Fig. 129 E) ....................................................................................................... Trichodagmia – Escudo con pelos dispuestos en grupos aparentando escamas, y con 1+1 mancha anterolateral o continuando submedianamente hasta el área grisácea posterior (Figs. 133 A, G); placa ventral distalmente con prolongaciones mediana y laterales (Figs. 133 D, I) .................................................................................................. Thyrsopelma 5(3) Gonostilo sin o con muy reducido espolón distal ............................. Chirostilbia......6 – Gonostilo con espolón distal ........................................................................................... 7 6(5) Gonostilo sin cresta lateral (Figs. 58 B, F, G, I, L, O) ........ PERTINAX sp. grupo – Gonostilo con cresta lateral (Figs. 61 D, F, H) .............................................................. ............................................................................. SUBPALLIDUM sp. grupo (en parte) 7(5) Escudo sin ornamentación; gonostilo achatado y curvo, con lóbulo interno subtriangular ............................................................................... Eusimulium, Nevermannia – Escudo generalmente ornamentado con manchas anteriores claras; gonostilo de variada forma ...................................................................................................................... 8 8(7) Gonostilo (visto en preparación microscópica) subcuadradado con concavidad distal y espolón subapical (Figs. 28 N; 30 R; 41 I) .......................... Pternaspatha......9 – Gonostilo (visto en preparación microscópica) de forma subtriangular a subtrapezoidal o cilíndrica y con espolón apical ....................................................... 10 9(8) Gonostilo con tubérculos (Figs. 28 N; 30 R; 31 S) ...................................................... .........................................................................................NIGRISTRIGATUM sp. grupo – Gonostilo sin tubérculos (Figs. 41 I; 47 K; 52 K) ........... NEMORALE sp. grupo 10(8) Gonostilo subtriangular o subtrapezoidal, con un espolón apical (Figs. 65 D, FH, J-N, P); placa ventral sin constricciones laterales (Figs. 65 C, E, I, O) ........... ...............................................................................................................................Inaequalium – Gonostilo subcilíndrico con suave curvatura distal y un espolón apical (Figs. 86 H-N) (excepto wolffhuegeli, donde es relativamente corto y con varios espolones apicales; placa ventral con suave constricción lateral (Figs. 87 A-H) ....................... ............................................ Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo) 11(2) Gonostilo notoriamente más corto que el gonocoxito ............................................ 12 – Gonostilo similar o más largo que el gonocoxito ..................................................... 19 12(11) Gonostilo sin espolón apical ......................................................................................... 13 – Gonostilo con espolón apical ........................................................................................ 14 13(12) Escudo sin ornamentación; gonostilo subovoidal, con borde reforzado (Figs. 54 N, S, T) .................................................................................................................. Notolepria

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Escudo con 1+1 manchas plateadas submedianas (Figs. 61 A, B, ); gonostilo subtrapezoidal con cresta sublateral (Figs. 61 D, F, H, I) ............................................ ......................................................................... Chirostilbia (SUBPALLIDUM sp. grupo) Escudo con 1+1 banda plateada submediana alcanzando o no área posterior grisácea; gonostilo subtriangular o subtrapezoidal ................................................... 15 Escudo sin 1+1 banda plateada submediana, pero presentando manchas plateadas anteriores subcuadradas o subtriangulares (Figs. 86 A-D; 91 A-D); gonostilo subrectangular o subcilíndrico ....................................................................................... 17 Gonostilo aproximadamente 2 veces más largo que ancho en la base; espolón apical espatulado (Figs. 81 C, G, H) ......................................................... Coscaroniellum Gonostilo más corto y solamente más largo que ancho en la base; espolón terminal subcónico ........................................................................................................... 16 Gonostilo subtriangular con espolón apical o subapical (Figs. 77 J, O; 81 L) ........ .................................................................................. Byssodon, Cerqueirellum, Coscaroniellum (QUADRIVITTATUM sp. grupo; Psaroniocompsa (SIOLII sp. grupo) Gonostilo subcuadrado o subtrapezoidal con espolón submediano (Figs. 72 F, H, I) .................................................................. Psaroniocompsa (INCRUSTATUM sp. grupo) Gonostilo generalmente subcilíndrico, con suave curvatura dorsal y un espolón apical (Figs. 86 H-N; 93 K; 118 F, H) ............................................................................. ............................................ Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo), Psilopelmia y Hemicnetha (OVIEDOI sp. grupo) Gonostilo subtrapezoidal con 2-15 espolones marginales ...................................... 18 Placa ventral más de 2 veces ancha que larga (Fig. 87 I); gonostilo con 7-15 espolones en el margen distal (Fig. 86 O) ........ Ectemnaspis (ROMANAI sp. grupo) Placa ventral tan ancha como larga; gonostilo con 2-4 espolones apicales ............. ...................................................................................................................................... Psilozia Gonostilo subcilíndrico y fino basalmente, generalmente más angosto que la mitad del gonocoxito, borde distal ancho y generalmente con una protuberancia basal ..................................................................................................................................... 20 Gonostilo basalmente más ancho que la mitad del gonocoxito, borde distal ancho y sin protuberancia basal ................................................................................................ 21 Placa ventral generalmente haciéndose más angosta medio distalmente (Figs. 106 M-P); gonostilo con protuberancia basal (Figs. 105 B, C, E-H, J-L) ......... Aspathia Placa ventral no haciéndose más angosta medio distalmente (Fig. 121 D, G ); gonostilo sin protuberancia basal pero con pequeña carena basal (Figs. 121 C, F) ...................................................................................................................................... Hearlea Gonostilo achatado, flexuoso y escasamente afinado distalmente (Figs. 114 A, E, G, I, M, N); endoparámeros con largos dientes y base bien esclerotizada (Fig. 114 K) ........................................................................................................ Hemicnetha......22 Gonostilo subtriangular afinándose distalmente y notoriamente curvo (Figs. 133 B, C, H, J); endoparámeros sin dientes y base no bien esclerotizada (Fig. 133 F) . .............................................................................................................................. Thyrsopelma Placa ventral notoriamente más ancha que larga, con una protuberancia mediana ancha (Figs. 114 B, C, F, H, L) .............. PAYNEI y BRACHYCLADUM sp. grupo

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Placa ventral aproximadamente tan larga como ancha, sin protuberancia mediana, pero con carena mediana (Figs. 114 J, O) ....................... MEXICANUM sp. grupo Gonostilo cerca de 2 veces más largo que el largo del gonocoxito ...................... 24 Gonostilo más corto o escasamente más largo que el largo del gonocoxito ...... 28 Gonostilo con 3-6 espolones o más (Fig. 129 L); endoparámeros sin dientes (Fig. 129 C) ............................................................................................. Trichodagmia (en parte) Gonostilo con 1 espolón o ausente; endoparámeros generalmente con dientes grandes (Figs. 61 I; 99 I) ................................................................................................. 25 Sector basal de R sin pelos; escudo gris opaco; gonostilo con borde interno recto (Figs. 98 V; 99 H) .................................................. Psilopelmia (BLANCASI sp. grupo) Sector basal de R con pelos (Fig. 60 C); escudo no gris opaco; gonostilo con borde interno curvo .................................................................................................................... 26 Gonostilo sin espolón apical, con cresta sublateral no curvada distalmente (Figs. 61 D, F, H, I) ................................................ Chirostilbia (SUBPALLIDUM sp. grupo) Gonostilo con espolón apical, sin cresta sublateral y suavemente curvada distalmente ......................................................................................................................... 27 Placa ventral generalmente más larga que ancha en la base (Figs. 87 J-L); gonostilo sin curvatura distal (Figs. 86 P, Q, R) .................. Ectemnaspis (PERFLAVUM sp. grupo) Placa ventral generalmente más corta que ancha en la base (Figs. 87 M-O; 94 AL); gonostilo generalmente con curvatura distal (Figs. 86 S; 93 H-R) ..................... ............................................................... Psilopelmia, Ectemnaspis (DINELLII sp. grupo) Escudo castaño oscuro con pelos agrupados simulando escamas, gonostilo relativamente grueso distalmente (Fig. 133 J) ............................................ Thyrsopelma Escudo castaño rojizo con pelos no agrupados; gonostilo relativamente fino distalmente (Figs. 129 B, D) .......................................................................... Trichodagmia

Pupas 1 – 2(2) – 3(2) – 4(3) –

Branquias con ramas relativamente gruesas con ápice agudo y bien esclerotizado en forma de espina, con 12-50 ramas (Figs. 134 B, E, G, K); capullo resistente y proyectado ventralmente hacia delante .................................... Thyrsopelma (en parte) Branquias de forma variable pero sin ápice agudo bien esclerotizado en forma de espina; capullo de forma variable ................................................................................... 2 Branquia notoriamente gruesas, asimétricas, generalmente con curvaturas .......... 3 Branquias con ramas generalmente filiformes, simétricas y sin curvaturas laterales ................................................................................................................................................ 6 Frontoclípeo y porción expuesta del tórax con tricomas simples (Fig. 121 H); branquias con estriaciones (Figs. 122 A-K; 123 A-O; 124 A-K); capullo sin prolongación dorsal .................................................................................................Hearlea Frontoclípeo y porción expuesta del tórax con tricomas multirramosos con 2-8 ramas; branquias sin estriaciones; capullo generalmente con una prolongación dorsal .................................................................................................................................... 4 Capullo generalmente sin prolongación dorsal, borde anterior recto (Figs. 66 A, B, D, L; 70 A, B, P, Q) ...........................................................................................Inaequalium Capullo prolongado dorsalmente (Figs. 73 H-K; 88 C, D, F; 107 I) ...................... 5

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– 9(7) – 10(9) – 11(10) – 12(11) – 13(6) – 14(13) – 15(14) – 16(15) –

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Tricomas del frontoclípeo y tórax bífidos (Figs. 73 O-Q) ................. Psaroniocompsa Tricomas del frontoclípeo y tórax con 3-8 ramas .................... Ectemnaspis, Aspathia Capullo con tejido resistente y porción anterior elevada, generalmente protegiendo la base de la branquia (Figs. 59 L, O, S; 115 A, D); filamentos de las branquias relativamente gruesos ..................................................................................... 7 Capullo con tejido no resistente y generalmente con la porción anterior no elevada protegiendo la branquia; filamentos de las branquias relativamente finos .......... 13 Abertura anterior del capullo con festones protegiendo la base de la branquia (Figs. 115 A-D) ................................................................................................................... 8 Abertura anterior del capullo sin festones .................................................................... 9 Branquia con 8 ramas, frecuentemente con el ápice romo y resistente (Figs. 116 A, B); frontoclípeo generalmente reforzado en la base y agrandado en el área facial, mostrando las hembras y machos similar ancho del área facial (Figs. 115 O-T) ... ........................................................................................ Hemicnetha (PAYNEI sp. grupo) Branquias con 10 ramas frecuentemente con el ápice romo no resistente (Fig. 59 N); frontoclípeo no reforzado en la base y no agrandado en área facial, mostrando el macho la base de la frente más angosta que la hembra ........... Chirostilbia (en parte) Branquia con 6 filamentos ............................................................................................. 10 Branquia con más de 6 filamentos ............................................................................... 11 Frontoclípeo y tórax con numerosos tubérculos, algunos de ellos aguzados como cortas espinas (Figs. 119 B, D, E); capullo escasamente elevado anteriormente (Fig. 119 A) ................................................................Hemicnetha (OVIEDOI sp. grupo) Frontoclípeo y tórax sin tubérculos (Fig. 115 R); capullo frecuentemente muy elevado anteriormente (Fig. 115 E) ............................................................................... ............................................................... Hemicnetha (BRACHYCLADUM sp. grupo) Branquia con 18-20 ramas ordenadas como un ramo (Fig. 130 L) ........................... .........................................................................................................Trichodagmia ( en parte) Branquias con 8-16 ramas .............................................................................................. 12 Branquia con 12 ramas ....................................................................................................... ................................. Hemicnetha (MEXICANUM sp. grupo), Trichodagmia (en parte) Branquias con 10 ramas ...................................................................................................... ................................................ Hemicnethha (BRACHYCLADUM sp. grupo en parte) Branquia con 4 ramas filamentosas .............................................................................. 14 Branquia con más de 4 ramas filamentosas ............................................................... 17 Tubérculos de frontoclípeo y tórax aguzados (Fig. 82 D) .................. Coscaroniellum Tubérculos de frontoclípeo y tórax no aguzados ...................................................... 15 Capullo reducido ventralmente (Figs. 98 X; 99 K); tricomas cefálicos y torácicos simples (Fig. 98 a-b) .............................................. Psilopelmia (BLANCASI sp. grupo) Capullo no reducido ventralmente; tricomas cefálicos y torácicos con 1-8 ramas .............................................................................................................................................. 16 Tricomas cefálicos y torácicos generalmente bífidos (Figs. 73 O, P, Q) .................. ................................................................................................. Psaroniocompsa, Cerqueirellum Tricomas cefálicos y torácicos con 1-8 ramas ............................................................... ................................................................... Eusimulium, Byssodon, Nevermannia Psilopelmia

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17(13) Branquia con 6 ramas filamentosas .............................................................................. 18 – Branquia con más de 6 ramas filamentosas ............................................................... 20 18(17) Tricomas torácicos en número sobrepasando 10 de cada lado y frecuentemente muy modificado (Figs. 40 U-V; 44 R; 53 S, T; 108 G-H; 109 A, G) ........................ ......................................................................................... Pternaspatha, Aspathia (en parte) – Tricomas torácicos en número de alrededor de 5 de cada lado ............................ 19 19(18) Tricomas cefálicos y torácicos bífidos (raramente simples o trífidos) ...................... ............................................................. Psaroniocompsa, Cerqueirellum, Aspathia (en parte) – Tricomas cefálicos y torácicos no bífidos generalmente con más de 3 ramas ....... ................................................. Byssodon, Inaequalium, Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo ), Notolepria, Psilopelmia 20(17) Branquias con 8-10 ramas .............................................................................................. 21 – Branquias con más de 10 ramas ....................................................................................... ....................... Psilozia, Chirostilbia, Aspathia Ectemnaspis (PERFLAVUM sp. grupo) 21(20) Tricomas torácicos sobrepasando en número de 10 de cada lado y frecuentemente muy modificados (Figs. 29 N, P; 31 Y, Z); branquias con 8 ramas ......... Pternaspatha – Tricomas torácicos en número de 5 de cada lado; branquias con 8-10 ramas .......... 22 22(21) Tejido del capullo esponjoso, con margen anterior elevado y carena dorsal (Figs. 88 B, H, I, J) ............................................................................................................. ....................... Ectemnaspis (BICOLORATUM y ROMANAI sp. grupo, en parte) – Tejido del capullo compacto, sin margen anterior elevado y sin carena dorsal (Figs. 62 C, D, G; 78 A, G, J) .............................. Chirostilbia (SUBPALLIDUM sp. grupo), Notolepria, Cerqueirellum, Coscaroniellum, Psilozia, Byssodon, Aspathia, Ectemnaspis (PERFLAVUM y DINELLII sp. grupo), Psilopelmia Larvas 1 – 2(1)

–

3(2)

–

Anillo anal con más de 150 hileras de ganchos ........................................................... 2 Anillo anal con menos de 120 hileras de ganchos ...................................................... 6 Cutícula del cuerpo con tricomas lanceolados (Figs. 135 G, L, S); hipostomio con diente mediano y dientes intermediarios no bien diferenciados (Figs. 135 D, I, N); abdomen no achatado dorsalmente en su porción distal (Fig. 135 A); escleritos cervicales agrandados, más de 2 veces más ancho que largo (Fig. 135 Q) ............. .............................................................................................................................. Thyrsopelma Cutícula del cuerpo sin tricomas lanceolados; hipostomio con diente mediano y generalmente los dientes intermediarios bien diferenciados (Figs. 117 N, Q; 126 J; 131 G); abdomen dorsalmente achatado distalmente (Figs. 127 G; 131 A); escleritos cervicales no agrandados menos de 2 veces más ancho que largo (Fig. 131 I) ..................................................................................................................................... 3 Esclerito anal frecuentemente con refuerzo esclerotizado mostrando en algunos casos ramas ventrales agrandadas completando un anillo (Figs. 127 B-J); margen anterior del hipostomio generalmente en forma de arco, con dientes laterales e intermediarios reducidos (Figs. 126 A, B, I, H, J) .............................................Hearlea Esclerito anal sin refuerzo esclerotizado ni alargadas las ramas ventrales completando un anillo; margen anterior del hipostomio de forma variable ....................................... 4

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Apotoma cefálico oscurecido en la mitad basal mostrando como un triángulo (Fig. 63 A) .................................................................................................................... Chirostilbia – Apotoma cefálico con diferente ornamentación ......................................................... 5 5(4) Bordes laterales del hipostomio descendiendo abruptamente desde los dientes laterales con aspecto de meseta (Figs. 117 P, Q; 131 G); mandíbula con diente preapical más corto que el apical (Figs. 117 L; 131 E, K) .......................................... ........................................... Hemicnetha (BRACHYCLADUM sp. grupo), Trichodagmia – Bordes laterales del hipostomio descendiendo suavemente desde los dientes laterales con aspecto de curva (Figs. 117 N, O, R); mandíbula con diente preapical tan largo o más que el apical (Figs. 117 K, M) .............................................................. ........................................................ Hemicnetha (PAYNEI; MEXICANUM sp. grupo) 6(1) Anillo anal con 100-120 hileras de ganchos; rayos del abanico cefálico con dientes de similar tamaño (Fig. 119 J); mandíbulas con un par de dientes marginales muy grandes (Fig. 119 K) ................................................ Hemicnetha (OVIEDOI sp. grupo) – Anillo anal con menos de 100 hileras de ganchos; rayos del abanico cefálico alternando un grupo de pequeños dientes con alguno más grande; mandíbula con solo un diente marginal, y si existen 2 el segundo es muy pequeño (Figs. 29 S; 79 B; 83 C, O, R; 90 Q) .......................................................................................................... 7 7(6) Apotoma cefálico sin ornamentación o reducida a una pequeña mancha en el área medio basal; con numerosos y cortos pelos sobre la cápsula cefálica (Fig. 79 C); generalmente con banda oscura alrededor del primer segmento abdominal; antena fina y sobrepasando el ápice del tallo del abanico cefálico (Fig. 79H) ................... 8 – Apotoma con evidente ornamentación; sin pelos o muy escasos en cápsula cefálica; antena generalmente gruesa y sobrepasando o no el ápice del tallo del abanico cefálico .................................................................................................................. 9 8(7) Mandíbulas con una hilera de cerdas supramarginal inferior (Fig. 79 B); artejo antenal mediano similar o más largo que el proximal (Figs. 79 D, E, I) .................. ............................................................................................................................ Cerqueirellum – Mandíbulas sin una hilera de cerdas supramarginal inferior; artejo antenal mediano aproximadamente 1/2 a 4/5 del largo del proximal (Figs. 83 B, N, Q) .. .......................................................................................................................... Coscaroniellum 9(7) Hendidura postgenal escasamente desarrollada, con puente postgenal más largo que el hipostomio (Figs. 112, A, B, D, G); papilas rectales sin divertículos en los lóbulos ................................................................................................................................ 10 – Hendidura postgenal bien desarrollada con puente postgenal tan largo o más corto que el largo del hipostomio; papilas rectales generalmente con divertículos en los lóbulos .................................................................................................................... 13 10(9) Antena con estriación transversal y sobrepasando el ápice del tallo del abanico cefálico (Figs. 111 A, B, H, L); artejo antenal mediano más largo que el proximal (Figs. 111 B, H, L, M) ........................................................................................... Aspathia – Antena sin estriación transversal y no sobrepasando el ápice del tallo del abanico cefálico; artejo mediano más corto que el artejo proximal ..................................... 11 11(10) Noveno esternito sin 1+1 tubérculos ............... Psilopelmia (BLANCASI sp. grupo) – Noveno esternito con 1+1 tubérculos bien evidentes ............................................. 12

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12(11) Antena sobrepasando el ápice del tallo del abanico cefálico ...................................... ....................................................................................................... Eusimulium, Nevermannia – Antena llegando aproximadamente al ápice del tallo del abanico cefálico .............. ...................................................................................................................................... Psilozia 13(9) Papilas rectales sin divertículos en los lóbulos .............................................................. ................................................................................... Psilopelmia (BLANCASI sp. grupo) – Papilas rectales con divertículos en los lóbulos ......................................................... 14 14(13) Apotoma cefálico con manchas aisladas postero medianas, antero-laterales y postero-laterales, generalmente positivas (Figs. 30 Y; 55 C; 56 K; 75 L) ............. 15 – Apotoma cefálico con ornamentación variable diferente a la señalada en el dilema opuesto ............................................................................................................................... 19 15(14) Esclerito anal sin escamas (Fig. 63 G) ......................................................................... 16 – Esclerito anal con escamas (Fig. 63 P) ........................................................................ 17 16(15) Antena sobrepasando el ápice del tallo del abanico cefálico (Figs. 75 D, L); artejo antenal mediano generalmente más corto que el distal (Figs. 75 A, E, M) ............. ......................................................................................................................... Psaroniocompsa – Antena no sobrepasando el ápice del tallo del abanico cefálico; artejo antenal mediano generalmente más largo que el distal (Figs. 90 M, N) ................................. ....................... Ectemnaspis (BICOLORATUM sp grupo; PERFLAVUM sp. grupo) 17(15) Largo máximo del cuerpo por debajo de 4,5 mm ....................................... Notolepria – Largo máximo del cuerpo por arriba de 5,0 mm ...................................................... 18 18(17) Artejo antenal mediano más corto que el apical (Fig. 63 N) ...................................... ......................................................................... Chirostilbia (SUBPALLIDUM sp. grupo) – Artejo antenal mediano más largo que el apical (Fig. 29 S) ........................................ .......................................................................................... Byssodon, Pternaspatha (en parte) 19(14) Apotoma cefálico oscurecido en la mitad basal, dejando una ventana clara sobre la porción medio basal (Figs. 90 E, F; 97 A-G); esclerito anales sin escamas ............ ........................................................................ Ectemnaspis (BICOLORATUM sp. grupo; ROMANAI sp. grupo; DINELLII sp. grupo), Psilopelmia – Apotoma cefálico con ornamentación variable, diferente al del dilema opuesto; esclerito anal con o sin escamas ................................................................................... 20 20(19) Esclerito anal con escamas; artejo antenal mediano más largo que el distal (Figs. 29 S; 31 b; 35 X) ............................................................................................. Pternaspatha – Esclerito anal sin escamas (Fig. 68 C); artejo antenal mediano más corto que el distal (Figs. 68 D, L) ........................................................................................................ 21 21(20) Apotoma cefálico con una mancha oscura difusa mediana basal que se vuelve más fina hacia adelante, delimitando 2 áreas claras alargadas (Figs. 68 B, M; 70 M) ...............................................................................................................................Inaequalium – Apotoma cefálico con fuerte pigmentación basal, generalmente con forma triangular (Fig.63 A) ............................................. Chirostilbia (PERTINAX sp. grupo)

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Simulium (Eusimulium) ROUBAUD Eusimulium ROUBAUD, 1906a: 521. Type-species: Simulium aureum FRIES, 1824: 16.

Only a short diagnosis is given. Female: Coloration brown to black, without scutum ornamentation; calcipala well developed; claws with wide basal tooth lanceolate; cibarium without teeth; paraprocts short; spermatheca sclerotization reaching spermathecal duct. Male: Gonocoxite larger than gonostylus; gonostylus with strong curvature, subtriangular and 1 terminal spur; ventral plate laterally compressed. Pupa: Cocoon slipper shaped; generally with 4 gill filaments; thoracic trichomes generally single. Larva: Hypostomium with well-developed teeth, with median tooth at same level as lateral teeth. Postgenal bridge nearly 2 times hypostomium length; ninth segment with 1+1 ventral tubercles, rectal papillae without diverticula. Distribution: This taxon is primarily Holarctic, with only one species in the Neotropics. Simulium (Eusimulium) donovani VARGAS, 1943b: 359-360; VARGAS & DÍAZ NÁJERA, 1957b: 164-166; DALMAT, 1955: 117; ADLER et al., 2004: 328-329. Simulium (Eusimulium) aureum (FRIES), 1824: 16; VARGAS et al., 1946: 166; VARGAS & DÍAZ NÁJERA, 1957b: 153, 164-166, 231; DALMAT, 1955: 116-121, 153, 164- 166, 231 (misidentification). Simulium diazi DE LEÓN, 1945: 70.

The species was treated by ADLER et al. (2004). Distribution: USA: Arizona, California, Colorado, Nevada, New Mexico, Oregon, Texas; Mexico: Chiapas, Durango, Guanajuato, Michoacan, Morelos, Veracruz; Guatemala. Simulium (Nevermannia) ENDERLEIN Nevermannia ENDERLEIN, 1921a: 199. Type-species: Simulium annulipes BECKER, 1908: 72-73 (= Simulium ruficorne MACQUART, 1838: 88).

Only a short diagnosis is given. Female: Coloration brown to black, without scutum ornamentation; calcipala well developed; claws with well-developed subbasal tooth; cibarium without teeth. Male: Gonostylus distally with medially directed, subtriangular flange. Paramere with 1 or 2 large, distal spines. Dorsal plate well developed. Pupa: Cocoon slipper shaped, with or without anterodorsal projection. Gill of 4-8 filaments. Larva: Hypostomium with well-developed teeth; lateral teeth moderately enlarged. Ninth segment with 1+1 ventral tubercles, rectal papillae with or without diverticula. Distribution: This taxon is primarily Holarctic and Oriental, with only one species in the Neotropics.

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Simulium (Nevermannia) carbunculum ADLER, CURRIE & WOOD, 2004 The species was treated by ADLER et al. (2004). Distribution: USA: Arizona, California, Colorado, Idaho, Nevada, New Mexico, Utah, Wyoming; Canada: British Columbia, Saskatchewan; Mexico: Durango. Discussion: Simulium pugetense was recorded from Durango, Mexico, by VARGAS & DÍAZ NÁJERA (1957b: 153, 164-166, 231), but is actually Simulium carbunculum ADLER, CURRIE & WOOD, 2004, as reported by ADLER et al. (2004). Simulium (Byssodon) ENDERLEIN Byssodon ENDERLEIN, 1925: 209. Type-species: Simulium forbesi MALLOCH, 1914: 63-65 (= Simulium meridionale RILEY, 1887: 513-514).

Female: Small species; scutum blackish with submedian silvery stripes in shape of lyre; claw generally with elongate basal tooth, as in Eusimulium; cibarium generally without teeth; radius without hair; paraproct short weakly acuminate ventrally. Male: Gonostylus shorter than gonocoxite; relatively thin and curved with one apical spur; ventral plate flat, wider than long; endoparameres with large teeth. Pupa: Gill with 2-6 or 22-26 branches; cocoon slipper or shoe shaped. Larva: Cephalic apotome without clear ornamentation, with spots positive and negative; postgenal bridge about half of hypostomium length; hypostomial teeth with median tooth beneath level of lateral teeth; ninth segment with or without ventral tubercles; rectal papillae without diverticula on lobes. Distribution: This taxon is present in the Holarctic, Oriental and Afrotropical Regions.

Simulium (Byssodon) meridionale RILEY, 1887: 513-514. Simulium tamaulipense TOWNSEND, 1897: 171-172 described from Mexico, is considered a synonym.

Distribution: Very well represented in Canada and USA; Mexico: Tamaulipas. Discussion: Simulium tamaulipense was described from Mexico but it is considered a synonym of Simulium meridionale. Simulium (Psilozia) ENDERLEIN Psilozia ENDERLEIN, 1936: 113-114. Type-species: Psilozia groenlandica ENDERLEIN, 1936: 114 (= Simulium vittata ZETTERSTEDT, 1838: 803. Simulium (Neosimulium) VARGAS et al., 1946: 103, 108, 160. Type-species Simulium vittata ZETTERSTEDT, 1838.

Female: Grayish pollinose; scutum black with 5 longitudinal silvery stripes; cibarium with small regular teeth; basal sector of R without hair; calcipala small; claws without basal tooth; paraprocts subtriangular, slightly longer than wide. Male: Scutum black with silvery spots anteriorly; gonostylus about half gonocoxite length; subtrapezoidal, with 2-3 apical spurs; ventral plate subtriangular with short basal branches.

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Pupa: Cocoon slipper shaped, relatively short; gills with 10-24 filaments: cephalic and thoracic trichomes simple. Larva: Cephalic apotome with positive spots; postgenal bridge nearly 1.5 times longer than hypostomium; ninth segment without ventral tubercles; rectal papillae generally without diverticula. Discussion: This subgenus has five species in the USA; three of them are also present in Mexico. Simulium (Psilozia) argus WILLISTON, 1893: 253-254; VARGAS & DÍAZ NÁJERA, 1957b: 263; PETERSON & KONDRATIEF, 1995: 36-37. Simulium kamloopsi HEARLE: VARGAS et al., 1946: 171.

Distribution: Mexico: Distrito Federal, Nueva León, Oaxaca, Puebla, San Luis Potosi, Sonora, Veracruz; Morelos by VARGAS & DÍAZ NÁJERA, 1948b, 1949, 1957b. Discussion: For more information see ADLER et al., 2004: 354-355. Simulium (Psilozia) encisoi VARGAS & DÍAZ NÁJERA, 1949: 292-295; 1957b: 180-182, 233, 263, 336. Material not revised. Distribution: Cited in Mexico: Hidalgo, Michoacan. Discussion: For more information see ADLER et al., 2004: 355. Simulium (Psilozia) vittatum ZETTERSTEDT, 1838: 803; VARGAS, 1942a: 243; VARGAS & DÍAZ NÁJERA, 1948b: 324; PETERSON & KONDRATIEFF, 1995: 37; ADLER et al., 2004: 356-361, 363-364. Material not revised. Distribution: Mexico: Distrito Federal, Tabasco. Discussion: For more information see ADLER et al., 2004: 356-364. Simulium (Pternaspatha) ENDERLEIN (Figs. 28-53; 137, 138.) Pternaspatha ENDERLEIN, 1930: 84, 88; EDWARDS, 1931: 151; ENDERLEIN, 1934b: 274; 1936: 115; SMART, 1945: 465, 488; VARGAS, 1945a: 102; VARGAS & DÍAZ, 1951: 143; STONE, 1962: 206, 208; 1963: 17. Type-species: Pternaspatha nigristrigata ENDERLEIN. Acropogon ENDERLEIN, 1934b: 276; 1936: 118; SMART, 1945: 489; VARGAS, 1945a: 85; STONE, 1962: 206; 1963: 1. Type: Acropogon barbatipes ENDERLEIN. Dasypelmoza ENDERLEIN, 1934b: 275; SMART, 1945: 488; VARGAS, 1945a: 90; VARGAS & DÍAZ, 1951: 146; STONE, 1962: 206; 1963: 5. Type-species: Simulium varipes PHILIPPI, 1865: 634.

Adults: Antennae 11 segmented; eyes of female with fronto-ocular triangle; row of hairs between eyes of male; hind border of cibarium of female unarmed; mandibles and lacinia of female toothed on both sides; mesonotum of female gray, with 1+1 sublateral

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and one submedian, black, longitudinal stripes, median stripe divided longitudinally by fine grayish line; scutum of male black, bordered with gray; anterior border with 1+1 small, white, submedian spots; delicate, white, central line often visible on anterior half of disc of mesonotum; pleural tuft present; setae lacking from pleural membrane and sternopleurum; costa with both hair and spinelike setae; radial sector not branched; basal portion of R bare, very rarely with hairs dorsally; R1 with both hair and spiniform setae; numerous long hairs on fore coxa, femur, and tibia of male; fore tarsi narrow; hind basitarsus of female narrow; of male, from narrow to conspicuously widened, spindleshaped; calcipala small to absent; pedisulcus well developed; claws of female with small subbasal tooth; basal fringe of abdomen of male exceptionally long; last tergites of female dull; gonostylus subquadrate, shorter than gonocoxite, with single apical spine; posterior surface glabrous or granulose; ventral plate transverse, in many instances with 1+1 translucent areas; basal arms short; median sclerite elongate, widened and deeply incised on distal half; spermatheca subglobular; outer surface not sculptured; inner surface with minute spicules arranged in groups of one to five; genital fork slender; stem strongly pigmented; arms not pigmented; apical expansions of arms partly translucent, partly pigmented; paraprocts short, rather rounded or subtriangular. Pupa: Cocoon from slipper shaped to shoe shaped, with distinct anteroventral bridge; close-woven, threads not conspicuous; rim of aperture slightly reinforced, not festooned; gills shorter than pupa, with six or eight slender filaments; their surface with granules forming spiral pattern; apex of filaments rounded; exposed portion of thorax with numerous trichomes, from approximately 10+10 to more than 100+100; cephalic and thoracic trichomes of greatly varied structure: hair like, spatulate, ribbon-shaped, simple or branched; all trichomes either of single type or various types intermixed; abdominal tergite II with 3+3, rarely with 4+4, simple spines; tergites III-IV with 4+4 simple spines; tergites VI-IX on anterior half with transverse rows of small, spinelike or scale-like tubercles; sternites V-VII with 2+2 strong, simple or bifid spines; sternites IIIVIII on anterior portion of disc with 1+1 fields of minute scales; apical hook of abdomen not developed. Larva: Thorax slightly swollen; abdomen gradually expanded, slightly clubbed posteriorly; no ventral tubercles at posterior of abdomen; body surface glabrous, or with only very sparse, simple setae; setae more numerous at base of anal sclerite; hypostomial setae arranged in two irregular series on each side, very rarely only in one series; postgenal cleft subtriangular, narrowed apically; postgenal bridge from well developed to extremely short, but postgenal cleft in no case completely attaining hypostomial groove; anal ring with 80-90 rows composed of 13 to 20 hooks each (Figs. 137, 138). Distribution: Cool and cold temperate western and extreme southern South America, north as far as central Ecuador. KEYS

TO

SIMULIUM (PTERNASPATHA)

SPECIES

Females The female of Simulium (Pt.) bordai is not known, and we do not have sufficient information to include Simulium (Pt.) schoenemanni in the key.

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Basal portion of R bare (Fig. 39 I) ................................................................................. 2 Basal portion of R setose (Fig. 51 D) ............................................ Simulium annulatum Abdomen (Fig. 36 C) dull silver-gray; each of terga II-V with relatively small, central, velvety black spot, and with lateral dark spots absent; calcipala very small (Fig. 36 G) ...........................................................................................Simulium strigidorsum – Abdominal terga II-V with more extensive dark markings; calcipala absent or present .................................................................................................................................. 3 3(2) Calcipala absent (Figs. 32 A; 50 J) .................................................................................. 4 – Calcipala present, although very small in some species ............................................. 9 4(3) Terga IV and V without 1+1 silver-white spots, entirely black, although in some specimens narrowly margined with white along hind borders (Fig. 50 A) .............. ....................................................................................................... Simulium prodexargenteum – Terga IV and V with 1+1 large silver-white spots (Fig. 28 A) ................................. 5 5(4) Claws without subbasal tooth (Fig. 32 B) ..................................... Simulium horcochuspi – Claws with subbasal tooth (Fig. 28 J) ............................................................................ 6 6(5) Abdominal tergites III-V with 1+1 well-developed lateral blackish spots meeting anteriorly with median spot (Figs. 28 A; 29 B); wing length 2.3-3.5 mm .............. 7 – Abdominal tergites III-V with 1+1 small isolated lateral blackish spots (Figs. 35 B; 37 B); wing length 3.3-3.7 mm ........................................................................................ 8 7(6) Abdominal tergites III-V with light grayish coloration extending to lateral area of tergites (Fig. 29 B); fore and mid femora light brown, not conspicuously darkened apically; wing length 2.8-3.5 mm ............................................................. Simulium limay – Abdominal tergites III-V with light grayish coloration extending or not to lateral area of tergites (Fig. 28 A); fore and mid femora conspicuously darkened at apex; wing length 2.3-2.7 mm ................................................................ Simulium nigristrigatum 8(6) Abdominal tergite VI with median black spot (Fig. 37 B); 1+1 lateral black spots on tergites III-V reaching to posterior border; length/width ratio of hind basitarsus = 7.2-7.6 ............................................................................. Simulium deagostinii – Abdominal tergite VI without median blackish spot (Fig. 35 B); 1+1 lateral black spots on tergites III-V not reaching posterior border; length/width ratio of hind basitarsus = 6.6 ................................................................................Simulium diamantinum 9(3) Tergum III without 1+1 light-colored spots, thus terga III-V entirely dark except in some species with narrow white line along lateral portions of hind margin (Fig. 51 A); mid-longitudinal light-colored line of scutum invisible or only faintly perceptible ......................................................................................................................... 10 – Tergum III with 1+1 gray or whitish spots (Figs. 47 A; 48 A; 49 A; 50 A), in some specimens small and not easily seen but in all cases perceptible; central lightcolored line of scutum invariably clearly visible ....................................................... 11 10(11) Bases of tibiae conspicuously pigmented, but apex of hind basitarsus lacking conspicuous pigment; terga III-V entirely dark (Fig. 52 A); paraprocts roundly salient; cerci relatively high (Fig. 52 H) .................................................. Simulium caprii – Bases of tibiae lacking distinct pigment, with hind basitarsus conspicuously pigmented on apical third (Fig. 53 H); posterior border of terga III-V narrowly

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bordered with gray (Fig. 53 A); paraprocts very short and truncate, cerci relatively low (Fig. 53 L) ....................................................................................... Simulium stelliferum Terga IV and V lacking 1+1 gray or white spots, only in some cases their hind margins narrowly bordered with white laterally (Figs. 47 A; 50 A); tibiae with distinct dark basal or subbasal annulus ....................................................................... 12 Terga IV and V, in some cases only tergum V, with 1+1 light-colored spots of varied sizes (Figs. 33 A; 38 C; 40 A; 41 A); tibiae with or without dark subbasal or basal annulus ..................................................................................................................... 16 Calcipala not well developed (Fig. 50 J) ................. (some)......Simulium prodexargenteum Calcipala well developed, distinct (Figs. 39 K; 48 C) ............................................... 13 Dark area separating white spots on abdominal tergum II very narrow, acuminate anteriorly (Fig. 48 A); tergum VI entirely silver-white ............... Simulium albicinctum Dark area separating white spots on abdominal tergum II wide (Figs. 39 B; 47 A; 49 A), widened anteriorly or as wide anteriorly as posteriorly; tergum VI dark at center and laterally ........................................................................................................... 14 Setae and spines on R1 arranged in several irregular rows (Fig. 39 I) ...................... .................................................................................................................... Simulium nemorale Setae and spines on R1 arranged in single row (similar to Fig. 42 P) ................... 15 Calcipala very small, narrowly triangular (Fig. 49 E) ...........................Simulium luchoi Calcipala larger, rounded (Fig. 47 F) ............................................Simulium hectorvargasi Frons very wide (Fig. 38 D); paraproct with ventral posterior angle elongated (Fig. 38 N); scutum black, with variable ornamentation in accordance with light incidence (Figs. 38 A, B) ....................................................................... Simulium cotopaxi Frons not wide (Fig. 40 B); paraproct with ventral posterior angle rounded (Fig. 40 L); scutum grayish, with median and 1+1 sublateral stripes blackish .................. 17 Tibiae not conspicuously darkened at base (Figs. 40 F, G, H) ............................... 18 Tibiae conspicuously darkened at base ....................................................................... 23 1+1 light-colored spots on terga III-V very small, in some cases difficult to perceive (Fig. 40 A; 41 A); wing length 2.9-3.9 mm ................................................. 19 1+1 light-colored spots on terga III-V larger, all distinct (Fig. 33 A); wing length 2.5-3.6 mm ......................................................................................................................... 20 Light submedian spots on tergites III-V crossing from anterior to posterior border of segment and with parallel sides (Fig. 40 A); claws with small subbasal tooth (Fig. 40 J) ................................................................................... Simulium barbatipes Light submedian spots on tergites III-V not reaching anterior border, and with enlarged base (Fig. 41 A); claws without subbasal tooth (Fig. 41 F) ........................ .................................................................................................................... Simulium philippii Calcipala well developed (Fig. 33 E) ............................................................................ 21 Calcipala minute (Fig. 31 I) ............................................................................................ 22 Wing length 3.2-3.5 mm; hind tibia with pigment often only at extreme apex ...... ......................................................................................................................... Simulium simile Wing length 2.6-2.8 mm; pigment of hind tibiae always more widely extended, on apical two-fifths of segment (Fig. 34 D) .......................................... Simulium pulchrum

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22(20) Fronto-ocular triangle longer than wide; wing length greater than 3.0 mm; Sc with approximately 10 hairs .............................................................................. Simulium dureti – Fronto-ocular triangle not longer than wide; wing length 3.0 mm or less; Sc without or with at most 1 or 2 hairs ........................................................ Simulium pichi 23(17) Setae and spines on R1 and setae on Rs arranged in several irregular rows (Fig. 39 I); white spots on terga IV and V faint, narrowly elongate (Fig. 39 C) ................... .................................................................................................................... Simulium nemorale – Setae and spines on R1 and setae on Rs arranged in single row, as in male (Fig. 42 P); white spots on terga IV and/or V generally distinct, rounded or rectangular (Fig. 42 A) .......................................................................................................................... 24 24(23) Paired white spots on tergum V as large as those on tergum III (Fig. 44 A) ......... .............................................................................................................. Simulium albilineatum – Paired white spots on tergum V distinctly smaller than those on tergum III (Figs. 42 A; 46 A) ........................................................................................................................ 25 25(24) Tooth of claws exceptionally small (Fig. 43 F); central area of eighth sternum lighter than lateral portions; paraprocts pointed (Fig. 43 G) .................... Simulium yacuchuspi – Tooth of claws normally developed (Fig. 46 A); central area of eighth sternum darker than lateral portions; paraprocts broadly rounded ....................................... 26 26(25) Abdomen (Fig. 42 A) with 1+1 silver-white spots of tergum III only slightly surpassing posterior half of tergum anteriorly; tergum IV with 1+1 small whitish spots; central whitish area of tergum VI not bisected longitudinally by black stripe ....................................................................................................................... Simulium herreri – Abdomen (Fig. 46 A) with paired whitish spots of tergum III coming close to anterior border of tergum; tergum IV lacking spots; light-colored whitish area of tergum VI bisected longitudinally by dark stripe ...................... Simulium quechuanum Males The males of Simulium (Pt.) bordai, Simulium (Pt.) schoenemanni and Simulium (Pt.) strigidorsum are not known and have, therefore, not been included in the keys. The male of Simulium (Pt.) horcochuspi is only known from a poorly preserved pharate specimen and also has not been included; if better known, it might key out as Simulium (Pt.) deagostinii. 1 – 2(1) – 3(2) – 4(3) – 5(4)

Basal portion of R bare (Fig. 39 I) ................................................................................. 2 Basal portion of R setose (Fig. 51 D) ............................................ Simulium annulatum Scape, pedicel and basal palpomeres with very long hair (Figs. 38 Q, R); scape and pedicel combined about as long as flagellum .................................. Simulium cotopaxi Scape, pedicel and basal palpomeres with short hair; scape and pedicel combined about 1/3 as long as flagellum ........................................................................................ 3 Calcipala absent (Figs. 50 P-R) ........................................................................................ 4 Calcipala present, although in some species very small ............................................. 8 Hind tibiae conspicuously darkened at base (Figs. 37 L; 50 P-Q) .......................... 5 Hind tibiae not conspicuously darkened at base ......................................................... 6 Abdomen with paired silvery spots on terga II-VII (Fig. 50 L); gonostylus with apical surface smooth (similar to Fig. 42 S) ........................ Simulium prodexargenteum

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Abdomen with paired silvery spots only on segments II, VI, and VII; apical surface of gonostylus granulose (similar to Fig. 37 M) .............. Simulium deagostinii Hind basitarsus 5.0 times as long as wide ............................................. Simulium limay Hind basitarsus 3.3-3.8 times as long as wide .............................................................. 7 1+1 silvery whitish spots not reaching anterior rim on tergites III-V; wing length 2.12.5 mm; length/width ratio of hind basitarsus = 3.3-3.5 .......... Simulium nigristrigatum 1+1 silvery whitish spots reaching anterior rim on tergites III-V; wing length 2.83.0 mm; length/width ratio of hind basitarsus = 3.0 ............. Simulium diamantinum Apical surface of gonostylus smooth (Fig. 47 K) ....................................................... 9 Apical surface of gonostylus tuberculate (Fig. 28 N) .............................................. 20 Base of hind tibia conspicuously darkened (Figs. 47 J; 49 L; 50 P, Q; 53 N) .... 10 Base of hind tibia not conspicuously darkened (Fig. 40 O) ...... Simulium barbatipes Hair and spines on R1 and Rs arranged in 1 or 2 irregular rows (as in female, Fig. 39 I); hind basitarsus narrower, about 4.0 times as long as wide .......................... 11 Hair and spines on R1 and Rs arranged in single row (Fig. 42 P); hind basitarsus wider, not more than 3.5 times as long as wide ........................................................ 14 Wing length > 3.5 mm .......................................................................... Simulium nemorale Wing length < 3.5 mm .................................................................................................... 12 Wing length 2.7-2.9 mm; hind basitarsus narrower (4.3-5.5 times as long as wide) (Fig. 53 N) .............................................................................................. Simulium stelliferum Wing length 3.1-3.3 mm; hind basitarsus wider (3.5-4.0 times as long as wide) .............................................................................................................................................. 13 Hind basitarsus 3.5 as long as wide .................................................... Simulium philippii Hind basitarsus 3.8-4.0 as long as wide .................................................. Simulium caprii Abdominal terga III-V entirely black, or III with 1+1 minute light-colored spots (Fig. 47 I) ........................................................................................................................... 15 Abdominal tergum III or V with distinct 1+1 silvery white spots (Figs. 42 M; 43 H; 50 L) .............................................................................................................................. 18 Basal 3/5 of posterior femur whitish, sharply contrasting with dark apex (Fig. 49 L); posterior basitarsus <3.0 times as long as wide (Fig. 49 L) .............................. 16 Basal three-fourths of hind femur light brown (Fig. 47 J); posterior basitarsus at least 3.0 times as long as wide (Fig.47 J) ..................................................................... 17 Calcipala very small, subtriangular (Fig. 49 M) .....................................Simulium luchoi Calcipala somewhat larger, distinctly rounded (Fig. 48 G) .......... Simulium albicinctum Calcipala very small (Fig. 46 C) ..................................................... Simulium quechuanum Calcipala larger (Fig. 47 F) .............................................................Simulium hectorvargasi Hair of abdomen silvery; tergum III uniformly dark, but V with 1+1 distinct white spots (Fig. 42 M); dark spot at bases of fore and mid femora short ............ ....................................................................................................................... Simulium herreri Hair of abdomen light brass colored; tergum III with 1+1 small but distinct silvery white spots at hind border, about half as high as height of segment; spot at bases of fore and mid femora narrowly elongate ..................................................... 19 Posterolateral margins of abdominal terga IV and V narrowly bordered with silver or white (Fig. 50 L) ................................................................... Simulium prodexargenteum

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Posterolateral margin of abdominal terga IV and V without light colored areas except rarely on V (Figs. 43 H; 44 I) ........ Simulium albilineatum, Simulium yacuchuspi 20(8) Calcipala very small (similar to Fig. 30 P) ................................................................... 21 – Calcipala well developed (similar to Fig. 33 E) .......................................................... 22 21(20) Wing length 3.0 mm; hind basitarsus wide, 3.2-3.4 times as long as wide; under certain angles of illumination, at least anterior 1/3 of scutum grayish ................... ........................................................................................................................ Simulium dureti – Wing length 2.3-2.6 mm; hind basitarsus narrower, 3.8 times as long as wide; only about 1/5 of anterior portion of scutum gray ..................................... Simulium pichi 22(20) Length of wing 2.8-3.0 mm; abdomen with small but distinct, paired, lightcolored spots on tergum III (Figs. 33 J, K) ........................................... Simulium simile – Length of wing 2.6-2.7 mm; tergum III lacking any trace of white spots .............. ................................................................................................................... Simulium pulchrum Pupae Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni, and Simulium (Pt.) strigidorswn are not included in the key below because their pupae are not known. 1 – 2(1) – 3(2) – 4(3) – 5(2) – 6(5) – 7(6)

–

Gill with 8 filaments .......................................................................................................... 2 Gill with 6 filaments ........................................................................................................ 16 Lower or both filaments of ventral primary branch of gill sharply diverging from remaining filaments, closely adhering to body of pupa (Figs. 42 X, U; 46 F; 49 O; 50 U) ..................................................................................................................................... 3 All filaments of gill approximately parallel, at least on apical half ......................... 5 Many trichomes of head and thorax shortly lanceolate, generally bifid (Figs. 49 Q-S) ................................................................................................................Simulium luchoi Trichomes of head and thorax not as above ............................................................... 4 Trichomes of thorax elongate-lanceolate, numerous, about 250+250 (Figs. 42 X, Y) .......................................................................................... (some)......Simulium herreri Trichomes of thorax fewer, about 12+12, simple or bifid (Fig. 50 W) ................... ....................................................................................................... Simulium prodexargenteum Trichomes of head and thorax very short, a few branched but most simple, from hair-like to spine-like, those of thorax concentrated in transverse row at base of exposed portion (Fig. 39 X) ................................................................. Simulium nemorale Trichomes of head and thorax much longer, their arrangement not as above .... 6 Trichomes of head and thorax very numerous, flattened, their branches ribbonlike (Figs. 33 R-T; 34 I) ..................................................................................................... 7 Trichomes of head and thorax not as above ............................................................... 8 Length of cocoon along base 5.2-6.8 mm; frontoclypeus (Fig. 33 R) with about 400 tubercles, with cephalic tubercles not surpassing area of facial trichomes, this region reinforced by sclerotized edge; thoracic trichomes not numbering more than 250+250, average 140+140 ............................................................. Simulium simile Length of cocoon along base 3.8-4.5 mm; frontoclypeus (Fig. 34 I) with about 800 tubercles; cephalic tubercles surpassing area of facial trichomes, this region

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–

14(13) – 15(14) – 16(1)

smooth, not reinforced at edges; thoracic trichomes numbering not more than 130+130, average 70+70 ..................................................................... Simulium pulchrum Disc of exposed portion of thorax without tubercles, or tubercles only along area adjacent to rim of aperture of cocoon (Figs. 39 X; 47 Q) ....................................... 9 Disc of exposed portion of thorax with numerous tubercles (Fig. 30 V) .......... 11 Frontal trichomes reduced to minute cones (Fig. 47 O); thorax with about 150+150 trichomes (47 Q) .............................................................Simulium hectorvargasi Frontal trichomes normally developed, hair-like; thorax with fewer than 100+100 trichomes ............................................................................................................................ 10 Thorax with about 70+70 trichomes (Figs. 52 Q-R), these trichomes either simple or, frequently, with 2 or 3 branches ........................................................ Simulium caprii Thorax with not more than 20+20 trichomes, these generally with 4 or more branches (Fig. 28 T) ....................................................................... Simulium nigristrigatum Trichomes of thorax lacking from anterior portion of sclerite ............................. 12 Trichomes of thorax practically covering all of exposed surface of sclerite (similar to figure 45 D) ................................................................................................... 13 Thorax with about 20+20 trichomes (Fig. 29 N); tubercles of frontoclypeus sparse, their surface minutely spinose; male pupa with tubercles only at base of frontoclypeus (Fig. 29 Q) .......................................................................... Simulium limay Thorax with about 60+60 trichomes (Fig. 30 V); tubercles of frontoclypeus more numerous, their surface smooth; male pupa with frontoclypeus entirely covered by tubercles (Fig. 30 U) ............................................................................... Simulium pichi Cocoon with leaden sheen; surface of tubercles of frontoclypeus smooth; frontal trichomes (Figs. 32 F, H) arranged in groups of 3 or 4, distinctly separated from facial trichomes which are single; frontal trichomes (Fig. 32 H) single or in groups of 2; trichomes of thorax most frequently with 1 to 3 branches (Fig. 32 I) .......... ................................................................................................................ Simulium horcochuspi Cocoon without leaden sheen; tubercles of frontoclypeus more or less distinctly tuberculate; frontal trichomes (Fig. 31 X) more numerous, contiguous with facial trichomes which number more than 1; ocular trichomes (Figs. 31 X; 37 P) numbering more than 2; trichomes of thorax most frequently with more than 3 branches (Fig. 31 Z) ......................................................................................................... 14 Number of thoracic trichomes on each side 35-80 (Fig. 37 Q); cocoon with large anteroventral projection thick, translucent (Fig. 37 O) ............... Simulium deagostinii Number of thoracic trichomes on each side 150-220; cocoon with large anteroventral projection thick, not translucent .......................................................... 15 Number of thoracic trichomes on each side 150 (Fig. 31 Y); trichome length 0.100.19 mm; cocoon maximum length at base 4.3 mm .......................... Simulium dureti Number of thoracic trichomes on each side 160-220; trichome length 0.24-0.35 mm; cocoon maximum length at base 5.2-6.0 mm ................. Simulium diamantinum One or both filaments of ventral primary branch of gill sharply diverging from remaining filaments, downward and backwardly directed (Figs. 42 U, X; 43 L, M; 46 F) .................................................................................................................................... 17

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Filaments of gill approximately parallel, or, if one or both filaments of ventral branch somewhat diverging, then not backwardly directed (Figs. 53 Q) ............. 19 All thoracic trichomes long, hair-like (similar to Figs. 43 O, P) or elongatelanceolate (Figs. 42 X, Y) ............................................................................................... 18 Thoracic trichomes either hair-like, simple or branched, and, most frequently, spatulate (Fig. 46 H) ........................................................................ Simulium quechuanum Both filaments of ventral primary branch of gill sharply diverging from remaining filaments (Fig. 42 U); 1+1 areas of tubercles at base of frontoclypeus (Fig. 42 V); trichomes of thorax elongate-lanceolate, subequal (Figs. 42 X, Y) ....................................................................................................................... Simulium herreri Generally only lower filament of primary ventral branch of gill diverging from remaining filaments (Fig. 43 M); no tubercles in area mentioned (Fig. 43 N); trichomes of thorax slender, hair-like, long and short (Figs. 43 O, P) ..................... ................................................................................................................. Simulium yacuchuspi Most thoracic trichomes roughly stellate (Figs. 53 S, T) ............. Simulium stelliferum Thoracic trichomes not stellate ..................................................................................... 20 Thoracic trichomes both long, hair-like, and shortly lanceolate (Figs. 40 U, V) .... ................................................................................................................. Simulium barbatipes All thoracic trichomes hair-like, simple or branched (Figs. 44 R; 45 D, F; 51 N-O) .............................................................................................................................................. 21 Practically all thoracic trichomes simple (Fig. 51 N) ................... Simulium annulatum Many thoracic trichomes branched (Figs. 44 R; 45 D-F) ........................................ 22 Thoracic trichomes (Fig. 44 R) simple or bifid, only rarely with more than 2 branches; their number 250-300 on each side ........................... Simulium albilineatum Thoracic trichomes (Figs. 45 D-F) practically never simple, generally with 2 or more branches, and their number 120-170 on each side ........................................ 23 Thoracic trichomes 5-6 on each side, very short and with 8 branches (Fig. 38 b); tubercles of frontoclypeus and thorax rugose ................................ Simulium cotopaxi Thoracic trichomes 5-170 on each side, relatively elongated with 1-6 branches; tubercles of frontoclypeus and thorax smooth ......................................................... 24 Thoracic trichomes of 60-80 on each side; trichomes mostly bifid with short branches (Figs. 41 M, N); frontoclypeus with about 1200 tubercles ........................ .................................................................................................................... Simulium philippii Thoracic trichomes of 120-170 on each side; trichomes mostly with 3 to 6 branches (Figs. 45 D-F); frontoclypeus without tubercles ................ Simulium bordai

Larvae Simulium (Pt.) albicinctum, Simulium (Pt.) bordai, Simulium (Pt.) luchoi, Simulium (Pt.) schoenemanni and Simulium (Pt.) strigidorsum are not included in this key because their larvae are not known. 1 –

Postgenal bridge extremely reduced, its length about 1/5 or less of depth of postgenal cleft (Figs. 29 V; 50 c) ..................................................................................... 2 Postgenal bridge much larger, about 1/2 of depth of postgenal cleft or larger (Figs. 42 e; 46 J; 51 T ) ...................................................................................................... 9

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Rectal papillae with 3 simple lobes, at most with 1 pair of secondary lobules .... 3 Rectal papillae with about 3 pairs of diverticula on each lobe ................................ 6 Rectal papillae with only 3 simple lobes (Figs. 30 b; 41 U) ...................................... 4 Rectal papillae with 1 secondary diverticulum close to base of lateral lobes, or with 2 pairs on middle lobe ............................................................................................. 5 4(3) Cephalic apotome very light colored, only slightly darkened at posterior margin (Fig. 34 J) .............................................................................. (some)......Simulium pulchrum – Cephalic apotome with overall light or medium brown pigment, also perceptibly darkened at hind border (Fig. 50 Y) ..................................... Simulium prodexargenteum 5(3) Rectal papillae with diverticula close to base of each of lateral lobe (Fig. 28 Y); first 2 antennal segments extremely light colored; hypostomial setae arranged in 1, rarely 2, irregular series, with 6-8 setae on each side (Fig. 28 V) .............................. ............................................................................................................ Simulium nigristrigatum – Rectal papillae with 2 diverticula on middle lobe; first 2 antennal segments distinctly pigmented (Fig. 33 W); hypostomial setae arranged in 1 to 3 rows, with about 14-18 setae on each side (Fig. 33 Y) ...................................... Simulium pulchrum 6(2) Maximum length about 6.7 mm; maximum width of cephalic capsule about 0.6 mm .......................................................................................................... Simulium limay – Maximum length greater than 7.5 mm, maximum width of cephalic capsule more than 0.7 mm ........................................................................................................................ 7 7(6) Maximum length 8.0 mm; maximum width of cephalic capsule 0.75 mm; postgenal cleft approximately as wide at base as deep .............................................. 8 – Maximum length 9.0 mm; maximum width of cephalic capsule 0.80 mm; postgenal cleft wider at base than deep ................................................. Simulium simile 8(7) Cephalic apotome light brown (Fig. 35 W); ratio of proximal, medial and distal antennal segments = 1:1.8-1.9:1.2-1.3 ........................................ Simulium diamantinum – Cephalic apotome dark brown (Fig. 37 T); ratio of proximal, medial and distal antennal segments = 1:1.3-1.5:0.8-1.0 ............................................ Simulium deagostinii 9(1) Lobes of rectal papillae each with 16-20 very long and slender diverticula (Fig. 43 X) ...................................................................................................... Simulium yacuchuspi – Lobes of rectal papillae with fewer or no diverticula .............................................. 10 10(9) Postgenal bridge about as long as postgenal cleft depth (Fig. 42 e) ..................... 11 – Postgenal bridge much shorter, about half as long as postgenal cleft depth (Figs. 46 J; 47 V; 53 a) ................................................................................................................ 12 11(10) Rectal papillae with 1+1 diverticula (Fig. 42 f); ratio of proximal, medial and distal antennal segments = 1:1.10-1.25:1.00-1.08 .......................................... Simulium herreri – Rectal papillae without diverticula (Fig. 41 U); ratio of proximal, medial and distal antennal segments = 1:1.0-1.2:.0.8 ..................................................... Simulium philippii 12(10) Lobes of rectal papillae simple, without diverticula ................................................. 13 – Lobes of rectal papillae with diverticula ..................................................................... 19 13(12) Base of anal sclerite with scale-like cuticular structures, from simple to trifid .................................................................................................................... Simulium nemorale – Base of anal sclerite generally without cuticular structures, or only with simple spicules ............................................................................................................................... 14

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14(13) Sclerotized area between anterior and posterior arms of anal sclerite extensive (Fig. 53 c); base of anal sclerite with simple spicules .............................................. 15 – Sclerotized area between anterior and posterior arms of anal sclerite comparatively narrow; base of anal sclerite lacking spicules .................................. 17 15(14) Anterior half of cephalic apotome conspicuously lighter than basal portion of sclerite (Figs. 53 V, W) ..................................................... (some)......Simulium stelliferum – Pigmentation of cephalic apotome not with strong contrasts ............................... 16 16(15) First marginal tooth of mandible about 4 times as long as second (Fig.46 I) .............................................................................................................. Simulium quechuanum – First marginal tooth of mandible about 2 times as long as second (Fig. 47 U) ..............................................................................................................Simulium hectorvargasi 17(14) Pattern of cephalic apotome positive; maximum size of mature larva 7.0 mm .......................................................................................................................... Simulium pichi – Pattern of cephalic apotome negative (Fig. 31 a); size of mature larva 8.0 mm or more .................................................................................................................................... 18 18(17) Distal antennal segment shorter than proximal (Fig. 31 b) .............. Simulium dureti – Distal antennal segment about as long as proximal (Fig. 32 J) .................................. ................................................................................................................ Simulium horcochuspi 19(12) Base of anal sclerite lacking spicules or scale-like cuticular structures ................ 20 – Base of anal sclerite with scale-like cuticular structures ......................................... 22 20(19) Hypostomium with anterior border depressed and median tooth below lateral teeth (Fig. 38 i); rectal papillae with 11-14 diverticula on each lobe (Fig. 38 n); medial antennal segment relatively elongated (Fig. 38 f), ratio of antennal segment lengths = 1:1.4-1.7:0.8 ........................................................................... Simulium cotopaxi – Hypostomium with anterior border not depressed and median tooth almost as prominent as lateral teeth; rectal papillae with 4-6 diverticula on each lobe; medial antennal segment not overly elongated, ratio of antennal segment lengths = 1:1.21.3:0.8-0.9 ........................................................................................................................... 21 21(20) Sclerotized area between anterior and posterior arms of anal sclerite comparatively small (similar to Fig. 32 K) ................................................. Simulium annulatum – Sclerotized area between anterior and posterior arms of anal sclerite extensive ............................................................................................... (some)......Simulium stelliferum 22(19) Anal ring with not more than 70 rows of hooks; distal antennal segment as long as proximal; pattern of cephalic apotome as shown in figure 44 S ....................... .............................................................................................................. Simulium albilineatum – Anal ring with 80 rows of hooks or more; distal antennal segment slightly but distinctly shorter than first; pattern of cephalic apotome different ..................... 23 23(22) Spicules at base of anal sclerite not numerous ........... (some)......Simulium stelliferum – Spicules at base of anal sclerite much more numerous (Fig. 40 b) ....................... 24 24(23) Pattern of cephalic apotome as shown in figure 40 W .............. Simulium barbatipes – Pattern of cephalic apotome difficult to perceive, as shown in figure 52 S ........... ......................................................................................................................... Simulium caprii

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CLAVES

DE ESPECIES DE

SIMULIUM (PTERNASPATHA)

Hembras Simulium (Pt.) bordai no se conoce la hembra y Simulium (Pt.) schoenemanni no tiene suficientes caracteres para incluirlo en la clave. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(6)

– 8(6) – 9(3)

Porción basal de R desnuda (Fig. 39 I) ......................................................................... 2 Porción basal de R con pelos (Fig. 51 D) ...................................... Simulium annulatum Abdomen (Fig. 36 C) gris plateado opaco; tergitos II-V cada uno con una mancha relativamente pequeña, central, negro aterciopelada y con manchas laterales oscuras ausentes; calcipala muy pequeña (Fig. 36 G) ................ Simulium strigidorsum Tergitos abdominales II-V con manchas oscuras más extensas; calcipala ausente o presente ................................................................................................................................ 3 Calcipala ausente (Figs. 32 A; 50 J) ................................................................................ 4 Calcipala presente, aunque en algunas especies muy pequeña ................................. 9 Tergitos IV y V sin 1+1 manchas blanco plateadas de coloración enteramente negra, aunque en algunos especimenes con angosta banda blanquecina a lo largo del borde posterior (Fig. 50 A) ............................................... Simulium prodexargenteum Tergitos IV y V con 1+1 manchas blancas plateadas (Fig. 28 A) ........................... 5 Uñas sin diente subbasal (Fig. 32 B) .............................................. Simulium horcochuspi Uñas con diente subbasal (Fig. 28 J) .............................................................................. 6 Tergitos abdominales III-V con 1+1 manchas laterales negruzcas uniéndose anteriormente con una mancha mediana (Figs.28 A; 29 B); largo del ala 2,33,5 mm ................................................................................................................................. 7 Tergitos abdominales III-V con 1+1 manchas laterales negruzcas pequeñas (Figs. 35 B; 37 B); largo del ala 3,3–3,7 mm ........................................................................... 8 Tergitos abdominales III-V con coloración gris claro extendiéndose al área lateral de los tergitos (Fig. 29 B); fémur anterior y mediana castaño claro, no conspicuamente oscurecidos apicalmente; longitud ala 2,8-3,5 mm ................... ......................................................................................................................... Simulium limay Tergitos abdominales III-V con coloración gris clara extendiéndose o no al área lateral de los tergitos (Fig. 28 A); fémures anterior y medio conspicuamente oscurecidos en el ápice; longitud del ala 2,3-2,7 mm ............. Simulium nigristrigatum Tergito abdominal VI con una mancha mediana negra (Fig. 37 B); tergitos III-V con 1+1 mancha negra lateral alcanzando al borde posterior; relación largo/ ancho del basitarso posterior = 7,2-7,6 .......................................... Simulium deagostinii Tergito abdominal VI sin una mancha mediana negra (Fig. 35 B); tergitos III-V con 1+1 manchas negras laterales no alcanzando el borde posterior; relación largo/ancho del basitarso posterior = 6,6 ................................. Simulium diamantinum Tergito III sin 1+1 manchas blanquecinas, con tergitos III-V enteramente oscuros, excepto en algunas especies con angosta línea a lo largo de bordes laterales del margen posterior (Fig. 51 A); línea medio longitudinal clara del escudo invisible o débilmente perceptible .................................................................. 10

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Tergito III con 1+1 mancha gris o blanquecina (Figs. 47 A; 48 A; 49 A; 50 A); difícil de observar en algunos especímenes pequeños pero en todos los casos perceptible; línea media longitudinal del escudo claramente visible ..................... 11 Bases de las tibias conspicuamente pigmentadas pero el ápice del basitarso posterior faltando pigmento; tergitos III-V enteramente oscuros (Fig. 52 A); paraproctos redondeados, salientes; cercos relativamente altos (Fig. 52 H) ........... ......................................................................................................................... Simulium caprii Bases de las tibias faltando evidente pigmento, con basitarso posterior bien pigmentado en el tercio apical (Fig. 53 H); borde posterior del tergito III-V angostamente pigmentado de gris (Fig. 53 A); paraproctos muy cortos y truncados; cercos relativamente bajos (Fig. 53 L) ......................... Simulium stelliferum Tergitos IV-V faltando las 1+1 manchas grises o blancas, solamente en algunos casos con margen posterior angostamente pigmentado con blanco lateralmente (Figs. 47 A; 50 A); tibia con nítido anillo oscuro basal o subbasal ....................... 12 Tergitos IV y V, en algunos casos solo tergito V con 1+1 manchas claras de variado tamaño (Figs. 33 A; 38 C; 40 A; 41 A); tibias con o sin anillo oscuro basal o subbasal .......................................................................................................................... 16 Calcipala no desarrollada (Fig. 50 J) .............. (algunos)......Simulium prodexargenteum Calcipala bien desarrollada (Figs. 39 K; 48 C) ........................................................... 13 Área oscura separando las manchas blancas del tergito abdominal II muy angosta (Fig. 48 A); tergito VI enteramente blanco plateado .................. Simulium albicinctum Área oscura separando las manchas blancas del tergito abdominal II ancha (Figs. 39 B; 47 A; 49 A), ensanchado anteriormente o tan ancho anterior como posteriormente; tergito VI oscuro en el centro y lateralmente .............................. 14 Cerdas y espinas sobre R1 dispuestas en varias hileras irregulares (Fig. 39 I) ........ .................................................................................................................... Simulium nemorale Cerdas y espinas sobre R1 dispuestas en una hilera simple (similar a figura 42 P) .............................................................................................................................................. 15 Calcipala muy pequeña, angostamente triangular (Fig. 49 E) ............Simulium luchoi Calcipala más grande, redondeada (Fig. 47 F) ...........................Simulium hectorvargasi Frente muy ancha (Fig. 38 D); paraprocto con ángulo ventral posterior alargado (Fig. 38 N); escudo negro, con variable ornamentación de acuerdo a la incidencia de la luz (Figs. 38 A-B) ......................................................................... Simulium cotopaxi Frente no ancha (Fig. 40 B); paraprocto con el ángulo ventral posterior redondeado (Fig. 40 L); escudo grisáceo, mostrando una banda mediana y 1+1 sublaterales negruzcas ..................................................................................................... 17 Tibia no conspicuamente oscurecida en la base (Figs. 40 F, G, H) ...................... 18 Tibia conspicuamente oscurecida en la base .............................................................. 23 Tergitos III-V con 1+1 manchas claras muy pequeñas, en algunos casos difíciles de percibir (Figs. 40 A; 41 A); longitud ala 2,9-3,9 mm .......................................... 19 Tergitos III-V con 1+1 manchas claras más grandes, bien distinguibles (Fig. 33 A); longitud ala 2,5-3,6 mm .................................................................................................. 20

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19(18) Manchas claras submedianas sobre tergitos III-V cruzando desde el borde anterior al posterior del segmento y de lados paralelos (Fig. 40 A); uñas con pequeño diente subbasal (Fig. 40 J) ................................................ Simulium barbatipes – Manchas claras submedianas sobre tergitos III-V no arribando al borde anterior y con base agrandada (Fig. 41 A); uñas sin diente subbasal (Fig. 41 F) ...................... .................................................................................................................... Simulium philippii 20(18) Calcipala bien desarrollada (Fig. 33 E) ........................................................................ 21 – Calcipala pequeña (Fig. 31 I) ......................................................................................... 22 21(20) Longitud ala 3,2-3,5 mm; tibia posterior con pigmento frecuentemente solo en el ápice ............................................................................................................... Simulium simile – Longitud ala 2,6-2,8 mm; tibia posterior con pigmento siempre más extendido, sobre los 2/5 del ápice del segmento (Fig. 34 D) .......................... Simulium pulchrum 22(20) Triángulo frontoocular más largo que ancho; longitud ala sobrepasando 3,0 mm; Sc con aproximadamente 10 pelos ......................................................... Simulium dureti – Triángulo frontoocular no más largo que ancho; longitud ala 3,0 mm o menos; Sc sin o con 1 ó 2 pelos cuando mas ............................................................ Simulium pichi 23(17) Cerdas y espinas sobre R1 y cerdas sobre Rs dispuestas en varias hileras irregulares (Fig. 39 I); manchas blancas sobre tergitos IV y V débiles angostamente alargadas (Fig. 39 C) ................................................................................................ Simulium nemorale – Cerdas y espinas sobre R1 y cerdas sobre Rs dispuestas en una hilera como en el macho (Fig. 42 P); manchas blancas sobre tergitos IV y V notorias redondeadas o rectangulares (Fig. 42 A) ................................................................................................. 24 24(23) Tergito V con 1+1 manchas blancas tan grandes como las del tergito III (Fig. 44 A) ......................................................................................................... Simulium albilineatum – Tergito V con 1+1 manchas blancas notoriamente más pequeñas que aquellas del tergito III (Figs. 42 A; 46 A) .......................................................................................... 25 25(24) Diente de las uñas excepcionalmente pequeño (Fig. 43 F); área central del octavo esternito más clara que las porciones laterales; paraprocto relativamente aguzado (Fig. 43 G) ............................................................................................. Simulium yacuchuspi – Diente de las uñas normalmente desarrollado (Fig. 46 A); área central del octavo esternito más oscura que las porciones laterales; paraprocto anchamente redondeado ........................................................................................................................ 26 26(25) Tergito III del abdomen (Fig. 42 A) con 1+1 manchas blanco plateadas solo sobrepasando ligeramente la mitad anterior del tergito; tergito IV con 1+1 manchas blanquecinas pequeñas; área central blanquecina del tergito VI no dividida longitudinalmente por una banda negra ............................... Simulium herreri – Tergito III del abdomen (Fig. 46 A) con 1+1 manchas blanquecinas, llegando muy cerca del borde anterior del tergito; tergito IV sin manchas; área blanquecina del tergito VI dividido longitudinalmente por una banda oscura Simulium quechuanum Machos Los machos de Simulium (Pt.) bordai, Simulium (Pt.)schoenemanni y Simulium (Pt.) strigidorsum no se conocen y Simulium horcochuspi no posee caracteres suficientes para incluirlo en la clave. Si se conociera mejor Simulium horcochuspi saldría en la clave como Simulium (Pt.) deagostinii.

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1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(4) – 7(6) – 8(3) – 9(8) – 10(9) – 11(10) – 12(11) – 13(12) – 14(10)

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Porción basal de R sin pelos (Fig. 39 I) ........................................................................ 2 Porción basal de R con pelos (Fig. 51 D) ...................................... Simulium annulatum Escapo, pedicelo y palpómeros basales con pelos muy largos (Figs. 38 Q, R); escapo y pedicelo combinados son tan largos como el flagelo ......................... .................................................................................................................... Simulium cotopaxi Escapo, pedicelo y palpómeros basales con pelos cortos; escapo y pedicelo combinados son 1/3 del largo del flagelo .................................................................... 3 Calcipala ausente (Figs. 50 P-R) ...................................................................................... 4 Calcipala presente, aunque en algunas especies muy pequeña ................................. 8 Tibia posterior conspicuamente oscurecida en la base (Figs. 37 L; 50 P-Q) ........ 5 Tibia posterior no conspicuamente oscurecida en la base ........................................ 6 Abdomen con 1+1 manchas plateadas en tergitos II-VII (Fig. 50 L); gonostilo con superficie apical lisa (similar a la figura 42 S) ..................... Simulium prodexargenteum Abdomen con 1+1 manchas plateadas solamente en tergitos II, VI y VII; superficie distal granulosa (similar a figura 37 M) ....................... Simulium deagostinii Basitarso posterior 5,0 veces tan largo como ancho ........................... Simulium limay Basitarso posterior 3,3-3,8 veces tan largo como ancho ........................................... 7 Tergitos abdominales III-V con 1+1 manchas blanco plateadas no llegando al borde anterior; largo ala 2,1-2,5; relación largo /ancho de basitarso posterior = 3,3-3,5 ............................................................................................... Simulium nigristrigatum Tergitos abdominales III-V con 1+1 manchas blanco plateadas llegando al borde anterior; largo ala 2,8-3,0 mm; relación largo/ancho del basitarso posterior = 3,0 .............................................................................................................Simulium diamantinum Superficie apical del gonostilo liso (Fig. 47 K) ............................................................ 9 Superficie apical del gonostilo con tubérculos (Fig. 28 N) ..................................... 20 Base de tibia posterior notoriamente oscurecida (Figs. 47 J; 49 L; 50 P-Q; 53 N) .............................................................................................................................................. 10 Base de tibia posterior no notoriamente oscurecida (Fig. 40 O) ......................... ................................................................................................................. Simulium barbatipes Pelos y espinas de R1 y Rs ordenados en 1 o 2 hileras irregulares (como en la hembra, Fig. 39 I); basitarso posterior angosto, alrededor de 4,0 veces tan largo como ancho ....................................................................................................................... 11 Pelos y espinas sobre R1 y Rs dispuestos en 1 hilera simple (Fig. 42 P ); basitarso posterior más ancho, no mayor de 3,5 veces más largo que ancho ...................... 14 Largo del ala > 3,5 mm ........................................................................ Simulium nemorale Largo del ala < 3,5 mm .................................................................................................. 12 Largo del ala 2,7-2,9 mm; basitarso posterior angosto (4,3-5,5 veces tan largo como ancho) (Fig. 53 N) .................................................................... Simulium stelliferum Largo del ala 3,1-3,3 mm; basitarso posterior más ancho (3,5-4,0 veces tan largo como ancho ) .................................................................................................................... 13 Basitarso posterior 3,5 veces tan largo como ancho ...................... Simulium philippii Basitarso posterior 3,8-4,0 veces tan largo como ancho .................... Simulium caprii Tergitos abdominales III-V enteramente negro, o III con 1+1 manchas pequeñas claras (Fig. 47 I) ................................................................................................................ 15

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– 15(14) – 16(15) – 17(15) – 18(14) – 19(18) – 20(8) – 21(20) – 22(20) –

Tergitos abdominales III-V con 1+1 manchas blancas plateadas (Figs. 42 M; 43 H; 50 L) .................................................................................................................................... 18 Fémur posterior con 3/5 basales blanquecinos, contrastando con el ápice oscuro (Fig. 49 L); basitarso posterior < 3,0 veces largo como ancho (Fig. 49 L) ......... 16 Fémur posterior con 3/4 basales castaño claro (Fig. 47 J); basitarso posterior por lo menos 3.0 veces tan largo como ancho (Fig. 47 J) .............................................. 17 Calcipala muy pequeña, subtriangular (Fig. 49 M) ...............................Simulium luchoi Calcipala algo más grande, redondeada (Fig. 48 G) ................... Simulium albicinctum Calcipala muy pequeña (Fig. 46 C) ............................................... Simulium quechuanum Calcipala más grande (Fig. 47 F) ..................................................Simulium hectorvargasi Pelos del abdomen plateados; tergito III uniformemente oscuro, pero el V con 1+1 mancha blanca (Fig. 42 M); mancha oscura del fémur anterior y mediano corto ............................................................................................................. Simulium herreri Pelos del abdomen bronceado claro; tergito III con 1+1 mancha blanca plateada pequeña en borde posterior, aproximadamente la mitad de alto del segmento; manchas en la base del fémur medio y anterior angostamente alargada ............. 19 Margen posterolateral de tergitos abdominales IV y V angostamente bordeado de plateado o blanco (Fig. 50 L) .................................................. Simulium prodexargenteum Margen posterolateral de tergitos abdominales IV y V sin áreas plateadas o blanquecinas (Figs. 43 H; 44 I), excepto raramente sobre el V ................................. ........................................................................... Simulium albilineatum, Simulium yacuchuspi Calcipala pequeña (similar figura 30 P) ....................................................................... 21 Calcipala bien desarrollada (similar figura 33 E) ....................................................... 22 Longitud ala 3,0 mm; basitarso posterior ancho 3,2-3,4 veces tan largo como ancho; bajo ciertos ángulos de iluminación, por lo menos el 1/3 anterior del escudo grisáceo ........................................................................................... Simulium dureti Longitud del ala 2,3-2,6 mm; basitarso posterior más angosto, 3,8 veces tan largo como ancho; solo alrededor de 1/5 de la porción anterior gris ........ Simulium pichi Largo del ala 2,8-3,0 mm; abdomen con 1+1 manchas claras pequeñas pero diferenciables sobre tergito III (Fig. 33 J K) ......................................... Simulium simile Largo del ala 2,6-2,7 mm; tergito III sin ninguna mancha blanquecina .................. ................................................................................................................... Simulium pulchrum

Pupas En Simulium (Pt.) albicinctum, Simulium (Pt.) schoenemanni y Simulium (Pt.) strigidorsum no se conoce la pupa. 1 – 2(1) –

Branquias con 8 filamentos .............................................................................................. 2 Branquias con 6 filamentos ............................................................................................ 16 El más bajo o los 2 filamentos de la rama ventral primaria de la branquia divergiendo de los restantes filamentos, disponiéndose muy cerca al cuerpo de la pupa (Figs. 42 X, U; 46 F; 49 O; 50 U) ........................................................................ 3 Todos los filamentos de la rama ventral primaria dispuestos aproximadamente paralelos, por lo menos en la mitad apical .................................................................... 5

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Mayoría de tricomas de cabeza y tórax ligeramente lanceolados, generalmente bífidos (Figs. 49 Q-S) .................................................................................Simulium luchoi – Tricomas de cabeza y tórax no como arriba ................................................................ 4 4(3) Tricomas de la cabeza lanceolados alargados, en número de alrededor de 250+250 (Figs. 42 X, Y) ..................................................................... (algunos)......Simulium herreri – Tricomas del tórax simples o bífidos (Fig. 50 W), en número de alrededor de 12+12 ........................................................................................... Simulium prodexargenteum 5(2) Tricomas de la cabeza y tórax muy cortos, algunos ramificados pero la mayoría simples, desde forma de pelo hasta de espina, presentando los del tórax concentrados en una hilera transversa en la base de la porción expuesta (Fig. 39 X) .................................................................................................................... Simulium nemorale – Tricomas de la cabeza y tórax más largos con una disposición diferente a la de arriba ..................................................................................................................................... 6 6(5) Tricomas de la cabeza y tórax numerosos, achatados, con ramas acintadas (Figs. 33 R-T; 34 I) ............................................................................................................................. 7 – Tricomas de la cabeza y tórax no como arriba ............................................................ 8 7(6) Largo del capullo en la base 5,2-6,8 mm; frontoclípeo (Fig. 33 R) con alrededor de 400 tubérculos, con los tubérculos cefálicos no sobrepasando el área de los tricomas faciales, presentando esta región reforzada por un borde esclerotizado; tricomas torácicos no sobrepasando 250+250, con un promedio de 140+140 ......................................................................................................................... Simulium simile – Largo del capullo en la base 3,8-4,5 mm; frontoclípeo (Fig. 34 I) con alrededor de 800 tubérculos; tubérculos cefálicos sobrepasando el área de tricomas faciales, presentando esta región lisa, no reforzada por bordes esclerotizados; tricomas torácicos no sobrepasando 130+130, con un promedio de 70+70 .......................... ................................................................................................................... Simulium pulchrum 8(6) Disco de porción expuesta del tórax sin tubérculos o con tubérculos solo a lo largo del área adyacente al borde de la abertura del capullo (Figs. 39 X; 47 Q) ............... 9 – Disco de porción expuesta del tórax con numerosos tubérculos (Fig. 30 V) ........ 11 9(8) Tricomas frontales reducidos a pequeños conos (Fig. 47 O); tórax con alrededor de 150+150 tricomas (Fig. 47 Q) .................................................Simulium hectorvargasi – Tricomas frontales normalmente desarrollados, en forma de pelo; tórax con alrededor de 100+100 tricomas .................................................................................... 10 10(9) Tórax con alrededor de 70+70 tricomas (Figs. 52 Q-R), estos tricomas desde simples a frecuentemente con 2 o 3 ramas ........................................... Simulium caprii – Tórax no con más de 20+20 tricomas, estos generalmente con 4 o más ramas (Fig. 28 T) .................................................................................................. Simulium nigristrigatum 11(8) Tricomas del tórax faltando en porción anterior del esclerito ............................... 12 – Tricomas del tórax cubriendo prácticamente toda la superficie de la porción expuesta del tórax (similar a figura 45 D) .................................................................. 13 12(11) Tórax con alrededor de 20+20 tricomas (Fig. 29 N); tubérculos del frontoclípeo esparcidos, su superficie espinosa, tubérculos del frontoclípeo del macho solamente presentes en la base (Fig. 29 Q) ........................................... Simulium limay

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Tórax con alrededor de 60+60 tricomas (Fig. 30 V); tubérculos del frontoclípeo más numerosos, su superficie lisa; tubérculos del clípeo del macho cubriéndolo en su totalidad (Fig. 30 U) ............................................................................... Simulium pichi Capullo con brillo plomizo; superficie de los tubérculos lisa; tubérculos frontales (Figs. 32 F, H) dispuestos en grupos de 3 o 4, bien separados de los tricomas faciales los cuales son simples; tricomas frontales (Fig. 32 H) simples o en grupos de 2; tricomas del tórax frecuentemente con 1 a 3 ramas (Fig. 32 I) ....................... ................................................................................................................ Simulium horcochuspi Capullo sin brillo plomizo; tubérculos del frontoclípeo con superficie más o menos tuberculada; tricomas frontales (Fig. 31 X) más numerosos, contiguos, con los tricomas faciales en número mayor de 1; tricomas oculares (Figs. 31 X; 37 P) en número superior a 2; tricomas del tórax muy frecuentemente con más de 3 ramas (Fig. 31 Z) .............................................................................................................. 14 Con 35-80 tricomas torácicos de cada lado (Fig. 37 Q); prolongación anteroventral del capullo gruesa y translúcida (Fig. 37 O) .................... Simulium deagostinii Con 150-220 tricomas torácicos de cada lado; prolongación antero-ventral del capullo gruesa, no translúcida ....................................................................................... 15 Con alrededor de 150 tricomas torácicos de cada lado (Fig. 31 Y); longitud de tricomas 0,10-0,19 mm; largo máximo del capullo en la base 4,3 mm ................ Simulium dureti Con 180-220 tricomas torácicos de cada lado; longitud de tricomas 0,24-0,35 mm; largo máximo del capullo en la base 5,2-6,0 mm ..................... Simulium diamantinum Uno o ambos filamentos de la rama primaria ventral de la branquias divergiendo fuertemente de los restantes filamentos, dirigidos hacia abajo y atrás (Figs. 42 U, X, 43 L, M; 46 F) ............................................................................................................. 17 Filamentos de branquias aproximadamente paralelos, o si 1o ambos filamentos de la rama primaria ventral son algo divergentes, no están dirigidos hacia atrás (Fig. 53 Q) ................................................................................................................................... 19 Todos los tricomas torácicos largos, en forma de pelo (similar Fig. 43 O-P) ó alargado y lanceolado (Figs. 42 X-Y) ........................................................................... 18 Tricomas torácicos en forma de pelo, simple o ramificado o más frecuentemente espatulados (Fig. 46 H) ................................................................... Simulium quechuanum Ambos filamentos de la rama primaria ventral de la branquias fuertemente divergentes de los restantes filamentos (Fig. 42 U); presencia de 1+1 áreas de tubérculos en la base del frontoclípeo (Fig. 42 V); tricomas del tórax alargados, lanceolados, subiguales (Figs. 42 X, Y) ................................................. Simulium herreri Generalmente solo el filamento más bajo de la rama primaria ventral divergiendo de los restantes filamentos (Fig. 43 M); sin tubérculos en el área mencionada (Fig. 43 N); tricomas del tórax alargados, en forma de pelo, largos y cortos (Figs. 43 O, P) ................................................................................................................. Simulium yacuchuspi Mayoría de tricomas torácicos de forma aproximadamente estrellada (Figs. 53 S, T) .................................................................................................................. Simulium stelliferum Tricomas cefálicos no de forma estrellada ................................................................. 20 Triocomas torácicos desde alargados en forma de pelo o más cortos lanceolados (Figs. 40 U, V) ...................................................................................... Simulium barbatipes

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Tricomas torácicos en forma de pelo simple o ramificado (Figs. 44 R; 45 D, F; 51 N-O) ................................................................................................................................... 21 Prácticamente todos los tricomas torácicos simples (Fig. 51 N) ............................... .................................................................................................................Simulium annulatum Muchos tricomas torácicos ramificados (Figs. 44 R; 45 D-F) ................................ 22 Tricomas torácicos (Fig. 44 R) simples o bífidos, raramente con más de 2 ramas, en número de 250-300 de cada lado ............................................ Simulium albilineatum Tricomas torácicos (Figs. 45 D-F) prácticamente nunca simples, generalmente con 2 o más ramas y en número de 120-170 de cada lado ............................................. 23 Tricomas torácicos en número de 5-6 de cada lado, muy cortos y con hasta 8 ramas (Fig. 38 b); tubérculos de frontoclípeo y tórax rugoso ...... Simulium cotopaxi Tricomas torácicos de 5-170 de cada lado, relativamente largos con 1-6 ramas; tubérculos del frontoclípeo y tórax lisos ..................................................................... 24 Tricomas torácicos en número de 60-80 de cada lado; tricomas mayormente bífidos, con ramas cortas (Figs. 41 M, N); frontoclípeo con alrededor de 1200 tubérculos ................................................................................................ Simulium philippii Tricomas torácicos en número de 120-170 de cada lado; tricomas mayormente con 3 a 6 ramas (Figs. 45 D-F); frontoclípeo sin tubérculos ............ Simulium bordai

Larvas En Simulium (Pt.) albicinctum, Simulium (Pt.) bordai, Simulium (Pt.) luchoi, Simulium (Pt.) schoenemani y Simulium (Pt.) strigidorsum sus larvas son desconocidas. 1 – 2(1) – 3(2) – 4(3) – 5(3)

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Puente postgenal extremadamente reducido, su longitud aproximadamente 1/5 o menos de la profundidad de la hendidura postgenal (Figs. 29 V; 50 c) ................. 2 Puente postgenal más largo, aproximadamente la mitad de la profundidad de la hendidura postgenal (Figs. 42 e; 46 J; 51 T) ................................................................. 9 Papilas rectales con 3 lóbulos simples, cuando mucho con 1 par de divertículos ................................................................................................................................................ 3 Papilas rectales con 3 pares de divertículos en cada lóbulo ...................................... 6 Papilas rectales con solo 3 lóbulos simples (Figs. 30 b; 41 U) ................................. 4 Papilas rectales con 1 divertículo próximo a la base de los lóbulos laterales o con 2 pares sobre el lóbulo mediano ..................................................................................... 5 Apotoma cefálico con coloración clara, solo ligeramente más oscuro sobre el margen posterior (Fig. 34 J) ......................................... (algunos)......Simulium pulchrum Apotoma cefálico todo claro o castaño en el medio y también oscurecido en el borde posterior (Fig., 50 Y) .................................................... Simulium prodexargenteum Papilas rectales con un divertículo próximo a la base de los lóbulos laterales (Fig. 28 Y); primeros 2 artejos antenales extremadamente claros; cerdas del hipostomio ordenadas en 1, raramente 2 series irregulares, con 6-8 cerdas de cada lado (Fig. 28 V) .................................................................... Simulium nigristrigatum Papilas rectales con 2 divertículos sobre el lóbulo mediano; primeros 2 artejos antenales notoriamente pigmentados (Fig. 33 W); cerdas del hipostomio dispuestas desde 1 a 3 hileras, con alrededor de 14-18 cerdas de cada lado (Fig. 33 Y) ......... Simulium pulchrum

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Longitud máxima alrededor de 6,7 mm; ancho máximo de la cápsula cefálica alrededor de 0,6 mm .................................................................................. Simulium limay Longitud máxima superior a 7,5 mm; ancho máximo de la cápsula cefálica mayor de 0,7 mm ............................................................................................................................ 7 Longitud máxima 8,0 mm; ancho máximo de la cápsula cefálica 0,75 mm; hendidura postgenal aproximadamente tan profunda como ancha en la base ..... 8 Longitud máxima 9,0 mm; ancho máximo de la cápsula cefálica 0,80 mm; hendidura postgenal menos profunda que ancha en la base ............. Simulium simile Apotoma cefálico castaño claro (Fig. 35 W); relación de los artejos antenales proximal, mediano y distal = 1:1,8-1,9:1,2-1,3 .......................... Simulium diamantinum Apotoma cefálico castaño oscuro (Fig. 37 T); relación de los artejos antenales proximal, mediano y distal = 1:1,3-1,5:0,8-1.0 ............................... Simulium deagostini Lóbulos de las papilas rectales cada uno con 16-20 divertículos largos y delgados (Fig. 43 X) ............................................................................................. Simulium yacuchuspi Lóbulos de las papilas rectales con pocos o sin divertículos .................................. 10 Puente postgenal aproximadamente tan largo como el de la hendidura postgenal (Fig. 42 e) ........................................................................................................................... 11 Puente postgenal más corto, aproximadamente la mitad del largo de la hendidura postgenal (Figs. 46 J; 47 V; 53a ) ................................................................................... 12 Papilas rectales con 1+1 divertículo sobre cada lóbulo (Fig. 42 f); relación de los artejos antenales proximal, mediano y distal = 1:1,10-1,25:1.00-1,08 ...................... ....................................................................................................................... Simulium herreri Papilas rectales sin divertículos (Fig. 41 U); relación de los artejos antenales proximal, mediano y distal = 1:1,0-1,2:0,8........................................ Simulium philippii Lóbulos de papilas rectales simples, sin divertículos ................................................ 13 Lóbulos de papilas rectales con divertículos .............................................................. 19 Base del esclerito anal con estructuras cuticulares como escamas desde simples a trífidas ....................................................................................................... Simulium nemorale Base del esclerito anal generalmente sin estructuras cuticulares como escamas o solamente como espículas simples ............................................................................... 14 Área esclerotizada entre ramas anteriores y posteriores del esclerito anal extensa (Fig. 53 c); base del esclerito anal con espículas simples ......................................... 15 Área esclerotizada entre ramas anteriores y posteriores del esclerito anal comparativamente angosta; base del esclerito anal sin espículas ........................... 17 Mitad anterior del apotoma cefálico conspicuamente más claro que la porción basal del esclerito (Figs. 53 V, W) .............................. (algunos)......Simulium stelliferum Pigmentación del apotoma cefálico sin fuertes contrastes ..................................... 16 Primer diente marginal de la mandíbula aproximadamente 4 veces más largo que el segundo (Fig. 46 I) ....................................................................... Simulium quechuanum Primer diente marginal aproximadamente 2 veces más largo que el segundo (Fig. 47 U) ...................................................................................................Simulium hectorvargasi Manchas del apotoma cefálico positivas; tamaño máximo de la larva 7,0 mm .......................................................................................................................... Simulium pichi

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Manchas del apotoma cefálico negativas (Fig. 31 a); tamaño de la larva madura 8,0 mm o más .......................................................................................................................... 18 Artejo distal de antena más corto que el proximal (Fig. 31 b) ......... Simulium dureti Artejo distal de antena aproximadamente tan largo como el proximal (Fig. 32 J) ................................................................................................................ Simulium horcochuspi Base del esclerito anal sin espículas o estructuras cuticulares como espinas ...... 20 Base del esclerito con estructuras cuticulares como escamas ................................ 22 Hipostomio con borde anterior deprimido y con diente mediano por debajo de los dientes laterales (Fig. 38 i); papilas rectales con 11-14 divertículos en cada lóbulo (Fig. 38 n); artejo antenal mediano relativamente alargado (Fig. 38 f); relación del largo de los artejos antenales proximal, mediano y distal = 1:1, 1.41,7:0,8 ........................................................................................................ Simulium cotopaxi Hipostomio con el borde anterior no deprimido y diente mediano casi tan prominentes como los dientes laterales; papilas rectales con 4-6 divertículos sobre cada lóbulo; artejo mediano de la antena no muy alargado, relación de los artejos antenales = 1:1,2-1,3:0,8-0,9 .......................................................................................... 21 Área esclerotizada entre las ramas anterior y posterior del esclerito anal comparativamente pequeña (similar a figura 32 K) ..................... Simulium annulatum Área esclerotizada entre las ramas anterior y posterior del esclerito anal comparativamente grande ........................................... (algunos)......Simulium stelliferum Anillo anal con no más de 70 hileras de ganchos; artejo distal de la antena tan largo como el proximal; ornamentación del apotoma cefálico como se muestra en figura 44 S .......................................................................................... Simulium albilineatum Anillo anal con 80 hileras de ganchos o más; artejo distal de la antena ligeramente más corto que el proximal; ornamentación del apotoma cefálico diferente ....... 23 Espículas en la base del esclerito anal no numerosos .................................................. ........................................................................................... (algunos)......Simulium stelliferum Espículas en la base del esclerito anal numerosos (Fig. 40 b) ................................ 24 Ornamentación del apotoma cefálico como se muestra en figura 40 W ................. ................................................................................................................. Simulium barbatipes Ornamentación del apotoma cefálico poco intensa, como en figura 52 S .............. ......................................................................................................................... Simulium caprii

NIGRISTRIGATUM species group Simulium (Pternaspatha) nigristrigatum (ENDERLEIN) (Fig. 28.) Pternaspatha nigristrigata ENDERLEIN, 1930: 88;1934b: 275; STONE, 1962: 208. Simulium nigristrigatum SMART, 1945: 509; VARGAS, 1945a: 170; STUARDO ORTIZ, 1946: 41; WYGODZINSKY, 1949: 304; VARGAS & DIAZ, 1953: 142; WYGODZINSKY, 1958: 132. Simulium (Simulium) nigristrigatum EDWARDS, 1931: 154; ORFILA, 1939: 1533. [Simulium] (Pternaspatha) nigristrigata STONE, 1963: 17. Simulium (Pternaspatha) nigristrigatum WYGODZINSKY & COSCARÓN, 1967: 69-70; COSCARÓN, 1991: 113. Acropogon edwardsi ENDERLEIN, 1934b: 279; VARGAS, 1945: 134.

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Simulium edwardsi STUARDO ORTIZ, 1946: 41. Simulium (Pternaspatha) bachmanni WYGODZINSKY & COSCARÓN, 1967: 58; COSCARÓN & WYGODZINSKY, 1972b: 209; COSCARÓN, 1991: 110; COSCARÓN & COSCARÓN ARIAS, 1996a: 703.

Female: Wing length 2.5-2.6 mm; head black; antennae, palpi, and labrum dark gray; occiput, frons, and clypeus dark gray pollinose; pilosity of antennae very short, that of palpi, clypeus, frons, and occiput somewhat longer, silver-gray; first and second antennal segments much lighter than remaining segments, brownish; scutum dark gray, central dark band divided by distinct, longitudinal, gray line; hair of thorax silver-gray; appressed setae of scutum very dense, somewhat obliterating pattern; scutellum, metanotum, pleura, and sterna dark gray; wings hyaline, veins light gray, hair and spines blackish, tufts at base of wing silver-gray; stem of halteres tinged with brown; knob white; legs light yellow; hair silvery gray to black; following regions darkened: all coxae, those of fore legs as intensely as others; trochanters; femora, their apices very strongly so on all pairs; apical three-fifths of mid and hind tibiae, fore tibiae almost entirely; entire fore tarsi; apical third of basal, and apical half of second, segment of mid and hind tarsi, and their entire third to fifth segments; exact proportions and relative intensity of pigmented areas as shown in figure F-H; abdomen (Fig. A) matte silver-gray; first tergite darkened; tergites II-V each with one large central and 1+1 lateral black spots connected by black line along anterior border of tergites; these spots narrowed posteriorly; distance between central and lateral spots about as large as width of central spot; tergite VI only very slightly darkened at middle anteriorly; tergite VII with one very large, black, central spot; tergites VIII and IX gray, VIII faintly darkened anteriorly at center; hair of abdomen from black to silver-gray; antenna as in figure B; distal palpomere as long as two anterior united (Fig. C); sensory vesicle of palp as in figure D; fronto-ocular triangle wider than high (Fig. E); wing: Sc with two or three hairs on its basal and three or four hairs on its central portion; basal portion of R glabrous; R1 with hair and setae arranged in one row, but occasional setae or spines out of line; Rs with one somewhat irregular row of setae, in some specimens a few out of line; shape and proportion of segments of legs as shown in figure F-H; calcipala not developed (Fig. I); claws with well-developed tooth (Fig. J); eighth abdominal sternite as shown in figure L; rather faintly pigmented, its central portion lighter than lateral areas; posterior border of central portion slightly sclerotized; paraprocts short, their free extremity truncate; genital fork as shown in figure K; median process distinctly sclerotized, including its terminal knob; spermatheca oval, its inner surface with minute spiculae, either single or arranged in groups of two. Male: Length of wing, 2.5 mm; head black; eyes deep purple; antennae and palpi piceous; clypeus gray pollinose; antennae with short pubescence only; setae of basal segments not distinctly longer than remainder; occiput, clypeus, and basal segment of maxillary palp with some long hair; isolated and rather short setae between eyes; scutum with usual color pattern; basic color from dark matte gray to velvety black, depending on the angle of view; appressed setae very light brass-colored; scutellum blackish, silvery gray pollinose; metanotum blackish; pleura blackish, silvery gray pollinose; pleural tuft silvery white; color of wings and halteres as in female; legs pale yellow to whitish, their hair from silvery white to black; following regions dark: all coxae and trochanters; base and apex of fore and mid femora, and apex of hind femur strongly pigmented, rest of femora slightly

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A

D

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C

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H

K

M N

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P Q

R T

S

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U

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Y Figure 28. Simulium (Pternaspatha) nigristrigatum. A-L: Female; A: abdomen dorsal view in different specimens; B: antenna; C: maxillary palp; D: sensory vesicle of maxillary palp; E: fronto-ocular triangle; F-H: fore, mid and hind leg, respectively; I: distal portion of hind basitarsus and pedisulcus; J: claw; K: genital fork; L: portion of eighth sternite and gonapophyses. M-P: Male; M: hind basitarsus; N: gonostylus; O: ventral plate; P: endoparameres and median sclerite. Q-T: Pupa; Q: general aspect; R: gill; S: frontoclypeus, with trichomes with high magnification at right; T: portion of thorax with trichomes. U-Y : Larva; U: teeth of mandible; V: hypostomium; W: hypostomium and postgenal cleft; X: anal sclerite; Y: rectal papillae.

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pigmented; fore and mid tibiae with apical fourth intensely pigmented, and with dark stripe running toward base, becoming evanescent in basal third; hind tibia intensely pigmented on apical third, faintly more basad, very faintly near base; entire fore tarsus, mid tarsus on apical half of basitarsus, and entire second through fifth segments; hind tarsus on apical third of basitarsus, apical half of second, and entire third through fifth segments; abdomen velvety black, its hair silvery white, including that of basal fringe; tergites II, VI, and VII with 1+1 silvery gray spots, those of segments II and VI very large, separated dorsally only by small black area on II, virtually confluent on VI; those of segment VII much smaller; last segment of maxillary palp more than twice as long as penultimate; diameter of sensory vesicle smaller than half diameter of third segment; chaetotaxy of wings like that of female; femur of forelegs with large number of long hairs; hind basitarsus 3.3-3.4 times as long as wide; calcipala not developed (Fig. M); gonocoxite wider than long; gonostylus subquadrate, distal surface heavily granulose (Fig. N); central portion of ventral plate somewhat salient; lateral projection and basal margin distinctly sclerotized, disc uniformly pigmented (Fig. O); bifid portion of median sclerite narrow and endoparameres with hard teeth (Fig. P). Pupa: Cocoon (Fig. Q) slipper shaped, its anteroventral bridge very short; color stramineous, surface smooth, closely woven, threads not distinct under medium magnification (X 60); rim of aperture not perceptibly reinforced; length of cocoon at dorsum along middle, 2.3-2.7 mm.; maximum length along base, 3.8 mm; length of pupal body 2.8-3.0 mm; length of gills, approximately 1.5 mm, viz., less than half as long as cocoon, and about half as long as pupa; gills each with eight filaments, forming a compact group (Fig. R); head and thorax of pupa light brown; head with 2+2 frontal and 1+1 facial trichomes (Fig. S); trichomes simple or with two or three branches; thorax with exposed portion without tubercles and not more strongly sclerotized than rest; region adjacent to aperture of cocoon with tubercles only near dorsal midline; disc of thorax (Fig. T) with about 10+10 long, simple or branched, hair like trichomes, latter with not more than four branches; length of trichomes, 0.10-0.16 mm. Larva: Maximum length, 5.2 mm; maximum width of head, 0.5 mm; general color whitish, with transverse bands of hypodermal pigment on first three or four abdominal segments dorsally; entire posterior portion of body pigmented; ventral surface pigmented only on anterior portion of body; head ivory-colored; cephalic apotome without perceptible pattern, only slightly darkened at posterior margin; body integument glabrous; antennae very pale, only distal segment slightly pigmented; ratio of proximal, medial and distal segments = 1:1.3:0.9; cephalic fan with 30-35 rays; toothing of mandibles as shown in figure U; median tooth of hypostomium about as prominent as lateral ones; serrations of lateral border of hypostomium exceptionally prominent (Fig. V); hypostomial setae arranged in one, rarely in two, irregular series, numbering about six in each group, in addition to two to four short, posterior setae; disc of hypostomium with one or two short, single, bifid hairs; postgenal cleft narrow and exceptionally deep, its apex almost attaining hypostomial groove (Fig. W); anal sclerite as shown in figure X, its base with numerous minute scales with from one to four rays; anal ring with approximately 80 rows of 15-17 hooks each; rectal papillae consisting of three primary lobes; two lateral ones close to their base, each with one secondary lobule (Fig. Y).

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Distribution: Argentina: Buenos Aires, Chubut, Neuquen, Rio Negro. Bionomics: Simulium nigristrigatum breeds in Patagonian rivers with fast flow over trees branches and leaves, especially of Salix humboldtiana, Cortaderia sp., or over stones. Females are slightly anthropophilic. Discussion: COSCARÓN and COSCARÓN ARIAS (1996: 709) established that Simulium bachmanni WYGODZINSKY & COSCARÓN is a synonym of Simulium nigristrigatum, based on variation of the female abdominal coloration of Simulium bachmanni. Simulium bachmanni is very well known in all stages, where the pupa provides the best characters to differentiate this species. In the topotypical area of Simulium nigristrigatum (known only from adults), pupae and larvae were found on several occasions and breeding males and females of Simulium bachmanni were also obtained. These are similar to the Simulium nigristrigatum female syntype and male lectotype from Zoologisches Museum of Humboldt University, Berlin. Simulium nigristrigatum is closely related to Simulium limay. The male of Simulium nigristrigatum differs from that of Simulium limay by its much wider basitarsus (at least four times as long as wide). The female of Simulium nigristrigatum with the apically truncate paraprocts are similar in shape to those of Simulium limay, but in the latter species tergites VIII and IX of the female are almost completely black, and neither the fore nor the mid femur is totally darkened. The pupa of Simulium nigristrigatum differs from that of Simulium limay by the absence of granulosity from the thoracic disc, by the smaller number of thoracic trichomes and the very long ventral primary branch of the gill. The very elongated postgenal cleft of the larva is remarkable in Simulium nigristrigatum. Simulium (Pternaspatha) limay WYGODZINSKY (Fig. 29.) Simulium limay WYGODZINSKY, 1958: 123; VULCANO, 1967: 15. Simulium (Pternaspatha) limay STONE, 1963: 16; WYGODZINSKY & COSCARÓN, 1967: 69-70; COSCARÓN & WYGODZINSKY, 1972b: 212; COSCARÓN, 1987: 18; 1991: 111. Simulium (Pternaspatha) huemul WYGODZINSKY & COSCARÓN, 1967: 67; COSCARÓN, 1991: 111. Simulium walterwittmeri WYGODZINSKY, 1958: 129.

Female: Wing length 2.8-3.5 mm; scutum gray pollinose with median and 1+1 sublateral bands blackish, covered by abundant gray-yellowish to silvery pilosity (Fig. A); abdomen gray pollinose with grayish dark brown spot medially on tergite I; another similarly median and 1+1 sublateral that don’t reach posterior border on tergites II-V, small median anterior spot on tergite VI and another more expanded on tergites VII-IX (Fig. B); legs grayish brown-yellow with apex of coxa, trochanter, apex of femur, tibia and tarsi more darkened; calcipala very small (Fig. C); claws with subbasal tooth evident (Fig. D); genitalia as shown in figures E-F; gonapophyses with abundant microtrichia and paraprocts with soft curvature on distal angles. Male: Wing length 2.5-3.0 mm; scutum black with 1+1 anterior, small subtriangular spots light gray continued laterally and posteriorly by thin grayish band bordering the scutum (Fig. G) and covered by grayish-tin pilosity; abdomen black with 1+1 silvery-gray spots on tergites II and VI, and very reduced on tergite III, and lateral borders of tergites VII; tergite IX grayish brown (Fig. H); hind leg with tibia and basitarsus scarcely stretched (Figs. I, J); length/width ratio of hind basitarsus = 5.2-5.6; genitalia as shown in figure K; gonostylus with tubercles on distal border (Fig. L).

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Figure 29. Simulium (Pternaspatha) limay. A-F: Female; A: scutum; B: abdomen, dorsal view; C: portion distal of hind basitarsus with small calcipala and pedisulcus; D: claw E: eighth sternite, gonapophyses, genital fork and spermatheca; F: paraproct and cercus. G-L: Male; G: scutum; H: abdomen dorsal view; I: hind leg; J: hind basitarsus; K: ventral view of gonocoxite, gonostylus, ventral plate, endoparameres and median sclerite; L: gonocoxite, gonostylus dorsal view. M-Q: Pupa; M: general view; N: portion of thorax with trichomes and gill; O: gill; P: portion of thorax with trichomes of different specimen than O; Q: frontoclypeus. R-V: Larva; R: cephalic apotome; S: antenna; T-U: mandible teeth of different specimens; V: hypostomium and postgenal cleft.

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Pupa: Cocoon length dorsally 3.0-3.8 mm, on base 3.3-4.2 mm, maximum 4.8-5.5 mm; cocoon light brown, with shoe-shaped strong basal projection, protecting gill base (Fig. M); body pupa length 4.0-4.5, gill length 1.5-2.0 mm; gill with 8 branches disposed as shown in figures N and O; frontoclypeus with 3 pairs of frontal trichomes, 1-2 pairs facials multibranched; thorax with 20-55 trichomes specially positioned on free posterior border; with 4-8 thin branches emerging close to base (Figs. N, P); frontoclypeus with tubercles on entire frons in female and reduced to basal area on male (Fig. Q); tubercles with smooth surface. Larva: Length of mature larva, 6.7-7.5 mm; maximum width of head capsule, 0.7 mm; general body shape much as in Simulium annulatum; color of larva dark brownish green, except unpigmented undersurface of posterior half of body; head dark brown, pigment pattern of cephalic apotome as shown in figure R; body integument with isolated short hair, more numerous in area of anal sclerite; antennae (Fig. S) with third segment darkest, second segment with one or two small unpigmented areas; ratio of length of segments proximal, medial and distal = 1:1.1:1.5:0.9; mouth brushes with 39-45 rays; toothing of mandibles as shown in figures T, U; anterior border of hypostomium strongly pigmented; hypostomial setae arranged in one or two irregular rows, with 14-16 setae in each group; postgenal cleft (Fig. V) very deep, five times as deep as length of hypostomial bridge; anal sclerite with some hair and without scales; anal ring with approximately 95 rows, each composed of 17 hooks; rectal papillae with three primary lobes, each with about six diverticula; median primary lobe simple in some specimens. Distribution: Argentina: Chubut, Mendoza, Neuquén, Río Negro, San Juan, Santa Cruz on precordilleran area. Bionomics: Simulium limay breeds in fast–flowing, clear rivers or creeks of large volume, with tree branches and leaves. Females could be collected several kilometers from breeding places on the Patagonian steppes. Discussion: The closest species to Simulium limay is S. nigristrigatum with adult abdomen ornamentation very similar, showing in the latter some variations. It can be separated from Simulium nigristrigatum by the absence in the pupa of tubercles on the frontoclypeus and dorsum of the thorax, scarce trichomes on the thorax (about 10), wider hind basitarsus in the male (length/width ratio = 3.2-3.4) and in larvae the rectal papillae with one lobule on the lateral lobes. Simulium walterwittmeri shows great similarity with S. nigristrigatum and S. limay (COSCARÓN 1991: 113) and lives in the same river. Based on pupal characters that are closest to Simulium limay, we consider S. walterwittmeri a synonym of S. limay. Simulium (Pternaspatha) pichi WYGODZINSKY and COSCARÓN (Fig. 30.) Simulium (Pternaspatha) pichi WYGODZINSKY & COSCARÓN, 1967: 70-74; COSCARÓN & WYGODZINSKY, 1972b: 213; COSCARÓN, 1991: 114-115.

Female: Wing length 2.4-2.9 mm; head blackish; color of appendages and hair like that of Simulium nemorale; scutum dark gray, with three longitudinal, light gray lines: a central very delicate one not attaining anterior border, and 1+1 wider submedian ones that meet central line posteriorly, posterior fourth of scutum light gray; lateral borders of scutum also light gray (Fig. A); scutum covered with very numerous appressed, silvery hair; scutellum light brown, with long silvery or dark brown hair; metanotum brown,

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tinged faintly with violaceous, delicately pollinose; pleural tuft silvery white; wing veins whitish; setae at base of wings whitish; base of halteres brownish, knob yellowish white; legs light brown; distribution of pigment on legs very similar to that of male; abdomen dull silvery gray; distribution of pigmentation in accordance with figure B; frons like that of Simulium nemorale; fronto-ocular triangle approximately as long as wide (Fig. C); shape and proportions of antennal segments as shown in figure D; last segment of maxillary palp (Fig. E) more than twice as long as penultimate, and approximately as long as two preceding segments combined; diameter of sensory vesicle as large as half of diameter of third segment; cibarium with well-sclerotized lateral projections (Fig. F); Sc in most cases without, rarely with one or two, setae on its center; setae and spines on R1 arranged in one row, in rare cases in irregular double row on basal portion of vein; setae on Rs arranged in one series; calcipala very small, wider than long; claws with small tooth (Fig. G); posterior portion of eighth sternite with 1+1 groups of approximately 35 long setae, their size diminishing toward central region (Fig. H); central portion of sternite slightly less pigmented than lateral areas; gonapophyses subtriangular, beset with microtrichia, their internal borders more strongly sclerotized; paraprocts and cerci as shown in figure I; cerci higher than wide, their distal border rounded; projecting portion of paraprocts subtriangular, extreme apex rounded; genital fork (Fig. J) with median process well sclerotized, its extreme apex widened; anterior projections narrow, with large lateral expansions at their tips, their external border pigmented; spermatheca like that in Simulium nemorale. Male: Wing length 2.3-2.6 mm; head, palpi, and labrum dark brown; eyes dark purple; antennae blackish, first and second segments slightly lighter; frons, clypeus, and labrum silvery pollinose; hair of palpi from silvery to black; antenna as in figure K; scutum velvety black, lateral margins and anterior and posterior declivities silvery pollinose, under certain angles with 1+1 submedian grayish spots adjacent to anterior gray border (Fig. L); hair of scutum very dense, silvery, decumbent, giving scutum overall grayish cast; metanotum dark brown, tinged with violaceous; pleura dark brown, grayish pollinose; pleural tuft silvery white; legs from yellowish to brownish; distribution and relative intensity of pigment of legs as shown in figures M, N, O; dorsal surface of abdomen velvety black, with 1+1 silvery gray spots on tergites II, VI, and VII; ventral surface of abdomen grayish brown; hair of abdomen, including that of basal fringe, silvery white; last segment of maxillary palp longer than two preceding segments combined; diameter of sensory vesicle about as large as half of diameter of third segment, this segment notably darker than remainder; wings like those of female, but no setae on Sc, and setae and spines on R1 invariably in single row, even at base of vein; hind basitarsus narrow, 3.8 times as long as wide; calcipala very small, wider at base than long (Fig. P); general structure of genitalia very similar to that of Simulium dureti; gonocoxite and gonostylus as shown in figures Q, R; ventral plate with 1+1 distinct, translucent areas. Pupa: Cocoon slipper shaped, with moderately extended anteroventral bridge (Fig. S); color of cocoon light brown, surface smooth, closely woven, but threads perceptible under moderate magnification; rim of aperture distinctly reinforced; gills covered by cocoon only at their extreme base; length of cocoon along dorsal surface 2.7-3.4 mm; along ventral surface 3.8-4.7 mm; length of body of pupa 2.8-3.0 mm; length of gill 1.2-1.6 mm; gills (Fig. T) each with eight filaments, slightly diverging on basal, and approximately parallel on apical,

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Figure 30. Simulium (Pternaspatha) pichi. A-J: Female; A: scutum; B: abdomen, dorsal view; C: fronto-ocular triangle; D: antenna; E: maxillary palp; F: cibarium basal portion; G: claw; H: eighth sternite; I: paraproct and cercus; J: genital fork. K-R: Male; K: antenna; L: scutum; M-O: fore, mid and hind leg respectively ; P: distal portion of hind basitarsus; Q: gonocoxite and gonostylus; R: gonostylus. S-W: Pupa; S: general aspect; T: gill; U: frontoclypeus; V: portion of thorax with trichomes and gill base; W: thoracic trichome enlarged. X-Z: Larva; X: general aspect; Y: cephalic apotome; Z: hypostomium; a: anal sclerite; c: rectal papillae.

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half; frontoclypeus (Fig. U) with 3+3 branched frontal and 1+1 branched facial trichomes; densely covered with small tubercles; exposed portion of thorax (Fig. V) covered with numerous tubercles and branched trichomes, latter limited to posterior half of exposed portion; trichomes slender, hair like, with two to five branches (Fig. W); average length of trichomes 0.12 mm; total number of trichomes of thorax approximately 60+60; chaetotaxy of abdomen like that in Simulium nemorale, but tubercles on tergites VI-VIII less numerous. Larva: Maximum length, 6.7 mm; maximum width of head capsule, 0.6 mm; general body shape as shown in figure X; color of larva light brown, with faint, dark, hypodermal pigment on dorsal surface and on underside of anterior third of body; cephalic apotome with faint brown pattern (Fig. Y); body integument smooth, hair present at base of anal sclerite; antennae light brown, distal segment darkest; medial segment with two slight constrictions, coinciding with unpigmented areas; ratio of lengths of proximal, medial and distal segments = 1:1.1-1.15:0.9; cephalic fan with about 26 rays in large fan; anterior border of hypostomium strongly pigmented (Fig. Z); median tooth almost as prominent as lateral and outermost intermediate teeth, remaining intermediate teeth smaller; sublateral tooth absent; lateral borders of hypostomium with seven or eight minute serrations; number of hypostomial setae from 7 to 11 in each group, arranged in one, rarely in two, irregular series; disc of hypostomium glabrous or with occasional simple short seta; postgenal cleft deep, about twice as deep as postgenal bridge; anal sclerite as shown in figure a; scales at base absent; anal ring with about 83 rows composed of about 18 hooks each; rectal papillae simple, composed of three simple lobes (Fig. b). Distribution: Argentina: Chubut, Santa Cruz. Bionomics: The larvae and pupae of Simulium pichi were collected in a crystal-clear stream about 2 m wide and 1 m deep. The specimens were attached to stems and leaves of Potamogeton sp. and Heleocharis sp., but mostly on reeds, generally on the downstream side of the leaves. Discussion: Simulium pichi resembles S. dureti, from which it differs by its smaller size, the smaller extension of the anterior gray portion on the scutum of the male, the absence of a median frontal blackish spot on tergite VI of the female, the absence of setae from Sc and the more salient cerci in the same sex, fewer and less-branched cephalic and thoracic trichomes of the pupa, and the different pigmentation of the larval head. Simulium limay also is similar, but has fewer trichomes on the thorax (20-55) and the frontoclypeal tubercles have more rugosity, and in the male pupa, the tubercles are only on the clypeal base. Simulium diamantinum also has a similar pattern on the abdomen, but is larger (wing length 3.4-3.9 mm), and has the calcipala absent, the claw subbasal teeth larger, the cocoon shoe shaped, more abundant frontal (5-15) facial (3-6), and thoracic (160-120) trichomes, a relatively shorter proximal antennal segment in larvae and more diverticula on the rectal papillae lobes (5-7). Simulium (Pternaspatha) dureti WYGODZINSKY & COSCARÓN (Fig. 31.) Simulium (Pternaspatha) dureti WYGODZINSKY & COSCARÓN, 1967: 62-67; COSCARÓN & WYGODZINSKY, 1972b: 209-210; COSCARÓN, 1991: 115-116.

Female: Wing length 3.6 mm; body with overall gray tinge; color of head like that of Simulium nemorale, hair of clypeus silvery; color of scutum very similar to that of Simulium

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nemorale, but grayish pattern more extended, occupying anteriorly one-third of surface of scutum; appressed setae of scutum silvery, very dense, giving scutum overall grayish aspect; scutellum grayish brown, its apical portion darker; pleural tuft silvery white; wing veins stramineous; hair at base of wings silvery white; stem of halteres brown; their knob yellowish white; legs light brown, their hair silvery to black; exact distribution and relative intensity of pigment of legs as in figures F-H; overall color of abdomen (Fig. K) light gray, dull; first segment darker; tergites II-V with well-developed, central, velvety black spot, separated from 1+1 slightly less intense, lateral, dark spots by gray areas as wide as central spots; tergite VI with very small, in some cases indistinct, black spot; lateral dark areas present; tergites VII and VIII with large, dark gray spots at center, in some cases entire tergites darkened; frons with obsolete median sulcus (Fig. A); fronto-ocular triangle slightly longer than wide (Fig. B); shape and proportions of antennal segments as in figure C; last segment of maxillary palp more than twice as long as penultimate and longer than two preceding segments combined (Fig. D); diameter of sensory vesicle larger than half diameter of third segment, its structure as shown in figure E; cibarium with wellsclerotized, lateral projections; lacinia with about 25 teeth, mandible with approximately 40 teeth; Sc with not more than 10 setae on its central portion; setae and spines on R1 arranged in one, or in some cases two, irregular rows; setae on Rs arranged in one series; shapes and proportions of segments of legs as shown in figures F, G, H; calcipala very small but distinct, somewhat wider at base than long (Fig. I); claws with small tooth (Fig. J); posterior portion of eighth sternite with 1+1 groups of long setae becoming progressively shorter toward center; about 30 setae in each group (Fig. L); central portion of eighth sternite less pigmented than lateral; gonapophyses subtriangular, covered with microtrichia; their internal border sclerotized; paraprocts and cerci as shown in figure M; cerci with apical border rounded, apex of paraprocts subrectangular; genital fork with median process distinctly sclerotized, its extreme apex widened; anterior projections with wide, lateral expansions; external border of distal portion pigmented; spermatheca like that in Simulium nemorale. Male: Wing length 3.0 mm; body with overall gray tinge; color of head and its appendages like that in Simulium nemorale; scutum with color pattern of the group, but all of anterior half gray under most angles except at middle, and posterior gray area also more widely extended than usual (Fig. N); scutum with very numerous silvery, appressed hair; pleura, halteres, and wings like those in female; legs light brown, their hair from silvery to black; abdomen (Figs. O, P) with dorsal surface velvety black, undersurface grayish; tergites II, VI, VII, and in some cases also VIII, with 1+1 light gray spots, smaller and with bluish sheen on posterior segments; hair of abdomen including that of basal fringe, silvery; last segment of maxillary palp approximately twice as long as penultimate; diameter of sensory vesicle slightly less than half diameter of segment; wings like those in female, but Sc devoid of hair, and spines and hair of R1 invariably in single row; shape, proportion and coloration of segments of legs very close to those of female and as shown on hind leg (Fig. Q); hind basitarsus 3.2-3.4 times as long as wide; calcipala very small but distinct, approximately as long as wide at base; gonocoxite and gonostylus as in figure R; gonostylus subquadrate, apical concavity beset with very numerous heavily pigmented tubercles (Fig. S); ventral plate as in figure T, without distinct translucent areas.

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Figure 31. Simulium (Pternaspatha) dureti. A-M: Female; A: frons; B: fronto-ocular triangle; C: antenna; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg, respectively; I: apex of hind basitarsus and tarsomere I; J: claw; K: abdomen, dorsal view; L: eighth sternite and gonapophyses; M: cercus and paraproct. N-T: Male; N: scutum; O-P: abdomen in dorsal and lateral views; Q: hind leg; R: gonocoxite and gonostylus, ventral view; S: gonostylus, dorsal view; T: ventral plate. U-Z: Pupa; U: lateral view; V-W: gill in different specimens; X: frontoclypeus; Y: exposed portion of thorax with trichomes and tubercles; Z: trichomes with high magnification. a-f: Larva; a: cephalic apotome; b: antenna; c: teeth of mandible; d: hypostomium; e: cephalic capsule in ventral view; f: anal sclerite and trichomes.

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Pupa: Pupae single or forming compact groups; cocoon shoe shaped, with large, anteroventral bridge; surface of all cocoons examined covered with foreign particles of various sizes (Fig. U); cocoon closely woven, threads not individualized; rim of aperture slightly reinforced; gills covered laterally on basal half by walls of cocoon, in some cases only extreme apex visible; length of cocoon at dorsum along middle to border of aperture, 3.3-4.0 mm; maximum length along base to anterior border of antero-ventral bridge, 4.3 mm; length of body of pupa 3.5-4.5 mm; length of gill 2.0-2.5 mm, viz., less than half as long as cocoon, and slightly more than half as long as pupa proper; gill with eight filaments (Figs. V, W), head and thorax of pupa light brown; clypeus covered with small tubercles; head with 1+1 groups composed each of approximately 17 branched frontal and facial trichomes (Fig. X) and 1+1 groups of three branched, ocular trichomes each; exposed portion of thorax more strongly sclerotized than remainder, covered with tubercles and long, branched trichomes; tubercles are abundant and those situated toward midline more or less pointed, spinelike; trichomes with as many as 8 branches (Figs. Y, Z); their average length, 0.2 mm; total number of thoracic trichomes approximately 150+150; chaetotaxy of abdomen like that in Simulium nemorale. Larva: Maximum length, 8.5 mm; width of head capsule, 0.7 mm; color yellowish, with dark greenish brown hypodermal pigment over whole body surface, especially intense on posterior half dorsally; cephalic apotome dark with negative spots (Fig. a); body integument smooth, with short setae at base of anal sclerite; antennae dark brown, pigment most intense on subapical segment (Fig. b); ratio of proximal, medial and distal segments = 1:1.3-1.45:0.7-0.8, viz., last segment distinctly shorter than first; cephalic fans with approximately 46 rays in large fan; toothing of mandible as shown in figure c; two external, one strong apical, three subapical, four internal, and two external teeth; anterior border of hypostomium strongly pigmented (Fig. d); median tooth as prominent as lateral ones, intermediate teeth gradually increasing in size from submedian to lateral ones; sublateral tooth barely perceptible; lateral borders of hypostomium with five to seven very small serrations; number of hypostomial setae about 14 in each group, arranged in two irregular series; disc of hypostomium with some short, simple setae; postgenal cleft deep (Fig. e), almost twice as deep as hypostomial bridge; anal sclerite as shown in figure f, without scales at its base; anal ring with approximately 90 rows, each composed of about 17 hooks; rectal papillae with three simple lobes. Distribution: Argentina: Jujuy, Salta; Bolivia: Potosí. Bionomics: Larval stages live in torrential creeks at 2000 to 4300 m altitude, flowing over stones and leaves. Discussion: Simulium dureti is close to Simulium pichi and S. deagostinii. The differences for S. pichi were treated under that species. Simulium deagostinii shows a more darkened abdominal tergite VI in the female and no calcipala In the pupa, the frontal and facial trichomes are fewer in number (5-6 on each side), and the larva has 6 diverticula on the median lobe and 4 on each lateral lobe of the rectal papillae. Simulium diamantinum is another similar species that can be differentiated by the absence of a blackish median anterior spot on female tergite VI, wider blackish spot on tergite VII, shorter blackish 1+1 lateral spots on tergites III-V, calcipala absent, thoracic trichomes more abundant (160-220), relatively shorter proximal segment of the larval antenna, and more diverticula on the rectal papillae lobes (5-7).

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Simulium (Pternaspatha) horcochuspi COSCARÓN & WYGODZINSKY (Fig. 32.) Simulium (Pternaspatha) horcochuspi COSCARÓN & WYGODZINSKY, 1972b: 210-212; COSCARÓN, 1991: 118.

Female: Wing length, 2.9-3.6 mm; color of head and thorax as in Simulium nemorale; appressed setae of scutum silvery, not very dense; overall color of dark portions of scutum distinctly blackish; scutellum and metanotum gray; pleural tuft silvery white; color of wings, halteres, legs and abdomen much as in Simulium dureti; head and its appendages and general shape of legs as in S. dureti; calcipala absent (Fig. A); claw without tooth, only with occasional slight angular projection in place of tooth (Fig. B); genitalia as in S. dureti. Male: Wing length about 2.8 mm; color of body and legs close to Simulium dureti; general structure as in S. dureti, including genitalia; apex of gonostylus granulose; length/ width ratio of hind basitarsus = 3.5; calcipala absent. Pupa: Pupae single or forming groups; cocoon with large anteroventral bridge (Fig. C); cocoon very thin, translucent, with conspicuous leaden sheen; cocoon closely woven; individual threads difficult to perceive; rim of aperture slightly reinforced; sides of cocoon covering basal half or two thirds of gills; length of cocoon at dorsum along middle to border of aperture 2.4-3.4 mm; maximum length along base to anterior border of anteroventral bridge, 4.6-5.2 mm; length of body of pupa 2.9-3.2 mm; length of gills 1.5-1.8 mm., viz., about half as long as pupa proper; gills with eight filaments, arising from three primary branches (Figs. D, E); frontoclypeus covered with small tubercles, more numerous in male (Fig. F) than in female (Fig. G); surface of tubercles smooth; head with 3(-4)+3(-4) frontal, 1+1 facial and 1-2+1-2 ocular trichomes; these trichomes hair like, with 1-4 branches (Fig. H); exposed portion of thorax covered with very numerous tubercles forming irregular groups (Fig. I) and long hair like trichomes; number of trichomes 75-85+75-85, each with from one to three, rarely four, branches, most frequently two; average length of trichomes 0.25 mm; abdomen as in Simulium dureti. Larva: Maximum length of mature larva, 8.0 mm; maximum width of head capsule, 0.75 mm; general body color whitish, but almost entire body with very intense dark brown hypodermal pigment; pattern of cephalic apotome negative; general body shape as in Simulium dureti; antennae as shown in figure J, similar to those of S. dureti, but distal segment about same length as proximal; cephalic fans, maxillary palp, mandibles, hypostomium, postgenal cleft, anal ring, and rectal papillae as in S. dureti; anal sclerite as figure K. Distribution: Argentina: Jujuy. Bionomics: Simulium horcochuspi was collected between 3900 and 4700 m of altitude in creeks of reduced volume and crystalline water flowing over stones. Discussion: Simulium horcochuspi is close to S. deagostinii and S. dureti but S. deagostinii has a median black spot on female tergite VI. The female of S. horcochuspi differs from the two related species mentioned, by the absence of a process on the claws; it differs further from S. dureti, with which it might possibly be found together, by the light-colored area of the anterior portion of the scutum, which is much less extensive than in S. dureti. The male of S. horcochuspi, known only from a pharate specimen, cannot be distinguished from that of S. deagostinii, but differs from that of S. dureti by the absence of a calcipala. The pupa of S. horcochuspi is characterized by the leaden sheen of its cocoon, which distinguishes it immediately from the two other species, and thoracic trichomes that in S. dureti number

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Figure 32. Simulium (Pternaspatha) horcochuspi. A-B: Female; A: apex of hind basitarsus and pedisulcus; B: claw. C-I: Pupa; C: pupa in lateral view; D-E: gill in different specimens; F-G: portion of frontoclypeus, showing different number of tubercles (F: male, G: female); H: frontoclypeus; I: trichomes of thorax. J-K: Larva; J: antenna; K: anal sclerite.

about 150. Further differential characters, mostly relating to the number and structure of its cephalic and thoracic trichomes, serve to distinguish S. horcochuspi from S. dureti and S. deagostinii, as shown in the key. In the larva, S. dureti and S. deagostinii have the distal segment of the antenna shorter than the proximal. Simulium (Pternaspatha) simile SILVA FIGUEROA (Fig. 33.) Simulium simile SILVA FIGUEROA, 1917: 33; PINTO, 1931: 730; VARGAS, 1945a: 196; BEQUAERT, 1945: 114; STUARDO ORTIZ, 1946: 41; WYGODZINSKY, 1949: 304; 1958: 133; STONE, 1962: 207. Simulium (Simulium) simile EDWARDS, 1931: 153; ORFILA, 1939: 1534. Acropogon simile ENDERLEIN, 1934b: 280. Simulium (Pternaspatha) simile WYGODZINSKY & COSCARÓN, 1967: 79; COSCARÓN & WYGODZINSKY, 1972b: 216-218; COSCARÓN, 1991: 119. [Simulium (Pternaspatha)] simile STONE, 1963: 17. Simulium figueroa SMART, 1944: 133; 1945: 505; VARGAS & DIAZ, 1953b: 141;VULCANO, 1967: 11. Simulium illiesi WYGODZINSKY & COSCARÓN, 1967: 75. Acropogon punctativentris ENDERLEIN, 1936: 118. Simulium punctativentris SMART, 1945: 512; VARGAS, 1945a: 188; STUARDO ORTIZ, 1946: 41; VULCANO, 1967: 19.

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Simulium (Pternaspatha) punctativentris WYGODZINSKY & COSCARÓN, 1967: 115; COSCARÓN & WYGODZINSKY, 1972b: 218. Simulium barbatipes VARGAS & DIAZ, 1953b: 141 (part).

Female: Wing length 3.2-3.5 mm; head blackish; antennae, palpi, and labrum piceous; scape and pedicel dark orange; frons and clypeus gray pollinose; pilosity of antennae very short, that of palpi, clypeus, frons, and occiput somewhat longer, silvery gray; scutum as in Simulium nemorale; hair of thorax silvery gray; appressed setae of scutum very dense, somewhat obliterating pattern; scutellum, metanotum, pleura, and sterna dark gray; pleura with silver pollinosity; legs similar to those of Simulium pulchrum; color of abdomen blackish with tergite II with 1+1 large, tergites III-V with 1+1 small, and tergites VI and VII again with 1+1 large, silvery white spots; tergites VIII and IX extensively silvery white (Fig. A); fronto-ocular triangle about as long as wide (Fig. B); maxillary palp as in figure C; diameter of sensory vesicle equal to half of width of third palpomere; structure of vesicle as shown in figure D; lacinia with 28-29 teeth, mandible with approximately 43 teeth; calcipala well developed (Fig. E); claws with relatively large tooth (Fig. F); eighth sternite as illustrated in figure G, its central portion more strongly sclerotized than lateral areas; gonapophyses triangular, beset with microchaetae, their inner border faintly sclerotized; cerci with distal border rounded; paraprocts salient, their free portion subtriangular (Fig. H); genital fork as illustrated (Fig. I). Male: Wing length more than 3.5 mm; color of head and cephalic appendages like those in Simulium nemorale; thorax black, with abundant, appressed, golden setae; abdomen blackish with ornamentation as shown in figures J-K; hind basitarsus 4.5 times as long as wide; calcipala well developed but shorter than wide at base; gonostylus with distal portion widely concave, heavily tuberculate (Fig. L); ventral plate as in figure M. Pupa: Cocoon shoe shaped (Figs. N-P), with large, anteroventral bridge; color of cocoon light brown; surface smooth, closely woven, but threads individualized; rim of aperture distinctly reinforced; gills protected laterally by wall of cocoon; length of cocoon at dorsum 4.0-4.5 mm; total length to border of anteroventral bridge, 5.5-7.0 mm; length of body of pupa 4.0-5.0 mm; length of gills 1.8-2.0 mm, slightly less than one-third of length of cocoon, and not quite half as long as pupa proper; gills whitish, each with eight filaments arranged in fan-like pattern (Figs. P, Q); head and thorax of pupa dark brown; frontoclypeus (Fig. R) with approximately 40 frontal and facial, and apparently 4+4 ocular, trichomes, their shape as described below for thoracic trichomes; clypeus heavily beset with tubercles; thorax (Fig. S) strongly sclerotized on exposed portion, latter heavily beset with tubercles and with large number (approximately 140+140) of trichomes; trichomes (Figs. T-V) narrow at extreme base, then flattened and band-shaped, apically again narrowed and pointed; trichomes simple or divided near base into two to four branches, latter closely parallel and difficult to individualize; tubercles of frontoclypeus at base not distinctly surpassing area of frontal trichomes; tubercles in many cases with sharply pointed protuberances (Fig. U); area of facial trichomes with conspicuously marked depression (Fig. R); borders of depression heavily sclerotized. Larva: Maximum length, 9.0 mm; maximum width of head capsule, 0.8 mm; body dark greenish gray; head dark brown; antenna as in figure W; ratio of proximal, medial and distal segments = 1:1.6:0.8; cephalic fans with 28-32 rays in large fan; toothing of

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Figure 33. Simulium (Pternaspatha) simile. A-I: Female; A: abdomen, dorsal view; B: fronto-ocular triangle; C: maxillary palp; D: sensory vesicle; E: hind basitarsus showing calcipala; F: claw; G: portion of eighth sternite and gonapophyses; H: cercus and paraproct; I: genital fork. J-M: Male; JK: abdomen in dorsal and lateral views; L: gonocoxite and gonostylus in ventral view; M: ventral plate. N-V: Pupa; N-P: lateral, dorsal and ventral view, respectively; Q: gill; R: frontoclypeus, showing trichomes and tubercles; S: portion of exposed portion of thorax, with abundant trichomes; T-V: different trichomes with high magnification. W-Z: Larva; W: antenna; Y: portion of hypostomium; X: teeth of mandible; Z: cephalic capsule in ventral view.

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mandibles as shown in figure X; pigment pattern of hypostomium as shown in figure Y; median tooth more pronounced than lateral ones; lateral serrations distinct; hypostomial setae arranged in two or three irregular rows 21-24 in each group; postgenal cleft pointed, very deep, its apex very close to hypostomial groove (Fig. Z); anal sclerite at its base with simple to trifid scales, few in number; anal ring with 95-118 rows, each composed of 1819 hooks; rectal papillae with three primary lobes, each bearing three pairs of long secondary lobules; median pair in some cases with four pairs of secondary lobules. Distribution: Argentina: Chubut, Neuquén, Rio Negro; Chile: from Tarapaca to Bio-Bio. Bionomics: Simulium simile is a medium- to high-altitude species which prefers colder, very fast-flowing streams. Discussion: Simulium simile is characterized by the unique chaetotaxy of the pupal thorax. In the shape of its cocoon it resembles Simulium nemorale, but the latter has a very different thoracic chaetotaxy. The male can be distinguished from that of similar species as shown in the key. The female is not well enough known to be compared with females of related species. Simulium simile is very close to S. pulchrum PHILIPPI redescribed below; the differences between these two species are indicated in our keys, and summarized in a table given under the heading of S. pulchrum. These species also differ in habitat preferences. Simulium (Pternaspatha) pulchrum PHILIPPI (Fig. 34.) Simulium pulchrum PHILIPPI, 1865: 633; KERTESZ, 1902a: 290; SILVA FIGUEROA, 1917: 30; PINTO, 1931: 729; VARGAS, 1945a: 187; SMART, 1945: 512; STUARDO ORTIZ, 1946: 42; VULCANO, 1967: 19. Simulium (Pternaspatha) pulchrum WYGODZINSKY & COSCARÓN, 1967: 115; COSCARÓN & WYGODZINSKY, 1972b: 204, 213-216; COSCARÓN, 1991: 118-119.

Female: Wing length 2.6-2.8 mm; color of head, thorax, and legs as in S. simile; legs brownish with apex of segments more darkened as shown in figures B-D; abdomen close to Simulium limay but sublateral blackish spots on tergites III-V wider (Fig. A); frontoocular triangle, slightly deeper than wide (Fig. E); shape and proportions of antennal segments as shown in figure F; diameter of sensory vesicle of third palpomere equal to half width of segment (Fig. G); lacinia with 25-26 teeth, mandible with 38-41 teeth; calcipala, claws and wings as in Simulium simile; general structure of genitalia as in Simulium simile, but paraproct less salient. Male: Wing length 2.6-2.7 mm; color of head and its appendages, thorax wings, and halteres as in Simulium simile; legs as in female, although somewhat darker; abdomen velvety black dorsally, except dark gray on tergum IX; terga II, VI and VII with 1+1 large, silvery white spots; terga III-V entirely black; antennae as shown in figure H; hind basitarsus five times as long as wide; genitalia as in Simulium simile. Pupa: Structure of cocoon as in Simulium simile; its length dorsally 2.2-2.7 mm, maximum length along its base 3.8-4.5 mm; length of body of pupa 2.5-3.2 mm, of gills 1.11.3 mm; head of pupa light brown; chaetotaxy of head (Fig. I) much as in S. simile, but tubercles of frontoclypeus more numerous (about 800) than in S. simile, distinctly surpassing level of frontal trichomes, attaining level of uppermost facial trichomes; protuberances of tubercles not pointed; thorax of pupa much as in S. simile, but trichomes less numerous

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Figure 34. Simulium (Pternaspatha) pulchrum. A-G Female; A. abdomen, dorsal view; B-D: fore, mid and hind leg, respectively ; E: fronto-ocular triangle; F: antenna; G: sensory vesicle of maxillary palp. H: Male, antenna. I: Pupa, frontoclypeus with abundant tubercles and trichomes. J: Larva, cephalic apotome of different specimens.

(normally 70-90, occasionally up to 130, on each side), and tubercles more numerous; length of trichomes 0.15-0.28 mm, much shorter than in S. simile; abdomen as in S. simile. Larva: Length of mature larva, 5.6-6.0 mm; maximum width of head capsule, 0.8 mm; general body shape as in Simulium simile; color light greenish brown, head light brown; cephalic apotome very light, dark pigment restricted to base (Fig. J); general aspect of antennae as in S. simile; ratio of proximal, medial and distal segments = 1:1.5:1.1; cephalic fan with 29-33 rays; maxillary palpi, hypostomium, postgenal cleft and anal sclerite as in S. simile; anal ring with 85100 rows of about 16 hooks each; rectal papillae in most specimens with three simple lobes, with one pair of diverticula in specimens from the province of Aconcagua, and two pairs diverticula on each lobe in specimens from southern part of range. Distribution: Chile: Aconcagua, Colchagua, Curico, Nuble, O’Higgins, Santiago, Talca. Bionomics: The larvae and pupae of this species were found attached to stems of grasses (such as Scirpus sp.) trailing in the water, or to branches of trees like Salix trailing in the stream, but only rarely to rocks. The species was collected mostly in large, fastflowing streams or irrigation ditches. It prefers clear streams, with only occasional turbidity. This is a lowland species in Central Chile, collected generally at 400-800 m; the highest elevation recorded was 1100 m. Discussion: Simulium pulchrum is closely related to Simulium simile, from which it can be distinguished as shown in table 1.

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Table 1. Differences between Simulium pulchrum and Simulium simile (All measurements are in millimeters.) Simulium pulchrum

Character

Simulium simile

ADULTS Wing length, female Wing length, male Darkened portion of posterior tibia of Female Tergum III of abdomen of male

2.6-2.8 2.6-2.7 Apical fifth Black

Terga IV and V of abdomen of male

Black

3.2-3.5 3.2-3.5 Apical two-fifths with 1 + 1 small whitish spots with 1 + 1 very small whitish spots

PUPAE Length of cocoon, dorsally Length of cocoon, ventrally Length of body of pupa Length of gills Number of tubercles of frontoclypeus Shape of projections of tubercles Area of facial trichomes

2.2-2.7 3.8-4.5 2.5-3.2 1.1-1.3 +800 Not pointed almost smooth 0.15-0.30 130+130 70+70

3.3-4.5 5.2-6.8 3.2-4.5 1.8-2.0 +400 Pointed Distinct impression limited by carinae 0.3-0.5 250+250 140 + 140

light brown light greenish brown 6.0 0.6 85-100 14-17 13-18 1 or 2

dark brown dark greenish brown 9.0 0.8 95-118 18-19 21-24 3, rarely 4

Length of thoracic trichomes Maximum number of trichomes of thorax Average number of trichomes of thorax LARVAE Color of head Color of body Maximum length Maximum width of head capsule Number of rows in anal ring Number of hooks per row Number of hypostomial setae Number of diverticula to each rectalPapillae

Simulium (Pternaspatha) diamantinum COSCARÓN & COSCARÓN ARIAS (Fig. 35.) Simulium (Pternaspatha) diamantinum COSCARÓN & COSCARÓN ARIAS, 1996a: 703.

Female: wing length 3.3-3.7 mm; general color gray pollinose; antennae, maxillary palp and proboscis blackish; head brown, dark gray pollinose; frons, clypeus and occiput blackish with gray pollinosity; occiput a little darker with hair grayish to black in accordance with light; scutum grayish black with silvery-gray, submedian, lyre-shaped pattern, lighter anteriorly and grayish laterally and posteriorly (Fig. A); with changing light

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position lyre-shaped pattern becomes thicker, and in some lights its anterior border becomes black, to silky bright light gray; scutum hair sparse and upright; scutellum blackish-gray; scutum and scutellum hair light gray; metanotum velvety blackish; pleura blackish gray; legs blackish with basal third of femur III, basal half of tibiae I-III, basal half of basitarsus II and 3/4 of basitarsus III and base of tarsomeres II-III light brownish; distribution of pigmentation of hind leg as in figure G; abdomen light grayish pollinose, tergite I dark grayish; black subtriangular median spots on tergites II-V, lighter and pollinose on VII, and 1+1 lateral, blackish, subrectangular spots not reaching posterior border on tergites IV-V (Figs. B, C); hair of pleural tuft light gray; frons as in figure D; fronto-ocular triangle about as long as wide (Fig. E); sensory vesicle of palp about 1/3-1/4 of third palpomere, its structure as in figure F; mandible with 25+8-30+9 teeth, and lacinia 23-25 teeth; Sc with 3-5 setae; base of R without setae; R1 and R with setae and spines arranged in one irregular row; shape and proportion of segments of hind leg as shown in figure G; without calcipala; ratio of length to width of hind basitarsus = 7.2-7.6; claws with well-developed basal tooth (Fig. H); eighth stemite well sclerotized, with 34-40 hairs on each side (Fig. I), gonapophyses subtriangular; paraproct and cercus as in figure J; genital fork as in figure K.; spermatheca with small internal spines (Fig. L). Male: Wing length 2.8-3.0 mm; head coloration as in female; scutum black-gray with 1+1 subtriangular spots anteriorly (Fig. M); with light anterior to specimen 1+1 anterior gray pollinose spots occupying anterior third; decumbent, silky grayish hair sparsely covering scutum; abdomen blackish with 1+1 large gray spots on tergite II, smaller on tergites VII-VIII, VI and IX mostly grayish except for small and short median darker spot; legs yellowish brown with light gray hair; blackish brown on coxae, apex of femur III, apex of tibiae, all of basitarsus I, distal third of basitarsi II and III and tarsi; hind legs as in figure N, basitarsus three times as long as wide; calcipala absent; genitalia very close to species of Simulium (Pternaspatha) NIGRISTRIGATUM species group; gonostylus with abundant tubercles (Fig. O). Pupa: Cocoon shoe shaped with well-developed antero-ventral bridge (Fig. P); surface generally smooth, but some specimens with small sand granules incorporated; cocoon closely woven; gill covered basally by walls of cocoon; length of cocoon 3.8-4.0 mm at dorsum, maximum length along base 5.2-6.0 mm, length of body 4.0-4.7 mm, of gills 2.2-3.3 mm; gills with eight filaments, position of basal branching as in figure Q; frontoclypeus and thorax with 500-1000 tubercles (Fig. R), those of thorax slightly pointed, especially on posterior border (Fig. S), others rounded with smooth or slightly rough surface (Fig. T); trichomes filiform with 4-8 branches, 5-15 frontal and 3-7 facial trichomes on each side in continuous group; length of frontal trichomes 0.19-0.23 mm; thoracic trichomes 0.24-0.35 mm, 160-220 in number, with 6-10 filiform branches (Fig. U); chaetotaxy of abdomen following general pattern shown in Simulium nemorale. Larva: Maximum length 7.5-8.5 mm; head width 0.7 mm; color dark, greenish brown, ornamentation and shape in accordance with figure V; integument smooth; cephalic apotome brown with negative spots without well defined borders (Fig. W); antennae dark brown; ratio of proximal, media and distal segments = 1:1.8-1.9:1.2-1.3 (Fig. X); cephalic fans with 35-39 combs; mandible with 4-6 secondary teeth arranged in several rows (Fig. Y); teeth of hypostomium border as in figure Z, with 5-6 lateral

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Figure 35. Simulium (Pternaspatha) diamantinum. A-L: Female; A: scutum; B-C: abdomen in dorsal and lateral views; D: frons; E: fronto-ocular triangle; F: second palpomere with sensory vesicle; G: hindleg; H: claw; I: eighth sternite and gonapophyses; J: cercus and paraproct; K: genital fork; L: spermatheca. M-O Male; M: scutum; N: hind leg; O: gonostylus. P-U : Pupa; P: lateral view; Q: gill; R: frontoclypeus; S-T: tubercles of thorax pointed and rounded; U: thoracic trichomes. V-Z, a-c : Larva; V: general aspect in lateral view; W: cephalic apotome; X: antenna; Y: teeth of mandible; Z: hypostomium and postgenal cleft; a: proleg sclerite; b: anal sclerite; c: rectal papillae.

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serrations and 15-17 lateral setae arranged in 1-2 rows; postgenal bridge short, ratio of hypostomium/postgenal cleft = 1.4-1.5; comb of thoracic proleg with about 43-54 teeth arranged in 12-14 groups (Fig. a); anal sclerite without scales but with isolated trichomes as in figure b; anal ring with about 94-100 rows, each composed of about 15-17 hooks; rectal papillae with 5-7 diverticula on each of the three well developed lobes (Fig. c). Distribution: Argentina: Mendoza, Neuquén, San Juan. Bionomics: Simulium diamantinum breeds in creeks with strong current, crystalline and cold waters on aquatic vegetation at 3000 m altitude. Females show strong anthropophily. Discussion: Simulium pichi, S. dureti, S. diamantinum, S. horcochuspi and S. deagostinii are very closely related species that can be distinguished by some morphological characters and different distributions. Simulium diamantinum can be distinguished from the other species by several distinctive characters such as the female abdomen with 1+1 small, narrow and short, black lateral spots on tergites III-V, well isolated from the median black spot and leaving a wide white transverse band before the posterior border; scutum hair not appressed; narrower posterior basitarsi; longer pupa gill; greater number of frontal and thoracic trichomes and longer thoracic trichomes. The larva has a longer medial antennal segment, more lateral setae on the hypostomium, and a shorter postgenal bridge in relation to the hypostomial length. Simulium (Pternaspatha) strigidorsum (ENDERLEIN) (Fig. 36.) Acropogon strigidorsum ENDERLEIN, 1934b: 279. Simulium strigidorsum SMART, 1945: 514; VARGAS, 1945a: 199. Simulium (Pternaspatha) strigidorsum WYGODZINSKY & COSCARÓN, 1967: 74-75; COSCARÓN & WYGODZINSKY, 1972b: 218-219; COSCARÓN, 1991: 121.

Female: Wing length 3.3-3.6 mm; head black; antennae and mouthparts piceous; occiput, frons, and clypeus gray pollinose; pilosity of antennae very short, of palpi somewhat longer; occiput with long hair; scutum dark gray to black, with 1+1 light gray, median, longitudinal stripes arising from 1+1 anterior white spots; narrow, median, longitudinal line separated from gray stripes on its central half by elongate blackish areas; stripes and midline confluent on posterior third of sclerite, connected to equally gray though somewhat darker lateral borders of scutum; width of submedian gray stripes each about one-eighth of total width of scutum; pubescence of scutum sparse and short, decumbent, silvery; scutellum and metanotum piceous, silvery pollinose; scutellum with long decumbent and erect silvery hair; pleural tuft silvery; pleura and sterna piceous, silvery pollinose; wing veins stramineous, hair and setae black, tufts of hair at base of wings whitish; coloration of legs of lectotype as in figure A, B, shows more darkened fore legs and femur of hind legs; coxae and trochanters blackish, gray pollinose; forelegs entirely dark brown (Fig. A); basal half of tibia, basal three-fourths of first and basal half of second tarsal segment yellowish; hind legs dark brown; extreme base of femur and proximal half of tibia yellowish, mid legs brown, basal half of tibia, basal three-fourths of first and basal half of second tarsal segments yellowish; hind leg dark brown, extreme base of femur and proximal half of tibia yellowish; basal two-thirds of first and proximal half of second tarsal segment whitish; very

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small dark spot on base of second tarsal segment (Fig. B); dorsal surface of abdomen dull silvery gray; first segment darkened, its posterior fringe whitish; tergites II-V each with median velvety black spot (Fig. C); lateral dark spots or areas absent; tergite VI entirely gray; tergite VII with distinct gray spot, VIII faintly darkened at center; frons wide; fronto-ocular triangle about as long as wide (Fig. D); shape and portion of antennal segments as shown in figure E; last segment of maxillary palp more than twice as long as penultimate and longer than two preceding segments combined (Fig. F); diameter of sensory vesicle about half as large as diameter of third segment of palp; wings as usual for subgenus; Sc and basal portion of R bare; R1 with setae and spines arranged in single row, setae of Rs in single row; calcipala (Fig. G) very small, in shape of small protrusion; claws with small tooth (Fig. H); genitalia as illustrated in figures I-K; central portion of eighth sternite less strongly pigmented than lateral areas, only its hind margin narrowly dark; setae of lateral portion of sternite not very numerous (Fig. I); gonapophyses membranous, subtriangular, beset with microtrichia; cerci rather short (Fig. J); paraprocts somewhat more intensely pigmented than cerci, their free extremity very shortly salient; spermatheca subglobular with spiculae on inner surface; genital fork as shown in figure K. Male, pupa and larva: Unknown. Distribution: (?Argentina: Chubut and Mendoza); Peru: Puno. Bionomics: Unknown. Discussion: The differences between the types from Peru and the Argentinean specimens are very slight, and we do not hesitate to consider all specimens as conspecific.

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Figure 36. Simulium (Pternaspatha) strigidorsum. A-K: Female; A: fore leg; B: hind leg; C: abdomen, dorsal view; D: fronto-ocular triangle; E: antenna; F: maxillary palp; G: apex of hind basitarsus and tarsomere I; H: claw; I: portion of eighth sternite and gonapophyses; J: cercus and paraproct; K: genital fork. (D-F taken from Mendoza specimens; A-C, G-J from type).

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Preimaginal stages from the type habitat could help to elucidate the identity. The color pattern of the abdomen of the female without blackish lateral areas is unique in the subgenus. Simulium diamantinum from the same place in Mendoza shows some similarities. The female abdomen of S. diamantinum has 1+1 blackish lateral spots on tergites III-V, and the legs are lighter. Several other species of the Simulium NIGRISTRIGATUM species group are similar in abdominal ornamentation, such as S. pichi, which is smaller (wing length 2.4-2.9 mm), and S. dureti, which has a small subbasal claw tooth. Simulium herreri has smaller light gray spots on tergites IV-VIII of the female abdomen and a subbasal claw. Simulium (Pternaspatha) deagostinii COSCARÓN & WYGODZINSKY (Fig. 37.) Simulium deagostinii COSCARÓN & WYGODZINSKY, 1962: 322-332; VULCANO, 1967: 9. Simulium (Pternaspatha) deagostinii WYGODZINSKY & COSCARÓN, 1967: 61-62; COSCARÓN & WYGODZINSKY, 1972b: 209; COSCARÓN, 1987: 18; 1991: 117.

Female: Wing length 3.4-3.5 mm; frons and clypeus blackish with gray pollinosity; scutum blackish gray with 1+1 submedian longitudinal light gray stripes defining dark bands (Fig. A); abdomen grayish, black dorsally with 1+1 submedian wide silvery gray spots on tergites II-VII, wider on II, VI-VII (Fig. B); pleural and ventral areas grayishbrown; legs brown with hair silvery to black, darkened on coxa, apex of femur, tibia and tarsi as in figures E, F, G; fronto-ocular triangle wider than high (Fig. C); distal palpomere length shorter than two before combined (Fig. D); calcipala absent (Fig. H), and claws with small subbasal tooth (Fig. I); eighth sternite homogeneously darkened (Fig. J), paraproct prominent distally (Fig. K). Male: Wing length 3.3-3.4 mm; scutum black bordered anteriorly, laterally, and posteriorly by grayish band; legs dark brown with darkened areas as shown on hind leg (Fig. L); abdomen velvety black dorsally, grayish ventrally; tergites II, VI, VIII with 1+1 sublateral light gray spots, more developed on tergite II and getting narrower gradually from VI to VIII; ratio of length/width of hind basitarsus = 3.2; calcipala not developed; gonocoxite and gonostylus as in figure M, the latter with abundant tuberosity; ventral plate as in figure N. Pupa: Cocoon length on dorsum 4.0 mm, at base 4.5-5.0 mm; color light brown, shoe shaped, strongly projected anteriorly, protecting base of gills (Fig. O); length of pupal body 3.0 mm, gill 2.4 mm; frontoclypeus and thorax dorsally with abundant tubercles; cephalic trichomes 5 or 6 in number on each side with 5 or 6 branches (Fig. P); thorax with 65-95 trichomes (Fig. Q) with 2-8 branches (Fig. R) covering dorsally all free area; gill with 8 branches (Fig. S). Larva: Maximum length on dorsum 8.0 mm; color generally light grayish brown; cephalic apotome light brown, mostly darkened basally (Fig. T); ratio of segments proximal, medial and distal = 1:1.4:0.85; cephalic fan with about 45 rays; hypostomium with median teeth at same level as lateral teeth; plate with 11-13 setae on each side, arranged in more than one row; postgenal cleft deep, postgenal bridge about 1/5 of hypostomium length (Fig. U); anal sclerite without scales; anal ring with 95 rows of 25-28 hooks each; rectal papillae with 6 diverticula on median lobe and 4 on each lateral lobe (Fig. V). Distribution: Argentina: from Tierra del Fuego to center of Chubut.

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Bionomics: Simulium deagostinii breeds in rivers or large-volume creeks, with fastflowing, crystalline waters. Females were captured biting humans and horses. Discussion: The closest species to S. deagostinii are S. dureti and S. pichi. Differences from the first species were explained under that species. Comparison with S. pichi shows that in S. pichi the female abdomen has a narrower median black stripe and tergite VI is totally light gray, the calcipala is small, the trichomes of the thorax number about 60 on each side, the tubercles are comparatively few and the lobes of the larval gill are without diverticula. F E 0.1

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Simulium (Pternaspatha) cotopaxi WYGODZINSKY & COSCARÓN (Fig. 38.) Simulium (Pternaspatha) cotopaxi WYGODZINSKY & COSCARÓN, 1979: 19; COSCARÓN, 1991: 103; COSCARÓN ARIAS, 2003: 198, 204, 208, 215, 219.

Female: Wing length 3.8-4.2 mm; head with eyes black; clypeus, frons and occiput dark brown, with silver gray pruinosity; antennae and palpi black, their setae from dark brown to black; scutum black, with variable color pattern according to incidence of light: in some instances (Fig. A) almost entirely black, with 1+1 triangular anterosubmedian and 1+1 submedian spots on posterior third silver gray, and 1+1 sublateral and one median longitudinal stripe of same color; with different illumination, scutum with 1+1 anterosubmedian triangular silver spots and 1+1 obliquely longitudinal silvery stripes beginning somewhat before middle of scutum and connected posteriorly by transverse band (Fig. B); with still different illumination central portion of scutum almost entirely silvery, with 1+1 anterior and 1+1 submedian spots appearing black; scutellum dark brown, narrowly margined with lighter along hind border; metanotum velvety grayish brown; setae of scutum and scutellum from brass to golden colored; pleura blackish; legs grayish brown, with coxae, apices of femora and tibiae, apex of basitarsi and remaining tarsomeres, darkened (Figs. H, I, J); setae of legs large and very numerous, producing overall lightcolored appearance of legs; abdomen black, with 1+1 silver gray spots on tergites II-VII, most conspicuous on segments II and Vl, reduced on Vll (Fig. C); terminal tergites from dull to slightly polished; ventral surface of abdomen blackish; frons without median sulcus (Fig. D); frontal angle 125 degrees; fronto-ocular triangle as deep as wide at base (Fig. E); shape of segments of antennae shown in figure F; setae not unusually elongate or numerous; maxillary palp as shown in figure G; sensory vesicle of maxillary palp slightly wider than half diameter of third segment; lacinia and mandibles serrate on both edges; lacinia with approximately 28 teeth, mandible with 42-47 teeth; Sc with 12-15 hair; R1 with hair and spiniform setae arranged in one or two irregular rows; shape of legs shown in figures H-J; calcipala (Fig. K) slightly less than half as wide as basitarsus; pedisulcus normal; claws with well-developed triangular process subbasally (Fig. L); eighth stemite more darkened on median area (Fig. M); gonapophyses divergent, their inner margin conspicuously sclerotized, their disc with very numerous microtrichia; cerci low; paraprocts narrowly and shortly projecting apically, their posterior border emarginated (Fig. N); genital fork with stem slender, heavily pigmented; spermatheca subglobular, pigmented, except circular area of insertion of spermathecal duct; inner surface of spermatheca with faintly visible slender spicules. Male: Wing length 3.6-3.8 mm; head black; eyes dark brown to black; clypeus black with silvery pruinosity; antennae and palpi black, their hair also black; scutum entirely black, with certain illumination anterior margin appearing gray with 1+1 short submedian posteriorly directed gray projections, or with narrow silver gray band along margins of entire sclerite; setae of scutum and scutellum colored as in female; color of metanotum, pleura, wings and legs as in female; abdomen (Fig. O) sooty black, with 1+1 silver gray spots each on segments II, VI, and VII, the one on II largest, the one on Vll smallest; head strongly hairy (Fig. P); row of hair between eyes; scape and pedicel (Fig. Q) extraordinarily enlarged, combined about as long as flagellum, and much wider than segments of latter;

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setae of scape and pedicel unusually numerous and long, surpassing level of middle of flagellum; maxillary palp with setae of third and to lesser degree also fourth palpomere unusually long and numerous (Fig. R); fifth segment slightly over twice as long as penultimate; sensory vesicle small; shape and pigment pattern of legs very close to those of female but more darkened and hind leg with basitarsus strongly widened (Fig. S), about three times as long as wide; calcipala small, one-fourth as wide as basitarsus apically (Fig. T); genitalia with gonocoxite and gonostylus subtrapezoidal; gonostylus with distal margin truncate and slightly concave and smooth; apical spine short, situated on slight projection of inner apical angle (Fig. U); median sclerite with apical half widened and deeply incised; endoparameres with numerous spinelike processes; membrane of aedeagus with numerous minute cuticular spicules; ventral plate (Fig. V) wider than long. Pupa: Cocoon (Fig. W) light brown, slipper shaped, translucent; fabric of cocoon very uniform, parchment-like, individual threads difficult to perceive; anterior border of cocoon reinforced, opening of cocoon subcircular; length of cocoon along dorsal surface 3.2-3.7 mm, along ventral surface 4.5-4.8 mm; body pupa 3.7-4.7, and gills 3.0-3.5 mm; gills brown, darkest on basal half or third, each consisting of six slender filaments; all six filaments forward directed, of subequal length, forming very tight bundle; head, thorax and abdomen light brown, with frontoclypeus, antennal sheaths and exposed portion of thorax dark brown, heavily sclerotized; tubercles of head and thorax delicately and abundantly spinulose; shape of frontoclypeus of female as in figure Y, and male as in figure Z; frontoclypeus with 1+1 facial and 2+2 frontal trichomes, from simple to bearing up to eight branches (Fig. a); thorax (Fig. X) with 5-6+5-6 trichomes, also very small, with up to eight branches (Fig. b); tergites III and IV with 4+4 simple hooks; apex of abdomen (Fig. c) without pair of hooks or denticles. Larva: Length 8.0-8.5 mm; width of head 0.8 mm; general body color light yellow brown, tinged with greenish; head dark brown; general body shape as shown in figure d, abdomen gradually widened posteriorly, not abruptly truncate at tip; cephalic apotome as shown in figure e; spots positive, feeble, darkest along posterior margin; antennae as shown in figure f, faintly pigmented, third segment darkest; ratio of proximal, medial and distal segments = 1:1.4-1.7:0.8; cephalic fan with approximately 40 rays; toothing of mandible as shown in figure g; maxillary palp as shown in figure h; hypostomium with median and lateral teeth equally prominent (Fig. i); hypostomial setae (Fig. j) in two or three irregular rows, each group consisting of 12-18 setae; postgenal cleft slightly deeper than length of postgenal bridge (Fig. k); lateral sclerite of proleg shown in figure l, setae arranged in three or four groups on prominent tubercles; body of larva glabrous, except some setae near anal region and isolated ones scattered over rest of body; anal ring with approximately 90 rows of 16 hooks each; anal sclerite as shown in figure m; rectal papillae with 11-14 diverticula on each lobe (Fig. n). Distribution: Ecuador: Cotopaxi, Pichincha, Tungurahua. Bionomics: The mouthparts of the female of this species are normally developed; therefore, females are presumed to be haematophagous. Larvae and pupae of Simulium cotopaxi live in streams 2-5 m wide, with many small rapids and falls; the water temperature was 11 ºC. Pupae were mostly found attached to the sides and undersurface of large stones and boulders, as well as on blades of grass trailing in fast-flowing water.

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Figure 38. Simulium (Pternaspatha) cotopaxi. A-N: Female; A-B: scutum with different illumination; C: abdomen, dorsal view; D: frons; E: fronto-ocular triangle; F: antenna; G: maxillary palp; H-J: fore, mid and hind leg, respectively; K: apex of hind basitarsus and tarsite I; L: claw; M: eighth sternite and gonapophyses; N: cercus and paraproct. O-V: Male; O: abdomen in lateral view; P: head; Q: antenna; R: maxillary palp; S: hind leg; T: apex of hind basitarsus and tarsomere I; U: gonocoxite and gonostylus; V: ventral plate. W-Z & a-c : Pupa; W: general aspect, lateral view; X: portion of exposed portion of thorax and gill; Y: female frontoclypeus; Z: male, frontoclypeus; a: frontoclypeus trichome; b: thoracic trichomes and tubercles; c: abdomen distal portion. d-n: Larva; d: lateral view; e: cephalic apotome; f: antenna; g: teeth of mandible; h: maxillary palp; i: teeth of hypostomium; j: hypostomium; k: hypostomium and postgenal cleft; l: proleg lateral sclerite; m: anal sclerite; n: rectal papillae.

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Discussion: Simulium cotopaxi is a peculiar species with a variety autopomorphic and plesiomorphic characters that help separate it from other species, but constitute a dilemma to place it in either one of the species groups of Pternaspatha. The gonostylus is without tubercles (plesiomorphic), the gill branches and the enlarged hairs on the antennae and maxillary palp of the male are characteristic of the NEMORALE group. On the other hand ,the pupa with multiple filamentous branched trichomes, reduced number of trichomes on the thorax (5-6 on each side) and female abdomen with 1+1 light gray stripes on tergites II-VII, are more common in the NIGRISTRIGATUM group. Some autopomorphies such as ornamentation of the female scutum, the ventral projection of the paraproct, the elongate male pedicel, hair on the basal segments of the antenna and maxillary palp, very wide hind basitarsus in the male, and the absence of terminal hooks on the pupal abdomen, show a specialization probably present in a line of Pternaspatha ancestors that evolved independently of the rest of the Pternaspatha. NEMORALE species group Simulium (Pternaspatha) nemorale EDWARDS (Fig. 39.) Simulium (Simulium) nemorale EDWARDS, 1931: 152; ORFILA, 1939: 1533. Simulium nemorale SMART, 1945: 509; VARGAS, 1945a: 168; WYGODZINSKY, 1949: 304; VARGAS & DIAZ, 1951: 138. Simulium (Pternaspatha) nemorale WYGODZINSKY & COSCARÓN, 1967: 83-88; COSCARÓN & WYGODZINSKY, 1972b: 222; COSCARÓN, 1991: 122-123.

Female: Wing length 3.2-4.0 mm; head black; eyes, antenna, maxillary palp, and labium dark brown; occiput, frons, and clypeus gray pollinose; pilosity of antenna very short, that of palp, clypeus, frons, and occiput somewhat longer; hair brass-colored to black; first and second segments of antennae slightly lighter colored than remainder, and with longer hair; scutum (Fig. A) black, with 1+1 silvery gray, submedian stripes arising from 1+1 anterior white spots; median dark stripe divided longitudinally by delicate white line; submedian stripes connected posteriorly to gray lateral borders of sclerite; appressed setae of scutum brass-colored, dense, somewhat obliterating pattern; scutellum grayish brown, spines and hair brass-colored to dark brown; metanotum brown to black, silvery pollinose; pleural tuft brass-colored; wings hyaline, veins light brown, spines and hair black; stem of halteres brownish, their knob yellowish; legs light brown with some regions darkened as shown in hind leg figure J, and male legs (Figs. O, P, Q); their hair silvery to black; abdomen (Fig. B) black; 1+1 large, silvery gray spots on sides of tergite II, and 1+1 smaller and more medially situated spots on tergite III; tergites IV and V entirely black except a narrow line laterally on hind margin, very rarely with traces of paired very narrow gray spots, as shown in figure C; tergite VI white except at sides and forward-pointing, triangular, central, posterior, dark gray spot; tergites VI-IX black, with lateral light-colored areas diminishing in size toward posterior segments; hair of abdomen, brass-colored, rarely with silvery tinge; frons as shown in figure D; fronto-ocular triangle approximately as long as wide (Fig. E); last segment of maxillary palp (Fig. F) more than twice as long as penultimate segment, and slightly longer than two preceding segments combined;

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diameter of sensory vesicle at least equal to half width of third segment; mandibles (Fig. G) with about 40 teeth; lacinia (Fig. H) with 25-29; scutum with isolated long hair and numerous short, appressed setae; Sc with few setae on its base, and from one to eight setae on its central portion; basal portion of R glabrous; R1 with hair and spiniform setae arranged in several rows; Rs with one or two irregular rows of setae (Fig. I); shapes and proportions of hind segments of leg as shown in figure J; calcipala distinct, approximately as long as wide at its base (Fig. K); claws with subbasal tooth well-developed (Fig. L); eighth sternite with about 25 long hairs on each side (Fig. M); hind border distinctly pigmented; gonapophyses subtriangular, beset with microchaetae, their inner border slightly sclerotized; paraproct and cercus as shown in figure N; genital fork with median process distinctly sclerotized except its apex which is widened; anterior projections narrow, with wide, medially directed expansions. Male: Wing length 3.8 mm; head blackish; eyes, occiput, antenna, and maxillary palp dark brown, almost black; clypeus black, gray pollinose; occiput, clypeus, and basal segments of maxillary palp with numerous long hair, latter also in row between eyes; scutum black, with appressed setae that appear to be golden to black; scutellum light brown, with brown to black hair; metanotum dark brown, golden pollinose; pleura dark brown, silvery pollinose; legs brown, with hair golden to black; dark regions distributed as in figures O, P, Q; abdomen velvety black, its hair golden to black; tergites II and VI with 1+1 silvery gray spots, those of second segment larger and more brilliant, those of sixth segment darker, with bluish tinge; last segment of maxillary palp more than twice as long as penultimate; diameter of sensory vesicle smaller than half of diameter of third segment; wings like those of female, but center of Sc only occasionally with setae, and hair and spiniform setae on R1 slightly less numerous; hind basitarsus 3.8 times as long as wide; calcipala well developed, about as long as wide at base (Fig. R); genitalia as shown in figures S, T; gonocoxite subtrapezoidal, slightly wider than long; gonostylus subquadrate, distal portion widely concave, smooth (Fig. S); central portion of ventral plate salient posteriorly at middle, with numerous curved setae; lateral projections and basal margin distinctly sclerotized (Fig. T); disc of ventral plate with 1+1 translucent areas; endoparameres membranous, with strongly pigmented spines. Pupa: Cocoon shoe shaped (Fig. U), with large, anteroventral bridge; color of cocoon light brown; surface smooth, closely woven; threads not individualized; rim of aperture slightly reinforced; gills protected laterally by walls of cocoon, only some of dorsal filaments free; length of cocoon at dorsum along middle, 4.2 mm; maximum length along base, 6.0 mm; length of body of pupa, 3.8 mm; length of gill 1.9 mm; gills (Fig. V) each with eight almost parallel filaments, slightly converging toward their apices; head and thorax of pupa light brown; head with 3+3 simple, short, frontal, 1+1 long, mostly branched facial, and 1+1 simple long, ocular trichomes (Fig. W); thoracic region adjacent to aperture of cocoon with band of tubercles and numerous interspersed, very short and stout trichomes (Fig. X); several smaller tubercles posteriorly on covered portion of thorax; disc of thorax on each side with approximately 20 additional trichomes; their size and exact distribution individually variable, in some cases bifid, in latter case generally longer than others; toward ventral surface, along band of tubercles, additional trichomes, similar to those described above, but more delicate; total number of thoracic trichomes,

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approximately 60+60; abdominal tergites light brown, intensity of pigment diminishing toward posterior segments; chaetotaxy as shown in figure Y. Larva: Maximum length, 6.0 mm; width of head capsule, 0.37 mm; color of larva whitish; cephalic apotome light-colored, darker along middle and at hind margin (Fig. Z); cuticle with isolated short hair, more numerous toward posterior end; antennae as shown in figure a, light brown, distal segment darkest; medial segment with two slight constrictions, regions of constrictions with unpigmented areas; ratio of proximal, medial and distal segments = 1:1.2:0.8; cephalic fan with 37-40 rays; toothing of mandibles as shown in figure b; two outer, one strong apical, three approximately identical subapical, five internal and two marginal teeth, first of which much larger than second; anterior border of hypostomium strongly pigmented (Fig. c); median tooth less prominent than lateral ones; intermediate teeth gradually increasing in size from II to IV; sublateral tooth barely perceptible; lateral borders rugose, with four to five very small serrations; hypostomial setae arranged in two irregular rows, about 14 in each group; disc of hypostomium glabrous; postgenal cleft (Fig. d) deep, pointed, its depth much larger than postgenal bridge; anal sclerite base with simple to trifid scales in large numbers; anal ring with about 85 rows, with approximately 15 hooks each; rectal papillae with three simple lobes. Bionomics: The pupae were found attached to roots of the bamboo Chusquea culeou DESVAUX (common name, “caña colihue”) trailing in the water, at about 10 cm under the surface. They live in torrential creeks at the base of the Andean mountains. Distribution: Argentina: Chubut, Neuquen, Río Negro, Tierra del Fuego; Chile: Aysen. Discussion: This species is very similar to S. barbatipes in the female abdominal ornamentation, but the 1+1 lighter spots of tergite III are smaller in S. nemorale, and in S. barbatipes the pupa has 6 branches and the thoracic trichomes number about 120. Simulium (Pternaspatha) barbatipes (ENDERLEIN) (Fig. 40.) Acropogon barbatipes ENDERLEIN, 1934b: 277; STONE, 1962: 206. Simulium barbatipes: SMART, 1945: 501; VARGAS, 1945a: 119; VARGAS & DIAZ; 1953b: 141. WYGODZINSKY, 1953b: 321; 1958: 133; STONE, 1962: 206; 1963: 17. [Simulium (Pternaspatha)] barbatipes STONE, 1963: 1. Simulium (Pternaspatha) barbatipes WYGODZINSKY & COSCARÓN, 1967: 99; COSCARÓN & WYGODZINSKY, 1972b: 220; COSCARÓN, 1991: 136-137.

Female: Wing length 3.1-3.9 mm; head black; occiput, frons, and clypeus gray pollinose; palpi and antennae black, latter with two basal segments more or less conspicuously orange-colored; scutum as usual in group; medium, narrow, longitudinal, white line distinct; appressed setae of scutum very pale brass-colored; scutellum and metanotum piceous, former with long, pale, brass-colored or dark hair; pleura gray pollinose; pleural tuft pale brass-colored; base of halteres dark, remainder yellowish white; legs light brown, their hair silvery to black; relative intensity of pigmented areas as shown in figures F, G, H; abdomen dark gray, tergites II-V deep black at center; tergite II with 1+1 large, silvery white spots; tergites III-V with 1+1 small spots, those on III largest, those of IV and V very small, in some cases difficult to perceive; tergite VI extensively silver-gray, black at lateral extremities, in some cases very faintly darkened at center; tergite

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VII dark, light gray at sides; tergites VIII and IX entirely dark; hair of abdomen, including that of basal fringe, silver-colored (Fig. A); frons as shown in figure B, with obsolete median sulcus; fronto-ocular triangle (Fig. C) slightly wider than long; last segment of maxillary palp (Fig. D) more than twice as long as penultimate segment; diameter of sensory vesicle approximately half that of third segment of maxillary palp; structure of vesicle as shown in figure E; maxillae with about 25 teeth, mandible with 40-47, teeth; Sc with few setae on central portion; setae and spines on R1 in single row, setae on R in single series; legs as in figures F, G, H; calcipala small, about as long as wide at base (Fig. I); claws with small but distinct tooth Fig. J); eighth sternite as in figure K; its lateral portions more heavily pigmented than central area, latter with posterior dark band; gonapophyses subtriangular, membranous, beset with microchaetae, inner border faintly sclerotized; paraprocts and cerci as shown in figure L; cerci much higher than wide, their distal border rounded; protruding portion of paraprocts subtriangular, their apex rounded; median process of genital fork strongly sclerotized, its apex widened, light-colored; anterior projections slightly darker than arms of fork; spermatheca globular, its inner surface with minute spicules arranged in groups. Male: Wing length 3.4-4.0 mm; head blackish; eyes, antennae, and palpi dark brown, almost black, first two and extreme base of third antennal segments lighter, all with silvery pilosity; palpi, basal two antennal segments, clypeus, line between eyes, and occiput all with long black hair; scutum as usual for group; appressed setae brass-colored; scutellum brown, with brass-colored hair; metanotum and pleura dark brown, silvery pollinose; pleural tuft brass-colored; legs whitish to brown, distribution and intensity of pigment as shown in figures M, N, O; abdomen velvety black; its hair, including that of basal fringe, silvery; segment II with 1+1 large, silvery white spots, segments VI and VII with 1+1 smaller ones (Fig. P); last segment of maxillary palp more than twice as long as penultimate; diameter of sensory vesicle approximately half that of third palpomere; wings like those of female, but Sc devoid of setae; hind basitarsus 3.4-3.9 times as long as wide; calcipala slightly shorter than wide at base (Fig. Q); genitalia very similar to those of Simulium nemorale. Pupa: Cocoon slipper shaped (Fig. R), with very narrow anteroventral bridge; rarely shoe shaped; color of cocoon brown; surface smooth, closely woven, threads not distinctly individualized; rim of aperture slightly reinforced; only extreme base of gills protected by lateral walls of cocoon, all filaments free; length of cocoon at dorsum, 3.03.6 mm; maximum length along base, 4.2-5.1 mm; length of pupa, 3.5-4.0 mm; length of gill, approximately 2.0 mm; gills (Fig. S) each consisting of six closely parallel filaments; three primary branches arising from short basal trunk, each branch dividing into two filaments not far from its base; exact level of branching somewhat variable; head and thorax of pupa dark brown; head with 3+3 branched frontal, 3+3 branched facial, and 1+1 branched ocular, trichomes (Fig. T); thorax showing zone adjacent to aperture of cocoon with numerous tubercles of various sizes and numerous trichomes of different structure; short, simple or bifurcate, spatulate, flattened, in some cases curved, in others hair like, with from one to six long and delicate filaments (Figs. U, V); total number of trichomes, 110-130+110-130; chaetotaxy of abdomen like that of S. nemorale. Larva: Maximum length, 8.0 mm; maximum width of head capsule, 0.7 mm; larva with extensive hypodermal pigment on dorsal and on anterior half of ventral surface of

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Figure 40. Simulium (Pternaspatha) barbatipes. A-L: Female; A: abdomen, dorsal view; B: frons; C: fronto-ocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg, respectively; I: apex of hind basitarsus and tarsomere I; J: claw; K: eighth sternite and gonapophyses; L: cercus and paraproct. M-Q: Male; M-O: fore, mid and hind leg, respectively; P: abdomen, dorsal view; Q: apex of hind basitarsus and tarsomere I. R-V: Pupa; R: lateral view; S: gill; T: frontoclypeus; U: exposed portion of thorax and trichomes; V: trichomes and tubercles of thorax. W-Z & a-c: Larva; W: cephalic apotome; X: antenna; Y: teeth of mandible; Z: hypostomium a: cephalic capsule in ventral view; b: anal sclerite; c: rectal papillae.

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body; cephalic apotome yellowish, with dark pattern as shown in figure W; body integument glabrous except some short setae at base of anal sclerite; antennae dark brown, distal segment darkest; ratio of proximal, medial and distal segments = 1:1.2:0.72 (Fig. X); cephalic fan with approximately 40 rays; toothing of mandible as shown in figure Y; hypostomium with median tooth a little more prominent than lateral ones; lateral borders with four or five small serrations; hypostomial setae arranged in two irregular series, their number about 10 in each group (Fig. Z); postgenal cleft (Fig. a) triangular, its depth larger than length of hypostomial bridge; anal sclerite as shown in figure b, its base with numerous small scales; anal ring with about 80 rows, each composed of 13-14 hooks; rectal papillae (Fig. c) consisting of three primary lobes, middle one with six, lateral ones with two diverticula each. Distribution: Argentina: Jujuy, Mendoza, Neuquén, Salta, Tucuman; Chile: Aconcagua, O’Higgins, Santiago; Bolivia: La Paz; Peru: Chocó, Cuzco, Junin, Puno. Bionomics: Simulium barbatipes breeds in torrential high-mountain streams of relatively low temperatures (10-12 °C). It is also found in irrigation ditches with fastflowing water. The larvae and pupae are attached to vegetation trailing in the current or to the surface of rocks. Discussion: Simulium barbatipes is one of the most wide-ranging species of the Pternaspatha group. Although there is some individual variation, no characters were found that would separate the various populations that were examined. Simulium (Pternaspatha) philippii COSCARÓN (Fig. 41.) Simulium (Pternaspatha) philippii COSCARÓN, 1976a: 29-33; 1991: 125-126.

Female: Wing length 2.9-3.4 mm; head blackish, antenna, palp and proboscis dark brownish; scutum as in Simulium nemorale; legs light brown with apex of segments and most of tarsi dark brown; abdomen grayish brown, blackish on median area of tergites, with 1+1 whitish gray submedian spots on tergites III-V, larger on tergite II and extending sublaterally on tergites VI-VIII (Fig. A); fronto-ocular triangle deep (Fig. B); lacinia with about 20 teeth and mandible with 30-34 teeth; maxillary palp as in figure C, with enlarged sensory vesicle with abundant sensilla (Fig. D); short calcipala (Fig. E); claw without subbasal tooth (Fig. F); eighth sternite and gonapophyses similar to those of Simulium stelliferum; paraprocts relatively elongated, not acuminate distally (Fig. G). Male: Wing length 3.1 mm; antenna and palp coloration as in female; thorax and legs with golden to grayish brown pilosity; abdomen with 1+1 silvery spots on tergites II, VI and VII, the two latter smaller; calcipala small (Fig. H); ratio of length/width of hind basitarsus = 3.5; gonostylus without tubercles (Fig. I). Pupa: Cocoon length on dorsum 2.5-2.8 mm; base 3.0-3.2 mm; gills 2.4-2.8 mm; pupa body 3.0 mm; cocoon slipper shaped, scarcely projected ventrally and with reinforced border (Fig. J); gill with 6 branches disposed as shown in figures L, M; head and thorax with abundant tubercles; on frontoclypeus with about 1200, those of thorax bigger than on head; frontoclypeus with 2+2 frontals and 1+1 facial trichomes relatively short, uni- or biramous (Fig. K); thorax with 60-85 trichomes on each side, filiform, with 1-3 branches. (Figs. M, N); abdominal chaetotaxy as in Simulium nemorale.

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Larva: Length 7.5 mm; color light grayish brown-green; head dark brown; cephalic apotome more darkened on base (Fig. O), central spots negative; antenna with distal segment more darkened (Fig. P); ratio of proximal, medial and distal segments = 1:1.21.3:0.8; cephalic fan with 42-52 rays; toothing of mandible as in figures Q, R; hypostomium with apical border well pigmented, with median teeth as high as lateral teeth; hypostomial setae 7-11 in number on each side and postgenal bridge as long as hypostomium length (Fig. S); anal sclerite without scales; anal ring with 83-93 rows with 13-15 hooks in each; rectal papillae with 3 single lobes (Fig. U); comb of thoracic proleg sclerite with about 17 groups of setae of different length (Fig. T). Distribution: Chile: O’Higgins, Tarapaca. Bionomics: Simulium philippii breeds in creeks with high current and cold waters (about 10 °C) in the southern and temperate areas of the northern region. The feeding habits of the females are unknown. E C

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Figure 41. Simulium (Pternaspatha) philippii. A-G: Female; A: abdomen, dorsal view B: fronto-ocular triangle; C: maxillary palp; D: sensory vesicle of maxillary palp; E: apical portion of hind basitarsus and tarsomere I; F: claw; G: cercus and paraproct. H-I: Male; H: apical portion of hind basitarsus and tarsomere I; I: gonostylus. J-N: Pupa; J: lateral view; K: trichomes and tubercles of frontoclypeus; L: basal portion of gill; M: exposed portion of thorax with trichomes and gill; N: trichomes and tubercles of thorax. O-U: Larva; O: cephalic apotome; P: antenna; Q-R: teeth of mandible in different specimens; S: hypostomium and postgenal cleft; T: lateral sclerite of proleg; U: rectal papillae.

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Discussion: On the basis of the abdominal ornamentation, the closest species to S. philippii is S. nigristrigatum, but in S. nigristrigatum the light spots on tergites III-V are larger, the calcipala is absent, the subbasal claw teeth are well developed, the pupal gill has 8 filaments, and the thoracic trichomes are multiramous. Simulium (Pternaspatha) herreri WYGODZINSKY & COSCARÓN (Figs. 42 A-Z, a-f.) Simulium (Pternaspatha) herreri WYGODZINSKY & COSCARÓN, 1967: 104-109; COSCARÓN, 1991: 130.

Female: Wing length 2.8-3.5 mm; head black; eyes piceous; antennae, palpi, and labium black, first two antennal segments tinged with gray; occiput, frons, and clypeus gray pollinose; pilosity of antennae very short; basal segments of palpi, clypeus, frons, and occiput with some long dark hair; scutum as usual for group; dark regions velvety black; appressed setae of scutum brass-colored, not very dense, not obliterating pattern; scutellum grayish brown, with long hair brass-colored to brown; metanotum, pleura, and sterna piceous, silvery pollinose; legs light brown, pigment areas as shown in figures F, G, H; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spots, 1+1 smaller ones on tergite III, 1+1 very small ones each on tergites IV and V; tergite VI with transverse, silvery white, posteriorly emarginated band on central two-fourths; tergites VII and VIII grayish or silvery white on sides; tergites II-VI narrowly banded with grayish on lateral portions of hind margin; hair of fringe of first segment yellowish white, that of rest of abdomen brass-colored to very dark; frons as shown in figure B; fronto-ocular triangle about as long as wide (Fig. C); last segment of maxillary palp (Fig. D) slightly more than twice as long as penultimate segment, and as long as third and fourth segments combined; diameter of sensory vesicle about equal to half diameter of third palpomere, its surface structure as shown in figure E; lacinia with 26 or 27 teeth, mandible with 36-40, teeth; scutum with appressed setae on disc; posterior declivity with erect or forwardly curved longer hair; Sc with three or four hairs on central portion; R1 with hair and spiniform setae arranged in one row, in some cases becoming double on apical portion of vein; leg as shown in figures F-H; calcipala (Fig. I) small but distinct, about as long as wide; claws with well-developed tooth (Fig. J); eighth sternite pigmented, central posterior portion as in figure K, gonapophyses subtriangular, rounded apically, their inner border slightly sclerotized, their disc with very numerous microtrichia; paraprocts and cerci as shown in figure L; projecting portion of paraprocts semi elliptical; cerci with distal border truncate. Male: Wing length 3.0-3.2 mm; head black; eyes purple; occiput, antennae, and palps black, clypeus black, gray pollinose; occiput, clypeus, and basal maxillary segments with numerous very long hair, latter also arranged in row between eyes; scutum as usual for group, deep velvety black, on its anterior three-fourths a very delicate, median, longitudinal line; hair brass-colored; scutellum black, its hair brass-colored; metanotum, pleura, sterna, as in female; legs yellowish brown, their hair brass-colored to black; relative intensity of pigment areas as shown in figures M, N, O; abdomen (Fig. M) velvety black, hair silvery gray; segment II with 1+1 very large, silvery white spots closely approaching but not meeting dorsally; segment V in some individuals very faintly iridescent, at certain angle showing small, silvery white area dorsally at center; VI entirely silvery white except small dark area dorsally at center posteriorly, VII with 1+1 medium-sized, silvery white

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Figure 42. Simulium (Pternaspatha) herreri. A-L: Female; A: abdomen, dorsal view B: frons; C: frontoocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg, respectively; I: apical portion of hind basitarsus and tarsomere I; J: claw; K: portion of eighth sternite and gonapophyses; L: cercus and paraproct. M-S: Male; M: abdomen in lateral view; N: antenna; O: maxillary palp; P: anterior veins of wing; Q: hind leg; R: apical portion of hind basitarsus and tarsite I; S: gonostylus. T-Y: Pupa; T: lateral view; U: gill; V: frontoclypeus; W: frontoclypeus tubercles with high magnification; X: exposed portion of thorax and trichomes; Y: thoracic trichomes with high magnification. Z, a-f: Larva; Z & a: cephalic apotome in different specimens; b: antenna; c: teeth of mandible; d: hypostomium; e: hypostomium and postgenal cleft; f: rectal papillae.

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spots widely separated above; IX faintly iridescent; eyes separated by series of long hairs; scape and pedicel with long hair (Fig. N) as also basal segments of maxillary palp (Fig. O); diameter of sensory vesicle of palp half that of third palpomere; wings without setae on central portion of R1 with one row of setae intermixed with hair (Fig. P); shapes and proportion of segments of hind legs as shown in figure Q; hind basitarsus 3.2 times as long as wide; calcipala (Fig. R) small but distinct, slightly wider than long; legs with numerous long setae; gonocoxite wider than long; gonostylus subquadrate, distal portion widely concave, not tuberculate (Fig. S); ventral plate with 1+1 distinct, translucent areas. Pupa: Cocoon (Fig. T) slipper shaped, with short anteroventral bridge; color light brown, surface smooth, closely woven, threads not individualized under medium magnification; rim of aperture not or only feebly reinforced; length of cocoon at dorsum, 2.5-3.0 mm; maximum length along base, 4.0 mm; average length of pupa, 3.5 mm; length of gills 2.5-3.0 mm; gills (Fig. U) each with six filaments, dorsal and median primary branches and their filaments forming tight bundle, ventral primary branch and its filaments sharply curved ventrally and backward, filaments almost entirely hidden by cocoon; head and thorax of pupa light brown; head with 1+1 fields of tubercles at base of clypeus (Figs. V, W), 2+2 frontal and 1+1 facial trichomes, all simple, facial trichomes much longer than frontal trichomes; thoracic region adjacent to aperture of cocoon with band of tubercles that extends to midline ventrally and attains base of gill; most of remainder of exposed surface of thorax covered with trichomes in shape of elongate, lanceolate setae (Figs. X, Y), their length, 0.07-0.10 mm, their total number approximately 250+250; few fine hair near insertion of gills; chaetotaxy of abdomen on tergite I with 2+2 lateral setae; tergite II with 3+3 relatively small, equidistant spines (acrostichal, dorsal and dorsocentral), and in some cases also dorsolateral one, one or two anterior dorsocentral and approximately 3+3 lateral setae also present; tergites III and IV with 4+4 large, spinelike setae, dorsolateral separated from dorsocentral by relatively wide space; tergites III and IV also with 1+1 anterior dorsocentral and 2+2 lateral setae; tergite V with 1+1 small, dorsal and 1+1 dorsolateral setae, as well as with 1+1 small, anterolateral groups of small, scale like structures; tergites VI-VIII at anterior border with 1+1 rows of spiniform, simple, bifid, or trifid tubercles, last transformed laterally into small, scale like structures arranged in several irregular rows and occasionally with 1+1 dorsocentral setae; rows of tubercles on tergites VI and VII widely separated from one another, those of VIII closely approximated, almost confluent; numbers of tubercles in rows 6-13+6-13 on tergites VI and VII, 10-13+10-13 on VIII; anterior border of tergite IX in some cases with 1+1 groups of one to three tubercles; sternite III with 1+1 lateral and 1+1 ventral setae, central region with transverse area of numerous small and very delicate, scale like structures; sternite IV with 3+3 or 4+4 ventral and lateral setae, and central area of scale like structures as on III; sternites V-VII with 2+2 spines before posterior margin; spines of each pair close to one another on fifth, separated by wider space on both sixth and seventh segments; spines of V generally bifid, but in some cases ventrolateral one simple; on VI and VII, generally ventral spine bifid and ventrolateral one simple, but in some both simple; sternites V-VII each with 1+1 suboval fields of scale like structures; VIII with one transverse group of scales near anterior margin. Larva: Maximum length, 7.0 mm; width of head capsule, 0.60-0.65 mm; color variable; dark hypodermal pigment generally present on dorsal and ventral of anterior

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half of body and on dorsal surface of posterior half; cephalic apotome with two different basic color patterns, as illustrated in figures Z, a; cuticle with isolated short hair, becoming more numerous toward posterior end; antennae as shown in figure b; ratio of proximal, medial and distal segments = 1:1.1-1.25:1.0-1.08; cephalic fan with approximately 40 rays; toothing of mandibles as shown in figure c; hypostomium with median tooth as prominent as lateral ones; central intermediate tooth smallest; lateral serration well developed, irregular in shape; hypostomial setae arranged in two irregular rows, with average of 11 setae in each group (Fig. d); postgenal cleft rather wide (Fig. e), its depth not much larger than length of postgenal bridge; anal sclerite with scales at its base small and few in number; anal ring with approximately 90 rows, of about 15 hooks each; central lobe of rectal papillae with 1+1, lateral lobes each with one secondary diverticula (Fig. f). Distribution: Peru: Lima. Bionomics: The larvae and pupae S. herreri live at 2300-2500 m altitude on trailing grass leaves and rocks in a small stream and irrigation ditches, at 1-3 cm below the water surface. Pupae frequently form small aggregations. The temperature in the watercourses was 15 °C. Discussion: Simulium herreri, S. albilineatum, and S. yacuchuspi are very similar. The male of S. herreri differs from males of the two other species by the absence of paired light-colored spots on tergite III. The female of S. herreri is distinguished by the shape of the paraprocts and the light-colored central portion of the eighth sternite. The pupa is characterized by the sharply divergent filaments of the ventral primary branch of the gill, the conspicuous areas of tubercles at the base of the frontoclypeus, and the lanceolate trichomes of the thorax. The larvae of S. herreri have the postgenal bridge about as long as the postgenal cleft and in S. albilineatum and S. yacuchuspi it is much shorter, about a half as long as the postgenal cleft. Simulium (Pternaspatha) yacuchuspi WYGODZINSKY & COSCARÓN (Fig. 43.) Simulium (Pternaspatha) yacuchuspi WYGODZINSKY & COSCARÓN, 1967: 110-115; COSCARÓN, 1991: 127-128.

Female: Wing length 3.5-3.8 mm; pilosity and color of head and thorax as in Simulium herreri; legs light brown, darkened on apices of femur of tibia and tarsi; their hair silvery to black; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spots, tergite III with 1+1 smaller ones, tergites IV and V each with 1+1 very small ones; tergite VI entirely silvery white, very faintly darkened posteriorly at center; VII black on central half, white on lateral fourths; tergite VIII black, silvery white only at lateral extremities; tergite IX grayish black; tergites IV and V narrowly banded with grayish on lateral portion of hind border; hair of basal fringe and of remainder of abdomen silvery; frons as shown in figure B; fronto-ocular triangle about as long as wide; maxillary palp like that of Simulium herreri, but tubercles of sensory vesicle (Fig. C) relatively larger and somewhat less numerous; lacinia with 22-27 teeth, mandible with 42-48, teeth; shape of hind leg as in figure D; calcipala distinct (Fig. E), about as long as wide; claws with very small tooth (Fig. F); eighth sternite and gonapophyses like those of Simulium albilineatum; paraprocts and cerci as shown in figure G; projecting portion of paraprocts rather narrowly subtriangular. Male: Wing length 3.0-3.5 mm; color of head and its appendages like that of Simulium herreri; color of thorax like that of S. herreri, but knob of halteres yellow; legs pale

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yellow-brown with darkened area as in female but more extended on hind basitarsus, their hair silvery white to black; abdomen (Fig. H) velvety black; hair, including that of basal fringe, generally silvery, rarely brass-colored; segment II with 1+1 very large, silvery white spots closely approaching but not meeting dorsally; segment III with 1+1 small, silvery white spots laterally at hind margin; segment VI almost entirely silvery white except small dark area at center behind; VII with 1+1 medium-sized, lateral, silvery white spots; segment IX faintly iridescent; shape and proportions of segments of hind legs as in figure I; hind basitarsus 2.7-3.0 times as long as wide; calcipala (Fig. J) very small but distinct, about as long as wide; genitalia much like those of Simulium herreri. Pupa: Cocoon slipper shaped (Fig. K) with well-developed anteroventral bridge; color light brown, translucent, surface smooth, texture closely woven, threads not perceptible under low magnification; rim of aperture feebly reinforced; length of cocoon at dorsum along middle, 2.8-3.4 mm; maximum length along base, 4.1 mm; length of body of pupa 3.2-3.4 mm; length of gills 2.2-2.5 mm, viz., half as long as cocoon, and twothirds as long as body of pupa; gills (Figs. L, M) each with six filaments arising from three primary branches; dorsal and median primary branches and their filaments lying closely together, ventral primary branch sharply ventrally bent, its ventral filament curved backward under cocoon, its dorsal filament generally curved forward, but not closely associated with remaining filaments; all primary branches with division into filaments not far removed from base of gill; division of dorsal primary branch closest to, and either that of median or ventral branch most remote from, base of gill; dorsal primary branch thickest, ventral primary branch thinnest; dorsal filament of dorsal primary branch conspicuously thicker on basal portion than any other; head and thorax of pupa dark brown; head (Fig. N) with (1-)2+2(-1) frontal and 1+1 facial trichomes, latter frequently longer, and generally bifid; thoracic region adjacent to aperture of cocoon with band of not very numerous tubercles, becoming evanescent toward sides of thorax; smaller tubercles also occupying large area of dorsal portion of part of thorax covered by cocoon; exposed surface of thorax with large and very variable number of simple (rarely bifid) trichomes, longest and most delicate near band of tubercles on lateral portion of thorax, diminishing in size toward dorsum and lateral disc of thorax (Figs. O, P); trichomes mostly slender, hair like (Fig. Q), but in some specimens trichomes on disc becoming very short and pointed, almost spinelike; length of trichomes varying from 0.02 to 0.16 mm., most frequently around 0.1 mm; total number of trichomes varying from 50+50 to 130+130; type and number of trichomes invariably closely similar on both halves of thorax in one individual; chaetotaxy of abdomen like that of Simulium herreri, but spiniform tubercles on tergite VIII generally forming continuous row. Larva: Maximum length, 7.0 mm; width of head capsule, 0.65-0.70 mm; larvae color brownish green (in alcohol) on whole body except posterior fifth of ventral surface; larvae appearing black in life; head very variable in color, extremes of pattern of cephalic apotome shown in figures R, S.; dark form prevalent; cuticle with few scattered short hair; antennae as shown in figure T, ratio of proximal, medial and distal segments = 1:1.11.3:0.75-0.9; cephalic fan with 40-46 rays in large fan; toothing of mandibles (Fig. U) much like that of Simulium herreri; structure and pigmentation of hypostomium as shown in figure V; lateral serrations well developed, irregular in shape; hypostomial setae

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Figure 43. Simulium (Pternaspatha) yacuchuspi. A-G: Female; A: abdomen, dorsal view B: frons; C: sensory vesicle of maxillary palp; D: hind leg; E: apical portion of hind basitarsus and tarsomere I; F: claw, in different position; G: cercus and paraproct. H-J: Male; H: abdomen in lateral view; I: hind leg; J: apical portion of hind basitarsus and tarsomere I. K-Q: Pupa; K: lateral view; L-M: gill; N: frontoclypeus; O-P: exposed portion of thorax with different number of trichomes; Q: thoracic trichomes with high magnification. R-X: Larva; R-S: cephalic apotome in different specimens; T: antenna; U: teeth of mandible; V: portion of hypostomium; W: hypostomium and postgenal cleft; X: rectal papillae.

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arranged in two irregular rows, with average of 15 setae in each group; region of insertion of setae distinctly less pigmented than remainder of disc of hypostomium; postgenal cleft relatively narrow, pointed apically, its depth larger than length of postgenal bridge (Fig. W); anal sclerite with small scales at its base and few in number; anal ring with 81-89 rows, each composed of 12-20 hooks; central and lateral lobes of rectal papillae each with 16-20 very long and slender secondary lobules arranged in two to three irregular rows (Fig. X). Distribution: Peru: Junin. Bionomics: The larvae and pupae of S. yacuchuspi were found at 3750 m altitude, attached to leaves and stems of grasses trailing in a small stream, near the surface; the stream was not more than 50 cm wide and not more than 10 cm deep, and the temperature of the water was 10 °C. Discussion: Simulium yacuchuspi shows great variation in the number and size of the trichomes of the pupal thorax. This species is closely related to S. herreri, S. albilineatum, and S. albicinctum. The larva differs from that of S. herreri and S. albilineatum (that of S. albicinctum is unknown) mainly by the large number of diverticula of the rectal papillae. The pupa of S. yacuchuspi differs from that of S. herreri (with which it shares the backwarddirected ventral filament of the ventral primary branch of the gill) by the thicker lanceolate-shaped thoracic trichomes in S. herreri, and from that of S. albilineatum by the virtual absence of bifid trichomes on the thorax and the presence of the abovementioned backward-directed ventral filament. The adults are more difficult to distinguish with certainty. The male of S. yacuchuspi differs from that of S. albicinctum by the presence of 1+1 white spots on abdominal tergite III, and a less strongly widened hind basitarsus. It differs from that of S. herreri by the presence of the spots and the absence of such spots from segment V, but it cannot be separated from the male of S. albilineatum. The female of S. yacuchuspi can be recognized by the structure of its paraprocts. It differs from that of S. albilineatum by the white spots on tergite V which are much smaller than those of tergite III, and from that of S. herreri also by the dark central portion of the eighth sternite, the poorly developed tooth of the claw, and the almost completely white tergite VI. The female of S. yacuchuspi is rather similar to that of S. albicinctum, but the latter lacks white spots on tergites IV and V, tergite VI is entirely white, the dark area at the base of the posterior tibia is more extended, and the calcipala is wider. Simulium (Pternaspatha) albilineatum (ENDERLEIN) (Fig. 44.) Pternaspatha albilineata ENDERLEIN, 1936: 115. Simulium albilineata SMART, 1945: 500. Simulium albilineatum VARGAS, 1945a: 109. Simulium (Pternaspatha) albilineatum WYGODZINSKY & COSCARÓN, 1967: 95-99; COSCARÓN, 1991: 126-127.

Female: Wing length, 3.5-3.7 mm; color of head and thorax like that of Simulium herreri; legs yellowish brown, their hair from silvery to black; color pattern, illustrated in figures C, D, E; abdomen (Fig. A) black; tergite II with 1+1 large, silvery white spot; tergites II and V with 1+1 smaller ones; tergite IV either entirely black or with 1+1 minute whitish spots; tergite VI silvery white darkened only at lateral extremities; tergite VII dark at middle, silvery white on lateral thirds, dark region widened posteriorly; tergite VIII black, silvery only

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at lateral extremities; tergite IX silvery to black; tergites II and III narrowly banded with grayish on lateral portions of hind margin (not shown in illustration); hair of fringe of first segment yellowish white, that of rest of abdomen brass-colored to very dark; fronto-ocular triangle (Fig. B) about as long as wide; lacinia with 22-28 teeth, mandible with 40-45 teeth; shape of legs as shown in figures C-E; calcipala (Fig. F) small but distinct, somewhat pointed, about as long as wide; subbasal projection of claw very small (Fig. G); eighth sternite with central portion lighter than lateral portions (Fig. H); gonapophyses subtriangular, rounded or pointed apically, their inner border slightly sclerotized; paraprocts with projecting portion angular; cerci with distal border rounded. Male: Wing length 3.5-3.6 mm; color of head, thorax and legs as in female but the latter more darkened; abdomen (Fig. I) velvety black; hair, including that of basal fringe, light brass-colored; segment II with 1+1 very large, silvery white spots closely approaching but not meeting dorsally; segment III with 1+1 small, silvery white spots laterally at hind margin, these spots in some cases faint; segment VI almost entirely silvery white except small dark area at center posteriorly; VII with 1+1 medium-sized, silvery white spots widely separated above; IX faintly iridescent; shape of legs as shown in figures J-L; hind basitarsus 3.0-3.2 times as long as wide; calcipala (Fig. M) small but distinct, slightly wider than long; legs with numerous long setae as illustrated; genitalia as for genus in general shape, but gonostylus distinctly wider than long (Fig. N). Pupa: Cocoon (Fig. O) like wall pocket or slipper shaped, with very short anteroventral bridge; color light brown, translucent; surface smooth, closely woven, threads not individualized under medium magnification; rim of aperture feebly reinforced; length of cocoon at dorsum 3.5-4.0 mm; maximum length along base, 5.0 mm; length of pupa, 3.0-3.5 mm; length of gill 2.5-3.0 mm; gills each with six filaments, filaments spreading and forming fanlike structure; ventral primary branch and its two filaments somewhat bent downward (Fig. P); head and thorax of pupa dark brown; frontoclypeus without tubercles; head with 2+2-3+3 short frontal, single or bifid, and 1+1 longer facial, trichomes (Fig. Q), latter generally bifid; thorax on region adjacent to aperture of cocoon with band of tubercles, becoming evanescent toward sides of thorax; tubercles also occupying large area of dorsal portion of that part of thorax covered by cocoon; exposed surface of cocoon with large number of simple or bifid trichomes (Fig. R), their length, 0.05-0.08 mm, their total number, 250-300+250-300; trichomes also present on region anterior to insertion of gill. Larva: Maximum length, 7.0 mm.; maximum width of head capsule, 0.7 mm; dark hypodermal pigment on dorsal and ventral surfaces of whole body; pattern of cephalic apotome as shown in figure S; cuticle with isolated short setae, slightly more numerous on caudal portion; antennae light brown, subapical segment darkest; ratio of proximal, medial and distal segments = 1:1.25:1; cephalic fan with about 40 rays; hypostomium median tooth as prominent as lateral ones; lateral serrations well developed, irregular in shape; hypostomial setae arranged in two irregular rows, with 10-12 setae in each group; postgenal cleft rather wide, much deeper than hypostomial bridge length (Fig. T); anal sclerite, with well-developed scales at base of sclerite; anal ring with approximately 65-70 rows, each composed of about 17 hooks; each lobe of rectal papillae (Fig. U) with diverticula arranged in two rows, from four to six diverticula on lateral, and from six to eight diverticula on central lobe.

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Distribution: Peru: Cuzco, Junin. Bionomics: The larvae and pupae of Simulium albilineatum were found on blades of heavy aquatic vegetation trailing in the stream. The temperature of the water was 10 °C, and the altitude was about 3700-4000 m. Discussion: Simulium albilineatum imagoes are similar to those of S. yacuchuspi and S. herreri but both have more reduced whitish areas on tergites III and V. The more convergent filaments of the gills and the structure of the thoracic trichomes distinguish C

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Figure 44. Simulium (Pternaspatha) albilineatum. A-H: Female; A: abdomen, B: fronto-ocular triangle; C-E: fore, mid and hind leg, respectively; F: apical portion of hind basitarsus and tarsomere I; G: claw; H: eighth sternite and gonapophyses. I-N: Male; I: abdomen dorsal view; J-L: fore, mid and hind leg, respectively; M: apical portion of hind basitarsus and tarsomere I; N: gonocoxite and gonostylus. O-R: Pupa; O: lateral view; P: gill; Q: frontoclypeus and trichomes; R: exposed portion of thorax and trichomes. S-U: Larva; S: cephalic apotome; T: hypostomium and postgenal cleft; U: rectal papillae.

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the pupa of S. albilineatum from the pupae of S. herreri and yacuchuspi; in the latter two, one or two filaments of the gills sharply diverge from the others, and the thoracic trichomes are thicker. Simulium bordai has very similar pupal characters but the thoracic trichomes have 2 to 6 branches and the number is lower (120-170) on each side. Simulium (Pternaspatha) bordai COSCARÓN & WYGODZINSKY (Fig. 45.) Simulium (Pternaspatha) bordai COSCARÓN & WYGODZINSKY, 1972b: 221-222; COSCARÓN, 1991: 124-125.

Female, Male and Larva: Unknown. Pupa: Cocoon as in Simulium albilineatum, viz., slipper shaped, with short anteroventral bridge, only slightly raised from substrate; cocoon light brown, its surface smooth, with threads not clearly perceptible; rim of aperture distinctly reinforced; length of cocoon dorsally, 4.5 mm; maximum length as seen from below, 5.6 mm; gill about 2.5 mm. long, with six filaments arising from three very short primary branches (Figs. A, B), with two filaments to each branch; filaments approximately parallel, only slightly diverging at base; dorsal filament of dorsal branch conspicuously wider on basal portion than remaining filaments; head and thorax dark brown; frontoclypeus (Fig. C) without tubercles; 3+3 frontal, 1+1 facial and 1+1 ocular trichomes, in shape of long bifid or trifid, very rarely simple, setae; facial trichomes inserted in distinct depression delimited by conspicuously sclerotized ridges; disc of thorax without tubercles, the latter restricted to area adjacent to aperture of cocoon and at insertion of gills (Fig. D); trichomes occupying disc of thorax, in shape of setae with three to six branches (Figs. E-F), from rather short and stout to long and slender, rather uniform or varied; number of trichomes approximately 120-170+120-170; their average length 0.09-0.11 mm; onchotaxy of abdomen as usual for subgenus. C

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E F Figure 45. Simulium (Pternaspatha) bordai. A-F: Pupa; A-B: gills; C: frontoclypeus; D: exposed portion of thorax; E: trichomes of disc of thorax; F: trichomes and tubercles of posterior margin of exposed portion of thorax.

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Distribution: Bolivia: La Paz to Yungas. Bionomics: Simulium bordai breeds in creeks of fast-flowing, crystalline and cold water, at about 4000 m altitude. Discussion: Simulium bordai is described and named even though only its pupa is known, because the latter is conspicuously different from that of any other described species. The pupa of S. bordai is similar in many ways with that of S. albilineatum, but the latter differs by the greater number of thoracic trichomes (250-300) that are simple or bifid, and the gill branches diverging immediately from the base. Simulium (Pternaspatha) quechuanum COSCARÓN & WYGODZINSKY (Fig. 46.) Simulium (Pternaspatha) quechuanum COSCARÓN & WYGODZINSKY, 1972b: 232-234; COSCARÓN, 1991: 135-136.

Female: Wing length 3.2-3.7 mm; color of head with appendages, thorax, wings, and halteres as in Simulium herreri; color pattern of legs as illustrated for S. hectorvargasi; abdomen (Fig. A) black, with 1+1 large silvery white spots on terga II, VI, and VII; terga III and V, and in some specimens also IV, with 1+1 smaller silver-white spots; posterior margins of these terga very narrowly white at sides; terga VIII and IX dark gray; frons and fronto-ocular triangle as in Simulium herreri; frontal angle 115 degrees; lacinia with 24-26 teeth, mandible with approximately 35 teeth; wings as in S. hectorvargasi, shapes and proportions of leg segments as in S. hectorvargasi; calcipala very small but distinct, wider than long as in male (Fig. C); claws with distinct tooth (Fig. B); genitalia similar to those of S. herreri. Male: Wing length 3.5 mm; color and structure extremely similar to those of Simulium hectorvargasi, but calcipala smaller (Fig. C); apex of gonostylus smooth. Intersex (= specimen with male and female characters): General aspect, color, and most morphological characters as in females, but genital fork shorter and stouter; legs with hind basitarsus distinctly widened, and claws of all legs as in male. Pupa: Cocoon (Figs. D, E) slipper shaped; anteroventral bridge distinct, rather salient; cocoon dark brown; surface smooth, but with individual threads distinctly perceptible with moderate magnification; rim of aperture distinctly reinforced; length of cocoon dorsally, 3.5 mm; maximum length along base, 5.0-5.5 mm; length of body 4.0 mm; of gills 2.0-2.2 mm; gills (Fig. F) each composed of six filaments arising from three main branches; dorsal and median primary branches and their filaments lying closely together, ventral primary branch sharply ventrally bent, its ventral filament curved backward under cocoon, its dorsal filament generally curved forward but not closely associated with remaining filaments; all primary branches with division close to base of gill, all about equidistant from base; head and thorax light brown; frontoclypeus (Fig. G) without tubercles; 2+2 frontal trichomes, extremely varied in shape, often different in same specimen, from very short, divided, spine-shaped, to elongate bristle-like, generally divided. 1+1 facial trichomes, also extremely varied in shape, from spinelike (although not as short as some frontal trichomes) to elongate hair like, with two branches, or simple; facial trichomes situated in depression delimited by distinct ridges; 1+1 ocular trichomes, hair like, with two branches; thorax with tubercles forming band along area adjacent to aperture of cocoon (Fig. F); trichomes of thorax very similar to those of Simulium

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barbatipes, from lanceolate, simple or divided, to elongate hair like (Fig. H), but most frequently lanceolate; number of thoracic trichomes from 120+120 to 180+180; onchotaxy of abdomen as usual for subgenus. Larva: Length of mature larva, 6.5-7.0 mm; width of head capsule, 0.6 mm; color much as in Simulium hectorvargasi, although somewhat darker; general structure of antennae as in S. hectorvargasi; ratio of proximal, medial and distal segments = 1:1.7:0.9-1.1; cephalic fan with 35-38 rays; toothing of mandibles generally with first marginal tooth much over twice as long as second (Fig. I), although in some individuals only twice as long as second; structure and pigmentation of hypostomium much as in S. hectorvargasi; hypostomial setae arranged in two or three irregular series, with 11-16 setae in each group; postgenal cleft (Fig. J) 2.0-2.5 times as deep as postgenal bridge; anal sclerite spicules or scales not perceptible; anal ring with 80-85 rows with about 14 hooks each; rectal papillae with three simple lobes. Distribution: Argentina: Jujuy, San Juan; Chile: Tarapaca. Bionomics: Simulium quechuanum is a species restricted to the puna or high areas (3000-4300 m altitude). It is found in streams having cold, crystal clear water, on leaves of trailing grasses, on aquatic plants such as Potamogeton, and occasionally on the surface of rocks. The intersexes mentioned in the above description are the first-ever reported from the subgenus Pternaspatha; they were bred from pupae collected west of Putre. Mermithids were here found to parasitize several larvae, and might explain the presence of intersexes. Discussion: Simulium (P.) quechuanum resembles S. barbatipes in the kind and arrangement of the thoracic trichomes of the thorax, but in S. barbatipes all branches of

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Figure 46. Simulium (Pternaspatha) quechuanum. A-C: Female; A: abdomen, dorsal view B: claw; C: apical portion of hind basitarsus and tarsite I. D-H: Pupa; D-E: dorsal and lateral views; F: gill and exposed portion of thorax with trichomes; G: frontoclypeus and trichomes; H: trichomes of thorax. I-J: Larva; I: marginal teeth; J: hypostomium and postgenal cleft.

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the gill lie close together. The diverging filaments of the ventral primary branch, as observed in S. quechuanum, are also found in S. herreri and S. yacuchuspi, but the thoracic trichomes are very different in these species. The adults of S. quechuanum are extremely similar to those of S. hectorvargasi; the two species are sympatric. The females can be distinguished by the presence in S. quechuanum of 1+1 white spots on abdominal tergum V, and the absence of these spots in S. hectorvargasi, and the slightly less-developed calcipala in S. quechuanum. This latter character is the only one to separate males of the two species. The very different pupae, distinguished easily by the number of filaments of gills and by the thoracic trichomes, are sufficient to recognize each species. There seem to be no clearcut characters to distinguish the larvae of both species. Simulium (Pternaspatha) hectorvargasi COSCARÓN & WYGODZINSKY (Fig. 47.) Simulium (Pternaspatha) hectorvargas COSCARÓN & WYGODZINSKY, 1972b: 222-225; COSCARÓN, 1991: 129-130.

Female: Wing length 3.3-3.5 mm; color of head with appendages, and thorax as in Simulium herreri; legs brown, their pattern as shown in figures C, D, E.; pattern of abdomen as shown in figure A; frons and fronto-ocular triangle much as in S. herreri; frontal angle 115 degrees; maxillary palp as illustrated (Fig. B); sensory vesicle half as wide as third palpomere, its structure as in S. herreri; lacinia with 25-29 teeth, mandible with 38-40 teeth; Sc glabrous or with one to four setae on central portion; basal section of R glabrous; setae and spines on R1 and setae on Rs arranged in one irregular row; shapes and proportions of leg segments as shown in figures C, D, E; calcipala (Fig. F) distinct but small, wider than high; claws with tooth well developed (Fig. G); genitalia much as in S. herreri; cercus and paraproct as shown in figure H. Male: Wing length 3.3-3.5 mm; color of head and its appendages and of thorax and its appendages very similar to that of Simulium herreri; abdomen (Fig. I) velvety black dorsally, especially intense on terga III-V; terga II, VI, and VII with 1+1 large sublateral silvery white spots; tergum III with 1+1 very small whitish spots, not easily perceptible in every specimen; terga III-V narrowly lighter along hind border; chaetotaxy of wings as in female, but Sc invariably lacking hair; shapes and proportions of segments of hind leg as shown in figure J; calcipala small as female; hind basitarsus 3.1-3.5 times as long as wide; genitalia much as in Simulium nemorale; apex of gonostylus smooth (Fig. K). Pupa: Cocoon (Figs. L, M) wall slipper shaped, with extremely short anteroventral bridge, not raised from substrate; cocoon dark brown, closely woven, surface smooth but with threads distinctly perceivable with moderate magnification; rim of aperture distinctly reinforced; length of cocoon dorsally 3.7-4.2 mm, maximum length along base, 4.5 mm; length of body pupa, 3.5-3.8 mm; length of gills 2.5-2.8 mm; gills (Fig. N) each with eight filaments arising from three main branches, more or less parallel-sided or spreading on basal half; ventral branch divided rather close to its base; median and dorsal branches divided at slightly larger distance from their base and again at considerable and rather variable distance; head and thorax light brown; frontoclypeus (Fig. O) without tubercles; 2+2 frontal trichomes, in shape of very short cones; 1+1 facial trichomes in shape of long setae; 1+1 ocular trichomes in shape of short spines (Fig. P); facial trichomes inserted

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basad of faint but perceptible transverse ridge; disc of thorax without tubercles, the latter restricted to area adjacent to aperture of cocoon and at insertion of gills (Fig. Q); trichomes occupying large part but not all of free surface of thorax, mostly simple hair like, very rarely bifid (Fig. R); their number approximately 150+150, their length, 0.9-0.12 mm; onchotaxy of abdomen as usual for subgenus. Larva: Maximum length of mature larva, 6.0-6.5 mm; width of head capsule, 0.50.6 mm; head light brown; cephalic apotome as shown in figure S; in some cases more G

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Figure 47. Simulium (Pternaspatha) hectorvargasi. A-H: Female; A: abdomen, dorsal view; B: maxillary palp; C-E: fore, mid and hind leg, respectively; F: apical portion of hind basitarsus; G: claw; H: cercus and paraproct. I-K: Male; I: abdomen, dorsal view; J: hind leg; K: gonostylus. L-R: Pupa; LM: cocoon in lateral and dorsal view; N: gill; O: frontoclypeus and trichomes; P: frontal trichome enlarged; Q: exposed portion of thorax with trichomes; R: trichomes of thorax. S-V: Larva; S: cephalic apotome; T: antenna; U: teeth of mandible; V: hypostomium and postgenal cleft.

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light colored; antennae (Fig. T) with distal segment darkest; medial with two constrictions accompanied by unpigmented areas; ratio of proximal, medial and distal segments = 1:1.2-1.4:0.6-0.7; cephalic fan with approximately 45 rays; toothing of mandible as shown in figure U; hypostomium with median tooth as prominent as lateral ones; hypostomial setae in one or two irregular rows, with 8-15 setae in each group; postgenal cleft (Fig. V), about twice as deep as length of postgenal bridge; anal sclerite with some simple spicules; anal ring with 75-80 rows, composed of 15-18 hooks each; rectal papillae with three simple lobes. Distribution: Chile: Tarapaca. Bionomics: Simulium hectorvargas larvae and pupae were collected on blades of grass and aquatic vegetation in fast-flowing streams with crystal-clear water. The temperature of the water was very low; ice formed in the mornings in shallow spots. The area in which this species was collected is typical puna. Discussion: Simulium albicinctum, S. herreri and S. yacuchuspi show female abdominal ornamentation very similar to that of S. hectorvargasi, but can be differentiated by the presence of the grayish tergite VI in the first species, and the other two have 1+1 small light spots on tergites IV and V. The latter species has a pupal gill with 6 branches. A distinctive character of S. hectorvargasi is to have vestigial frontal trichomes. Simulium (Pternaspatha) albicinctum (ENDERLEIN) (Fig. 48.) Acropogon albicinctus ENDERLEIN, 1934b: 277. Simulium albicinctus SMART, 1945: 500; VARGAS, 1945a: 109. Simulium (Pternaspatha) albicinctum WYGODZINSKY & COSCARÓN, 1967: 94-95; COSCARÓN, 1991: 128-129.

Female: Wing length, 3.6 mm; head black; antennae and mouthparts piceous; occiput, frons, and clypeus gray pollinose; long hair on occiput, frons, clypeus, basal antennal segments, and basal segments of maxillary palpi; scutum black, median and submedian stripes dull gray, not contrasting strongly with black areas; pubescence relatively dense; a few suberect to decumbent, brass-colored, long hairs present; scutellum, metanotum, pleura, and sterna like those in male; legs brown to yellowishbrown with coxae very dark, and distal portion of femur, tibia and tarsi darkened; dorsal surface of abdomen (Fig. A) velvety black; tergite II with 1+1 silvery white spots, dorsal dark area very small; tergite III with 1+1 smaller spots, their distance from each other about subequal to their diameter; tergites IV and V entirely black; tergite VI entirely silvery white; tergite VII black at center, extensively silvery white at sides; tergite VIII light-colored at sides, otherwise dark but not black; tergite IX entirely dark; hair of fringe of first segment whitish, hair of posterior segments dark; frons wide; fronto-ocular area about as long as wide; hind legs as shown in figure B; calcipala small but distinct (Fig. C); claws with small tooth (Fig. D); genitalia not well known. Male: Wing length, 3.1 mm; head black; eyes, antennae, and palpi piceous; clypeus black, silvery pollinose; conspicuous dark long hair on occiput, between eyes, on clypeus, basal two segments of antennae, and basal segments of maxillary palpi; scutum velvety black; median longitudinal line not clearly perceptible; pre-scutellar region and sides grayish at certain angle; anterior border with 1+1 distinct, submedi-

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an, silvery white spots; scutum with rather sparse, long, suberect and decumbent, brass-colored setae; scutellum dark brown, slightly shining, with long and very long, erect and decumbent, dark or light-colored hair; metanotum black, slightly shining; pleura and sterna piceous, with gray pollinosity; pleural tuft silvery; coxae black, grayish pollinose; trochanter of forelegs dark, femur stramineous, with apical fifth piceous, tibiae and tarsi piceous, with central portion of tibiae somewhat lighter; pattern of mid legs like that of fore legs, but first tarsal segment somewhat lighter at base; pattern of hind legs as shown in figure F; dorsal surface of abdomen (Fig. E) velvety black; long hair of fringe of first segment, and hair of other segments, whitish; tergites II and VI each with 1+1 large, silvery white spots, dark dorsal portion small; at certain angle entire sixth tergite silvery white; tergite VII with 1+1 smaller, silvery white spots; dorsal dark spot large; tergite VIII dark, with small, lateral, silvery spots visible only at certain angle; wings: Sc and basal section of R glabrous; setae and spines on R1 arranged in single row, setae on Rs in single row with numerous long hairs on dorsal surface of most segments; basitarsus slightly less than three times as long as wide; calcipala small but distinct (Fig. G); genitalia not examined in detail; gonostylus apparently smooth apically. Distribution: Peru: High Peru. Bionomics: Unknown. Discussion: Simulium albicinctum is only known from the type material, a female and a male; both were designated types by ENDERLEIN. The female is close to S. yacuchuspi, but can be differentiated by the presence of 1+1 small submedian light grayish spots on tergites IV and V, and a median posterior small darker grayish spot on tergite VI. The males in both species are also characterized by an unusually wide posterior basitarsus but S. yacuchuspi has 1+1 small light grayish spots on tergite III. B

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C E Figure 48. Simulium (Pternaspatha) albicinctum. A-D: Female; A: abdomen, dorsal view; B: hind leg; C: apical portion of hind basitarsus and tarsomere; D: claw. F-G: Male; F: hind leg; G: apical portion of hind basitarsus. and tarsomere I.

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Simulium (Pternaspatha) luchoi COSCARÓN & WYGODZINSKY (Fig. 49.) Simulium (Pternaspatha) luchoi COSCARÓN & WYGODZINSKY, 1972b: 225-228; COSCARÓN, 1991: 134.

Female: Wing length approximately 3.5 mm; color of head, thorax and wings as described for Simulium herreri; color pattern of fore and mid legs as illustrated for male, of hind legs as shown in figure D; abdomen (Fig. A) dark gray; first tergum black; terga IIVIII velvety black at center, extension of black areas as illustrated, that on second tergum widened anteriorly; terga II, III, VI, VII, and VIII with 1 +1 sublateral silver-white spots, large on II and VI, smaller on III, VII, and VIII; fronto-ocular triangle as shown in figure B; frontal angle 115 degrees; sensory vesicle of maxillary palp as shown in figure C; lacinia with 22-23 teeth; mandibles not examined; wings as usual for subgenus; chaetotaxy of wings as in Simulium hectorvargasi; hind legs as shown in figure D; calcipala distinct but small, subtriangular (Fig. E); claws with distinct tooth (Fig. F); central portion of eighth sternum darker than peripheral areas; paraproct relatively elongated ventrally (Fig. G). Male: Wing length 3.3 mm; color of head, thorax, and abdomen much as in Simulium herreri; color pattern of legs as shown in figures J, K, L; structure of head as in S. herreri, antennae (Fig. H) and maxillary palp (Fig. I) with long setae; last palpomere more than twice as long as penultimate; shapes and proportions of segments of legs as in figures J, B

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Figure 49. Simulium (Pternaspatha) luchoi. A-G: Female; A: abdomen, dorsal view; B: fronto-ocular triangle; C: second palpomere with sensory vesicle; D: hind-leg; E: apical portion of hind basitarsus, with small calcipala; F: claw; G: cercus and paraproct. H-N: Male; H: antenna; I: maxillary palp; J-L: fore, mid and hind leg, respectively; M: apical portion of hind basitarsus, with small calcipala; N: gonocoxite and gonostylus. O-S: Pupa; O: gill; P: frontoclypeus; Q: exposed portion of thorax, with trichomes and tubercles; R: tubercles, high magnification; S: trichomes, high magnification.

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K, L; calcipala present but very small (Fig. M); hind basitarsus slightly less than three times as long as wide; abdomen with numerous long hairs; gonocoxite and gonostylus as in figure N; the latter smooth. Pupa: Cocoon slipper shaped, anteroventral bridge distinct, slightly elevated from substrate; general aspect of cocoon much as in Simulium quechuanum; cocoon light brown, translucent; surface smooth, but with individual threads distinctly perceptible; rim of aperture faintly reinforced; length of cocoon dorsally, 3.3 mm, maximum length at base, 4.04.5 mm; length of body, 3.5 mm; length of gill 2.0 mm; gill composed of eight filaments arranged as shown in figure O; all filaments lying close together except ventral most filament, which is sharply ventrally and backwardly bent, situated inside cocoon; head and thorax light brown; frontoclypeus (Fig. P) without tubercles, with 2+2 frontal, 1+1 facial, and 1+1 ocular trichomes; facial and ocular trichomes much longer than frontal trichomes; all trichomes slender, hair like, simple, or with one or two branches; facial trichomes situated in depression delimited by conspicuous ridges; thorax (Fig. Q) with disc smooth; tubercles forming band adjacent to aperture of cocoon; these tubercles frequently somewhat pointed (Fig. R); thorax with 30+30 to 50+50 trichomes (Fig. S), most shortly lanceolate, simple or divided, with some simple or branched hair like setae interspersed. Larva: Unknown. Distribution: Chile: Tarapaca. Bionomics: Simulium luchoi has been collected at 4100 m altitude. Discussion: The female of Simulium luchoi is very similar to that of S. hectorvargasi, but S. luchoi has smaller 1+1 light grayish spots on abdominal tergite III and the darkened area of tergites III-V is homogeneous. It also can be separated by the larger, rounded calcipala present in S. hectorvargasi. The males of S. luchoi are very close in structure and color to the males of S. albicinctum, S. hectorvargasi, and S. quechuanum. Minor characters of the structure and coloring of the hind legs, such as that in S. luchoi showing 3/5 of the hind femur whitish and in the other species light brown, may make specific identification possible. The pupa of S. luchoi can be distinguished by the structure and number of its thoracic trichomes from the other species with 8 gill branches; that in S. luchoi has one or two of these branches sharply diverging. Simulium (Pternaspatha) prodexargenteum (ENDERLEIN) (Fig. 50.) Pternaspatha prodexargenteum ENDERLEIN, 1936: 116. Simulium (Pternaspatha) prodexargenteum WYGODZINSKY & COSCARÓN, 1967: 109-110; COSCARÓN & WYGODZINSKY, 1972b: 228-232; COSCARÓN, 1991: 123-124.

Female: Wing length 2.9-3.2 (3.3-3.7) mm; color of head and thorax and their appendages much as in Simulium nemorale; pattern of legs as shown in figures G-I; general color of abdomen gray, showing terga II-V velvety black at center; terga II and III with 1+1 large, sublateral, silver-white spots; IV and V at sides of posterior border narrowly silver-white; VI almost entirely white dorsally, slightly darker at center and at sides, silvergray sublaterally; VIII and IX dark gray (Fig. A); frons as shown in figure B; frontal angle 115 degrees; fronto-ocular triangle as shown in figure C; shapes and proportions of antennal segments as shown in figure D; maxillary palp (Fig. E), sensory vesicle over half as wide as third palpomere, with abundant tubercles (Fig. F); lacinia with 28-29 teeth,

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mandible with 37-43 teeth; Sc with two to five scattered setae; setae and spines on R1 and setae on Rs arranged in single row; shapes and proportions of leg segments as shown in figures G-I; calcipala obsolescent or absent (Fig. J); claws with distinct tooth (Fig. K); eighth sternum, gonapophyses, genital fork, and spermatheca as in Simulium dureti. Male: Wing length 3.0 (3.1) mm; scutum velvety black with very narrow median longitudinal, white line; color pattern of legs very close to female, hind legs as in figures O, P; abdomen velvety black, with silvery spots as shown in figure L; antenna showing scape and pedicel with very long hair (Fig. M); maxillary palp with very long setae on basal segments (Fig. N); diameter of sensory vesicle less than half width of third segment of palp (Fig. O); chaetotaxy of wing veins as in female; shapes and proportions of hind leg segments as shown in figures P-Q showing wide basitarsus; calcipala obsolescent (Fig. R) or absent; hind basitarsus 3.0-3.6 times as long as wide; genitalia as in Simulium herreri; apex of gonostylus smooth. Pupa: Cocoon (Figs. S, T) wall-pocket shaped, but with very short anteroventral bridge; color light brown; when observed in liquid, shows leaden sheen; cocoon closely woven, threads not individualized, texture somewhat parchment-like; surface of cocoon smooth, in some specimens with adhering foreign particles; rim of aperture slightly reinforced; length of cocoon dorsally along middle, 2.5-3.0 (3.2-3.5) mm; maximum length along base 3.2-3.8 (4.2-4.5) mm; length of body 2.5-2.7 (3.6-3.7) mm; length of gills 1.8 (2.5) mm, viz., distinctly shorter than both cocoon and pupa proper; gills from light to dark brown, each with eight filaments (Fig. U); filaments of dorsal and median branches and dorsal filament of ventral branch diverging at base, approaching near apex; lower filament of ventral branch sharply diverging from rest of filaments, ventrally and backwardly curved; head and thorax of pupa from light to dark brown; frontoclypeus (Fig. V) without tubercles, and with 3+3 simple, hair like, short frontal, 1+1 in some cases bifid facial, and 1+1 simple hair like ocular trichomes; facial trichomes inserted in, or basad of, area of low irregular ridges; thorax with tubercles restricted to margin of exposed area (Fig. W), attaining base of gill; trichomes numbering approximately 10+10 (14+14), in shape of short simple or bifid setae. Larva: Length 4.5-4.7 (6.7) mm; width of head capsule, 0.48 (0.70) mm; general body shape as shown in figure X; color of larva greenish brown, head light brown; cephalic apotome with pattern very faint, and distinctly darkened near hind margin (Fig. Y); antennae dark brown (Fig. Z); ratio of proximal, medial and distal segments = 1:1.41.8:0.9-1.0; cephalic fan with 30-40 (39-50) rays; toothing of mandible as shown in figure a; anterior border of hypostomium strongly pigmented; median tooth as prominent or more prominent than lateral teeth; marginal serrations distinct; hypostomial setae arranged in one or two very irregular rows (Fig. b), with 8-12 (14-17) setae in each group; postgenal cleft (Fig. c) pointed, deep, its depth equal to five times length of postgenal bridge; anal sclerite much as in Simulium dureti; perianal hair few in number; scales at base of anal sclerite from inconspicuous to distinct; anal ring with approximately 78 (95) rows, with 14-15 (16-17) hooks each; rectal papillae with three simple lobes. Distribution: Argentina: Salta; Bolivia: La Paz; Peru (High Peru). Bionomics: Larvae of Simulium prodexargenteum live in small creeks with strong current, crystalline and cold waters at great altitudes (3200-4700 m), flowing over aquatic vegetation and stones.

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Discussion: The material studied is homogeneous regarding morphological features but heterogeneous regarding some meristic characters. All the known localities that correspond to the “puna” area are close together. No geographical pattern of variability emerges; it may be

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Figure 50. Simulium (Pternaspatha) prodexargenteum. A-K: Female; A: abdomen, dorsal view; B: frons; C: fronto-ocular triangle; D: antenna; E: maxillary palp; F: sensory vesicle of maxillary palp; G-I: fore, mid and hind leg, respectively; J: apical portion of hind basitarsus and tarsomere I. K: claw; LR: Male; L: abdomen in lateral view; M: antenna; N: maxillary palp; O: sensory vesicle of maxillary palp;. S-W: Pupa; S-T: dorsal and lateral views; U: portion of thorax with gill; V: frontoclypeus; W: portion of P-Q: hind legs; R: apical portion of hind basitarsus thorax with trichomes and tubercles. X-Z, a-c: Larva; X: lateral view; Y: cephalic apotome; Z: antenna; a: teeth of mandible; b: portion of hypostomium; c: hypostomium and postgenal cleft.

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significant, though, that the largest measurements were obtained from the lot collected at the highest altitude. The differences between the populations examined are only meristic, and such differences by themselves do not necessarily indicate specific differences. The abdominal ornamentation of females shows similarities with S. nemorale but this species has 1+1 whitish submedian longitudinal stripes on tergites IV and V, the calcipala well developed, tubercles on the frontoclypeus and gill branches not much opened basally. Simulium (Pternaspatha) annulatum PHILIPPI (Fig. 51.) Simulium annulatum PHILIPPI, 1865: 634; KERTESZ, 1902a: 285; KNAB, 1914a: 18; SILVA FIGUEROA, 1917: 30; PINTO, 1931: 727; EDWARDS, 1931: 153; SMART, 1945: 500; VARGAS, 1945a: 112; STUARDO ORTIZ, 1946: 42. Simulium (Pternaspatha) annulatum WYGODZINSKY & COSCARÓN, 1967: 89-94; COSCARÓN & WYGODZINSKY, 1972b: 219-220; COSCARÓN, 1991: 133-134. Simulium varipes PHILIPPI, 1865: 634; KERTESZ, 1902a: 291; SILVA FIGUEROA, 1917: 30; PINTO, 1931: 730; SMART, 1945: 515; VARGAS, 1945a: 205; VARGAS & DIAZ, 1951: 139; STUARDO ORTIZ, 1946: 41; STONE, 1963: 5. Simulium (Simulium) varipes EDWARDS, 1931: 152. Friesia varipes ENDERLEIN, 1929: 327; 1930: 92. Dasypelmoza varipes ENDERLEIN, 1934b: 275.

Female: Wing length 3.2 mm; color of head and its appendages, and of thorax, halteres, and wings like that in Simulium nemorale; legs light brown, with darker regions: mid and hind coxae; apical halves of trochanters; apices of all femora and tibiae; entire fore tarsus, apex of first tarsal segment of mid tarsus as well as all other segments entirely, apical third of hind basitarsus, apical two-thirds of second and entire third, fourth, and fifth segments of hind tarsus; abdomen (Fig. A) black on tergites I-V, dark gray on remainder; silvery gray areas distributed as follows: narrow line on posterior border of tergite I; 1+1 large lateral spots on tergite II; 1+1 very delicate lines on lateral portions of posterior borders of tergites III-V; 1+1 large lateral spots, in some cases meeting medially and thus occupying entire tergite, on tergites VI-IX; ventral surface of abdomen almost entirely gray pollinose; hair of abdomen, including that of basal fringe, silvery; frons as shown in figure B, median sulcus obsolete; fronto-ocular triangle slightly wider than long; last segment of maxillary palp (Fig. C) almost twice as long as penultimate segment; base of cibarium with well-sclerotized lateral projections; lacinia with approximately 25 teeth, mandible with about 48 teeth; scutum with abundant decumbent setae, longer erect ones on posterior declivity; Sc with 15-30 setae and several rows of setae also on basal sector of R; setae and spines on R1 arranged in several rows; hair on Rs, in single row (Fig. D); calcipala (Fig. E) well developed, about as long as wide; claws with well-developed tooth (Fig. F); eighth sternite (Fig. G) with central portion lighter than lateral areas; gonapophyses rounded apically, beset with microchaetae, their inner margins more strongly sclerotized; cercus with distal margin truncate to slightly rounded; projecting portion of paraprocts subtriangular (Fig. H); median process of genital fork strongly sclerotized, extreme apex widened. Male: Wing length 3.0 mm; scutum black, with golden-colored, decumbent setae; fore legs dark brown, mid and hind legs light brown, darkened area almost black, distributed very

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similar to female; setae brown to black; abdomen velvety black dorsally, with golden-colored hair; tergites II, VI, and VII with 1+1 lateral silvery gray spots; tergite IX dark gray pollinose; sternites grayish to dull brown; last segment of maxillary palp (Fig. I) more than twice as long as penultimate segment, and longer than two preceding segments combined; diameter of sensory vesicle approximately half that of third segment; wings as in female, but hairs on basal portion of R slightly less numerous; basitarsus of hind legs 4.2 times as long as wide, calcipala well developed, slightly wider than long; genitalia similar to those of Simulium nemorale, but the 1+1 translucent, areas of basal plate larger.

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Figure 51. Simulium (Pternaspatha) annulatum. A-H: Female; A: abdomen, dorsal view; B: frons; C: maxillary palp; D: anterior veins of wing; E: apical portion of hind basitarsus and tarsite I; F: claw; G: eighth sternite and gonapophyses; H: cercus and paraproct. I: Male, maxillary palp. J-O: Pupa; J: lateral view; K-L: gills of different specimens; M: frontoclypeus and trichomes; N: exposed portion of thorax and trichomes; O: thoracic trichomes. P-U: Larva; P: cephalic apotome; Q: antenna; R: teeth of mandible; S: hypostomium; T: cephalic capsule, ventral view; U: rectal papillae.

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Pupa: Cocoon slipper shaped (Fig. J), with elongate anteroventral bridge; color of cocoon light to dark brown; closely woven, individual threads perceptible; rim of aperture slightly reinforced; gills protected laterally by cocoon at their extreme base only; length of cocoon at dorsum 3.3 mm; at base, 4.5 mm; length of pupa body 3.6 mm; length of gill 2.5 mm; gills (Fig. K) with six (exceptionally eight as Fig. L) closely parallel filaments; head and thorax of pupa distinctly pigmented, more strongly so on frontal region of head and anterior dorsal portion of thorax; trichomes of head and thorax simple, hair like; head (Fig. M) with 3+3 frontal, 1+1 facial, and 1+1 ocular, trichomes; region of thorax adjacent to aperture of cocoon with band of small tubercles, dorsal exposed portion of thorax covered with long trichomes (Fig. N), approximately 150+150 in number, their length (Fig. O), 0.14-0.17 mm; abdomen of pupa much like that of Simulium nemorale, spines on tergite II relatively smaller. Larva: Maximum length, 6.5 mm; maximum width of head capsule, 0.66 mm; color greenish brown; head brown; cephalic apotome darkened on median line and 1+1 lateral, sometimes darkened also on anterior area (Fig. P); body integument with isolated delicate hairs, latter numerous at base of anal sclerite; antennae brown, pigment most intense on third segment (Fig. Q); ratio of proximal, medial and distal segments = 1:1.3:0.9; cephalic fan with approximately 40 rays; toothing of mandibles as shown in figure R; anterior border of hypostomium (Fig. S) distinctly pigmented, median tooth less prominent than lateral ones; intermediate teeth gradually increasing in size from inner to outer ones; sublateral teeth very small; lateral borders of hypostomium serrated, with five small teeth; hypostomial setae with 7-13 setae in each group; postgenal cleft deep subtriangular, longer than postgenal bridge (Fig. T); anal sclerite without scales at base; anal ring with about 85 rows, each composed of 13-14 hooks; rectal papillae consisting of three primary lobes, each bearing three pairs of long secondary lobules (Fig. U). Distribution: Argentina: Chubut over Andean area, Neuquen, Rio Negro; Chile: Central and southern region from Santiago to Llanquihue. Bionomics: Simulium annulatum breeds in rivers with fast-flowing, sometimes torrential crystalline waters. Females are anthropophilic and are known by the regional inhabitants as “petros”, the araucanian name. Discussion: Simulium annulatum differs from all other species of Pternaspatha by the presence of hair on the basal portion of the radial vein, but agrees with Pternaspatha in all other essential characters. The pupa shows great similarity with that of S. caprii, but has fewer trichomes on the thorax (about 100). The female has similar ornamentation but the posterior rim of tergites III-V has no narrow grayish band as in S. caprii. Simulium (Pternaspatha) caprii WYGODZINSKY & COSCARÓN (Fig. 52.) Simulium (Pternaspatha) caprii WYGODZINSKY & COSCARÓN, 1967: 79-83; COSCARÓN & WYGODZINSKY, 1972b: 222; COSCARÓN, 1991: 132.

Female: Wing length 3.1-3.5 mm; head blackish; antenna, maxillary palp, and labium brown, with eyes and basal segment of palp darkest; first and second antennal segments somewhat more light-colored than remainder; antenna with short hair, that on two basal segments distinctly longer; hair of frons, clypeus, occiput, and palpi silvery to black; color

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pattern of scutum very similar to that of Simulium nemorale, with decumbent, brass-colored to black hair. Median longitudinal light-colored line formed by hair only, not by gray pigment as in remaining species of the group; scutellum grayish brown, with long silvery to black hair. Metanotum and pleura dark brown, with velvety violaceous pollinosity; pleural tuft pale brass-colored; legs brown, with silvery to black hair; following regions dark: all coxae and trochanters, small spot on base and large one on apex of all femora, base and apex of all tibiae, entire fore tarsus, apical half of basitarsus and all remaining segments of mid tarsus, extreme base and apex of basitarsus and second segment, as well as entire third to fifth segments, of hind tarsus; abdomen (Fig. A) piceous; tergites II-V velvety black at center, dark gray at sides; tergite II with 1+1 large, silvery white spots; tergites VI-IX dark, with 1+1 sublateral, silvery white spots decreasing progressively in size toward posterior segments; ventral surface of abdomen brown; frons as shown in figure B, with obsolete median sulcus; fronto-ocular triangle approximately as wide as long; last segment of maxillary palp (Fig. C) much longer than two preceding segments combined, and three times as long as penultimate segment; diameter of sensory vesicle approximately equal to half of diameter of segment; structure of vesicle as shown in figure D; lacinia with about 27 teeth, mandible with about 45 teeth; Sc with approximately 10 setae on central and apical regions; spines and setae on R1 in two very irregular rows; setae on Rs in one row, in some places in two irregular rows; calcipala well developed, slightly wider at base than long (Fig. E); claws with short tooth (Fig. F); eighth sternite with about 30 long setae on each side (Fig. G); gonapophyses subtriangular, with abundant microtrichiae; inner borders more sclerotized; paraproct and cercus as shown in figure H with median process well sclerotized. Male: Wing length 3.3 mm; color of head, thorax, wings, and halteres like that of Simulium nemorale; legs light brown; darkened in some areas as hind leg (Fig. I); following dark: all coxae brown; trochanters except base; spot at base of fore and mid, and on apex of all femora; base and apex of tibiae of all legs; entire fore tarsus, tarsus of mid legs except basal two-fifths of basitarsus, base and apex of hind basitarsus, basal scale and apical half of second segment, and entire third to fifth segments, of hind tarsus; abdomen black, with 1+1 silvery gray spots on tergites II, VI, VII, and VIII, most conspicuous on second, becoming progressively smaller and fainter on posterior segments; last segment of maxillary palp longer than two preceding segments combined and sensory vesicle smaller than half diameter of third segment; wings like those of female, but Sc with not more than seven setae, and spines and hair on R1 and Rs slightly less numerous; segments of hind legs as shown in figure I; hind basitarsus narrow, 3.8-4.0 times as long as wide; calcipala distinct, about as long as wide at its base (Fig. J); gonostylus subquadrate; apical border widely concave, glabrous (Fig. K). Pupa: Cocoon (Fig. L) with relatively small, anteroventral bridge, nearly shoe shaped; color of cocoon dark brown; surface relatively rough, threads individualized; rim of aperture distinctly reinforced; gills protected by walls of cocoon at their extreme base only. Maximum length of cocoon dorsally along middle, 4.2 mm; maximum length along base, 5.2 mm; length of body of pupa 2.5 mm, of gill 2.1 mm; gill (Fig. M) with eight filaments; head and thorax of pupa dark brown; trichomes of head (Fig. N) exceptionally long; simple or bifid (Fig. O), until trifid in facial (Fig. P); clypeus without tubercles; thorax

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with band of tubercles adjacent to aperture of cocoon, with numerous interspersed trichomes (Fig. Q); disc of thorax lacking tubercles but possessing about 100 slender, hair like, simple or bifid trichomes; latter (Fig. R) with average length, 0.14 mm; chaetotaxy of abdomen like that in Simulium nemorale. Larva: Maximum length, 7.5 mm; maximum width of head capsule, 0.7 mm; color of larva dark greenish brown; cephalic apotome brown, with darker spots (Fig. S); some curved, short setae at base of anal sclerite; antennae as shown in figure T; brown, subapical segment darkest; ratio of proximal, medial and distal segments = 1:1.3:0.9; cephalic fan with 35-40 rays in large fan; toothing of mandible as shown in figure U: two J C

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Figure 52. Simulium (Pternaspatha) caprii. A-H: Female; A: abdomen, dorsal view B: frons; C: maxillary palp; D: sensory vesicle of maxillary palp; E: apical portion of hind basitarsus and tarsomere I; F: claw; G: eighth sternite and gonapophyses; H: cercus and paraproct. I-K: Male; I: hind leg; J: apical portion of hind basitarsus and tarsomere I; K: gonostylus. L-R: Pupa; L: lateral view; M: gill; N: frontoclypeus; O-P: frontal and facial trichomes, respectively; Q: exposed portion of thorax with trichomes and tubercles; R: thoracic trichomes. S-W: Larva; S: cephalic apotome; T: antenna; U: teeth of mandible; V: hypostomium; W: rectal papillae.

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external, one strong apical, three subapical, six internal, and two marginal teeth, internal ones variable in size; anterior border of hypostomium strongly pigmented (Fig. V); median tooth about as prominent as lateral ones; intermediate teeth gradually increasing in size from center to side; sublateral tooth hardly perceptible; lateral border with six to seven very small serrations; hypostomial setae in groups of 12-15, arranged in two irregular series; disc of hypostomium with few short and simple bifid setae; postgenal cleft deep, pointed, about twice as deep as postgenal bridge; anal sclerite with numerous scales at base; anal ring with approximately 85 rows, with 13-15 hooks each; rectal papillae (Fig. W) with three primary lobes, lateral ones each with three or four and median one with six to eight, diverticula. Distribution: Argentina: Chubut. Bionomics: The larvae and pupae of S. caprii were found attached to stems of grasses trailing in clear streams and on branches of trees that had fallen into the water, as well as on stones. Discussion: Simulium caprii is similar to S. nemorale but it is much smaller. The female of S. caprii differs from that of S. nemorale by the more conspicuous pilosity on the scutum which also forms a line along the middle of the sclerite (this line is formed by lightcolored pigment in S. nemorale as in all other species of the subgenus), and the central portion of the eighth sternite, which is darker than the lateral areas. The most obvious differences are found in the chaetotaxy of the head and thorax of the pupae, with longer and more abundant trichomes present in S. caprii. Simulium (Pternaspatha) stelliferum COSCARÓN & WYGODZINSKY (Fig. 53.) Simulium (Pternaspatha) stelliferum COSCARÓN & WYGODZINSKY, 1972b: 234-239; COSCARÓN, 1991: 137-139. Simulium (Pternaspatha) schoenemanni WYGODZINSKY & COSCARÓN, 1967: 89 (misidentification).

Female: Wing length 2.9-3.1 mm. (mode, 3.1 mm.); color of head and its appendages blackish, antennae, palpi and labrum piceous; frons and clypeus grayish pollinose; scutum as in Simulium nemorale; legs brownish with darker area distributed as shown in figures F-H; general color of abdomen gray; pattern as shown in figure A; terga II-V velvety black at center; tergum II with 1+1 large sublateral silver-white spots; terga III-V narrowly silver-white along hind margin except on median black area; terga VI and VII each with 1+1 large silver-white spots, those of VII separated by larger dark median area than those of VI; terga VIII and IX silver-gray; frons as shown in figure B; frontal angle 115 degrees; fronto-ocular triangle very slightly deeper than wide (Fig. C). Maxillary palp as illustrated in figure D; sensory vesicle (Fig. E) half as wide as third palpomere; lacinia with 26-28 teeth, mandible with 42-45 teeth; Sc with 12-14 hairs occupying most of length of vein; basal section of R glabrous; setae and spines on R1 and setae on Rs arranged in one or two irregular rows; shapes and proportions of leg segments as shown in figures F, G, H; calcipala well developed (Fig. I); claws with tooth well developed (Fig. J); eighth sternum (Fig. K) with central and lateral areas well pigmented; gonapophyses with inner margins distinctly pigmented; cerci and paraprocts as shown in figure L. Male: Wing length 2.7-2.9 mm. (mode, 2.8 mm); color of head and its appendages much as in Simulium nemorale, thorax black, legs as female but slightly darker; abdomen

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(Fig. M) grayish, velvety black dorsally except on segments VIII and IX which are dark gray; terga II, V, and VI with 1+1 large sublateral silver-white spots; terga III-VII narrowly lighter along hind margin; scape and pedicel with conspicuous long hair. Maxillary palp with basal segments with long hair; chaetotaxy of wing veins as in female, but Sc only with 8-10 hairs; shapes and proportions of segments of hind legs as shown in figure N; hind basitarsus narrow, 4.3-5.5 times as long as wide; calcipala well developed, not quite so long as wide; gonostylus smooth distally (Fig. O). Pupa: Cocoon slipper shaped, but with very short anteroventral bridge (Fig. P); cocoon brown, its surface smooth, closely woven, individual threads not perceptible; rim of aperture slightly reinforced; length of cocoon dorsally, 3.7-3.8 mm; along base, 4.5-4.7 mm; length of pupa 3.5-3.7 mm; gills 2.5-2.8 mm; gills (Fig. Q) each with six filaments arising from three main branches; ventral and dorsal primary branches divided very close to their bases; median primary branch slightly longer; dorsal and median branches and filaments closely parallel to each other; ventral branch diverging somewhat from remaining branches, its filaments much narrower than remaining, ventrally but not backwardly directed; surface structure of filaments as usual for subgenus; head and thorax of pupa brown, darkest on frontal region and exposed portion of thorax; frontoclypeus (Fig. R) with very numerous smooth tubercles, and with 2+2 frontal, 1+1 facial and 1+1 ocular trichomes; shape of these trichomes extremely variable, even on opposite sides in one individual; frontal trichomes generally rather short, with two or three branches, but occasionally longer; facial trichomes generally hair like, bifid; ocular trichomes short, inconspicuous; facial trichomes inserted in distinct depression delimited by irregular carinae; exposed portion of thorax covered with very numerous tubercles similar to those of head, and several hundred trichomes (Fig. S). Most thoracic trichomes roughly stellate, as shown in figure T, with three to six branches, the latter short, flattened, pointed apically, slightly curved; some trichomes long, hair like, branched; such trichomes more numerous in lower portion of exposed area; onchotaxy of abdomen as usual for subgenus. Larva: Length 8.5-8.8 mm; width of head capsule, 0.70-0.75 mm; general body shape of larva as in figure U; color of larva light greenish brown; head brown; anterior half or two-thirds of cephalic apotome (Fig. V) conspicuously lighter than rest of head, which is dark brown; in some specimens, pigment on cephalic apotome reduced to narrow dark band along basal margin (Fig. W); antenna (Fig. X) light brown, third segment darkest; ratio of proximal, medial and distal segments = 1:1.2:0.8; cephalic fan with 37-39 rays; toothing of mandible as shown in figure Y; anterior border of hypostomium strongly pigmented. Median tooth of hypostomium (Fig. Z) almost as prominent as lateral teeth; marginal serration distinct but not very pronounced; hypostomial setae arranged in one or two irregular rows, with 12-15 setae in each group; postgenal cleft pointed or narrowly rounded apically (Fig. a); postgenal bridge shorter than hypostomium length; sclerite of thoracic proleg with about 15 setal groups of different length (Fig. b); anal sclerite as shown in figure c, characterized by very extensive sclerotized areas between anterior and posterior arms; base of anal sclerite lacking scale like cuticular structures, at most with small number of spicules; anal ring with 81-83 rows composed of 14-15 hooks each; rectal papillae (Fig. d) with three primary lobes, either simple or each with not more than five lobules.

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Figure 53. Simulium (Pternaspatha) stelliferum. A-L: Female; A: abdomen, dorsal view; B: frons; C: fronto-ocular triangle; D: maxillary palp; E: sensory vesicle of maxillary palp; F-H: fore, mid and hind leg, respectively; I: apical portion of hind basitarsus and tarsomere I; J: claw; K: eighth sternite and gonapophyses; L: cercus and paraproct. M-O: Male; M: abdomen, dorsal view; N: hind leg; O: gonostylus. P-T: Pupa; P: lateral view; Q: gill; R: frontoclypeus and trichomes; S-T: thoracic trichomes and tubercles. U-Z, a-d: Larva; U: lateral view; V-W: cephalic apotome in different specimens; X: antenna; Y: teeth of mandible; Z: anterior portion of hypostomium; a: hypostomium and postgenal cleft; b: lateral sclerite of thoracic proleg; c: anal sclerite; d: rectal papillae.

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Distribution: Argentina: Río Negro isolated on the east low Patagonian steppe; Chile: from Santiago to Tarapaca on the lowlands. Bionomics: Simulium stelliferum has a comparatively wide ecological spectrum. It is found mainly in fast-flowing streams but also occasionally in irrigation ditches, at altitudes from 200 to 2500 m, and at water temperatures of 6-17 °C. The larvae and pupae were found attached to vegetation, such as grass or small branches of other plants trailing in the current, but also, in some cases, to the surface of rocks. The streams in which specimens were collected were mostly clear, but sometimes S. stelliferum was obtained in slightly turbid waters containing abundant diatoms and other organic matter; in the latter case, the current was exceedingly fast. Discussion: The adults of S. stelliferum are most similar to those of S. caprii from which they can be distinguished by tergites III-V being entirely black in the latter. The pupa differs from all other species of the subgenus by the unique structure of the trichomes of the thorax. The larva of S. stelliferum lacks peculiarities, and varies in some key characters; consequently, it had to be placed in the key more than once. Species inquirenda Simulium (Pternaspatha) schoenemanni (ENDERLEIN) Acropogon schoenemani ENDERLEIN, 1934b: 277. Simulium schoenemanni SMART, 1945: 513; VARGAS, 1945a: 195; STUARDO ORTÍZ, 1946: 141. Simulium (Pternaspatha) schoenemanni WYGODZINSKY & COSCARÓN, 1967: 89.

WYGOZINSKY & COSCARÓN (1967) tentatively identified certain Chilean specimens of Pternaspatha as Simulium schoenemanni. Careful re-examination led them to conclude that the females fail to agree with the characters given by ENDERLEIN (1934b) for the female type, the only specimen known of S. schoenemanni, especially as to the color pattern of the abdomen. The type of S. schoenemanni could not be found at the Zoological Museum in Berlin, where it was deposited, according to ENDERLEIN (1934b), nor at the Staatliches Museum fûr Tierkunde, Dresden, where the type might have been deposited by error. The species formerly identified by WYGODZINSKY & COSCARÓN as S. schoenemanni was described later as Simulium(Pt.) stelliferum COSCARÓN & WYGODZINSKY, 1972b, but S. schoenemanni itself continues to be an enigma. Simulium (Notolepria) ENDERLEIN (Figs. 54-56, 139.) Type-species: Simulium exiguum ROUBAUD, 1906b: 108.

Female: Species of small size, wing length 2.2-2.5 mm; coloration blackish; scutum black, bordered laterally and posteriorly by a silvery pollinose area; scutum and scutellum generally with bright golden hair grouped and simulating scales; abdomen black, tergites V-VIII with cereous sheen; fronto-ocular triangle absent or reduced; cibarium with basal portion smooth, scarcely sclerotized, without reinforcement; wings with Sc and R bare; legs with petaloid hair; claws with or without subbasal teeth; calcipala well developed,

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about as long as wide; hind basitarsus with normal width; gonapophyses subtriangular with internal border concave, and well sclerotized; paraprocts about as long as wide with posterior border acuminate and generally with thin elongated process; genital fork with branches well sclerotized with stem about two times longer than posterior branches and scarcely widened apically; spermatheca ovoidal. Male: Wing length 2.0-2.3 mm; coloration black; scutum velvety black, bordered by pollinose grayish area; hind basitarsus not inflated; abdomen with II, VI, VII with 1+1 light grayish spots, smaller on VIII tergite; gonostylus half as long as gonocoxite, subtriangular and with reduced obsolescent apical spine; ventral plate rather acuminate distally, nearly as long as wide, sometimes with median constriction. Pupa: Cocoon slipper shaped, generally slightly projected basally, weave not too thick, with visible threads; gill with 6-8 branches; frontoclypeus and exposed portion of thorax with abundant tubercles; frontoclypeus with 2 frontal and facial trichomes on each side, with one to 3-6 branches; thorax with 5-6 trichomes on each side with 3-6 branches; abdomen with apical spur on last tergite. Larva: Cephalic apotome darkened on base, generally with median and lateral spots not well defined; antennal medial segment longer than proximal and this about 1/3 hypostomium length. Mandible with several rows of teeth, and two marginal teeth thin, curved and with second tooth small; latero-mandibular process not evident; thoracic proleg lateral sclerite with teeth arranged in about 9 groups; anal ring with 52-76 rows of about 15 hooks each; rectal papillae with 0-12 lobules on each lobe. Distribution: Northern Argentina to Mexico, primarily in the intertropical occidental area (Fig. 139). Discussion: This subgenus is made up of 10 species: 5 with scarce information and 3 of which are considered species inquirendae. KEYS

TO

SIMULIUM (NOTOLEPRIA)

SPECIES

Females 1 –

2(1) – 3(2)

Scutum and scutellum with cylindrical hairs distributed homogeneously, without bright metallic scales; with some light, showing 1+1 anterior subtriangular light grayish spots; paraproct with small distal process (Fig. 56 E) ........ Simulium llutense Scutum and scutellum with flattened hairs (Fig. 5 B) arranged in groups (Fig. 5 A) appearing as bright metallic scales; with some light, not showing 1+1 anterior grayish subtriangular spots; paraproct with distal process generally well developed (Figs. 54 I, Q, X) ................................................................................................................ 2 Paraproct with reduced distal process (Fig. 54 X); posterior branches of genital fork long and thin and scarcely sclerotized .................................................................... ................................................. Simulium gonzalezi, Simulium subexiguum, Simulium sp. A Paraproct with well-developed distal process (Figs. 54 I, Q); posterior branches of genital fork relatively short and wide, and well sclerotized ...................................... 3 Paraproct distal process relatively short and hooked; eighth sternite well sclerotized medially and distally ................................................... Simulium cuasiexiguum

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– 4(3) –

Paraproct distal process relatively long (Figs. 54 I, Q); eighth sternite not well sclerotized medially and distally (Fig. 54 F) ................................................................. 4 Paraproct process thick at base (Fig. 54 I); genital fork stem comparatively shorter in relation to posterior branches (Fig. 54 H) ................................... Simulium exiguum Paraproct process thin at base (Fig. 54 Q); genital fork stem comparatively longer in relation to posterior branches (Fig. 54 R) ............................. Simulium paraguayense

Males 1 – 2(1) – 3(2) –

Scutum with normal hairs arranged homogeneously; ventral plate very short (Fig.56 G) ................................................................................................... Simulium llutense Scutum with flattened hairs arranged in groups appearing as bright scales; ventral plate more elongated (Figs. 54 O, P, U, W) .................................................................. 2 Gonostylus subtriangular (Figs. 54 M, N); ventral plate without (Fig. 54 O) or with (Fig. 54 P) lateral constriction ........................................................................................... .................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A Gonostylus subquadrate (Figs. 54 S, T); ventral plate with well-defined lateral constriction .......................................................................................................................... 3 Ventral plate with strong mediolateral constriction appearing like wide median spatulate process (Figs. 54 U-W) ................ Simulium gonzalezi, Simulium paraguayense Ventral plate with lateral constriction appearing as narrow distomedial process .. .............................................................................................................Simulium cuasiexiguum

Pupae 1 – 2(1) – 3(1) – 4(4) –

Gills with 6 branches (Fig. 55 L) .................................................................................... 2 Gills with 8 branches (Fig. 55 K) .................................................................................... 3 Frontal and facial trichomes simple ............................................ Simulium cuasiexiguum Frontal trichomes with 1-3 branches, facial trichomes with 3-5 branches .............. ................................................................................................................... Simulium gonzalezi Cocoon extended ventrally and closed anteriorly (Fig. 56 H) ........ Simulium llutense Cocoon slightly extended ventrally and open anteriorly (Figs. 55 A, I) ................. 4 Cocoon not extended ventrally (Fig. 55 I) ................................. Simulium paraguayense Cocoon with short ventral projection (Fig. 55 A) .................................................... .................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A

Larvae Larvae of Simulium cuasiexiguum, Simulium subexiguum and Simulium sp. A are unknown. Simulium gonzalezi larvae are not included due to lack of information. – –

Cephalic apotome with well-developed positive spots (Fig. 56 K); rectal papillae without diverticula; anal sclerite without scales ................................. Simulium llutense Cephalic apotome with negative or scarcely evident spots (Fig. 55 C); rectal papillae with 4-12 diverticula on each lobe (Fig. 55 G); anal sclerite with abundant scales (Fig. 55 H) ............................................. Simulium exiguum, Simulium paraguayense

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SIMULIUM (NOTOLEPRIA)

Hembras 1 –

2(1) – 3(2) – 4(3) –

Pelos del escudo y escutelo cilíndricos distribuidos uniformemente sin brillo metálico; con cierta iluminación presenta 1+1 manchas grisáceo claro; paraprocto con pequeña prolongación distal (Fig. 56 E) ..................................... Simulium llutense Pelos del escudo y escutelo achatados (Fig. 5 B) dispuestos en grupos (Fig. 5 A) con aspecto de escamas de brillo metálico; con cierta iluminación no presenta 1+1 manchas anteriores grisáceas; paraproctos con prolongación distal generalmente bien evidente (Figs. 54 I, Q, X) ...................................................................................... 2 Paraprocto con prolongación distal reducida (Fig. 54 X); ramas posteriores de la horquilla genital larga, delgada y poco esclerotizadas .................................................. ................................................. Simulium gonzalezi, Simulium subexiguum, Simulium sp. A Paraprocto con prolongación distal bien desarrollada (Fig. 54 I, Q); ramas posteriores de la horquilla genital relativamente cortas y bien esclerotizadas ...... 3 Prolongación distal del paraprocto relativamente corto y en forma de gancho; octavo esternito fuertemente esclerotizado mediano y distalmente ......................... ............................................................................................................. Simulium cuasiexiguum Prolongación distal del paraprocto relativamente larga (Fig. 54 I, Q); octavo esternito no muy esclerotizado mediano y distalmente (Fig. 54 F) ......................... 4 Prolongación del paraprocto grueso en la base (Fig. 54 I); tallo de la horquilla genital comparativamente más corto en relación a las ramas posteriores (Fig. 54 H) ........... .................................................................................................................... Simulium exiguum Prolongación del paraprocto fino en la base (Fig. 54 Q); horquilla genital comparativamente más larga en relación con las ramas posteriores (Fig. 54 R) .... ............................................................................................................. Simulium paraguayense

Machos 1 – 2(1) – 3(2) –

Escudo con pelos normales dispuestos homogéneamente; placa ventral muy corta (Fig. 56 G) .................................................................................................. Simulium llutense Escudo con pelos achatados dispuestos en grupos con aspecto de escamas; placa ventral más alargada (Figs. 54 O, P, U, W) .................................................................... 2 Gonostilo subtriangular (Figs. 54 M, N); placa ventral sin(Fig. 54 O) ó con (Fig. 54 P) contracción lateral ........... Simulium exiguum, Simulium subexiguum, Simulium sp. A Gonostilo de forma subcuadrada (Figs. 54 S, T); placa ventral con notoria contracción lateral .............................................................................................................. 3 Placa ventral con fuerte contracción medio-lateral, mostrándose como una prolongación espatulada mediana (Figs. 54 U-W) ........................................................ ............................................................................ Simulium gonzalezi, Simulium paraguayense Placa ventral con lateral contracción, mostrándose como una prolongación medio-distal ...................................................................................... Simulium cuasiexiguum

Pupas 1

Branquias con 6 ramas (Fig. 55 L) ................................................................................. 2

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Branquias con 8 ramas (Fig. 55 K) ................................................................................. 3 Tricomas frontales y faciales simples .......................................... Simulium cuasiexiguum Tricomas frontales con 1-3 ramas, faciales con 3-5 ramas .......... Simulium gonzalezi Capullo bien extendido ventralmente y cerrado anteriormente (Fig. 56 H) ........... ...................................................................................................................... Simulium llutense Capullo escasamente extendido ventralmente y abierto anteriormente (Figs. 55 A, I) ..... 4 Capullo no extendido ventralmente (Fig. 55 I) ......................... Simulium paraguayense Capullo con una corta prolongación ventral (Fig. 55 A) ............................................. .................................................. Simulium exiguum, Simulium subexiguum, Simulium sp. A

Larvas Larvas de Simulium cuasiexiguum, Simulium subexiguum y Simulium sp. A no se conocen y la de Simulium gonzalezi no se incluye por escasa información. – –

Apotoma cefálico con manchas positivas bien evidentes (Fig. 56 K); papilas rectales sin divertículos; esclerito anal sin escamas ........................................... Simulium llutense Apotoma cefálico con manchas negativas o escasamente evidentes (Fig. 55 C); papilas rectales con 4-12 divertículos sobre cada lóbulo (Fig. 55 G); esclerito anal con abundantes escamas (Fig. 55 H) .......... Simulium exiguum, Simulium paraguayense

Simulium (Notolepria) exiguum ROUBAUD (Figs. 5D, 54 A-P; 55 A-H.) Simulium exiguum ROUBAUD, 1906b: 108-109; WYGODZINSKY, 1951a: 214-217; TIDWELL et al., 1980: 119, 121, 126. Simulium (Notolepria) exiguum DALMAT, 1955: 112; COSCARÓN & WYGODZINSKY, 1975: 227-287; COSCARÓN, 1987: 20; SHELLEY et al., 1989: 89; COSCARÓN, 1991: 147-148; SHELLEY et al., 2000: 177. COSCARÓN ARIAS, 2003: 195, 198, 204, 209, 215, 219. Wilhelmia exigua ENDERLEIN, 1921a: 200. Simulium delpontei PATERSON & SHANNON, 1927: 739. Simulium glaucophthalmun KNAB, 1914c: 123. Simulium urubambanum ENDERLEIN, 1934b: 288.

Female: Wing length 1.8-2.5 mm; general color black; antennae dark brown; frons nacreous; scutum and scutellum with bright golden to greenish scales, formed by flat, grouped hairs (Figs. 5 A, B); pleurae with gray pruinosity; legs yellowish brown with darkened areas on first and hind coxae, 3/4 of hind femora, apex of tibiae, tarsi, as shown in male figures 54 J-L; abdomen blackish matte with tergites II-IV brown blackish and V-VII with cereous sheen; fronto-ocular triangle not developed (Fig. 54 A); cibarium with basal portion as in figure 54 B; wing chaetotaxy of anterior veins as in figure 54 C. Median claw with subbasal tooth variable on fore leg (Figs. 5 E, 54 D), present on median, and hind legs (Fig. 54 E); legs with cylindrical and flattened hair (Fig. 5 D); eighth sternite with median area slightly darkened and with about 15 setae on each side (Fig. 54 F); gonapophyses without setae, and sclerotized on internal margin; spermatheca ovoidal with insertion to spermathecal ducts membranous (Fig. 54 G); genital fork with well-sclerotized branches ovoidal (Fig. 54 H); cerci hemispherical distally, paraprocts subtriangular with acuminate distal posterior process (Fig. 54 I).

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Male: Wing length 1.5-2.2 mm; scutum velvety black, bordered posteriorly by gray pollinosity with pilosity similar to scales, as in female; legs as shown in figures 54 J-L; length/width ratio of hind basitarsus = 7.1-7.4; abdomen velvety black with silvery tergites II and VI and 1+1 lateral spots of this color on tergite VII; genitalia as shown in figure M; gonostylus subtriangular with distal spur obsolescent (Fig. 54 N); ventral plate subtriangular without lateral decolletage (Figs. 54 O, P). Median sclerite with median depression and endoparameres with large hooks (Fig. 54 M). Pupa: Cocoon length on dorsum 2.0-2.3 mm; on base 2.0-2.6 mm; body of pupa 1.9-2.2 mm; gills 1.2-1.8 mm; color brown, slipper shaped (Fig. 55 A); eight gill branches emerging from three primary branches, very similar to those of Simulium paraguayense (Fig. 55 K); frontoclypeus and thorax with abundant platelets; frontal and facial trichomes with 2-3 branches, thoracic with 2-6 branches; abdominal chaetotaxy as in figure 55 B. Larva: Maximum length 3.6-4.4 mm; color light gray-greenish, head light brown; cephalic apotome light brown with basal median portion darkened (Fig. 55 C); antenna as in figure 55 D, ratio of proximal, medial and distal segments = 1:1.1:1; mandible with thin and curved marginal teeth, as in Simulium paraguayense (Fig. 55 E); cephalic fan with about 40 rays; hypostomium well pigmented on anterior margin; setae 3 or 4 in number disposed in one row; postgenal cleft deep, rounded anteriorly (Fig. 55 F), postgenal bridge about 1/ 3 of hypostomium length; thoracic proleg sclerite with 9-10 groups of spines; rectal papillae with 4-8 diverticula on each lobe (Fig. 55 G); anal sclerite with abundant scales arranged in two to four rows (Fig. 55 H); anal ring with 70-76 rows with 13-15 hooks each. Distribution: Argentina: Jujuy, Salta, Tucumán; Bolivia: Beni, Cochabamba, Tarija; Brazil: Amazonas, Distrito Federal, Goiás, Mato Grosso, Minas Gerais, Roraima, São Paulo; Colombia: Cauca, Magdalena, Meta, Valle; Ecuador: Bolivar, Bucay, Cañar, Cotopaxi, El Oro, Esmeraldas, Guayas, Imbabura, Loja, Los Rios, Manabi, Napo, Pastaza, Pichincha, Puyo; Guatemala; Mexico: Morelos; Panama: Darien; Peru: Huanuco, Junin, Lima; Venezuela: Amazonas, Anzoategui, Aragua, Barinas, Bolivar, Carabobo, Cojedes, Distrito Federal, Guarico, Lara, Mérida, Miranda, Monagas, Sucre, Tachira, Trujillo, Yaracuy, Zulia. Bionomics: Simulium exiguum breeds in creeks and rivers with fast currents, in crystalline and temperate water flowing over aquatic vegetation and branches of trees or rocks, sometimes up to 2 m deep. Females bite humans and animals, both domestic and wild, such as tapirs. Simulium exiguum is a vector of Onchocerca volvulus in Ecuador, Colombia, Venezuela, and Brazil, and of Mansonella ozzardi in Argentina. Discussion: At least five species, Simulium paraguayense, S. cuasiexiguum, S. subexiguum, S. sp. A and S. gonzalezi, are very close to Simulium exiguum. The first species shows a few differences, such as in the female paraprocts, gonostylus and ventral plate, and some variations in ornamentation, size of legs, and proportion of larval antennae, as explained for that species. Simulium gonzalezi also shows differences in paraproct shape and especially in the number of pupal gill branches. Simulium cuasiexiguum is also differentiated by the 6 gill branches of the pupa. Simulium subexiguum has a short distal posterior process on the paraproct (similar to S. gonzalezi). As shown in Simulium sp. A, some of these differences are considered by SHELLEY et al. (1989) as possible variants of Simulium exiguum. It is also possible to see great variation in size and female feeding habits (SHELLEY et al., 2000),

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Figure 54. A-P: S. (Notolepria) exiguum; A-I: Female; A: fronto-ocular triangle; B: Basal portion of cibarium; C: wing anterior portion of chaetotaxy; D: claw without subbasal tooth; E: claw with subbasal tooth; F: eighth sternite and gonapophysis; G: spermatheca; H: genital fork; I: cercus and paraproct. J-P: Male; J-L: fore, mid and hind leg, respectively; M: genitalia showing gonocoxite, gonostylus, ventral plate, endoparameres and median sclerite; N: gonostylus; O, P: ventral plate of different specimens. Q-W: S. (N.) paraguayense; Q-R: Female; Q: cercus and paraproct; R: genital fork; S-W: Male; S-T: gonostylus in different position; U-W: ventral plate in different position. X: S. (N.) gonzalezi, cercus and paraproct.

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common in species with large geographic distributions. As indicated by the presence of at least 4 cytospecies in Ecuador (CHARALAMBOUS et al., 1993), it is probable that there are more morphological differences in the range of S. exiguum, and these small variations could correspond to sibling species. We prefer to maintain S. exiguum as a species, pending further information, such as cytological or molecular, to clarify its identity. Simulium (Notolepria) paraguayense SCHROTTKY (Figs. 54 Q-W; 55 I-K.) Simulium paraguayense SCHROTTKY, 1909: 63; LUTZ, 1909: 144-145; 1910: 235; SURCOUF & GONZÁLEZ RINCONES, 1911: 291-292; LUTZ et al., 1918: 165; LUTZ & NUÑEZ TOVAR, 1928: 43; WYGODZINSKY, 1953a: 298-301. Simulium (Eusimulium) paraguayense LUTZ, 1909: 144; ORFILA, 1939: 1531. Eusimulium paraguayense PINTO, 1931: 678; LANE & PORTO, 1939: 169, 175; BRICEÑO IRAGORRI, 1941: 114; LANE & VULCANO, 1943: 432. Simulium (Notolepria) paraguayense COSCARÓN & WYGODZINSKY, 1975: 283-287; COSCARÓN, 1987: 20; 1991: 148-150; SHELLEY et al., 2001: 485.

Female: Size, ornamentation and morphology very similar to those of Simulium exiguum but differentiated by the following: claw subbasal tooth always present on all legs; hind leg with half of distal portion of femur, apex of basitarsus, and second and third tarsomeres more darkened; paraproct distal process thinner acuminate apically and more elongated (Fig. 54 Q); genital fork stem proportionally more elongated and posterior branches more sclerotized (Fig. 54 R). Male: Very similar to Simulium exiguum. Maximum wing length 2.5 mm; gonostylus subquadrate to subovoidal with very small spur (Figs. 54 S, T); ventral plate with lateral constriction at about half length (more or less evident in accordance with position, (Figs. 54 U, V) and with small median elevation in lateral view (Fig. 54 W). Pupa: The small differences from Simulium exiguum are the more translucent cocoon, permitting the pupal body to be seen(Fig. 55 I) and more abundant platelets on the frontoclypeus and thorax (Figs. 55 J-K). Larva: Greatly similar to S. exiguum; showing only small differences in antennal segment ratios: 1:1.2:1.4; anal ring with 52-62 rows 11-13 hooks each, and presence of one latero-mandibular process. Distribution: Argentina: Misiones; Brazil: Bahia, Minas Gerais, Rio de Janeiro, São Paulo (cited by LUTZ); Paraguay; Venezuela: Aragua, Distrito Federal (cited by LUTZ & NUÑEZ TOVAR). Bionomics: Simulium paraguayense breeds in swift, large river up to 1 m deep, flowing over Podostemaceae. Females show great anthropophily and their bites are very painful. Discussion: Simulium paraguayense has only small differences, as shown in the description, that permit it to be separated from S. exiguum. These could be considered population differences, but S. exiguum has been known until now from 4 cytotypes only in Ecuador, showing the presence of several species in the same morphotypic species. This is the motive for not synonymizing S. paraguayense with S. exiguum. Simulium incertum LUTZ, with 8 pupal gill branches, probably is a synonym of S. paraguayense, which LUTZ also suspected. Material collected by us very close to the type area corresponds to S. paraguayense.

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I J E

A D B L C

K

G F

H

Figure 55. A-H: S. (Notolepria) exiguum; A-B: Pupa; A: general aspect; B: abdominal chaetotaxy. CH: Larva; C: cephalic apotome; D: antenna; E: dental portion of mandible; F: hypostomium and postgenal cleft; G: rectal papillae; H: anal sclerite with rows of scales on superior area and some hair on inferior. I-K: S. (N.) paraguayense; I-K: Pupa; I: general aspect; J: frontoclypeus; K: dorsal portion of thorax with trichomes and gill. L: S. (N.) gonzalezi; Pupa, dorsal portion of thorax showing tubercles, trichomes and gill.

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Simulium (Notolepria) gonzalezi VARGAS & DÍAZ NÁJERA (Figs. 54 X, 55 L.) Simulium gonzalezi VARGAS & DÍAZ NÁJERA, 1953b: 235; SHELLEY & ARZUBE, 1985: 329. Simulium (Notolepria) gonzalezi VARGAS & DÍAZ NÁJERA, 1957b: 158, 338; COSCARÓN, 1987: 20; SHELLEY et al., 1989: 88; IBÁÑEZ-BERNAL, 1992: 263; CHARALAMBOUS et al., 1997: 19; SHELLEY et al., 2002: 135; COSCARÓN ARIAS, 2003: 195, 198, 204, 208, 215, 219. Simulium gonzalesi VULCANO, 1967: 12 (incorrect spelling). Simulium exiguum KNAB, 1913: 154; VARGAS, 1941a: 115; 1941b: 213; 1942a: 229; 1943b: 363-365. Simulium (Notolepria) exiguum VARGAS et al., 1946: 171; DALMAT, 1955: 112-116.

Female: Wing length 1.5-2.2 mm; color and morphology of head, scutum, abdomen and appendages as in Simulium exiguum; claws without basal tooth, sometimes restricted to hind leg; paraproct with distal process more reduced than in S. exiguum (Fig. 54 X). Male: Wing length 1.1-2.0 mm; color and morphology very similar to that of Simulium exiguum; gonostylus subquadrate, very close to that of S. paraguayense; ventral plate very close to that of S. exiguum; some males dichoptic with external morphology similar to that of female, with narrow frons and more elongated antennal segments; abdominal ornamentation similar to that of S. exiguum. Pupa: Cocoon length dorsally 1.4-2.0 mm, along base 1.4-2.5 mm, pupa body 1.9-2.4 mm, gills 1.0-1.8 mm. Cocoon slipper shaped, with reinforced rim; gill with 6 branches (Fig. 55 L); chaetotaxy of cephalothorax and abdomen as in Simulium exiguum. Larva: Maximum length 4.2 mm; coloration and morphology as in Simulium exiguum; negative cephalic apotome pattern of Simulium gonzalezi also found in some specimens of S. exiguum (CHARALAMBOUS et al., 1997). Distribution: Belize: Cayo, Stan Creek, Toledo; Ecuador: Esmeraldas; Guatemala; Mexico: Colima, Chiapas, Guerrero, Hidalgo, Jalapa, Michoacan, Nayarit, Oaxaca, Potosi, San Luis, Tabasco, Tamaulipas, Veracruz. Bionomics: Simulium gonzalezi breeds in fast-flowing small creeks to rivers, from 1 to 1500 m in altitude in water of about 20 °C, on aquatic vegetation. Females are haematophagous, biting humans, domestic mammals and birds. This species is recognized as a vector of Onchocerca volvulus in Mexico and Guatemala. Discussion: Several Simulium (Notolepria) species, such as S. exiguum, S. paraguayene, S. cuasiexiguum, S. subexiguum and Simulium sp. A, are very similar to S. gonzalezi; it is possible to differentiate the first two by the 8 gill branches and more elongate distal process on the paraproct. The small differences in the ventral plate and gonostylus, as well as the negative spots on the cephalic apotome of larvae of S. gonzalezi, are included in the variation of characters of the other two species. All compared species (except S. cuasiexiguum) have pupal gills with 8 branches. Simulium (Notolepria) cuasiexiguum SHELLEY, LUNA-DIAS, MAIA-HERZOG & LOWRY Simulium (Notolepria) cuasiexiguum SHELLEY et al., 2000: 178; SHELLEY et al., 2001: 483; SHELLEY et al., 2002: 154.

Female: Wing length 1.5-2.1 mm; coloration and morphology similar to Simulium exiguum, S. paraguayense and S. gonzalezi; the great differences are in the bigger amplitude of

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sclerotized area of eighth sternite, the thinner and curved distal paraproct process, and the insertion of spermathecal duct to sclerotized spermatheca well, that can reach to the basal portion of spermathecal duct. Male: Wing length 1.4-1.8 mm; as in the female, it is very similar to the other tree species in this subgenus; it shows differences in the more acuminate ventral plate distal process and with Simulium paraguayense and S. gonzalezi by the subquadrate gonostylus. Pupa: Cocoon length dorsally 2.1-2.7 mm, ventrally 2.2-3.1 mm; body pupa 2.0-2.1 mm; gills 1.3-1.5 mm; gill has 6 branches with similar arrangement as Simulium gonzalezi. Larva: Unknown. Distribution: Brazil: Goiás, Mato Grosso. Bionomics: Simulium cuasiexiguum breeds in fast flowing rivers, over aquatic vegetation, in grasslands and gallery forests. Females have zoophilic haematophagous habits. Discussion: As was shown in the description, S. cuasiexiguum is very similar to S. exiguum, S. paraguayense and S. gonzalezi. The pupal gills with 8 branches allow S. cuasiexiguum to be separated from the first two species. The shorter distal process of the paraproct and the subtriangular gonostylus differentiate S. gonzalezi from S. cuasiexiguum. Simulium (Notolepria) subexiguum FIELD Simulium (Notolepria) subexiguum FIELD, 1967: 189; IBÁÑEZ-BERNAL, 1992: 286.

Female: Similar to Simulium exiguum; differs in paraproct, which has very short distal posterior process; also very similar to Simulium gonzalezi, S. llutense and Simulium sp. A. Male: Similar to Simulium exiguum in external aspects; gonostylus and ventral plate included in variations of S. exiguum. Pupa: Gill with 8 branches, as in Simulium exiguum, S. paraguayense, S. llutense and Simulium sp. A; without any difference from those species. Larva: Unknown. Distribution: Mexico: Chiapas; Panama: Canal Zone. Bionomics: The species has been collected, according to the species author, on “trailing grass”. Discussion: The paraproct morphology of S. subexiguum allows it to be differentiated from S. exiguum. The male subtriangular gonostylus and pupal gills with 8 branches separate it from S. gonzalezi. Simulium (Notolepria) sp. A IBÁÑEZ-BERNAL Simulium (Notolepria) sp. A IBÁÑEZ-BERNAL, 1992: 279.

IBÁÑEZ-BERNAL gave a detailed description of this species based on pharate adults. It has similarities with S. cuasiexiguum, S. exiguum and S. subexiguum in the adults and also the pupal gill with 8 branches. It can be differentiated from S. exiguum by the short paraproct posterior process and other small anatomical differences, some of them included among the variation in S. exiguum. Compared with S. gonzalezi, it shows differences in the paraproct distal elongation, which is greater in Simulium sp. A, and in the pupal gill with 8 branches. It is practically the same as S. subexiguum. As explained by IBÁÑEZ-BERNAL, it will

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be necessary to study more material of all stages. Maybe Simulium sp. A is a variation of S. gonzalezi with 8 branches, as suggested by DALMAT (1955) in Guatemala for S. exiguum with 8 and 6 branches (IBÁÑEZ-BERNAL, 1992: 286). We have seen the same situation in Simulium (Pternaspatha) annulatum with 6 and 8 gill branches. Distribution: Mexico: Chiapas. Bionomics: Simulium sp. A lives in the same places as S. gonzalezi. Simulium (Notolepria) llutense COSCARÓN & MATTA (Fig. 56.) Simulium (Notolepria) llutense COSCARÓN & MATTA, 1982: 49; COSCARÓN, 1987: 20; 1991: 150.

Female: Wing length 2.1-2.5 mm; head blackish, frons and clypeus dark brown; antennae, palpi and proboscis dark grayish-brown with scape and pedicel brown; scutum, scutellum black with abundant pilosity brownish or greenish-gray; with some light incidence 1+1 anterior submedian subtriangular grayish spots can be observed, changing light position these spots appear blackish. Metanotum velvety black; pleurae blackish with gray pollinosity; wing veins yellowish brown with hair and setae dark brown; legs yellowish brown with fore tarsomere and distal portion of median and posterior tarsomeres blackish; abdomen matte black, with cereous sheen on VI-IX tergites; fronto-ocular triangle slightly wider than long (Fig. A). Maxillary palp sensory organ about half palpomere length; cibarium basally as in figure B; legs without petaloid hair; calcipala well developed; claws without subbasal teeth; basal portion of R and Sc without hair or setae; eighth sternite (Fig. C) with about 70 setae; gonapophyses with internal border well sclerotized; genital fork with well-sclerotized branches (Fig. D); paraproct with truncate distal border and short distal posterior process (Fig. E). Male: Wing length 1.8 mm; head and scutum black, the latter with golden hair, and with some light with anterior fourth grayish; leg coloration as in female, but darker, especially hind legs; pleurae blackish with gray pollinosity; abdomen black with 1+1 light silvery spots on tergites II, VI-VIII; tergite IX grayish black; length/width ratio of hind basitarsus = 7; gonostylus subtriangular, smoothly emarginated apically with apical spur obsolescent (Fig. F); ventral plate shorter than wide at base (Fig. G); endoparameres with strong hooks. Pupa: Cocoon length at base 2.7-3.0 mm, at dorsum 1.8-2.5 mm; gills slipper shaped with thick weave strong, projected ventrally (Fig. H); frontoclypeus (Fig. I) and exposed area of thorax with regular quantity of smooth tubercles; frontoclypeus with 1+1 facial and 2+2 frontal trichomes simple or bifid, thorax dorsally with 7 multirramous trichomes; gill with 8 branches, primary branches with short basal trunk (Fig. J). Larva: Maximum length 4.6-4.8 mm; color light grayish-green, more intense on dorsum; head yellowish brown; cephalic apotome with isolated median, sublateral and subbasal positive spots (Fig. K); antenna longer than stem of cephalic fan (Fig. L); ratio of proximal, medial and distal segments = 1:1.2:1.1. Median tooth of hypostomium at same level as lateral teeth; with 4 setae on each side (Fig. M); postgenal cleft deep; with postgenal bridge shorter than half of hypostomium length. Mandible with 6 internal teeth, second marginal teeth shorter than first tooth by half; cephalic fan with about 25

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D B A E

G F

C H L

K

I

J

M

Figure 56. S. (Notolepria) llutense. A-E: Female; A: fronto-ocular triangle; B: cibarium, basal portion; C: eighth sternite and gonapophysis; D: genital fork; E: cercus and paraproct. F-G: Male; F: gonocoxite and gonostylus; G: ventral plate. H-J: Pupa; H: general aspect; I: frontoclypeus; J: portion of thorax with trichomes and gill. K-M: Larva; K: head dorsal view; L: antenna; M: hypostomium and postgenal cleft.

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rays; anal sclerite with some hair, scales not evident; anal ring with about 74 rows with 1314 hooks each; rectal papillae with 3 simple lobes. Distribution: Chile: Tarapacá. Bionomics: Simulium llutense breeds in small creeks of the Chilean desert area, which flow over aquatic vegetation. The feeding habits of the females are unknown. Discussion: Similarity in adult genitalia, cibarial shape, pupal gill and chaetotaxy, as well as in the larvae, allow S. llutense to be included in Notolepria, but the species shows some differences from other species of this subgenus, such as the absence of grouped hairs simulating scales on the scutum, 1+1 subtriangles for the anterior silvery spots on the female scutum; a small fronto-ocular triangle in the female and shorter ventral plate paraprocts, which are very similar to those of S. gonzalezi. Simulium spinifer KNAB from Peru, of uncertain position, shows some similarities in the female description, but the male genitalia, as well as the other stages, do not allow us to decide if they are the same species; therefore, we consider S. spinifer species inquirenda. Species inquirendae Simulium (Notolepria) blantoni FIELD Simulium (Notolepria) blantoni FIELD, 1967a: 192.

This species is known only from a female that is very close to Simulium gonzalezi and S. subexiguum. The absence of a male and pupa do not allow it to be adequately differentiated from the two previously mentioned species. Distribution: Panama: Bocas del Toro. Simulium incertum LUTZ Simulium incertum LUTZ, 1910: 259; PINTO, 1931: 755.

LUTZ described this species without illustrations. It is not easy to identify, and he says at the end of his description that it might be S. paraguayense SCHROTTKY. PINTO (1931) published a pupal gill (Fig. 8 j) from the type area, and considered it to be that of S. incertum, but it is not similar to the PINTO description of S. paraguayense that agrees with our interpretation of this species. It is not possible to be sure what S. incertum might be. Distribution: Brazil: Minas Gerais, São Paulo. Simulium paranense SCHROTTKY Simulium paranense SCHROTTKY, 1909: 63; VULCANO, 1967: 18. Simulium (Notolepria) paranense COSCARÓN, 1987: 20; CROSSKEY & HOWARD, 1997: 53.

The description is not sufficient to permit identification of this species. Distribution: Paraguay: Alto Paraná.

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Simulium (Chirostilbia) ENDERLEIN (Figs. 57-63, 140.) Type-species Chirostilbia flavifemur ENDERLEIN, 1921b: 79 = Simulium pertinax KOLLAR, 1832: 117.

Female: Species of medium size, wing length 2.4-3.6 mm; general coloration blackish with soft pollinosity gray blackish, but can be black-grayish to greenish yellow; some illumination shows 1+1 silvery submedian vittae changing to black with other lighting; scutellum brownish, metanotum blackish; abdomen blackish to grayish green to brown yellowish; frons medium wide; fronto-ocular triangle variable from deep to flat; cibarium basally with median area smooth and cornuae branches well sclerotized; wing generally with hair on basal sector of R; claws with or without basal tooth; gonapophysis subtriangular, nearly two times longer than wide at base; genital fork with anterior branch about one quarter longer than each posterior branch. Male: Wing length 2.2-3.5; scutum black to brown yellowish or reddish, bordered laterally and posteriorly by silver band, enlarged on pronotal angles; abdomen blackish to gray greenish with 1+1 grayish lateral spots on tergites II, V-VIII; gonocoxite subtrapezoidal to subquadrate, longer than gonostylus; gonostylus subtriangular to subtrapezoidal, curved, acuminate distally, without apical spur, or very reduced (like small seta); ventral plate wider than long, with slight curvature in lateral view and not acuminate apically. Median sclerite wide with strong median concavity; parameres with robust hooks. Pupa: Cocoon slipper to shoe shaped, compact with resistant weave, evident threads compact or with apertures and festoons; frontoclypeus and exposed portion of thorax with abundant to no tubercles; head with 2+2 frontal and 1+1 facial trichomes; thorax dorsally with 5-6 trichomes on each side; gills each with 8-10 to 120 branches, sometimes relatively thick; terminal spines relatively short. Larva: Body cuticle without hair or scales; cephalic apotome variable, generally with subtriangular darker medium basal area with median longitudinal, submedian and subbasal spots positive or negative; antenna proximal segment shorter than medial segment and this shorter than distal. Mandible with 8-9 internal teeth; second marginal tooth generally about half of first; cephalic fan with about 45 rays; postgenal cleft deep with postgenal bridge about half of hypostomium length; lateral sclerite of thoracic proleg with about 25 teeth, arranged in groups; anal sclerite with single hairs and sometimes with scales; anal ring with 65-190 rows with 12-24 hooks each; rectal papillae with 4-24 diverticula on each lobe. Distribution: Simulium (Chirostilbia) is a South American taxon, distributed predominantly in the intertropical non-Andean region (Fig. 140). Discussion: This subgenus is composed of two species groups: · PERTINAX species group: imagoes blackish; female claw with subbasal tooth; male gonostylus subquadrate, about as long as wide at base and without median distal diagonal crest; ventral plate about as long as wide, generally curved lateroapically; pupa generally with abundant tubercles on frontoclypeus and thorax, and cocoon slipper to shoe shaped; larval cephalic apotome basally with darker subtriangular. · SUBPALLIDUM species group: imagoes brown grayish to green yellowish especially females; female claw generally without subbasal tooth; male gonostylus

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subtriangular, longer than wide at base with median distal diagonal crest; ventral plate wider than long not curved apically; pupal frontoclypeus and thorax without tubercles or these very scarce; cocoon slipper shaped; larval cephalic apotome without basally darker subtriangular area. KEYS

TO

SIMULIUM (CHIROSTILBIA)

SPECIES

Females Not included: Simulium friedlanderi (unknown female) and S. empascae (scarce information). 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(4) – 8(7) – 9(1) –

General coloration blackish; scutum and abdomen black to dark reddish brown; claw with subbasal tooth (Fig. 57 M) ...................... PERTINAX species group......2 General coloration grayish brown to yellowish green; scutum yellowish brown to lead gray; abdomen yellowish, grayish brown to greenish gray; claw without subbasal tooth (Fig. 60 O) ................................ SUBPALLIDUM species group......9 Scutum blackish, without submedian silvery vittae joined to posterior grayish area .................................................................................................................... Simulium pertinax Scutum blackish to reddish brown, with 1+1 submedian silvery anterior subrectangular spots or vittae joining to posterior grayish area in some incident light (Figs. 57 F, G, J, O) ................................................................................................... 3 Scutum with 1+1 submedian silvery anterior subrectangular small spots ............... ............................................................................................................. Simulium bifenestratum Scutum with 1+1 submedian silvery vittae joining posterior grayish area (Figs. 57 F, G, J, O) ............................................................................................................................. 4 Scutum blackish; wing length 2.0-3.0 mm .................................................................... 5 Scutum dark reddish brown to light yellowish brown; wing length 2.5-3.6 mm ................................................................................................................................................ 7 Sc without hair; wing length 2.0-2.2 mm; scutum with pile silvery .......................... .....................................................................................................................Simulium serranus Sc with hair; wing length 2.2-3.0 mm; scutum with pile golden or with lead gray tones ...................................................................................................................................... 6 Scutum with abundant pile, with lead gray tones ................. Simulium spinibranchium Scutum with scarce pile, with golden tones ................................... Simulium laneportoi Scutum light yellowish brown, with light grayish to yellow pile ................................ ...............................................................................................................Simulium riograndense Scutum dark grayish brown or reddish brown, with golden to silvery pile .......... 8 Cercus with curved distal border; paraproct about 1.5 times longer than wide at base (Fig. 57 N); scutum dark grayish brown ............................... Simulium distinctum Cercus with straight distal border; paraproct about 2.5 times longer than wide at base (Fig. 57 U); scutum dark reddish brown .................................... Simulium obesum Scutum light yellowish brown with light brown posterior area; pleurae yellowish brown .................................................................................................. Simulium subpallidum Scutum dark grayish leaden, with blackish brown posterior area; pleurae dark grayish ................................................................................................................................. 10

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10(9) –

Wings with hair on basal sector of R; abdomen greenish with dark brown tergal plates (Fig. 60 E) ................................................................................. Simulium acarayense Wings without hair on basal sector of R; abdomen brownish with dark grayish brown tergal plates ............................................................................... Simulium papaveroi

Males Simulium empascae and S. friedlanderi are not included due to lack of information. 1

– 2(1) – 3(2) – 4(2) – 5(1) – 6(5) –

Gonostylus subquadrate, about as long as wide at base and without median distal diagonal crest (Figs. 58 B, F, G, I, J, L, O); ventral plate about as long as wide, generally curved apicolaterally (Figs. 58 A, C, H, K, M, P) ........................................ ......................................................................................... PERTINAX species group......2 Gonostylus subtriangular, longer than wide at base, with median distal diagonal crest (Figs. 61 D, F, H, I) ventral plate wider than long, not curved apically (Figs. 61 E, J, K) ............................................................ SUBPALLIDUM species group......5 Scutum dark grayish brown ............................................................................................. 3 Scutum black ....................................................................................................................... 4 Wing length 2.4-3.3 mm; scutum blackish; gonostylus with relatively short process on external distal side (Fig. 58 J); ventral plate relatively elongated (Fig. 58 K) .... ................................................................................................................. Simulium distinctum Wing length 3.3-3.5 mm; scutum reddish brown to yellowish; gonostylus with well-developed process on external distal side (Fig. 58 L), ventral plate relatively short (Fig. 58 M) ...............................................................................Simulium riograndense Wing length 3.2-3.5 mm; gonostylus with lateral external and posterior sides concave (Fig. 58 O) .......................................... Simulium obesum, Simulium bifenestratum Wing length 2.0-3.1 mm, gonostylus with lateral external and posterior sides from convex to straight (Figs. 58 B, F, G) ................................................................................ ....... Simulium pertinax, Simulium laneportoi, Simulium serranus, Simulium spinibranchium Scutum yellowish brown to reddish brown ................................ Simulium subpallidum Scutum black ....................................................................................................................... 6 Wings with hair on basal sector of R; gonostylus only a little longer than wide at base (Fig. 61 D) ................................................................................... Simulium acarayense Wings without hair on basal sector of R; gonostylus nearly 2 times longer than wide at base (Fig. 61 F) ....................................................................... Simulium papaveroi

Pupae 1 – 2(1) –

Gill with 8 filaments; cocoon generally not covering gill branches and with anterior portion smooth, without festoons (Figs. 59 A, D, G, J; 62 A, C, D, G) ....................... 2 Gill with 10-120 filaments; cocoon covering gill and anterior portion with festoons (Figs. 59 L, O, S) ................................................................................................ 8 Cocoon covering branches (Fig. 59 J); primary and secondary branches emerging as a bundle (Fig. 59 K) ......................................................................... Simulium empascae Cocoon not covering gill (Figs. 59 A, D, G; 62 A, C, D, G); primary and secondary branches emerging isolated, not as bundle (Figs. 59 C, F, H; 62 B, F, I) ............... 3

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Gill filaments thick basally (Fig. 59 H) ................................... Simulium spinibranchium Gill filaments not thick basally ........................................................................................ 4 Frontoclypeus and exposed portion of thorax with abundant tubercles (Figs. 59 B, E) ........................................................................................................................................... 5 – Frontoclypeus and exposed portion of thorax without or with few tubercles (Figs. 62 E, H) ................................................................................................................................ 6 5(4) Gill length about as long as pupal body length (Fig. 59 D) and arranged in 1 plane (Fig. 59 F); frontoclypeus and thorax with only moderate abundance of tubercles (Fig. 59 E) ................................................................................................ Simulium serranus – Gill length about half of pupal body length (Fig. 59 A), and arranged in more than 1 plane (Fig. 59 C); frontoclypeus and thorax with very abundant tubercles (Fig. 59 B) .............................................................. Simulium pertinax, Simulium laneportoi 6(4) Gill branches slightly grouped basally, especially dorsal branches (Fig. 62 I); primary and secondary dorsal branches thicker than ventral branches; cephalic and thoracic trichomes simple or bifid (Fig. 62 H) ...................... Simulium papaveroi – Gill branches open (Figs. 62 B, F); primary and secondary dorsal branches no thicker than ventral branches; cephalic and thoracic trichomes with 3-6 branches (Fig. 62 E) ............................................................................................................................ 7 7(6) Gill filaments approximately straight and disposed in 1 plane (Figs. 62 C, D, F); sometimes rounded tubercles present on frontoclypeus and thorax dorsally (Fig. 62 E) ...................................................................................................... Simulium acarayense – Gill filaments with slight curvature and arranged like basket (Figs. 62 A, B); without tubercles on frontoclypeus or thorax, or if present, with conical shape .............................................................................................................. Simulium subpallidum 8(1) Gill with about 120 branches; thorax covered totally with trichomes (Fig. 59 U) ...................................................................................................................... Simulium obesum – Gill with 10-11 branches; thorax with 5-6 trichomes on each side ......................... 9 9(8) Gill with 11 branches very thick and with strong curvature (Fig. 59 R) .................. ................................................................................................................ Simulium friedlanderi – Gill with 10 branches filamentous and without strong curvature (Figs. 59 N, Q) .............................................................................................................................................. 10 10(9) Cocoon with 1+1 anterolateral apertures; frontoclypeal margin elevated as a crown ................................................................................................. Simulium bifenestratum – Cocoon without 1+1 anterolateral apertures (Figs. 59 L, O); frontoclypeal margin not elevated ....................................................................................................................... 11 11(10) Frontoclypeus and thorax with regular quantity of tubercles uniformly distributed (Fig. 59 M); cocoon length basally 3.0 mm, dorsally 2.8 mm, gills 1.0 mm ............ ................................................................................................................. Simulium distinctum – Frontoclypeus and thorax with scarce quantity of tubercles, distributed on frontoclypeus near facial trichomes (Fig. 59 P) and on exposed portion of thorax; cocoon length basally 4.2-4.7 mm; dorsally 4.5-4.7 mm; gills 1.7-2.0 mm .............. ...............................................................................................................Simulium riograndense

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Larvae Simulium obesum larva not included (unknown). 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(2) – 7(6) – 8(6) – 9(8) –

Postgenal cleft subtriangular (Fig. 63 F) to dome shaped (Fig. 63 C); anal ring with 85-204 rows of hooks; maximum length 6.0-11.8 mm ............................................. 2 Postgenal cleft subovoidal (Fig. 63 Q); anal ring with 60-81 rows of hooks; maximum length 5.0-5.8 mm ............................................................................................. ......................................... Simulium subpallidum, Simulium acarayense, Simulium papaveroi Anal ring with 145-204 rows of hooks; maximum length 8.1-11.8 mm ................ 3 Anal ring with 76-122 rows of hooks; maximum length 5.6-7.5 mm .................... 6 Anal ring with 201-204 rows of hooks; rectal papillae with 4-6 diverticula on each lobe ........................................................................................................... Simulium empascae Anal ring with 145-190 rows of hooks; rectal papillae with 7-26 diverticula on each lobe .............................................................................................................................. 4 Postgenal cleft dome shaped; cephalic apotome darkened totally on basal 1/3 and continued anteriorly as triangle nearly reaching frontal area ...... Simulium bifenestratum Postgenal cleft subtriangular with blunt apex; cephalic apotome darkened only on basal 1/3 .............................................................................................................................. 5 Anal ring with 170-190 rows of hooks; maximum length 8.5-9.0 mm .................... ................................................................................................................. Simulium distinctum Anal ring with 145-152 rows of hooks; maximum length 9.5-10.5 mm .................. ...............................................................................................................Simulium riograndense Cephalic apotome darkened totally on basal 1/3 and continued anteriorly as triangle nearly reaching frontal area (Fig. 63 A) .......................................................... 7 Cephalic apotome darkened only on middle of base (Fig. 63 D) or basal 1/4 and slightly continued on midlongitudinal area ................................................................... 8 Postgenal cleft dome shaped (Fig. 63C); anal ring with 88-100 rows of hooks .................................................................................................................... Simulium pertinax Postgenal cleft subtriangular; anal ring with 76-86 rows of hooks ........................... ..........................................................................................................Simulium spinibranchium Cephalic apotome darkened on basal 1/4 and slightly continued on midlongitudinal area, median longitudinal and submedian spots positive............... .................................................................................................................. Simulium laneportoi Cephalic apotome darkened only at mid base, median longitudinal and submedian spots negative (Fig. 63 D) ................................................................................................ 9 Anal ring with 85-88 rows of hooks; rectal papillae with 15-18 diverticula on each lobe .............................................................................................................Simulium serranus Anal ring with 105-122 rows of hooks; rectal papillae with 20-24 diverticula on each lobe .............................................................................................. Simulium friedlanderi

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SIMULIUM (CHIROSTILBIA)

Hembras Simulium friedlanderi (hembra desconocida) y S. empascae (escasa información) no se incluyen. 1 –

2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(4) – 8(7) – 9(1) –

Coloración general negruzca; escudo y abdomen negro a castaño rojizo oscuro; uña con diente subbasal (Fig. 57 M) .............................. PERTINAX sp. grupo......2 Coloración general castaño grisáceo a verde amarillento; escudo castaño amarillento a gris plomizo; abdomen desde amarillo, castaño grisáceo a gris verdoso; uña sin diente subbasal (Fig. 60 O) ........................................................... ........................................................................................SUBPALLIDUM sp. grupo......9 Escudo negruzco, no mostrando con ninguna luz bandas plateadas submedianas llegando al área posterior grisácea ...................................................... Simulium pertinax Escudo negruzco a castaño rojizo presentando con alguna luz 1+1 manchas plateadas submedianas subrectangulares o bandas llegando al área posterior grisácea (Figs. 57 F, G, J, O) ............................................................................................ 3 Escudo con 1+1 mancha plateada subrectangular pequeña anterior submediana ............................................................................................................. Simulium bifenestratum Escudo con 1+1 banda plateada submediana, uniéndose al área grisácea posterior (Figs. 57 F, G, J, O) ............................................................................................................ 4 Escudo negruzco; longitud alas 2,0-3,0 mm ................................................................ 5 Escudo castaño rojizo oscuro a castaño amarillento claro; longitud de las alas 2,53,6 mm .................................................................................................................................. 7 Sc sin pelos; longitud ala 2,0-2,2 mm; pilosidad del escudo plateada ...................... .....................................................................................................................Simulium serranus Sc con pelos; longitud ala 2,2-3,0 mm; pilosidad del escudo dorado a gris plomizo ................................................................................................................................................ 6 Escudo con pilosidad abundante con tonalidad gris plomizo ................................... ..........................................................................................................Simulium spinibranchium Escudo con pilosidad relativamente escasa, con tonalidad dorada ........................... .................................................................................................................. Simulium laneportoi Escudo castaño amarillento claro, con pilosidad grisácea clara a amarillenta ........ ...............................................................................................................Simulium riograndense Escudo castaño grisáceo oscuro o castaño rojizo con pilosidad dorada a plateada ................................................................................................................................................ 8 Cerco con borde distal curvo; paraprocto aproximadamente 1,5 veces más largo que ancho en la base (Fig. 57 N); escudo castaño grisáceo oscuro .......................... ................................................................................................................. Simulium distinctum Cerco con borde distal recto; paraprocto aproximadamente 2,5 veces más largo que ancho en la base (Fig. 57 U); escudo castaño rojizo oscuro .............. Simulium obesum Escudo castaño amarillento claro, con fondo castaño claro; pleuras castaño amarillento ......................................................................................... Simulium subpallidum Escudo gris plomizo oscuro con fondo castaño negruzco; pleuras grisáceo oscuro .............................................................................................................................................. 10

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Alas con pelos en sector basal de R; abdomen verdoso con placas tergales castaño oscuro (Fig. 60 E) ............................................................................... Simulium acarayense Alas sin pelos en sector basal de R; abdomen castaño con placas tergales castaño grisáceo oscuro ..................................................................................... Simulium papaveroi

Machos Simulium empascae y S. friedlanderi no se incluyen por escasa información. 1

– 2(1) – 3(2) – 4(2) – 5(1) – 6(5) –

Gonostilo subcuadrado, aproximadamente tan largo como ancho en la base y sin cresta mediana distal (Figs. 58 B, F, G, I, J, L, O); placa ventral aproximadamente tan larga como ancha, generalmente curvada latero apicalmente (Figs. 58 A, C, H, K, M, P) ................................................................................ PERTINAX sp. grupo......2 Gonostilo subtriangular, más largo que ancho en la base, con una cresta medio longitudinal (Figs. 61 D, F, H, I); placa ventral más ancha que larga, no curvada apicalmente (Figs. 61 E, J, K) ..................................SUBPALLIDUM sp. grupo......5 Escudo castaño grisáceo oscuro ..................................................................................... 3 Escudo negro ...................................................................................................................... 4 Longitud ala 2,4-3,3 mm; escudo negruzco; gonostilo con relativamente corta prolongación sobre el costado distal externo (Fig. 58 J); placa ventral relativamente alargada (Fig. 58 K) ................................................... Simulium distinctum Longitud ala 3,3-3,5 mm; escudo castaño rojizo a amarillento; gonostilo con una prolongación bien desarrollada sobre el costado distal externo (Fig. 58 L); placa ventral relativamente corta (Fig. 58 M) ........................................Simulium riograndense Longitud ala 3,2-3,5 mm; gonostilo con el costado lateral externo y posterior cóncavos (Fig. 58 O) ....................................... Simulium obesum, Simulium bifenestratum Longitud ala 2,0-3,1 mm; gonostilo con el costado lateral externo y posterior desde convexo a recto (Figs. 58 B, F, G) ........................................................................ ....... Simulium pertinax, Simulium laneportoi, Simulium serranus, Simulium spinibranchium Escudo castaño amarillento a castaño rojizo ............................. Simulium subpallidum Escudo negro ...................................................................................................................... 6 Alas con pelos en sector basal de R; gonostilo solamente un poco más largo que ancho en la base (Fig. 61 D) ............................................................. Simulium acarayense Alas sin pelos en sector basal de R; gonostilo cerca 2 veces más largo que ancho en la base (Fig. 61 F) ........................................................................... Simulium papaveroi

Pupas 1 – 2(1)

Branquias con 8 filamentos; capullo generalmente no cubriendo las ramas de las branquias y con porción anterior lisa y sin festones (Figs. 59 A, D, G, J; 62 A, C, D, G) ........................................................................................................................................... 2 Branquias con 10-120 filamentos; capullo cubriendo las branquias; porción anterior con festones (Figs. 59 L, O, S) ......................................................................... 8 Capullo cubriendo las ramas (Fig. 59 J); ramas primarias y secundarias emergiendo agrupadas como un ramo (Fig. 59 K) ............................................... Simulium empascae

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Capullo no cubriendo la branquia (Figs. 59 A, D, G; 62 A, C, D, G); ramas primarias y secundarias emergiendo aisladas no como un ramo (Figs. 59 C, F, H; 62 B, F, I) ............................................................................................................................. 3 Filamentos branquiales gruesos basalmente (Fig. 59 H) ........ Simulium spinibranchium Filamentos branquiales no gruesos basalmente .......................................................... 4 Frontoclípeo y porción expuesta del tórax con abundantes tubérculos (Figs. 59 B, E) ................................................................................................................................................ 5 Frontoclípeo y porción expuesta del tórax sin o con escasos tubérculos (Figs. 62 E, H) ..................................................................................................................................... 6 Largo de la branquia aproximadamente como el largo del cuerpo (Fig. 59 D) y ordenadas en un plano (Fig. 59 F); frontoclípeo y tórax con solamente una moderada abundancia de tubérculos (Fig. 59 E) ..............................Simulium serranus Largo de las branquias aproximadamente la mitad del largo del cuerpo (Fig. 59 A) y ordenadas en más de un plano (Fig. 59 C); frontoclípeo y tórax con muy abundantes tubérculos (Fig. 59 B) ...................... Simulium pertina, Simulium laneportoi Ramas de las branquias relativamente agrupadas basalmente, en especial las ramas dorsales (Fig. 62 I); ramas primaria y secundaria dorsal más gruesas que la ventral; tricomas cefálicos y torácicos simples o bífidos (Fig. 62 H) .................... .................................................................................................................. Simulium papaveroi Ramas de la branquia abiertas (Figs. 62 B, F); ramas primaria y secundaria dorsal no más gruesas que la ventral; tricomas cefálicos y torácicos con 3-6 ramas (Fig. 62 E) ..................................................................................................................................... 7 Filamentos de la branquia aproximadamente rectos y dispuestos en un plano (Figs. 62 C, D, F); tubérculos redondeados a veces presentes en frontoclípeo y dorsalmente en tórax (Fig. 62 E) ..................................................... Simulium acarayense Filamentos de la branquia con una pequeña curvatura y dispuestos en conjunto como un cesto (Figs. 62 A, B); sin tubérculos en frontoclípeo y tórax, si existe alguno tiene forma de cono ............................................................ Simulium subpalidum Branquia con alrededor de 120 ramas; tórax totalmente cubierto con tricomas (Fig. 59 U) .................................................................................................. Simulium obesum Branquia con 10-11 ramas; tórax con tricomas de 5-6 ramas sobre cada costado ................................................................................................................................................ 9 Branquia con 11 ramas muy gruesas y con fuerte curvatura (Fig. 59 R) ................. ................................................................................................................ Simulium friedlanderi Branquia con 10 ramas filamentosas y sin fuerte curvatura (Figs. 59 N, Q) ...... 10 Capullo con 1+1 agujeros anteriores; áreas marginales del frontoclípeo elevadas semejando una corona ................................................................... Simulium bifenestratum Capullo sin 1+1 agujeros anteriores (Figs. 59 L-O); área marginal del frontoclípeo no elevada .......................................................................................................................... 11 Frontoclípeo y tórax con regular cantidad de tubérculos uniformemente distribuidos (Fig. 59 M); largo del capullo en la base 3,0 mm, en el dorso 2,8 mm, branquias 1,0 mm ................................................................................ Simulium distinctum Frontoclípeo y tórax con escasa cantidad de tubérculos distribuidos en frontoclípeo cercanos a los tricomas faciales (Fig. 59 P) y en la porción expuesta

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del tórax; largo del capullo en la base 4,2-4,7 mm, en el dorso 4,5-4,7 mm, branquias 1,7-2,0 mm .......................................................................Simulium riograndense Larvas La larva de Simulium obesum desconocida. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(2) – 7(6) – 8(6) – 9(8) –

Hendidura postgenal subovoidal (Fig. 63 Q); anillo anal con 60-81 hileras de ganchos; longitud máxima 5,5-8,0 mm ............................................................................ ......................................... Simulium subpallidum, Simulium acarayense, Simulium papaveroi Hendidura postgenal subtriangular (Fig. 63 F), con forma de domo (Fig. 63 C); anillo anal con 85-204 hileras de ganchos; longitud máxima 6,0-11,8 mm ........... 2 Anillo anal con 145-204 hileras de ganchos; longitud máxima 8,1-11,8 mm ....... 3 Anillo anal con 76-122 hileras de ganchos; longitud máxima 5,6-7,5 mm ............ 6 Anillo anal con 201-204 hileras de ganchos; papilas rectales con 4-6 divertículos sobre cada lóbulo .................................................................................. Simulium empascae Anillo anal con 145-190 hileras de ganchos; papilas rectales con 7-26 divertículos en cada lóbulo ..................................................................................................................... 4 Hendidura postgenal en forma de domo; apotoma cefálico oscurecido totalmente sobre 1/3 basal y continuando adelante como un triángulo llegando cerca del área frontal ................................................................................................ Simulium bifenestratum Hendidura postgenal subtriangular pero con ápice romo; apotoma cefálico oscurecido solo en el 1/3 basal ....................................................................................... 5 Anillo anal con 170-190 hileras de ganchos; longitud máxima 8,5-9,0 mm ............ ................................................................................................................. Simulium distinctum Anillo anal con 145-152 hileras de ganchos; longitud máxima 9,5-10,5 mm ......... ...............................................................................................................Simulium riograndense Apotoma cefálico oscurecido totalmente sobre 1/3 basal y continuando hacia delante como un triángulo, llegando cerca del área frontal (Fig. 63 A) ................. 7 Apotoma cefálico oscurecido solo en la mitad de la base (Fig. 63 D) o en 1/4 basal y continuando suavemente sobre el área medio longitudinal ................................... 8 Hendidura postgenal en forma de domo (Fig. 63 C); anillo anal con 88-100 hileras de ganchos ............................................................................................... Simulium pertinax Hendidura postgenal subtriangular; anillo anal con 76-86 hileras de ganchos ....... ..........................................................................................................Simulium spinibranchium Apotoma cefálico oscurecido en el 1/4 basal y suavemente continuado sobre el área mediana longitudinal; manchas positivas medio longitudinales y submedianas .................................................................................................................. Simulium laneportoi Apotoma cefálico oscurecido solo en la mitad basal; manchas negativas medio longitudinales y submedianas (Fig. 63 D) ..................................................................... 9 Anillo anal con 85-88 hileras de ganchos; papilas rectales con 15-18 divertículos en cada lóbulo ..........................................................................................Simulium serranus Anillo anal con 105-122 hileras de ganchos; papilas rectales con 20-24 divertículos en cada lóbulo ..................................................................................... Simulium friedlanderi

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PERTINAX species group Simulium (Chirostilbia) pertinax KOLLAR (Figs. 57 A-E; 58 A-C; 59 A-C; 63 A-C.) Simulium pertinax KOLLAR, 1832: 117; LUTZ, 1910: 228-230, 266; LUTZ et al., 1918: 107-109; D’ANDRETTA & D’ANDRETTA, 1950: 193-213; SHELLEY et al., 1984: 149. Simulium (Chirostilbia) pertinax COSCARÓN, 1987: 26; 1991: 215-217; GIL ACEVEDO et al., 2004: 433-437. Simulium pertinax PINTO, 1931: 678-680, 722-724, 725, 727, 742, 743, 760; LANE & PORTO, 1939: 169171; LANE & VULCANO, 1943: 432. Trichodagmia pertinax ENDERLEIN, 1936: 128. Simulium infuscatum LUTZ, 1910: 26-29. Simulium venustum (nec SAY, 1922): LUTZ, 1909: 132, 136-138. Chirostilbia flavifemur ENDERLEIN, 1921b: 79. Trichodagmia lutziana ENDERLEIN, 1934b: 291; 1935: 195. Simulium distinctum (nec LUTZ, 1910): PINTO, 1931: part: 670, 669-701, part: 724. Simulium inexorabile SCHROTTKY, 1909: 63; WYGODZINSKY, 1953a: 302. Simulium septentrionale CERQUEIRA & BARBOSA DE ALMEIDA, 1970: 1-11. Simulium cerqueirai BARBOSA DE ALMEIDA, 1979: 789. Chirostilbia pertinax PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2002: 133-135; 2004: 126-127.

Female: Wing length 2.2-2.8 mm; general coloration blackish with soft gray pollinosity and gray to golden pile; with some frontal illumination, black with median wide stripe; changing light to latero-posterior position shows grayish with 1+1 black submedian stripes nearly reaching blackish posterior border, similar as shown for laneportoi but wider and without contrast with gray area; in specimens preserved for long time, coloration is dark grayish brown homogeneous and without evident vittae; scutellum dark brown gray; metanotum velvety black with gray pollinosity; pleurae brown to violaceous with silvery pruinosity; legs brown blackish on third distal area of tibia and basitarsus of hind leg and tarsomeres of all legs; abdomen black with shiny tergal plates V-IX; cibarium with median area light and smooth (Fig. 57 A); fronto-ocular triangle deep (Fig. 57 B); basal sector of R with hair arranged in 2-3 rows, and Sc with 18 setae; claw with well developed subbasal tooth; eighth sternite well sclerotized, with about 20 setae on each side (Fig. 57 C) Gonapophysis subtriangular, divergent with rounded apex and slightly sclerotized internally; paraproct about 2.5 times longer than wide at base and without basal lobe (Fig. 57 D); anterior branch of genital fork with strong curvature (Fig. 57 E). Male: Wing length 2.2-2.5 mm; scutum black with golden pile and gray pollinosity showing with some light, a wide and lighter lateral stripe joined anteriorly with 1+1 fronto-lateral spots and posteriorly with gray prescutellar area, similar as shown for Simulium serranus (Fig. 58 E); frontolateral spots with antero-lateral illumination are continuous anteriorly as in Simulium laneportoi (Fig. 58 D); scutellum and metanotum black; abdomen black with 1+1 silvery pruinose spots on tergites II, V-VII and in some specimens on IX; gonostylus about 3/4 of gonocoxite length with curved apex (Figs. 58 A, B); ventral plate about as long as wide at base apically slightly acuminate and curved dorsally (Figs. 58 A, C); median sclerite with slight median cleft and endoparameres with strong hooks (Fig. 58 A).

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H

M O

G

F

J A

B

E

I

C N

Q

D

R

T S U

L

V

P

K Figure 57. Female. A-E: S. (Chirostilbia) pertinax; A: cibarium, B: fronto-ocular triangle; C: genitalia ventral view showing eighth sternite, gonapophysis, cercus, paraproct, genital fork and spermatheca; D: cercus and paraproct mounted on slide; E: genital fork. F-G: S. (C.) laneportoi; F: scutum with frontal illumination; G: scutum with posterior illumination. H-I: S. (C.) serranus; H: frons; I: cercus and paraproct. J-L: S. (C.) spinibranchium; J: scutum with frontal illumination; K: eighth sternite and gonapophysis; L: cercus and paraproct. M-N: S. (C.) distinctum; M: claw; N: cercus and paraproct. O-V: S. (C.) obesum; O: scutum with frontal illumination, P: sensory vesicle of maxillary palp and palpomere; Q-S: anterior, median and hind leg, respectively; T: eighth sternite and gonapophysis; U: cercus and paraproct; V: genital fork.

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Pupa: Cocoon slipper shaped, weave resistant with visible threads and reinforced anterior border (Fig. 59 A), length at base 4.0-4.5 mm, at dorsum 3.0-3.5 mm, gills 1.5-2.0 mm; frontoclypeus and exposed portion of thorax with numerous tubercles (Fig. 59 B), cephalic trichomes with 1-3 branches, thorax with 4-6 branches; gill with 8 filamentous branches, with short primary and secondary branches (Fig. 59 C). Larva: Maximum length 6.5-7.0 mm; body without hair or spines; cephalic apotome with dark subtriangular basal spot reaching half of length (Fig. 63 A); ratio of proximal, medial and distal antennal segments = 1:1.2:1.4 (Fig. 63 B); cephalic fan with about 40 rays; postgenal cleft dome shaped (Fig. 63 C); postgenal bridge a little longer than half of hypostomium length; anal ring with about 88-100 rows of 15-21 hooks each; rectal papillae with 15 diverticula on central lobe and 12 on each lateral lobe. Distribution: Argentina: Corrientes, Entre Rios, Misiones; Brazil: Bahia, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Rondônia, Santa Catarina, São Paulo; Paraguay. Bionomics: Simulium pertinax breeds in creeks with fast current and clear water. Larvae attach to aquatic plants, trees, submerged branches and leaves. Females are very anthropophilic and the bite is uncomfortable. This species is the most dangerous in southern Brazil and northeastern Argentina and Paraguay. Discussion: Simulium pertinax is a very peculiar species and easy to identify. The closest species is S. laneportoi with a very similar pupa, having gill branches only a little thinner. The female of S. laneportoi has a scutum with narrower black submedian stripes, and when seen with posterior light, the 1+1 grayish submedian vittae are more evident than in S. pertinax. Simulium (Chirostilbia) laneportoi VARGAS (Figs. 57 F, G; 58 D.) Simulium laneportoi VARGAS, 1941a: 118 (subst. name). Simulium pilosum LANE & PORTO, 1940: 190 (preocup). Eusimulium laneportoi LANE & VULCANO, 1943: 437. Simulium (Chirostilbia) laneportoi PY-DANIEL, 1987: 336-341; COSCARÓN, 1987: 26, 1991: 217-219. Simulium (Chirostilbia) dekeyseri SHELLEY & PY-DANIEL, 1981: 23-32. Chirostilbia laneportoi PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

Female: Wing length 2.2-2.9 mm; general coloration black; scutum black with gray pollinosity, pile relatively scarce with golden tones; with frontal light, median and 1+1 sublateral grayish areas appear on black scutum (Fig. 57 F); with posterior illumination, coloration reverses (Fig. 57 G). Metanotum and abdomen black; abdominal tergite II with 1+1 lateral silvery spots and tergites V-VIII shiny; basal sector of R with hair and Sc with 10-20 setae; claw with large subbasal tooth; eighth sternite with 15-17 setae; gonapophysis and paraprocts as in Simulium spinibranchium; genital fork as in S. pertinax. Male: Wing length 2.1-2.7 mm; scutum velvety black with anterior lateral and posterior border gray pollinose similar to Simulium serranus; with frontal light, anterior border gray, wide and continuous (Fig. 58 D); gonocoxite, gonostylus and ventral plate as in S. spinibranchium. Pupa: Cocoon shape, numerous tubercles on frontoclypeus and thorax, and gill with 8 branches practically indistinguishable from those of Simulium pertinax, with only small

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E D F

N H

B C

I G J

K

O

L M Figure 58. Male. A-C: S. (Chirostilbia) pertinax; A: genitalia ventral view showing gonocoxite, gonostylus, ventral plate, endoparameres and median sclerite; B: gonocoxite and gonostylus in ventral view; C: ventral plate. D: S. (C.) laneportoi, scutum with frontal illumination. E-F: S. (C.) serranus; E: scutum with lateral view; F: gonocoxite and gonostylus. G-H: S. (C.) spinibranchium; G: gonocoxite and gonostylus, dorsal view; H: ventral plate. I: S. (C.) empascae, gonocoxite and gonostylus. J-K: S. (C.) distinctum; J: gonocoxite and gonostylus, K: ventral plate. L-N: S. (C.) riograndense; L: gonocoxite and gonostylus; M: ventral plate; N: endoparamere and aedeagal membrane. O-P: S. (C.) obesum; O: gonocoxite and gonostylus, ventral view; P: ventral plate.

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thin gill branches; size slightly smaller with cocoon length on base 2.4-3.9 mm, on dorsum 2.8-3.8 mm and gills 1.5-2.4 mm. Larva: Maximum length 5.6-7.5 mm; cephalic apotome darkened on mid longitudinal and 1/4 basal area, with discontinuous positive spots on median line and 1+1 on submedian and subbasal areas; postgenal cleft subtriangular with blunt apex; postgenal bridge a little shorter than hypostomium length; ratio of proximal, medial and distal antennal segments = 1:1-1.3:1.3-1.7; cephalic fan with 38-45 rays; anal ring with 90-115 rows with 15-18 hooks each; rectal papillae with 19-30 diverticula on each lobe. Distribution: Brazil: Goiás, Mato Grosso, Minas Gerais, Rio de Janeiro, São Paulo. Bionomics: Simulium laneportoi breeds in fast-flowing creeks, where the larvae and pupae attach themselves to stones and aquatic vegetation. Females bite humans. Discussion: Simulium laneportoi’s similarity to S. pertinax was explained in the description. Simulium spinibranchium is very similar and the adults can be differentiated only by the more abundant pilosity on the scutum. The pupa with its shoe-shaped cocoon and thicker gill branches allow it to be separated easily. Simulium serranus is also very similar and can be separated only by its small size and lower number of head and thoracic tubercles. Comparative cytological or molecular studies would be convenient to assure the identity of these species. Simulium (Chirostilbia) serranus COSCARÓN (Figs. 57 H, I; 58 E, F; 59 D-F; 63 D-G.) Simulium serranus COSCARÓN, 1981a: 158-160. Simulium (Chirostilbia) serranus COSCARÓN, 1987: 26; 1991: 219-220. Chirostilbia serranus PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

Female: Wing length 2.0-2.2 mm; scutum black with gray pollinosity, with ornamentation as in Simulium laneportoi; legs darkened on tibial apex and tarsi of anterior leg, distally on tarsi, medially and apically on tibia and terminal portion of tarsi of hind leg; scutellum light brown; metanotum blackish; pleurae and abdomen brown grayish; frons with parallel sides (Fig. 57 H); Sc without hair; eighth sternite with about 30 setae, gonapophysis and genital fork as in Simulium pertinax; paraproct subtriangular, two times longer than wide at base (Fig. 57 I). Male: Wing length 2.0-2.1 mm; scutum black bordered by grayish pollinose 1+1 anterior spots joining lateral stripe to posterior gray area (Fig. 58 E); changing light position shows continuous grayish anterior band as in Simulium laneportoi; scutellum brown; metanotum blackish; length/width ratio of hind basitarsus = 5.4; gonostylus subtrapezoidal, distally truncate with acute apex (Fig. 58 F); ventral plate as in Simulium pertinax. Pupa: Cocoon slipper shaped, weakly projected basally (Fig. 59 D); weave compact, sometimes with threads surpassing anterior border and covering basal portion of gills; length at base 2.9-3.2 mm, at dorsum 2.8-3.0 mm, gills 2.0-2.5 mm; frontoclypeus and exposed portion of thorax with tubercles relatively abundant (Fig. 59 E) but more scarce than in Simulium pertinax; cephalic trichomes with 1-3 branches and thoracic with 3-8 branches; gill with 8 branches, with primary ventral branch longer than primary dorsal; secondary median branch two times longer than secondary dorsal branch (Fig. 59 F). Larva: Maximum length 6.0 mm; cephalic apotome scarcely pigmented (Fig. 63 D); antenna as long as cephalic fan stem; ratio of proximal, medial and distal antennal segments = 1:1-1.3:1.6-2; cephalic fan with 45-50 rays. Mandible with second marginal

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Figure 59. Pupa. A-C: S. (Chirostilbia) pertinax; A: lateral view; B: frontoclypeus; C: portion of thorax and gill. D-F: S. (C.) serranus; D: lateral view; E: frontoclypeus; F: portion of thorax and gill. G-I: S. (C.) spinibranchium; G: lateral view; H: gill; I: thoracic trichome. J-K: S. (C.) empascae; J: lateral view; K: gill. L-N: S. (C.) distinctum; L: lateral view; M: frontoclypeus; N: portion of thorax and gill. O-P: S. (C.) riograndense; O: lateral view; P: frontoclypeus, Q: gill. R: S. (C.) friedlanderi, gill, lateral view (extracted from original description). S-U: S. (C.) obesum; S: lateral view; T: frontoclypeus; U: portion of thorax with trichomes and gill.

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tooth very short (Fig. 63 E); postgenal cleft subtriangular with acute apex; postgenal bridge a little longer than half hypostomium length (Fig. 63 F); anal sclerite with single hairs (Fig. 63 G); anal ring with 85-88 rows with 13-15 hooks each; lateral sclerite of proleg with about 30 teeth; rectal papillae with about 15-18 diverticula on each lobe. Distribution: Brazil: Mato Grosso, Minas Gerais. Bionomics: Simulium serranus larvae live in small creeks with moderate current and crystalline water, flowing over aquatic vegetation. The females are haematophagous and very aggressive in Chapada dos Guimarães (MT), Brazil. Discussion: Simulium serranus is very close to S. laneportoi. The differences were explained in the description of that species. Simulium spinibranchium is similar but has more abundant pilosity on the scutum and the pupal gill is thicker and has 8 or 9 branches. The pupal cocoon is shoe shaped. Simulium pertinax can be distinguished by the absence of submedian silvery vittae on the scutum. Simulium (Chirostilbia) spinibranchium LUTZ (Figs. 57 J-L; 58 G, H; 59 G-I; 63 H.) Simulium spinibranchium LUTZ, 1910: 248, 261, 263-264; PINTO, 1931: 712-713, 726-727, 756-762; PYDANIEL & SHELLEY, 1980: 213-223. Simulium (Chirostilbia) spinibranchium COSCARÓN, 1987: 26; 1991: 220-223; SHELLEY et al., 2000: 175-176. Chirostillbia spinibranchia PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 128.

Female: Wing length 2.7-3.0 mm; scutum blackish gray pollinose with abundant gray pile; with frontal light, 1+1 submedian gray pollinose stripes wider anteriorly and posteriorly, last portion joining with grayish, prescutellar area (Fig. 57 J); with another light position this submedian vittae changes to black with median and sublateral gray lead coloration; scutellum brown grayish, metanotum velvety brown grayish; legs brown yellowish with darker apical portion of segments blackish; abdomen brown blackish with tergite I and 1+1 lateral spots on tergite II light grayish brown; tergites V-VIII shiny; fronto-ocular triangle absent or very reduced; eighth sternite well sclerotized with about 30 setae on each side, gonapophysis with internal border well sclerotized (Fig. 57 K); paraproct about two times longer than wide at base (Fig. 57 L); genital fork with anterior branch well sclerotized. Male: Wing length 2.4-2.7 mm; scutum black with pilosity gray, and bordered anteriorly, laterally and posteriorly by silvery pruinose stripe; with frontal light, grayish anterior border very wide as in Simulium laneportoi; scutellum and metanotum as in female; abdomen black with 1+1 grayish pollinose spots on tergites II, VI-VII, decreasing in size from anterior to posterior segments; gonostylus apically acuminate (Fig. 58 G); ventral plate body wider than high (Fig. 58 H). Pupa: Cocoon shoe-shaped, with reinforced anterior border and protecting gill base (Fig. 59 G); weave thick, with visible threads; frontoclypeus and exposed portion of thorax with numerous tubercles, similar to Simulium pertinax; cephalic and thoracic trichomes with 6-8 branches (Fig. 59 I); gills each with 8-9 branches emerging from 4 short primary branches from short basal trunk; branches thick at base and slightly thinning to blunt apex (Fig. 59 H). Microstructure of gill branches with thin porosity. Larva: Maximum length 6.0-6.7 mm; cephalic apotome darkened basally as in Simulium pertinax, antennae a little shorter than cephalic fan stem; ratio of proximal,

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medial and basal segments = 1:1-1.1:1.7-1.9. Mandible with latero-mandibular process; postgenal cleft subtriangular with blunt apex; postgenal bridge about 2/3 of hypostomium height; cephalic fan with 30-37 rays; lateral sclerite of proleg with 20 teeth arranged in 9-10 groups (Fig. 63 H); anal sclerite with hair and some scales like spines; anal ring with 76-86 rows with 12-14 hooks each; rectal papillae with 11-13 diverticula on each lobe; ninth segment with 1+1 very small ventral tubercles; body cuticle without hair. Distribution: Brazil: Bahia, Goiás, Mato Grosso, Minas Gerais, Rio de Janeiro, Rio Grande do Sul, São Paulo; Venezuela. Bionomics: Simulium spinibranchium larvae live in small creeks with slight discharge, deep, clear water and regular current velocity, from 400 to 1200 m in altitude. The feeding habits of the females are unknown. Discussion: The closest species to S. spinibranchium is S. laneportoi, whose adults are practically inseparable when the scutum pilosity is absent; the peculiar cocoon and gill shape permit separation from similar species. Simulium (Chirostilbia) empascae PY-DANIEL & MOREIRA (Figs. 58 I ; 59 J, K; 63 I-J.) Simulium (Chirostilbia) empascae PY-DANIEL & MOREIRA, 1988: 77-86; COSCARÓN, 1991: 222-223. Chrostillbia empascae PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 126.

(Based on original description). Female: (Pharate) General coloration black to dark brown; Sc and basal sector of R with hair; genitalia similar to those of Simulium spinibranchium and S. obesum. Male: (Pharate) Gonostylus wide basally, blunt apically, with very small subapical spine (Fig. 58 I); ventral plate as in Simulium distinctum. Pupa: Cocoon shoe shaped, protecting gills (Fig. 59 J); frontoclypeus without tubercles; exposed portion of thorax with short and acute tubercles on posterior area; cephalic trichomes single and short, thoracic single but more enlarged; gills with 8 branches emerging from short basal trunk and with primary and secondary branches also very short (Fig. 59 K). Larva: Maximum length 8.1-11.8 mm; cephalic apotome similar to that of Simulium pertinax; antenna relatively thick (Fig. 63 I); ratio of proximal, medial and distal segments = 1:1.3-1.6:1-1.3; cephalic fan with 53-54 rays. Mandible with latero-mandibular process (Fig. 63 J); postgenal cleft subtriangular with acute apex; postgenal bridge about half of hypostomium length; lateral sclerite of proleg with about 28 teeth; anal ring with 201-204 rows with 19-26 hooks each; rectal papillae with 4-6 diverticula on each lobe. Distribution: Brazil: Santa Catarina, Rio Grande do Sul. Bionomics: Simulium empascae breeds in fast-flowing creeks in areas with direct sunlight. Feeding habits of the females are unknown. Discussion: The peculiar cocoon shape covering all the gills and the gill branching allow the differentiation of this species from similar species in the subgenus. The gonostylus with a blunt apex and with a spine and the abundant rows of hooks on the anal ring also help to separate this species from others.

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Simulium (Chirostilbia) distinctum LUTZ (Figs. 57 M, N; 58 J, K; 59 L-N; 63 K, L.) Simulium distinctum LUTZ, 1910: 241-243; PINTO, 1931: 699-701; D’ANDRETTA & D’ANDRETTA, 1950: 197-198; EDWARDS, 1933: 210; D’ANDRETTA, 1954: 63-66; SHELLEY et al., 1984: 145. Simulium (Simulium) distinctum ORFILA, 1939: 1533. Simulium (Chirostilbia) distinctum COSCARÓN, 1987: 26; 1991: 223-225; PY-DANIEL et al., 1988: 43-48. ?Trichodagmia distincta ENDERLEIN, 1934a: 194; 1936: 128. Trichodagmia spadicidorsum ENDERLEIN, 1934a: 194. Simulium pertinax LANE & PORTO, 1929: 170; VARGAS, 1945a: 182. Simulium prumirimense COSCARÓN, 1981a: 160-164. Chirostilbia distincta PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 133-135; 2004: 125-126.

Female: Wing length 2.5-3.4 mm; general coloration dark grayish brown; scutum brown grayish with decumbent golden to silvery pilosity, with posterior border darker pollinose, giving dark tones, one median and 1+1 sublateral stripes gray pollinose not reaching anterior and posterior borders; scutellum brown; metanotum reddish brown with grayish pollinosity; pleura dark grayish brown with gray pollinosity; legs grayish brown with tarsi, distal portion of femur, tibia and basitarsus of hind leg blackish; abdomen dark brown grayish with tergite I, center of II-IV brown with dark pile; VI-IX tergites shiny; tergite II with 1+1 silvery pruinose lateral spots; ventrally light brown grayish; frontoocular triangle higher than wide; basal sector of R hairy and Sc with 10 setae; claw with subbasal tooth (Fig. 57 M); eighth sternite with about 60 setae; gonapophysis with internal margin more sclerotized and without microtrichia; paraprocts acuminate distally (Fig. 57 N) and genital fork with anterior branch strongly sclerotized. Male: Wing length 2.4-3.3 mm; scutum blackish with yellow greenish pile; abdomen blackish with 1+1 silvery spots on tergites II, V-VI and smaller spots on VII-VIII and IX; length/width ratio of hind basitarsus = 6; gonostylus subquadrate, with smooth external distal process (Fig. 58 J); ventral plate elongated with blunt apex (Fig. 58 K). Pupa: Cocoon shoe shaped with thick weave dark brown leaving holes anteriorly and protecting gill branches laterally (Fig. 59 L); length at base 3.0 mm; at dorsum 2.8, gills 1.0 mm; frontoclypeus and thorax with moderate quantity of tubercles (Fig. 59 M); cephalic trichomes with 2-3 branches, and thoracic with 3-6 branches; gills each with 10 short branches, with 3 short primary branches emerging from short basal trunk; gill filaments relatively thick, curved and sclerotized distally, terminal portion frequently lost (Fig. 59 N). Larva: Maximum length 8.5-9.0 mm; general aspect as in figure 63 K; cephalic apotome brown with median basal area darkened but not like triangle as in Simulium pertinax; slightly darkened antero-lateral and posterior spots; antennae as long as or surpassing cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.51.6:1.2-1.3; cephalic fan with 45-50 rays. Mandibles with second marginal tooth nearly as long as first (Fig. 63 L); postgenal bridge longer than half of hypostomial length; lateral sclerite of proleg with 32-38 teeth arranged in 10-13 groups; anal ring with 170-190 rows of 18-20 hooks each; body integument with scarce, short hair; rectal papillae with 7-9 diverticula on each lobe. Distribution: Brazil: Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, São Paulo.

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Bionomics: Simulium distinctum breeds in creeks of fast-flowing clear water, attached to stones. Feeding behavior of the females is unknown. Discussion: The closest species to S. distinctum is S. riograndense, but it can be separated by the light brown yellowish scutum, and very sparse pupal tubercles reduced on the frontoclypeus to 1+1 groups on the base at the sides of the facial trichomes and on the posterior area of the thorax. Simulium (Chirostilbia) riograndense PY-DANIEL, SOUZA & CALLAS (Figs. 58 L, M; 59 O-Q.) Simulium (Chirostilbia) riograndense PY-DANIEL et al., 1998: 38-43; CROSSKEY & HOWARD, 2004: 20. Chirostilbia riograndense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2002: 133-135; 2004: 127.

Female: Wing length 3.2-3.6 mm; scutum light brown-yellowish with light grayish to yellowish pile; with frontal light, 1+1 sublateral grayish pruinose stripes, becoming a little darker with posterior light and darker than prescutellar area; scutellum brown and metanotum and pleurae dark brown with gray pruinosity; legs yellowish with tibiae and tarsi of anterior leg, apices of basitarsi and tarsi of middle and posterior legs and apex of hind tibiae blackish; abdomen brown with 1+1 grayish spots on tergite II; eighth sternite with about 26-38 setae on each side, gonapophysis, paraproct and genital fork as in Simulium distinctum. Male: Wing length 3.3-3.5 mm; scutum brown-reddish to yellowish, more darkened than in female, with silvery to yellowish pile; with postero-lateral light, silvery pollinose band bordering posterior and sides; scutellum and metanotum as in female; abdomen blackish with 1+1 silvery pollinose spots on tergites II-VIII, more developed on tergite II; genitalia very similar to those of Simulium distinctum but gonostylus more sclerotized with well-developed process on external distal side, and microscopic subapical spur (Fig. 58 L); ventral plate not elongated and more rounded apically (Fig. 58 M). Pupa: Cocoon shoe shaped, weave resistant with abundant holes and protecting branches laterally (Fig. 59 O); length at base 4.2-4.7 mm, at dorsum 4.6-4.7 mm; gills 1.72.0 mm; frontoclypeus and exposed portion of thorax with scarce tubercles reduced to 1+1 small areas at frontoclypeal base (Fig. 59 P) and to posterior dorsal area of thorax, those mostly acuminate; cephalic trichomes short, single or bifid; thoracic trichomes with 1-4 branches; gills each with 10 branches emerging from short basal trunk and very short primary branches (Fig. 59 Q). Larva: Maximum length 9.5-10.5 mm; cephalic apotome with median basal area brownish; spots positive, median longitudinal and 1+1 submedian subbasally positioned; antennae scarcely surpassing cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.8-2.1:0.9-1.3; cephalic fan with 43-50 rays; mandible very similar to that of Simulium distinctum; postgenal cleft subtriangular, blunt apically; postgenal bridge more than half hypostomium length; anal ring with 145-152 rows with 19-25 hooks each; rectal papillae with 9-11 diverticula on each lobe. Distribution: Brazil: Paraná, Rio Grande do Sul. Bionomics: Simulium riograndense breeds in slow, acidic water in temperatures of 1920 °C, attached to stones. Feeding behavior of females is unknown. Discussion: The closest species to S. riograndense is S. distinctum in whose species description the differences are mentioned.

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Simulium (Chirostilbia) bifenestratum HAMADA & PEPINELLI Simulium (Chirostilbia) bifenestratum HAMADA & PEPINELLI, 2004: 45-52.

(Based on original description). Female: Wing length 3.3 mm; scutum dark gray with 1+1 submedian small subrectangular silver spots, pilosity pale golden; scutellum light brown, metanotum brown; pleurae dark brown; legs light brown with darkened black portion; abdomen dark brown with membranous areas light brown; tergite II with 1+1 silver pruinose spots, VI-IX shiny; cibarium smooth with well sclerotized external cornuae branches; basal sector of R and Sc with hair; claw with subbasal tooth; genitalia very similar to those of Simulium distinctum. Male: Wing length 3.1 mm; scutum black with dense golden pile; scutellum dark brown, metanotum black; legs coloration as in female; genitalia as in Simulium distinctum. Pupa: Cocoon slipper shaped, with strong and thick weave, showing 1+1 anterior sublateral subovoidal holes, and protecting most gill branches; length at base 1.8-2.2 mm; at dorsum 3.9-4.5 mm; frontoclypeus with rounded tubercles on thoracic middle area larger, with protuberances and sometimes acuminate, frontoclypeus with marginal area elevated, in special cases lateral border with a crown; cephalic trichomes with 1-3 branches, thoracic with 3-9 branches; gill with 10 branches, thicker at base and surrounding head. Larva: Maximum length 8.4-9.0 mm; cephalic apotome darkened on mid to posterior area, and postgenal cleft shaped as in Simulium pertinax; antenna as long as cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.8-1.9:0.8-.9; cephalic fan with 45-48 rays; anal ring with 152-158 rows with 15-19 hooks each; rectal papillae with 20-26 elongated diverticula in total; body cuticle with single-branched hairs. Distribution: Brazil: São Paulo. Bionomics: Simulium bifenestratum breeds in creeks with rocky bottoms, about 10 m wide, at an altitude of 1500 m. Feeding habits of the females are unknown. Discussion: Similar to species of the PERITNAX group, in particular to S. distinctum, S. bifenestratum can be separated by the presence on the female scutum of small anterior submedian silvery spots and 1+1 anterior sublateral apertures in the thick cocoon. Simulium (Chirostilbia) friedlanderi PY-DANIEL (Fig. 59 R.) Simulium (Chirostilbia) friedlanderi PY-DANIEL, 1987: 331-336; COSCARÓN, 1991: 227-228.

(Based on original description). Female: Unknown. Male: Scutum dark brown with yellowish pile; lateral illumination shows whitish pollinosity bordering scutum, frontal light shows 1+1 lateral and antero-dorsal pruinose spots; genitalia similar to those of Simulium obesum. Pupa: Cocoon shoe shaped very similar to Simulium riograndense and S. obesum, and very elevated anteriorly; weave with abundant holes and protecting all gill filaments laterally; length at base 3.1-3.9 mm; at dorsum 3.4-5.0 mm, gills 1.2 mm; frontoclypeus and exposed portion of thorax with numerous small tubercles; cephalic and thoracic trichomes with 3-7 branches; gill with 11 very thick filaments, thinning to acute apices with strong curvature (Fig. 59 R).

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Larva: Maximum length 6.2-6.4 mm; cephalic apotome darkened on mid basal area and with median longitudinal and median sublateral positive spots; antennae surpassing cephalic fan stem; ratio of proximal, medial and distal segments length = 1:1-1.2:1.3-1.5; cephalic fan with 43-47 rays; postgenal cleft subtriangular; with blunt apex; postgenal bridge a little longer than half hypostomium length. Mandible with second marginal tooth surpassing half of anterior length; latero-mandibular process present; lateral sclerite of proleg with about 23 teeth in 7-8 groups; anal ring with 105-122 rows of 16-20 hooks each; rectal papillae with 20-24 diverticula on each lobe. Distribution: Brazil: Mato Grosso. Bionomics: Larval stages of S. friedlanderi live in flows with rocky bottoms. Feeding behavior of the females is unknown. Discussion: The known stages are very close to those of S. distinctum, S. riograndense and S. obesum, but the peculiar thick gill of S. friedlanderi permits separation. Simulium (Chirostilbia) obesum VULCANO (Figs. 57 O-V; 58 O, P; 59 S-U.) Simulium obesum VULCANO, 1959: 351-360; LUTZ, 1922: 89-92 (nomen nudum); PINTO, 1931: 753, 762 (nomen nudum). Simulium (Chirostilbia) obesum COSCARÓN, 1987: 26; 1991: 228-229. Chirostilbia obesa PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

Female: Wing length 3.0-3.6 mm; scutum dark brown reddish to blackish with soft gray pollinosity and golden decumbent pilosity, lateral and posterior edges with silvery pollinose band; frontal illumination shows 1+1 silvery submedian and sublateral vittae separated by one median and 1+1 sublateral black stripes that becomes wider with lateral light (Fig. 57 O); scutellum dark brown reddish; metanotum velvety black and pleural areas blackish with gray pollinosity; legs light brown with darkened areas dark brown to black, as in figures 57 Q-S; abdomen blackish with tergal posterior border dark gray; tergite II with 1+1 lateral silvery spots; tergites I-V velvety; VI-IX shiny. Maxillary palpal sensory organ small, shorter than half of palpomere length (Fig. 57 P); claw with subbasal tooth well developed; Sc with about 30 setae; eighth sternite with about 40 setae on each side; gonapophysis with rounded apex and without microtrichia (Fig. 57 T); paraprocts two times longer than width at base (Fig. 57 U); genital fork with strong anterior branch (Fig. 57 V); spermatheca subovoidal without sclerotized neck and light circular areas between spiculae. Male: Wing length 3.2-3.5 mm; general coloration as in female; scutum velvety black, bordered laterally and posteriorly by silvery band; frontal light shows 1+1 anterior sublateral silver spots; legs as in female but more darkened especially on anterior legs; abdomen velvety black, tergites II and VI with 1+1 silvery spots, bigger on II; gonostylus about half gonocoxite length (Fig. 58 O); ventral plate subtriangular, a little wider than high (Fig. 58 P). Pupa: Cocoon with threads evident, leaving large apertures, anteriorly elevated, projected anteriorly and protecting all gill filaments (Fig. 59 S); length at base 3.1-4.0 mm; at dorsum 4.0 mm; gills 1.3 mm; frontoclypeus and exposed portion of thorax with abundant tubercles (Fig. 59 T); cephalic trichomes with 2-5 branches; thorax dorsally

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covered with abundant trichomes with 1-3 branches; gills filiform, arborescent, each with about 120 terminal branches; primary branches about 7 in number, relatively short, emerging form base without evident basal trunk (Fig. 59 U); abdomen with two strong terminal spines. Larva: Undescribed. Distribution: Brazil: Rio de Janeiro, Santa Catarina, São Paulo. Bionomics: Simulium obesum breeds in torrential creeks with crystalline, relatively cold water. The female feeding habits are unknown. Discussion: Coloration of the adults of S. obesum is similar to that of S. distinctum, but is lighter. The peculiar pupa with its many gill branches and abundant trichomes on the thorax allows separation of S. obesum from other species. SUBPALLIDUM species group Simulium (Chirostilbia) subpallidum LUTZ (Figs. 60 A-C; 61 H-K; 62 A, B; 63 M-P.) Simulium subpallidum LUTZ, 1910: 247-248; PINTO, 1931: 714, 742, 755; COSCARÓN, 1982: 66, 67, 73-76. Simulium (Chirostilbia) subpallidum COSCARÓN, 1987: 26; 1991: 230-232; SHELLEY et al., 2000: 176-177. Simulium subviride LUTZ & MACHADO, 1915: 46 (nomem nudum). Simulium guarani COSCARÓN & WYGODZINSKY, 1972a: 210-216. Chirostilbia subpallida PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 128.

Female: Wing length 2.0-2.8 mm; scutum and scutellum light brown yellowish with gray yellowish pile, some specimens with 1+1 small dark gray postero-lateral spots; posterior illumination shows submedian light gray stripes joined posteriorly with lyre pattern; pleurae gray yellowish with basal area silvery gray. Metanotum velvety brown blackish; legs yellowish with dark portion brown, nearly black on anterior tarsi; abdomen with tergites I-II yellowish, III-V yellow grayish with tergal plate brown dark gray not reaching posterior border and in III-IV another 1+1 lateral dark spot; VI-IX with cereous sheen on tergal plates and yellowish laterally; frons slightly narrowed (Fig. 60 A); frontoocular triangle as high as wide (Fig. 60 B); basal portion of cibarium smooth with cornuae well sclerotized; basal portion of R with hair in 1-2 rows and Sc generally without hair (Fig. 60 C); claws with or without small tooth; eighth sternite, gonapophyses and genital fork as in Simulium acarayense; spermatheca with abundant microtrichia arranged in rows; paraproct about two times longer than wide at base and with small basal lobe. Male: Wing length 1.8-2.5 mm; scutum and scutellum brown yellowish to reddish, with 1+1 blackish spots postero-laterally; laterally and posteriorly bordered by silvery gray band; with changing light position, lateral band can increase in size, covering one quarter of antero-lateral area; scutum pilosity golden; pleurae gray, silvery and shiny basally; legs brown yellowish with dark areas larger than in female; abdomen black with 1+1 silvery spots on tergites II, VI-VIII; gonostylus as long as gonocoxite, longer than wide, slightly curved and with lateral crest (Figs. 61 H, I); ventral plate wider than high, with thin hair (Figs. 61 J, K). Median sclerite and endoparameres as in Simulium papaveroi. Pupa: Cocoon slipper shaped, with straight anterior border and slightly projected anteriorly (Fig. 62 A); compact weave with evident threads; length at base 3.6-4.0 mm, at

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Figure 60. Female. A-C: S. (Chirostilbia) subpallidum; A: frons; B: fronto-ocular triangle; C: veins of anterior portion of wing. D-K: S. (C.) acarayense; D: basal portion of cibarium; E: abdomen; F: eighth sternite and gonapophysis; G: genital fork; H: cercus and paraproct; I-K: fore, mid and hind leg of male, respectively. L-P: S. (C.) papaveroi; L-M: scutum with lateral and frontal illumination; N: sensory vesicle of maxillary palp and palpomere; O: claw; P: cercus and paraproct.

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dorsum 2.6-3.5 mm, gills 1.5-2.3 mm; frontoclypeus and thorax without tubercles or these restricted to basal area on frontoclypeus, area of gill and postero-dorsally on thorax with conical shape; cephalic trichomes large with 3-5 branches and thoracic trichomes with 46 branches; gills with 8 branches emerging from short basal trunk and with short primary branches (Fig. 62 B). Larva: Maximum length 5.0 mm; color light green grayish; cephalic apotome light brown with positive spots (Fig. 63 M); antennae as long as or shorter than stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.5 (Fig. 63 N). Mandible with second marginal tooth half length of first tooth (Fig. 63 O); cephalic fan with 32-39 rays; postgenal cleft subovoidal; postgenal bridge half of hypostomium length; anal sclerite with single hairs, and scales with 3-5 branches (Fig. 63 P); anal ring with 70-80 rows with 12-14 hooks each; body cuticle without hairs; segment nine with 2 small ventral tubercles; rectal papillae with 4-7 diverticula on each lobe. Distribution: Argentina: Misiones; Brazil: Bahia, Espírito Santo, Goiás, Minas Gerais, Pará, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Simulium subpallidum breeds in small creeks, with moderate current, attached to aquatic vegetation. The feeding habits of the females are unknown. Discussion: The closest species to S. subpallidum is S. acarayense. Females can be separated by the gray-lead scutum, and males by the blackish scutum and shorter gonostylus. Simulium papaveroi also is similar but can be separated in the female by the dark grayish brown scutum, and more elongated paraproct; in the male by the blackish scutum; in the pupa by the more grouped branches and cephalic trichomes with 1-2 branches. Karyological or molecular studies are needed to assure the species identity, particularly of S. subpallidum and S. acarayense, which are very close. Simulium (Chirostilbia) acarayense COSCARÓN & WYGODZINSKY (Figs. 60 D-K; 61 A-E; 62 C-F.) Simulium acarayense COSCARÓN & WYGODZINSKY, 1972a: 216-223; COSCARÓN, 1982: 67-68. Simulium (Chirostilbia) acarayense COSCARÓN, 1987: 26; 1991: 232-233. Chirostilbia acarayense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 125.

Female: Wing length 2.0-2.2 mm; scutum gray-lead with dark background and gray pollinosity, pile yellowish; scutellum light brown with gray pollinosity; metanotum velvety dark brown; pleurae brown grayish; legs light yellow with dark areas blackish (Figs. 60 I-K); abdomen gray greenish with tergal plates of segments III-VI dark brown and more dilute laterally on I-VI (Fig. 60 E); fronto-ocular triangle higher than wide; cibarium with median portion light (Fig. 60 D); basal sector of R with hair; eighth sternite scarcely sclerotized with about 20 setae; gonapophysis longer than wide (Fig. 60 F); genital fork with anterior branch well sclerotized (Fig. 60 G); paraproct about 3 times longer than wide at base (Fig. 60 H). Male: Wing length 2.2 mm; scutum blackish with silvery band laterally and posteriorly and enlarged antero-medially (Figs. 61 A, B); scutellum, metanotum and pleurae as in female; legs yellowish with distribution of pigmentation and relative proportion of size similar to female; abdomen gray greenish, with large light areas on tergites II, and smaller ones on III-VI (Fig. 61 C); gonocoxite, gonostylus and ventral plate as shown in figures 61 D, E.

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B C A

D H

F

I

G J

E

K

Figure 61. Male. A-E: S. (Chirostilbia) acarayense; A-B: scutum in dorsal and lateral views; C: abdomen; D: gonocoxite and gonostylus; E: ventral plate. F-G: S. (C.) papaveroi; F: gonocoxite and gonostylus; G: median sclerite and endoparameres. H-K: S. (C.) subpallidum; H-I: gonocoxite and gonostylus in dorsal and ventral view; J-K: ventral plate in different position.

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Pupa: Cocoon slipper shaped with straight edge and slightly projected anteriorly (Figs. 62 C, D); shorter than in Simulium subpallidum; weave compact with threads obvious; length at base 2.6-2.7 mm; at dorsum 2.2-2.3 mm; gills 1.1-1.2 mm; frontoclypeus and thorax with scarce tubercles (Fig. 62 E); cephalic and thoracic trichomes with 3-5 branches; gills with 8 branches, open from base (Fig. 62 F). Larva: Maximum length 5.3-5.8 mm; cephalic apotome with more pigmentation at center and base; ratio of proximal, medial and distal antennal segments = 1:1.1-1.2:1.11.6; cephalic fan with 44-50 rays; postgenal cleft subovoidal, postgenal bridge a little shorter than half of hypostomium length; anal ring with 50-70 rows with 14-15 hooks each; rectal papillae with 8-10 diverticula on each lobe. Distribution: Argentina: Entre Ríos, Misiones; Brazil: Ceará, Mato Grosso, Minas Gerais, Rio Grande do Sul, Santa Catarina, São Paulo; Paraguay; Uruguay. Bionomics: Simulium acarayense breeds in creeks with much discharge and fastflowing, crystalline water, attached to leaves and branches. The female feeding habits are unknown. Discussion: Similarities of S. acarayense with S. subpallidum were treated in the species description. Simulium (Chirostilbia) papaveroi COSCARÓN (Figs. 60 L-P; 61 F, G; 62 G-I; 63 Q.) Simulium papaveroi COSCARÓN, 1982: 67-73. Simulium (Chirostilbia) papaveroi COSCARÓN, 1987: 26; 1991: 233-235. Chirostilbia papaveroi PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

Female: Wing length 1.8-2.0 mm; general coloration dark brown grayish; scutum brown with gray pollinosity and pilosity silver to golden; laterally and posteriorly bordered by silvery band; with lateral light, scutum shows 1+1 submedian silvery vittae, joined posteriorly to grayish posterior area (Fig. 60 L); with frontal illumination, vittae and border change to black (Fig. 60 M); scutellum light brown; metanotum velvety blackish with scarce darker portion; abdomen brown grayish with tergite I light brown grayish; tergite II with 1+1 silvery lateral spots; tergites III-VI black with silvery narrow posterior margin; tergites VII-IX green grayish; fronto-ocular triangle short; sensory vesicle of maxillary palp about half of palpomere length and with abundant tuberosity (Fig. 60 N); basal sector of R and Sc without hair; claw without basal tooth (Fig. 60 O); eighth sternite with about 28 setae; gonapophysis subtriangular, without microtrichia; paraproct nearly two times longer than wide at base (Fig. 60 P); genital fork with anterior branch well sclerotized. Male: Wing length 1.7-1.8 mm; general coloration blackish; scutum black, bordered laterally and posteriorly by silvery pruinose band; with lateral illumination, silvery area enlarged antero-laterally as in Simulium acarayense; scutellum, metanotum and pleurae as in female; abdomen blackish with 1+1 silvery spots on tergites II, V-IX; length/wide ratio of hind basitarsus = 6; gonostylus relatively thin, about two times longer than wide at base, with median diagonal crest (Fig. 61 F); ventral plate very similar to that of Simulium subpallidum; median sclerite with deep incision; endoparameres with big teeth (Fig. 61 G). Pupa: Cocoon slipper shaped, weave thick, with evident threads, frequently with thin weave covering thorax dorsally and gills (Fig. 62 G); length at base 2.5-2.8 mm, at dorsum

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A

E B I

F

G

C D

H

Figure 62. Pupa. A-B: S. (Chirostilbia) subpallidum; A: lateral view; B: anterior portion of thorax and gill. C-F: S. (C.) acarayense; C-D: lateral and dorsal views; E: frontoclypeus; F: anterior portion of thorax and gill. G-I: S. (C.) papaveroi; G: lateral view; H: frontoclypeus; I: anterior portion of thorax and gill.

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D

K

M

A N

B

H

I

E

L

J

O

Q F

C G

P

Figure 63. Larva. A-C: S. (Chirostilbia) pertinax; A: cephalic apotome; B: antenna, C: postgenal cleft and hypostomium. D-G: S. (C.) serranus; D: head, dorsal view; E: teeth of mandible; F: postgenal cleft and hypostomium; G: anal sclerite. H: S. (C.) spinibranchium; lateral sclerite of proleg. I-J: S. (C.) empascae; I: antenna; J: teeth of mandible. K-L: S. (C.) distinctum; K: lateral view; L: teeth of mandible. M-P: S. (C.) subpallidum; M: head, dorsal view; N: antenna; O: teeth of mandible; P: anal sclerite. Q: S. (C.) papaveroi, postgenal cleft and hypostomium.

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2.0-2.3 mm, gills 1.5-1.7 mm; frontoclypeus (Fig. 62 H) and exposed portion of thorax without tubercles, or only very scarce on thorax; cephalic and thoracic trichomes single or bifid and very short; gills each with 8 branches with short basal trunk, primary and secondary branches (Fig. 62 I); abdomen with short terminal spines. Larva: Maximum length 5.5-6.0 mm; head brownish, body gray green-bluish; cephalic apotome scarcely pigmented and light on anterior area and lateral borders; ratio of proximal, medial and distal antennal segments = 1:1.2:1.5-2; postgenal cleft rounded anteriorly with postgenal bridge about half of hypostomium length (Fig. 63 Q); second marginal tooth about 1/4 of first; latero-mandibular process present; cephalic fan with about 40-46 rays; anal sclerite with single hairs; anal ring with 70-81 rows with 12-14 hooks each; lateral sclerite of thoracic proleg with about 25 teeth arranged in 8 groups; body cuticle without hair. Distribution: Brazil: Bahia, Minas Gerais. Bionomics: Simulium papaveroi breeds in torrential creeks with large discharge and crystalline water. The female feeding habits are unknown. Discussion: Imagoes of the PERTINAX species group have coloration similar to that of S. papaveroi, in particular S. serranus. Differences in gonostylus morphology, presence of frontoclypeal and thoracic tubercles and disposition of the gill branches allow separation of S. papaveroi from other species. Simulium acarayense is also very similar but has a relatively shorter gonostylus and very open pupal gill branches. Species inquirenda Simulium striginotum (ENDERLEIN) Thyrsopelma striginotum ENDERLEIN, 1934b: 284.

Female type material from the Dresden Museum is very similar to S. laneportoi. Lack of knowledge of the genitalia morphology and other stages, do not permit its comparison with other species; we, therefore, maintain S. striginotum as a species inquirenda. Simulium (Inaequalium) COSCARÓN & WYGODZINSKY (Figs. 64-70; 141.) Type-species: Eusimulium inaequale PATERSON & SHANNON, 1927.

Female: Species medium sized and blackish; frons and clypeus gray pollinose; without bright opalescence; scutum black with grayish pollinosity, generally with gray band around lateral and posterior edges and in some species with 1+1 submedian anterior light gray stripes; scutum with decumbent homogeneous silvery to yellowish pilosity; scutellum, metanotum and pleurae black with gray pilosity; abdomen black with 1+1 silvery spots on tergite II; tergites I-IV velvety and V-VIII with cereous sheen; frons slightly convergent; fronto-ocular triangle deep; cibarium with basal portion projected medially and covered with robust and well-sclerotized spines; legs with petaloid hair; calcipala well developed, longer than wider; basal sector of R with hair; Sc with about 20 hairs; claws generally with well-developed subbasal tooth; gonapophysis divergent, subtriangular, with sclerotized internal edge; eighth sternite with about 30-40 hairs on

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each side; cerci rounded apically; paraprocts generally shorter than wide, apically rounded or truncated; genital fork elongated, about 2 times length of each anterior branch. Male: Scutum velvety black, grayish silvery on lateral and posterior borders; with some illumination, 1+1 submedian gray pollinose spots evident on anterior third; scutum pilosity yellowish to golden; abdomen blackish with tergites II, VI, VII and 1+1 on VIII with small anterior portion whitish-silvery; basal sector of R with hair arranged in one row; Sc with 3-4 hairs; calcipala as in female; gonocoxite subtrapezoidal, about as long as wide; gonostylus slightly shorter than gonocoxite, subtriangular to subtrapezoidal, with large apical or subapical spur; ventral plate subovoidal to subtriangular, wider than long; endoparameres with robust spines. Pupa: Cocoon slipper shaped with anterior edge generally straight and reinforced; weave strong with threads obvious; gill generally with 6 filiform branches or with robust 2-3 irregular shaped branches; cephalothorax with obvious tubercles; frontoclypeal trichomes with 3-5 branches, and thoracic trichomes with 3-8 branches; abdominal spines, hooks and terminal spines well developed. Larva: Body without hair or scales; cephalic apotome generally with darkened median posterior and basal area, margin with light spots, or with more diffuse ornamentation; postgenal cleft generally deep with postgenal bridge smaller or about as long as hypostomium. Median hypostomium tooth lower than lateral teeth; hypostomial setae about 6 on each side; antennae surpassing stem of cephalic fan; terminal segment longer than each preceding segment. Mandible with 2 marginal teeth, anterior tooth over two times length of second; with robust latero-mandibular tooth; cephalic fan with 35-40 rays; anal sclerite with single hairs; anal ring with 65-90 rows with 12-15 hooks in each; ninth segment with two small ventral tubercles, sometimes not evident; rectal papillae with 2-13 diverticula on each lobe. Distribution: Simulium (Inaequalium) is a peculiarly South American taxon (although reaching Panama) from Venezuela to northern Argentina, present in all the countries, except Chile; the greatest diversity is in the southeastern Brazil mountain areas (Fig. 141). Discussion: Simulium (Inaequalium) has 14 known species; adults are very similar, but pupae have good characters to differentiate the species, especially by gill morphology;. Two species groups are evident, as shown through cladistic analysis (COSCARÓN & COSCARÓN ARIAS (1997) and STRIEDER (1997): · The INAEQUALE species group is characterized as follows: basal portion of cibarium pyramidal, acuminate apically; paraprocts longer than high, not acuminate distally; gonostylus shorter than gonocoxite; ventral plate wider than long; pupal gill composed of 6 filamentous branches and postgenal bridge shorter than hypostomium length. Simulium inaequale, S. clarki, S. subnigrum, S. nogueirai, S. leopoldense, S. beaupertuyi, S. hauseri, S. clavibranchium, S. subclavibranchium, S. nahimi, S. mariavulcanoae, S. diversibranchium, S. rappae, S. travassosi.. · The BOTULIBRACHIUM species group is characterized as follows: basal portion of cibarium with very low pyramidal median process to relatively depressed medially; paraprocts higher than long and acuminate distally; gonostylus generally as long as gonocoxite; ventral plate as long as wide; pupa gill composed of 2-6 thick irregular filaments; larval postgenal bridge as long as hypostomium length. Simulium botulibranchium, S. souzalopesi, S. petropoliense.

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TO

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SPECIES

Females Simulium hauseri and S. leopoldense females are unknown. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(1) – 7(6) –

Paraprocts longer than high (Figs. 64 G, L, P) ............................................................ 2 Paraprocts higher than long, acuminate distally (Figs. 69 D, Q) ............................. 6 Scutum without 1+1 submedian anterior band in certain lighting ........................... ............................................................ Simulium mariavulcanoae, Simulium diversibranchium Scutum with 1+1 submedian anterior band in certain light (Fig. 64 A) ................ 3 Coxae I blackish; femora and tibiae with strong blackish coloration; paraprocts proportionally low (Fig. 64 L) ................................................... Simulium clavibranchium Coxae I light grayish-brown, femora and tibiae with slight blackish coloration (Fig. 64 C); paraprocts proportionally higher (Figs. 64 G, P) ............................................ 4 Paraproct relatively high with apical ovoidal edge (Fig. 64 P); genital fork with stem relatively longer and posterior branches proportionately thicker (Fig. 64 O) ...................................................................................................................Simulium travassosi Paraproct relatively low with apical rounded edge (Fig. 64 G); genital fork with stem relatively shorter and posterior branches proportionately thinner (Fig. 64 N) ................................................................................................................................................ 5 Wing length 1.8-2.0 mm ................ Simulium rappae, Simulium clarki, Simulium nahimi Wing length 2.3-2.8 mm ..................................................................................................... ....... Simulium inaequale, Simulium nogueirai, Simulium subclavibranchium, Simulium subnigrum Gonapophyses with parallel inner margins (Fig. 69 T) ............ Simulium petropoliense Gonapophyses with divergent inner margins (Fig. 69 P) .......................................... 7 Basal portion of cibarium with subconical medial elevation (Fig. 69 C) ................. ........................................................................................................ Simulium botulibranchium Basal portion of cibarium slightly elevated, with small median depression (Fig. 69 K) ................................................................................................................ Simulium souzalopesi

Males Simulium leopoldense, S. hauseri and S. petropoliense males are unknown. 1 – 2(1) – 3(2)

Gonostylus short, about 1/2 as long as gonocoxite (Figs. 65 C, G, M), and not more than 2 times longer than wide at base (Figs. 65 C, D, F, G, H, J, K, L, N, P); ventral plate wider than long (Figs. 65 C, E, I, O) ..................................................... 2 Gonostylus elongated, about same length as gonocoxite (Fig. 69 R), and about 3 times longer than wide at base (Fig. 69 I); ventral plate about as long as wide (Figs. 69 H, S) ................................................................................................................................ 6 Gonostylus with 3-8 spurs on dorsal external edge to apex (Fig. 65 K) ................. .......................................................................................................................Simulium nahimi Gonostylus with 1 spur at apex (Figs. 65 D, F, H, J, L, N, P) .................................. 3 Gonostylus not acuminate apically and with subapical spur (Figs. 65 J, L) ............ ............................................................ Simulium mariavulcanoae, Simulium diversibranchium

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Gonostylus acuminate and with apical spur (Figs. 65 D, F, G, M, N) .................... 4 Ventral plate rounded apically (Fig. 65 I) ........................... Simulium subclavibranchium Ventral plate acuminate apically (Figs. 65 E, O) ......................................................... 5 Gonostylus thinning from distal half of its length (Figs. 65 D, P) ........................... ..................................................................................... Simulium inaequale, Simulium rappae Gonostylus thinning from distal third of its length (Figs. 65 F, G, H, N) .............. ........ Simulium clavibranchium, Simulium nogueirai, Simulium subnigrum, Simulium travassosi Ventral plate well expanded basally (Fig. 69 H) .................. Simulium botulibranchium Ventral plate not well expanded basally (Fig. 69 S) .................... Simulium souzalopesi

Pupae 1 – 2(1) – 3(2) – 4(3) – 5(3) – 6(5) – 7(5) – 8(7) – 9(7) –

Gill with 6 branches generally filiform (Figs. 66 A B, D, G; 67 A-L) .................... 2 Gill with 2-3 thick branches, frequently asymmetrical and with diverticula (Figs. 66 N; 70 A-C, G-I, P-R) ................................................................................................. 12 Gill branches thick and about half of cocoon length; secondary branches about as long as primaries (Fig. 66 I) .................................................................... Simulium rappae Gill branches filiform, as long as or longer than cocoon length; secondary branches longer than primary branches (Figs. 66 A, B, D, G; 67 A-L) .................. 3 Gill filaments thicker distally (Figs. 67 G, H, I) .......................................................... 4 Gill filaments not thicker distally (Figs. 67 A-C, F, J, K) ........................................... 5 Gill filaments greatly thickened distally and thinning abruptly (Figs. 67 G, H); secondary dorsal and median branches bifurcating far from primary ventral bifurcation; gill length 3.5-4.2 mm ........................................... Simulium clavibranchium Gill filaments slightly thickened distally and thinning gradually (Fig. 67 I); secondary dorsal and median branches bifurcated close to primary ventral branch; gill length 4.8-5.5 mm ............................................................ Simulium subclavibranchium Gill filaments branching very close to base and at about same distance, with very short primary and secondary branches (Fig. 67 A) .................................................... 6 Gill filaments branching at variable and different distances from base, with variable lengths for primary and secondary branches (Figs. 67 B-F, J, K) ............ 7 Frontoclypeus and exposed portion of thorax with abundant tubercles ................ ....................................................................................... Simulium inaequale, Simulium clarki Frontoclypeus and exposed portion of thorax with scarce tubercles ...................... ...................................................................................................................... Simulium hauseri Gill filaments with dorsal branch thicker than others (Figs. 67 K, L) .................... 8 Gill filaments of about same diameter ......................................................................... 9 Primary and secondary dorsal branches much thicker and longer than secondary ventral branches; secondary ventral branches directed downward and posteriorly (Fig. 67 L) ................................................................................... Simulium diversibranchium Primary and secondary dorsal branches only slightly thicker than and about same length as secondary ventral branches (Fig. 67 K); secondary ventral branches directed anteriorly (Fig. 67 K) .................................................. Simulium mariavulcanoae Gill longer than cocoon .................................................................................................. 10 Gill about same length as cocoon ................................................................................ 11

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Primary ventral branch relatively short; secondary dorsal and median branches bifurcated at about same level (Fig. 67 B) ..................................... Simulium subnigrum – Primary ventral branch relatively long, reaching at least to 1/3 of secondary ventral branch; secondary dorsal and median branches bifurcated at different levels ....................................................................................................... Simulium leopoldense 11(9) Secondary ventral branch bifurcated at median distance between ventral primary and secondary dorsal bifurcation (according to D’ANDRETTA & GONZALEZ, Fig. 67 C), or sometimes at same level (PY-DANIEL & MOREIRA, 1989; & STRIEDER, 1998, Fig. 67 D) .................................................................................................Simulium nogueirai – Secondary dorsal and ventral branches bifurcated at greater distance than primary ventral bifurcation and at about same distance between them (Fig. 67 J) ............... .......................................................................................................................Simulium nahimi 12(1) Gill with 3 branches of about uniform diameter, symmetrical, without diverticula or branches (Figs. 66 L-N) ..................................................................Simulium travassosi – Gill with 2 primary branches of variable diameter, asymmetrical, with diverticula or terminal branches ........................................................................................................ 13 13(12) Gill integument with abundant and large microtrichiae; dorsal branch with 4 relatively thin branches (Figs. 70 P-R); very abundant tubercles on head and thorax (Fig. 70 S) .............................................................................. Simulium petropoliense – Gill integument with sparse spiculae and tubercles; dorsal branch with different apical branches; head and thorax with few tubercles ............................................... 14 14(13) Dorsal branch with 4 terminal diverticula with blunt apices (Fig. 70 C) ................. ........................................................................................................ Simulium botulibranchium – Dorsal branch with 6 apically acuminate branches (Fig. 70 I) ................................... ................................................................................................................ Simulium souzalopesi CLAVES

DE ESPECIES DE

SIMULIUM (INAEQUALIUM)

Hembras Simulium hauseri y S. leopoldense con hembras desconocidas. 1 – 2(1) – 3(2) – 4(3)

Paraproctos aproximadamente más largos que altos (Figs. 64 G, L, P) ................. 2 Paraproctos más altos que largos, aguzados distalmente (Figs. 69 D, Q) .............. 6 Escudo sin 1+1 banda submediana anterior observada con cierta luz .................... ............................................................ Simulium mariavulcanoae, Simulium diversibranchium Escudo con 1+1 banda submediana anterior observada con cierta luz (Fig. 64 A) ................................................................................................................................................ 3 Coxa I negruzca; fémur y tibia con fuerte coloración negruzca; paraproctos proporcionalmente bajos (Fig. 64 L) ........................................ Simulium clavibranchium Coxa I castaño grisáceo claro; fémures y tibias con escasa coloración negruzca (Fig. 64 C); paraproctos proporcionalmente más altos (Figs. 64 G, P) .................. 4 Paraprocto relativamente alto y de borde apical ovoidal (Fig. 64 P); horquilla genital con el tallo relativamente largo y ramas posteriores proporcionalmente gruesas (Fig. 64 O) ................................................................................Simulium travassosi

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Paraprocto relativamente bajo y de borde apical redondeado (Fig. 64 G); horquilla genital con el tallo relativamente corto, y ramas posteriores proporcionalmente más finas (Fig. 64 N) ......................................................................................................... 5 Longitud alas 1,8-2,0 mm ............... Simulium rappae, Simulium clark, Simulium nahimi Longitud alas 2,3-2,8 mm ................................................................................................... ....... Simulium inaequale, Simulium nogueirai, Simulium subclavibranchium, Simulium subnigrum Gonapófisis con borde interno paralelo (Fig. 69 T) ................. Simulium petropoliense Gonapófisis con bode interno divergente (Fig. 69 P) ................................................ 7 Porción basal del cibario medianamente elevado subcónico (Fig. 69 C) ................. ........................................................................................................ Simulium botulibranchium Porción basal del cibario escasamente elevada, con pequeña depresión mediana (Fig. 69 K) ............................................................................................ Simulium souzalopesi

Machos Simulium hauseri, S. leopoldense y S. petropoliense con machos desconocidos. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(1) –

Gonostilo corto, aproximadamente la mitad del largo del gonocoxito (Figs. 65 C, G, M) y no más de 2 veces más largo que ancho en la base (Figs. 65 C, D, F, G, H, J, K, L, N, P); placa ventral más ancha que larga (Figs. 65 C, E, I, O) .................. 2 Gonostilo alargado, aproximadamente del mismo largo que el gonocoxito (Fig. 69 R), y unas 3 veces más largo que ancho en la base (Fig. 69 I); placa ventral aproximadamente tan larga como ancha (Figs. 69 H, S) ........................................... 6 Gonostilo con 3-8 espolones en el borde dorsal externo hasta el ápice (Fig. 65 K) .......................................................................................................................Simulium nahimi Gonostilo con 1 espolón sobre el ápice (Figs. 65 D, F, H, J, N, L, P) ................... 3 Gonostilo romo apicalmente con espolón subapical (Figs. 65 J, L) ......................... ............................................................ Simulium mariavulcanoae, Simulium diversibranchium Gonostilo aguzado y con espolón apical (Figs. 65 D, F, G, M, N) ......................... 4 Placa ventral redondeada apicalmente (Fig. 65 I) ............ Simulium subclavibranchium Placa ventral aguzada apicalmente (Figs. 65 E, O) ..................................................... 5 Gonostilo afinándose desde la mitad distal de su largo (Figs. 65 D, P) ................... ..................................................................................... Simulium inaequale, Simulium rappae Gonostilo afinándose desde el tercio distal de su largo (Figs. 65 F, G, H, N) ....... ........... Simulium clavibranchium, Simulium nogueirai, Simulium subnigrum, Simulium travassosi Placa ventral bien expandida basalmente (Fig. 69 H) ........ Simulium botulibranchium Placa ventral no expandida basalmente (Fig. 69 S) ..................... Simulium souzalopesi

Pupas 1 – 2(1)

Branquias con 6 ramas generalmente filiforme (Figs. 66 A, B, D, G; 67 A-L) ..... 2 Branquias con 2-3 ramas gruesas, frecuentemente asimétricas y con divertículos (Figs. 66 N; 70 A-C, G-I, P-R) ...................................................................................... 12 Branquias de ramas gruesas, aproximadamente la mitad del largo del capullo; ramas secundarias aproximadamente tan largas como las primarias (Fig. 66 I) ..... ....................................................................................................................... Simulium rappae

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Branquias de ramas filiformes, tan largas o más que el largo del capullo; ramas secundarias más largas que las primarias (Figs. 66 A, B, D, G; 67 A-L) ................ 3 Filamentos de las branquias más gruesos distalmente (Figs. 67 G-I) ..................... 4 Filamentos de las branquias no más gruesos distalmente (Figs. 67 A-C, F, J, K) ................................................................................................................................................ 5 Filamentos de las branquias muy engrosados distalmente y afinándose abruptamente (Figs. 67 G-H); ramas secundaria dorsal y mediana bifurcadas lejos de la bifurcación primaria ventral; longitud de la branquia 3,5-4,2 mm .................. .......................................................................................................... Simulium clavibranchium Filamentos de la branquia moderadamente engrosados distalmente y afinándose suavemente (Fig. 67 I); ramas secundaria dorsal y mediana bifurcadas cerca de la rama primaria ventral; longitud de las branquias 4,8-5,5 mm .................................... ..................................................................................................... Simulium subclavibranchium Filamentos de las branquias ramificados muy cerca de la base y aproximadamente a la misma distancia, mostrando ramas primarias y secundarias muy cortas (Fig. 67 A) ................................................................................................................................................ 6 Filamentos de las branquias ramificados a diferente distancia de la base, mostrando ramas primarias y secundarias de variable longitud (Figs. 67 B-F, J, K) ................................................................................................................................................ 7 Frontoclípeo y porción expuesta del tórax con abundantes tubérculos .................. ....................................................................................... Simulium inaequale, Simulium clarki Frontoclípeo y porción expuesta del tórax con escasos tubérculos .......................... ...................................................................................................................... Simulium hauseri Filamentos de las branquias con rama dorsal más gruesa que las otras (Figs. 67 K, L) ........................................................................................................................................... 8 Filamentos de las branquias aproximadamente del mismo diámetro ..................... 9 Ramas primaria dorsal y secundaria dorsal notoriamente gruesas y más gruesas que la secundaria ventral; ramas ventrales secundarias dirigidas hacia abajo y atrás (Fig. 67 L) ................................................................................... Simulium diversibranchium Ramas primaria y secundaria dorsales solamente un poco más gruesas y aproximadamente de la misma longitud que la secundaria ventral (Fig. 67 K); ramas ventrales secundarias dirigidas en dirección frontal (Fig. 67 K) .................... ......................................................................................................... Simulium mariavulcanoae Branquia más larga que el largo del capullo ............................................................... 10 Branquia aproximadamente del mismo largo que el capullo .................................. 11 Rama primaria ventral relativamente corta; ramas secundarias dorsal y media bifurcadas aproximadamente al mismo nivel (Fig. 67 B) ........... Simulium subnigrum Rama primaria ventral relativamente larga, llegando por lo menos a 1/3 de la secundaria ventral; ramas dorsales y medias bifurcadas a diferentes niveles .......... ................................................................................................................. Simulium leopoldense Rama secundaria ventral bifurcada a media distancia entre la bifurcación de la primaria ventral y la bifurcación de la secundaria dorsal (según D’ANDRETTA & GONZÁLEZ, 1964), Fig. 67 C), o a veces al mismo nivel (PY-DANIEL & MOREIRA, 1989; STRIEDER, 1998; Fig. 67 D) ........................................................Simulium nogueirai

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Rama secundaria dorsal y ventral bifurcada a mayor distancia de la bifurcación de la primaria ventral y al mismo nivel entre ellas (Fig. 67 J) ...............Simulium nahimi 12(1) Branquias con 3 ramas de aproximadamente diámetro uniforme, simétricas, sin divertículos o ramas (Figs. 66 L-N) ................................................... Simulium travassosi – Branquias con 2 ramas primarias de diámetro variable, asimétricas, con divertículos o ramas terminales ..................................................................................... 13 13(12) Tegumento de la branquia con largas y abundantes microtriquias; rama dorsal con 4 ramas relativamente finas (Figs. 70 P-R); cabeza y tórax con abundantes tubérculos (Fig. 70 S) ....................................................................... Simulium petropoliense – Tegumento de la branquia con espículas y escasos tubérculos; rama dorsal presenta diferentes ramas apicales; cabeza y tórax sin o con escasos tubérculos .............................................................................................................................................. 14 14(13) Rama dorsal con 4 divertículos terminales con ápice romo (Fig. 70 C) .................. ........................................................................................................ Simulium botulibranchium – Rama dorsal con 6 ramas terminales con ápice aguzado (Fig. 70 I) ......................... ................................................................................................................ Simulium souzalopesi INAEQUALE species group Simulium (Inaequalium) inaequale (PATERSON & SHANNON) (Figs. 64 A-G; 65 A-E; 66 A, 67 A; 68 B, C.) Eusimulium inaequalis PATERSON & SHANNON, 1927: 738-740; PINTO, 1931: 677; VULCANO, 1967: 13. Simulium inaequalis WYGODZINSKY, 1949: 305; 1953a: 304. Simulium inaequale VARGAS, 1945a: 147; COSCARÓN & WYGODZINSKY, 1973b: 142. Simulium (Inaequalium) inaequale COSCARÓN & WYGODZINSKY, 1984: 80-84; COSCARÓN, 1987: 21; 1991: 157158; COSCARÓN & COSCARÓN ARIAS, 1997: 111; COSCARÓN & MIRANDA ESQUIVEL, 1998b: 337. Inaequalium inaequale PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 44; 2004: 118-119; STRIEDER & PY-DANIEL, 1999: 51; 2000: 34; 2002: 66. Trichodagmia manicata ENDERLEIN, 1934b: 288. Simulium jundiaense D’ANDRETTA & GONZÁLEZ, 1964: 108-110. Simulium baiensis PINTO, 1931: 685. Thyrsopelma argentata ENDERLEIN, 1936: 125.

Female: Wing length 2.4-2.7 mm; frons and clypeus grayish pollinose; scutum black with grayish pollinosity and pilosity gray to tin colored; with some light directions, 1+1 submedian gray pollinose band (Fig. 64 A) that can be thinner and extended to posterior edge; (these bands can be easier to see if there is no hair); cibarium with basal portion well elevated medially; sensory vesicle of maxillary palp about 1/3 of palpomere length (Fig. 64 B); legs gray yellowish, with darkened areas dark brown to blackish as shown in figures 64 C, D, E; calcipala well developed, about as long as wide (Fig. 64 F); genitalia as shown in figure 64 G; paraproct longer than high with abundant microtrichiae and rounded distal edge. Male: Wing 2.3-2.6 mm; scutum black with golden pilosity; with some illumination. 1+1 submedian light gray pollinose subtriangular spots (Fig. 65 A) and band of gray pollinosity laterally and posteriorly (Fig. 65 B); legs with coloration as in female but more

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K Figure 64. Female. A-G: S. (Inaequalium) inaequale; A: scutum, dorsal view; B: third palpomere and sensory vesicle; C-E: fore, mid and hind leg, respectively; F: calcipala and pedisulcus; G: genitalia in ventral view: eighth sternite, gonapophyses, spermatheca, genital fork, paraproct and cercus (the latter in two positions). H: S. (I.) subnigrum, chaetotaxy of wing, anterior border. I-L: S. (I.) clavibranchium; I: frons; J: fronto-ocular triangle; K: claw; L: paraproct and cercus. M-N: S. (I.) subclavibranchium; M: basal portion of cibarium; N: genital fork. O-P: S. (I.) travassosi; O: genital fork; P: paraproct and cercus.

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darkened; genitalia as shown in figure 65 C; gonostylus elongated, subtriangular and robust spur at apex (Fig. 65 D); ventral plate subtriangular (Fig. 65 E); endoparameres with large teeth. Pupa: Cocoon length on base 3.3-3.5 mm, on dorsum 3.0-3.2 mm, gill length 4.0-4.2 mm; aspect of pupa in accordance with figure 66 A; gill with 6 similar filamentous branches emerging very close to base from 3 short primary branches (Fig. 67 A); frontoclypeus and exposed portion of thorax with abundant and obvious tubercles; head with 2+2 frontal and 1+1 facial trichomes with 2-4 branches; thoracic trichomes with 4-6 branches; abdomen chaetotaxy as in Simulium subnigrum. Larva: Maximum length 5.5-7.0 mm; coloration light yellowish-gray, darkened and greenish gray dorso-laterally; head brown, with light median subbasal window on cephalic apotome (Fig. 68 B); antennae longer than stem of cephalic fan; ratio of proximal, medial and distal antennal segments = 1:1.1-1.3:1.2-1.3. Mandible with teeth as explained for subgenus; anal sclerite with abundant single hairs (Fig. 68 C); anal ring with 62-68 rows of 10-12 hooks each; rectal papillae with 11-16 diverticula on each lobe. Distribution: Argentina: Jujuy, Misiones, Salta, Tucumán; Bolivia: La Paz; Brazil: Amapá, Bahia, Espírito Santo, Mato Grosso, Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo; Paraguay: from the Cordillera and Alto Paraná. Bionomics: Immature stages of S. inaequale live in small shallow creeks, mostly on aquatic vegetation in fast-flowing areas. Females can be anthropophilic and sometimes pestiferous, as in Santo Inácio (Bahia, Brazil). Discussion: Arrangement of the gill branches helps differentiate the most similar species, such as S. subnigrum, with branching at further distances and in different positions. The most similar species to S. inaequale is S. clarki, with similar pupal gill branching, and adult ornamentation and morphology, but it is smaller in size. Simulium inaequale could represent subspecific variation in Panama and Venezuela. Simulium hauseri STRIEDER has very similar pupal gill branching, and small morphological differences in the number of frontoclypeal and thoracic tubercles and abdominal chaetotaxy. As the adults and larva are unknown, we do not have enough information to consider it a variation of S. inaequale or S. subnigrum. Simulium nogueirai, in accordance with the original illustrations, is also very close to S. inaequale. Simulium (Inaequalium) clarki FAIRCHILD Simulium clarki FAIRCHILD, 1940: 703; VARGAS & DÍAZ NÁJERA, 1948a: 75; RAMÍREZ PÉREZ, 1971: 286; 1983: 42; RAMÍREZ PÉREZ et al., 1982: 31. Simulium (Inaequalium) clarki COSCARÓN, 1983: 21; 1991: 159; SHELLEY et al., 2004: 449, 453, 460. Inaequalium clarki PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 29; STRIEDER & PYDANIEL, 2000: 26.

This species is very close to S. inaequale; females and males of the type material are morphologically and ornamentally similar to S. inaequale, only a little smaller (wing length 1.8-1.9 mm) and with lighter legs, more yellowish, and lighter distal blackish areas on the hind femur, tibia and tarsi. The pupa illustrated by FAIRCHILD of S. clarki shows gill branching similar to that of S. inaequale. Material from Venezuela determined by RAMÍREZ PÉREZ (1971-1973) as S. clarki, is similar to the type material, but is larger.

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Figure 65. Male. A-E: S. (Inaequalium) inaequale; A: scutum, dorsal view; B: scutum, lateral view; C: genitalia in ventral view: gonocoxite, gonostylus, ventral plate, endoparameres and median sclerite; D: gonostylus; E: ventral plate in another position than C. F: S. (I.) subnigrum, gonostylus. G-H: S. (I.) clavibranchium; G: gonocoxite and gonostylus; H: gonostylus in different position. I: S. (I.) subclavibranchium, ventral plate. J: S. (I.) mariavulcanoae, gonostylus. K: S. (I.) nahimi, gonostylus (from PY-DANIEL, 1984). L: S. (I.) diversibranchium, gonostylus. M-O: S. (I.) travassosi; M: gonocoxite and gonostylus; N: gonostylus in different position; O: ventral plate P: S. (I.) rappae, gonostylus; N: Gonostylus in different position; O: Ventral plate.

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Figure 66. Pupa. A: S. (Inaequalium) inaequale, lateral view. B-C: S. (I.) clavibranchium; B: lateral view; C: frontoclypeus. D-E: S. (I.) subclavibranchium; D: lateral view; E: frontoclypeus. F: S. (I.) mariavulcanoae, thoracic trichomes. G-H: S. (I.) diversibranchium; G: lateral view; H: thoracic trichomes. I-K: S. (I.) rappae; I: lateral view; J: frontoclypeus; K: cephalic trichomes. L-N: S. (I.) travassosi; L-M: dorsal and lateral view, respectively; N: gill.

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Female: Wing length 2.3 mm. Male: Wing length 2.2 mm. Pupa: Cocoon length on base 3.0 mm, dorsum 2.8 mm, gill filaments 3.3 mm. Distribution: Panama: Coclé, Darien; Venezuela: Amazonas, Aragua, Bolivar, Barinas, Carabobo, Cojedes, Guárico, Lara, Miranda, Trujillo. Bionomics: Breeds in small creeks in temperate to warm water from 150 to 1900 m altitude. Females bite humans. Discussion: It will be necessary to study the immature stages at the type locality to decide if S. clarki is a synonym of S. inaequale or a subspecies of S. inaequale from Central America, as suggested by WYGODZINSKY (1953), with variation characteristic of a species with a large distribution. Simulium (Inaequalium) subnigrum Lutz (Figs. 64 H; 65 F; 67 B, 68 D-F.) Simulium subnigrum LUTZ, 1910: 239-241; LUTZ & NUNEZ TOVAR, 1928: 43-44; LANE & VULCANO, 1943: 138; VULCANO, 1967: 21. Simulium (Inaequalium) subnigrum COSCARÓN & WYGODZINSKY, 1984: 90; COSCARÓN, 1987: 21; 1991: 162-163; STRIEDER et al., 1992: 60; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium subnigrum PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 90; 2004: 135; 2004: 120-121; STRIEDER & PY-DANIEL, 1999: 58; 2000: 66; 2002: 66. Simulium diversifurcatum LUTZ, 1910: 258-263; LUTZ & MACHADO, 1915: 46. Simulium mbarigui COSCARÓN & WYGODZINSKY, 1973a: 142-149.

Female: Wing length 2.3-2.8 mm; color similar to that of Simulium inaequale but with scutum pilosity silky grayish and femur of anterior and middle legs lighter without blackish apex; cibarium elevated medially with strong teeth covering median area, similar to Simulium subclavibranchium (Fig. 64 M); sensory vesicle of maxillary palp shorter than half length of palpomere; wing with one row of hair on Sc and more abundant on basal sector of R (Fig. 64 H); claw with or without subbasal tooth; genitalia as in Simulium inaequale, but paraprocts with distal edge shorter as in Simulium clavibranchium (Fig. 64 L). Male: Wing length 2.2-2.7 mm; coloration and morphology as for Simulium inaequale; gonostylus thicker and wider subapically than in S. inaequale (Fig. 65 F), and ventral plate subovoidal with apex well curved , similar to Simulium subclavibranchium. Pupa: Cocoon length at base 3.3-4.5 mm, dorsally 2.8-3.8 mm, body o f pupa 2.8-3.6 mm, gill filaments 3.8-5.0 mm; cocoon weave thick, with obvious threads, with anterior reinforcement and without dorsal projection; gill branching at different distances from base: dorsal and median secondary branches bifurcated at same distance or a little far than ventral branch (Fig. 67 B) with relative variability of distances in different specimens; frontoclypeus and exposed portion of thorax with abundant tubercles of medium size; cephalic trichomes with 2-3 and thoracic with 3-5 branches; abdominal chaetotaxy as shown in Fig. 68 A. Larva: Maximum length 5.3-6.0 mm; cephalic apotome darkened in basal and median areas; ratio of proximal, medial and distal antennal segments = 1:1.1:1.3-1.5 (Fig. 68 D). Mandible with one long latero-mandibular process surpassing inferior mandible rim (Fig. 68 E); cephalic fan with 35-40 rays; rectal papillae with 10-15 diverticula on each lobe (Fig. 68 F); anal ring with 67-75 rows with 12-14 hooks each.

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D Figure 67. Pupa. Gill. A: S. (Inaequalium) inaequale. B: S. (I.) subnigrum. C-E: S. (I.) nogueirai; C: extracted from D’ANDRETTA & GONZÁLEZ, 1964, D: from STRIEDER, 1998; E: from PY-DANIEL & MOREIRA, 1989. F: S. (I.) leopoldense. G-H: S. (I.) clavibranchium (H: gill, terminal portion with higher magnification). I: S. (I.) subclavibranchium. J: S. (I.) nahimi ( PY-DANIEL, 1984). K: S. (I.) mariavulcanoae. L: S. (I.) diversibranchium.

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C Figure 68. A: Pupa; S. (Inaequalium) subnigrum, chaetotaxy of abdomen. B-M: Larva; B-C: S. (I.) inaequale; B: head, dorsal view showing cephalic apotome; C: anal sclerite and trichomes; D-F: S. (I.) subnigrum; D: antenna; E: teeth of mandible; F: rectal papillae; G-J: S. (I.) diversibranchium; G: general aspect, lateral view; H: head, dorsal view; I: hypostomium and postgenal area; J: gill histoblast; K-L: S. (I.) rappae; K: general aspect; L: antenna; M: S. (I.) travassosi, head dorsal view.

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Distribution: Argentina: Corrientes, Misiones; Brazil: Bahia, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo; Colombia: Meta; Paraguay: Caaguzu; Trinidad; Venezuela: Bolivar, Distrito Federal, Miranda; Bionomics: Simulium subnigrum breeds in creeks with clear water and fast current in open areas. Female feeding habits are unknown. Discussion: A close species to S. subnigrum is S. inaequale, which occurs in the same habitats. It shows small differences in adult genitalia but the best character is the branching of the pupal gill; that of S. inaequale is very close to the base and the emergence of filaments is at the same distance. Simulium leopoldense has a very similar pattern of gill branching but the primary ventral branch reaches about the basal third of the secondary ventral branch, and the secondary dorsal and median branches are bifurcated at different distances. In S. subnigrum the primary ventral branch is shorter and is bifurcated near the base of the secondary branches, and the secondary dorsal and median branches are bifurcated at the same distance. Simulium hauseri also has the gill bifurcation closer to the base and sparser tubercles on the frontoclypeus and thorax than does S. subnigrum. Simulium nogueirai also is very similar, but the female scutum pilosity is golden and the pupal gill is shorter with a length about that of the venter of the cocoon; that of S. subnigrum is longer. The pupa of S. beaupertuyi has the primary ventral branch longer than the primary dorsal branch, the bifurcation of the secondary, median and dorsal branches proportionately lower than in S. subnigrum, and the thoracic tubercles larger in diameter than in S. subnigrum. Simulium (Inaequalium) nogueirai D’ANDRETTA & GONZÁLEZ (Figs. 67 C-E.) Simulium nogueirai D’ANDRETTA & GONZÁLEZ, 1964: 103-106; MOREIRA et al., 1994: 525. Simulium (Inaqualium) nogueirai COSCARÓN, 1987: 21; PY-DANIEL & MOREIRA, 1989: 55; STRIEDER et al., 1992: 55. Inaqualium nogueirai PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 65; 2004: 133-135; 2004: 119-120; STRIEDER & PY-DANIEL, 1999: 54; 2000: 46; 2002: 66.

Adults: Morphology and ornamentation as for INAEQUALE species group, without peculiar characters that permit separation; based on detailed redescription by PY-DANIEL & MOREIRA (1989) of Santa Catarina material (type material from São Paulo is lost), female genitalia as in Simulium inaequale, male gonostylus as in S. diversibranchium and S. clavibranchium, ventral plate as in S. travassosi; material from Santa Catarina studied by PY-DANIEL & MOREIRA does not show differences in adults described by D’ANDRETTA & GONZÁLEZ. Pupa: Cocoon slipper shaped, similar to Simulium subnigrum, but smaller, and proportionally with shorter gill in relation to cocoon basal length. PY-DANIEL & MOREIRA (1989), STRIEDER (1998) and STRIEDER & PY-DANIEL (2000) redescribed S. nogueirai from material of Rio Grande do Sul, Santa Catarina and São Paulo; it shows different levels of bifurcation of secondary dorsal and median branches, (Fig. 67 D, STRIEDER, 1998 or Fig. 67 E, PY-DANIEL & MOREIRA, 1989) in which median branch is shorter than dorsal; this is not in accordance with original descriptions, as in figure 67 C, showing approximately same level of bifurcation. Larva: Maximum length a little shorter than that of Simulium subnigrum; antennal segment ratio with medial segment shorter than proximal and distal; anal sclerite with

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single hairs and scales; rectal papillae with 3-6 diverticula on each lobe, anal ring with 7783 rows with 11-15 hooks each. Distribution: Brazil: Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: According to PY-DANIEL & MOREIRA (1989), immature stages were collected in small creeks with or without direct sunshine at 100-300 m altitude. Females feed on cows, horses and pigs (MOREIRA et al., 1994: 525, STRIEDER et al., 1992: 55). Discussion: Simulium nogueirai is very close to some other species of the INAEQUALE species group, especially to S. subnigrum, but can be separated by characters of the pupal gill and larva. It will be useful to study more material and with morphological and other methods to determine species validity, because the size of the pupa and the gill branching can be interpreted as variation in S. subnigrum, which frequently shows variation of gill bifurcation even in the same specimen. Simulium (Inaequalium) leopoldense STRIEDER & PY-DANIEL (Fig. 67 F.) Inaequalium leopoldense STRIEDER & PY-DANIEL, 2000: 14-19; STRIEDER, 2004: 119.

Adults: Unknown Pupa: Simulium leopoldense is very close to Simulium subnigrum but can be differentiated by the small size of Simulium leopoldense (cocoon length dorsally 2.4-3.0 mm, basally 2.9-3.2 mm), and gill branching; primary ventral branch three times longer than primary dorsal branch (Fig. 67 F) (about two times longer in Simulium subnigrum) and primary ventral branch reaching 1/3 of secondary ventral branch length (in S. subnigrum, about 1/5). Larva: Small differences also permit Simulium leopoldense to be differentiated from Simulium subnigrum; in S. leopoldense antenna surpasses length of cephalic fan stem, rectal papillae with 10-14 diverticula (in S. subnigrum 15-16) and body coloration with darker dorsal band relative to S. subnigrum. Distribution: Brazil: Rio Grande do Sul. Bionomics: Simulium leopoldense lives on aquatic vegetation in small, shallow creeks in open areas. Discussion: Simulium leopoldense shows few differences from S. subnigrum and they are difficult to separate. Simulium nogueirai is also similar, and studies to separate S. nogueirai from S. subnigrum could be used to separate it from S. leopoldense. Simulium (Inaequalium) beaupertuyi RAMÍREZ PÉREZ, Rassi & RAMÍREZ Simulium (Simulium) beaupertuyi RAMÍREZ PÉREZ et al., 1977: 165-177; RAMÍREZ PÉREZ et al., 1982: 29-31. Simulium beaupertuyi RAMÍREZ PÉREZ, 1983: 29. Simulium (Inaequalium) beaupertuyi COSCARÓN & WYGODZINSKY, 1984: 84; COSCARÓN, 1987: 21; 1991: 163-164. Inaequalium beaupertuyi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 20; STRIEDER & PY-DANIEL, 1999: 47; 2000: 19; 2002: 66.

Female: Very similar to S. subnigrum, but a little smaller (wing length 2.2 mm) and with golden pilosity on scutum. Male and Pupa: Very similar to Simulium subnigrum, having secondary central and dorsal gill branches bifurcated at about same distance.

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Larva: Very similar to Simulium subnigrum, but with proportionally (a little) shorter medial antennal segment. Distribution: Brazil: Roraima; Venezuela: Amazonas. Bionomics: Simulium beaupertuyi breeds in fast-flowing areas of small creeks with clear water. Discussion: As previously explained, S. beaupertuyi is very similar to S. subnigrum and it is very difficult to separate the two, similar to the case of S. inaequale and S. clarki, having few morphological differences and living in different regions from the other two wellknown southern species. A comparative study with cytological or molecular methods could help to resolve the identity of this species. Simulium (Inaequalium) hauseri (STRIEDER) Inaequalium hauseri STRIEDER, 1998: 41.

Known only from pupa. Pupa: Cocoon with gill branching and number of filaments as in Simulium inaequale; cocoon length ventrally 4.1 mm, on dorsum 3.7 mm, gills 4.2 mm; frontoclypeus and exposed portion of thorax with scarce rounded tubercles, on head mostly present on basal area around facial trichomes; on thorax irregularly arranged; cephalic trichomes with 2-4 branches, thoracic trichomes with 4-6 branches. Distribution: Brazil: Rio Grande do Sul. Bionomics: This species has been collected in a small creek. Discussion: Simulium hauseri is very similar to S. inaequale but can be differentiated by the large number of tubercles on the head and thorax and more basal gill branching. Simulium (Inaequalium) clavibranchium LUTZ (Figs. 64 I-L; 65 G, H; 66 B, C; 67 G, H.) Simulium clavibranchium LUTZ, 1910: 257-258; PINTO, 1931: 699; VARGAS, 1945a: 127; VULCANO, 1967: 9. Simulium (Inaequalium) clavibranchium COSCARÓN & WYGODZINSKY, 1984: 84-87; COSCARÓN, 1987: 21; 1991: 159-160; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium clavibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 31; 2004: 133135; 2004:117-118; STRIEDER & PY-DANIEL, 1999: 49; 2000: 27; 2002: 66.

Female: Wing length 2.3-2.8 mm; head blackish, palp and proboscis brown blackish, antenna base light brown; scutum and scutellum black with grayish pollinosity and hair yellowish-gray with copper tones, homogeneously arranged and not abundant. Metanotum velvet black; legs light brown with darkened areas as in Simulium inaequale; frons slightly convergent (Fig. 64 I); basal portion of cibarium as in Simulium subclavibranchium; fronto-ocular triangle wider than high (Fig. 64 J); Sc with about 10 hairs; claws with obvious subbasal tooth (Fig. 64 K); eighth sternite with about 80 setae and gonapophyses divergent; paraproct short with slight curvature distally (Fig. 64 L). Male: Wing length 2.3 mm; coloration as in Simulium inaequale but with abdominal silvery spots reaching segment IX; legs as in S. inaequale but more darkened with fore coxa blackish and also more darkened femora and tibiae; gonocoxite subtrapezoidal (Fig. 65

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G), gonostylus (Fig. 65 H) relatively shorter than in Simulium inaequale and S. subnigrum; ventral plate acuminate distally. Pupa: Cocoon with anterior edge reinforced and perpendicular to base (Fig. 66 B); weave with open threads obvious; length at base 3.2-3.5 mm, on dorsum 2.8-3.0 mm, gills 3.5-4.2 mm; gill with primary ventral branch bifurcated at half length of secondary central branch, and these latter bifurcated at half of secondary dorsal branch (Fig. 67 G ); all filaments enlarged apically (Fig. 67 H); cephalothoracic trichomes with 3-5 branches; frontoclypeus and exposed portion of thorax with tubercles in regular quantity and well darkened (Fig. 66 C). Larva (based on STRIEDER, 1998): Color yellowish-gray; cephalic apotome with negative spots; antenna longer than stem of cephalic fan; ratio of proximal, medial and distal antennal segments = 1:0.8-0.9:1.3; cephalic fan with about 35 rays; postgenal cleft subtriangular; latero-mandibular process approximately straight, not surpassing inferior rim of mandible; anal ring with 67-76 rows with 11-14 hooks each; rectal papillae with 1012 diverticula on each lobe. Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Rondônia, Santa Catarina, São Paulo. Bionomics: Simulium clavibranchium breeds in swift creeks of crystalline water flowing over aquatic vegetation or submerged tree branches. Feeding behavior of the females is unknown. Discussion: Adults of S. clavibranchium are similar to those of S. inaequale, S. subclavibranchium and S. subnigrum, although S. clavibranchium has the femora, tibiae and coxae of the anterior legs darker, the paraproct with a curved distal edge and the gonostylus with a more flattened distal edge. The subapically inflated gill filaments permit differentiation from the closest species, S. subclavibranchium, which also has enlarged gill filaments distally, but with smaller diameter and longer filaments. Another difference from this species is the proportional length of the secondary dorsal gill branch bifurcation; that in S. clavibranchium, in relation to the primary ventral branch, is more than four times the length; the gills also are more sclerotized. Simulium (Inaequalium) subclavibranchium LUTZ (Figs. 64 M, N; 65 I; 66 D, E; 67 I.) Simulium subclavibranchium LUTZ, 1910: 260-261; PINTO, 1931: 713; VULCANO, 1967: 21. Simulium (Inaequalium) subclavibranchium COSCARÓN & WYGODZINSKY, 1984: 87-90; COSCARÓN, 1987: 21; 1991: 160-162; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium subclavibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 85; 2002: 133-135; 2004:120; STRIEDER & PY-DANIEL, 1999: 57; 2000: 61; 2002: 66.

Female: Wing length 2.4-2.5 mm; coloration similar to that of Simulium inaequale with scutum pilosity from grayish to tin color; basal portion of cibarium as in figure 64 M; fronto-ocular triangle as in Simulium clavibranchium; cercus short, paraproct shorter than high with rounded edge; genital fork as in figure 64 N; eighth sternite with about 70 setae; gonapophyses scarce sclerotized on internal edge. Male: Wing length 2.5 mm; coloration as in Simulium inaequale with variation in dark color of legs; gonocoxite and gonostylus as in S. clavibranchium; ventral plate not acuminate apically (Fig. 65 I).

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Pupa: Cocoon similar to Simulium clavibranchium but with thicker weave and threads more evident; aspect as in figure 66 D, with gill much longer than body of pupa and cocoon; cocoon length on base 3.2-3.5 mm; on dorsum 2.8-3.2 mm; gill 4.8-5.5 mm; gill with lighter base than in S. clavibranchium, with secondary dorsal and ventral branches bifurcating nearer to base than in S. clavibranchium (Fig. 67 I); gill filaments slightly widened distally but gradually thinning toward apex; frontoclypeal and thoracic trichomes with 3-6 branches; tubercles abundant, about same as in S. clavibranchium, but a little lighter (Fig. 66 E). Larva: Maximum length 6.0-6.5 mm; coloration light yellowish-gray, darkened dorsolaterally in greenish-gray bands; head dark-brown; cephalic apotome darkened on basal and median area but more darkened than in Simulium inaequale; ratio of proximal, medial and distal antennal segments = 1:1.1-1.3:1.4-1.5; mandible as in Simulium diversibranchium; cephalic fan with about 40 rays; anal sclerite as in Simulium inaequale; rectal papillae with 6-9 diverticula on each lobe; anal ring with 74-86 rows of 13-15 hooks each. Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Rondônia, Santa Catarina, São Paulo. Bionomics: Simulium subclavibranchium breeds in small creeks, with clear water, on aquatic vegetation and stones. Feeding behavior of the females is unknown. Discussion: Imagoes of S. subclavibranchium are similar to those of S. inaequale, S. subnigrum and S. clavibranchium. The first two species have more pilosity and more grayish coloration on the scutum; the latter has darkened legs especially on coxa I, more elongated paraprocts and a shorter gonostylus. A great difference is shown in the pupal gill: Simulium inaequale and S. subnigrum do not have the distal expansion, and in S. clavibranchium the distal enlargement is bigger, the gills are proportionally shorter, and the bifurcation of the secondary dorsal and ventral branches are at a greater distance from the base. Also, the frontoclypeal and thoracic tubercles of S. clavibranchium are more darkened and abundant. It is difficult to separate S. subnigrum and S. subclavibranchium pupae when the former has very long gills that are slightly enlarged distally. Simulium (Inaequalium) nahimi PY-DANIEL (Figs. 65 K; 67 J.) Simulium nahimi PY-DANIEL, 1984: 125-137. Simulium (Inaequalium) nahimi COSCARÓN, 1987: 21; 1991: 164; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium nahimi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 60; STRIEDER & PYDANIEL, 1999: 53; 2000: 42; 2002: 66.

(Diagnosis based on PY-DANIEL, 1984 and STRIEDER, 1998). Female: Wing length 1.9-2.0 mm; coloration and morphology similar to that of Simulium inaequale and S. subnigrum, although scutellum in Simulium nahimi is brown yellowish. Male: General coloration and morphology similar to that of Simulium inaequale and S. subnigrum, but Simulium nahimi with gonostylus bearing 3-8 spurs positioned on distal edge (Fig. 65 K from PY-DANIEL, 1984). Pupa: Cocoon length on base 3.2-4.0 mm, on dorsum 2.1-3.2 mm; maximum length of gills 3.6-3.7 mm; primary dorsal branch shorter than primary ventral branch; secondary dorsal and median branches bifurcated at about same distance, at greater distance than bifurcation of dorsal primary branch (Fig. 67 J, from PY-DANIEL, 1984).

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Larva: Maximum length 5.0; antenna reaching apex of cephalic fan stem; ratio of proximal, medial and distal antennal segments = 1:0.9-1.03:1.2-1.5. Mandible with lateromandibular process; rectal papillae with 15-19 diverticula on each lobe; anal sclerite with small spiniform setae or filiform hair. Distribution: Brazil: Mato Grosso. Bionomics: Breeds in temporary swift springs or creeks in open areas, flowing over aquatic vegetation or rocks. Female feeding habits are unknown. Discussion: It is difficult to separate S. nahimi from similar species. The small size and gonostylus with a greater number of spurs than only one other species allows it to be readily identified. The number of diverticula on the rectal papillae is the highest for the species group and is another character that can help distinguish this species. Simulium (Inaequalium) mariavulcanoae COSCARÓN & WYGODZINSKY (Figs. 65 J; 66 F; 67 K.) Simulium (Inaequalium) mariavulcanoae COSCARÓN & WYGODZINSKY, 1984: 96; COSCARÓN, 1987: 21; 1991: 166; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium mariavulcanoae PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 55; STRIEDER & PY- DANIEL, 1999: 52; 2000: 40; 2002: 66.

Female: Wing length 2.4-2.6 mm; coloration and morphology similar to that of Simulium diversibranchium; paraproct slightly shorter is the only appreciable morphological difference. Male: Wing length 2.3-2.6 mm; similar to Simulium diversibranchium; gonostylus wider (Fig. 65 J) is the only difference. Pupa: Cocoon shape and weave structure as in Simulium diversibranchium; length at base 3.2-3.5 mm, on dorsum 3.0-3.2 mm, gills 3.2-3.6 mm; gills with short basal trunk; ventral primary branch thin and short, dorsal primary branch also is short but little thicker; secondary dorsal branch always a little thicker (Fig. 67 K); frontoclypeus and thorax with abundant and obvious tubercles; trichomes with 2-7 branches with basal portion curved (Fig. 66 F). Larva: Unknown. Distribution: Brazil: São Paulo. Bionomics: Simulium mariavulcanoae breeding sites and female feeding habits are unknown. Discussion: Adults of S. mariavulcanoae are similar to those of S. diversibranchium, but the more elongated paraproct and narrower gonostylus of this species help differentiate it. The bifurcation of the pupal gill secondary branches at a greater distance, the thicker dorsal branch and the more elongated tertiary dorsal branch, as well as the first ventral secondary branch directed downward and backward, help identify this species. Simulium (Inaequalium) diversibranchium LUTZ (Figs. 65 L; 66 G, H; 67 L; 68 G-J.) Simulium diversibranchium LUTZ, 1910: 238-239, 263; PINTO, 1931: 701; VULCANO, 1967: 10. Simulium (Inaequalium) diversibranchium COSCARÓN & WYGODZINSKY, 1984: 93-96; COSCARÓN, 1987: 21; 1991: 164-166; COSCARÓN & COSCARÓN ARIAS, 1997: 111.

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Inaequalium diversibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 36; 2004: 118; STRIEDER & PY-DANIEL, 1999: 50; 2000: 31; 2002: 66. Simulium missionum COSCARÓN, 1976b: 151-154.

Female: Wing length 2.3-2.7 mm; coloration blackish with ornamentation similar to that in Simulium inaequale species group; frons as in Simulium clavibranchium; fronto-ocular triangle about as wide as high; scutum pilosity golden to tin colored; metanotum black, velvety; legs yellowish, with darkened areas blackish and pilosity yellowish to black in accordance with position of lighting; abdomen brownish-gray with blackish tergal plates on segments II-IV; paraprocts relatively shortened, with truncate apical edge; eighth sternite with about 72 setae; gonapophyses slightly sclerotized internally and scarcely divergent. Male: Wing length 2.3-2.6 mm; color generally as in female; scutum similar to that of Simulium inaequale; abdominal segments I-II brownish, subsequent segments blackish, with 1+1 silvery spots on tergites II, V, VI; sternites grayish pollinose; gonostylus subtrapezoidal, with strong spur near inner corner (Fig. 65 L). Pupa: Cocoon with thin weave, with few evident threads, reinforced edge and slowly projected ventrally (Fig. 66 G); cocoon length on base 3.2-4.0 mm, on dorsum 2.6-3.5 mm, gills 2.8-3.5 mm; gills with 5-6 filaments emerging from two short primary branches: ventral branch thinner and bifurcated at short distance; secondary ventral branch curved downward and posteriorly; primary dorsal branch bifurcated at short distance, with secondary dorsal branch thicker; secondary dorsal and median branches bifurcated at same distance (Fig. 67 L); frontoclypeus and thorax with abundant tubercles; trichomes of frontoclypeus with 2-6 branches, thoracic trichomes with 5-8 branches and with basal trunk curved (Fig. 66 H). Larva: Maximum length 6.0-6.8 mm; aspect as in figure 68 G; coloration yellowish gray; head yellowish brown; cephalic apotome darkened in median basal area (Fig. 68 H); antenna as long as or slightly shorter than stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1:1.4; mandible teeth with latero-mandibular process; hypostomium with 4-6 setae on each side; postgenal cleft deep (Fig. 68 I); pupal gill histoblast as in figure 68 J; anal ring with about 84 rows of 12-13 hooks each; rectal papillae with about 20 diverticula. Distribution: Brazil: Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Simulium diversibranchium breeds in small, clear creeks flowing over submerged aquatic vegetation or trees branches. Female feeding habits are unknown. Discussion: Adults are difficult to separate from those of the other INAEQUALE species group. Simulium diversibranchium shows only small differences such as the paraproct being shorter and gonostylus subtrapezoidal. The asymmetrical gill of S. diversibranchium permits it to be identified. Simulium (Inaequalium) rappae PY-DANIEL & COSCARÓN (Figs. 66 I-K; 68 K, L.) Simulium rappae PY-DANIEL & COSCARÓN, 1982: 155-162. Simulium (Inaequalium) rappae COSCARÓN, 1987: 21; 1991: 167-168; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium rappae PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 77; STRIEDER & PY-DANIEL, 1999: 55; 2000: 55; 2002: 66.

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Female: Wing length 1.8-1.9 mm; scutum blackish with pilosity yellowish; with anterior light, showing median and 1+1 submedian grayish pollinose bands; with posterior illumination, grayish spots become blackish and black area becomes grayish; Sc with 9-10 hairs; eighth sternite with about 40 setae on each side; gonapophyses sclerotized on internal edge and divergent; paraproct short as in Simulium clavibranchium and genital fork as in Simulium travassosi. Male: Wing length 1.8 mm; scutum black with yellowish pilosity; with frontal illumination, median third of scutum dark; Sc without setae; length/width ratio of hind basitarsus = 5.7; gonostylus subtriangular; ventral plate similar to that of Simulium inaequale. Pupa: Cocoon with thick weave, and reinforced anterior edge (Fig. 66 I); length at base 2.8-3.6 mm, on dorsum 2.4-3.0 mm, gills 1.6-2.1 mm; gills light brown, with 6 thick branches emerging from 3 long primary branches bifurcated at about same distance; frontoclypeus and thorax with scarce tubercles (Fig. 66 J); cephalic trichomes of 2-3 long branches and thoracic trichomes with 3-5 branches (Fig. 66 K). Larva: Maximum length 4.2-5.0 mm; general aspect as in figure 68 K; postgenal cleft deep, with postgenal bridge shorter than hypostomium length; antenna as in figure 68 L, ratio of proximal, medial and distal antennal segments = 1:0.7-1:1.2-1.7; anal ring with 64 rows with 12-15 hooks each; rectal papillae with 13-17 diverticula on each lobe. Distribution: Brazil: Goiás, Minas Gerais, Rio de Janeiro, São Paulo. Bionomics: Simulium rappae breeds in clear creeks, on aquatic vegetation in fast current and in open areas. Feeding habits of the females are unknown. Discussion: Simulium rappae adults are very similar to those of the INAEQUALE species group, with very few differences seen with different directions of illumination. The pupal gill of S. rappae is unique and allows the species to be identified. Simulium (Inaequalium) travassosi D’ANDRETTA & D’ANDRETTA (Figs. 64 O, P; 65 M, O; 66 L-N; 68 M.) Simulium travassosi D’ANDRETTA & D’ANDRETTA, 1947: 161-179; VULCANO, 1967: 22. Simulium (Inaequalium) travassosi COSCARÓN & WYGODZINSKY, 1984: 96-98; COSCARÓN, 1987: 22; 1991: 168-169; STRIEDER et al., 1992: 59; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium travassosi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 97; 2004: 121; STRIEDER & PY-DANIEL, 1999: 59; 2000: 71; 2002: 66. Simulium botulibranchium LUTZ, 1910: 256; PINTO, 1931: 675.

Female: Wing length 2.2 mm; scutum blackish with golden dispersed hair; abdomen blackish, tergite I bright silvery with yellowish setae on posterior edge; tergite II blackish brown, bright silvery laterally; tergites III and V black matte, remaining tergites with cereous sheen; legs brown with darkened area blackish; fronto-ocular triangle deep; genital fork with elongate stem (Fig. 64 O); paraproct slightly higher than long with convex edge (Fig. 64 P); gonapophysis as in Simulium inaequale. Male: Wing length 2.1 mm; coloration as in Simulium inaequale; Sc with 3-4 hairs; gonocoxite about two times longer than gonostylus (Fig. 65 M); gonostylus short, subtrapezoidal, distally elongated, with strong apical spur (Fig. 65 N); ventral plate wider than long, with distal edge subacute (Fig. 65 O).

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Pupa: Cocoon with thin weave, threads evident, with straight reinforced anterior edge (Figs. 66 L, M); length at base 3.2-3.6 mm, on dorsum 2.9-3.2 mm, gills 1.5-1.8 mm; gills each of three branches with microscopical granulosity; dorsal branch largest, showing shape variation, with 1 or 2 small apical processes (Fig. 66 N); frontoclypeus and exposed portion of thorax with abundant tubercles; cephalic and thoracic trichomes with 2-4 branches. Larva: Maximum length 5.0-5.2 mm; color yellowish brown with tones greenish-gray dorso-laterally on abdomen; cephalic apotome with diffuse ornamentation (Fig. 68 M); antennal apex surpassing cephalic fan stem; ratio of proximal, medial and distal antennal segments = 1:0.9:1.6; mandible with latero-mandibular process surpassing inferior edge; cephalic fan with about 35 rays; hypostomium with 4 setae on each side; anal ring with about 78 rows with 15 hooks each; anal sclerite dorsally with 3-4 rows of scales with 2.5 spines each; rectal papillae with 3-7 diverticula on each lobe. Distribution: Argentina: Misiones; Brazil: Goiás, Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Simulium travassosi breeds on aquatic vegetation in small fast-flowing creeks. Female feeding habits are unknown. Discussion: Simulium travassosi adults are very difficult to separate from those of similar species such as S. inaequale, S. subnigrum and S. subclavibranchium, showing differences only in the genital fork and gonostylus. However, the characteristic pupal gill morphology allows it to be identified. BOTULIBRANCHIUM species group Simulium (Inaequalium) botulibranchium LUTZ (Figs. 69 A-I; 70 A-F.) Simulium (Inaequalium) botulibranchium LUTZ, 1910: 256-257, 263; D’ANDRETTA & D’ANDRETTA, 1947: 145; VULCANO, 1967: 7; 19; COSCARÓN, 1980: 293-295. Simulium (Inaequalium) botulibranchium COSCARÓN, 1987: 22; 1991: 170-171; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Simulium tulibranchium LUTZ, 1922: 92 (error). Inaequalium botulibranchium PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 24; 2004: 117; STRIEDER & PY-DANIEL, 1999: 48; 2000: 22; 2002: 66.

Female: Wing length 2.1-2.6 mm; scutum blackish with abundant pilosity tin-colored to golden; scutellum dark brown; metanotum and pleurae blackish with nacreous pollinosity; abdomen black, tergite I laterally with grayish pollinosity; tergites II-III black matte, showing 1+1 postero-lateral whitish spots on tergite II; remaining tergites with cereous sheen; frons convergent and relatively narrow (Fig. 69 A); fronto-ocular triangle deep (Fig. 69 B); cibarium with basal portion elevated and with thick spines (Fig. 69 C); Sc with about 20 setae; paraprocts subtriangular, about two times higher than wide and with basal posterior lobe (Fig. 69 D); gonapophysis divergent, with edge slightly sclerotized; genital fork with elongated stem, longer than two posterior branches combined (Fig. 69 E). Male: Wing length 2.1-2.4 mm; general coloration as in female, but more darkened, especially legs, with more abundant black hair; scutum black, with some directions of

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B A

J

C

D

K

G

Q P

F

T

L N

M

O

S H E I R

Figure 69. A-I: S. (Inaequalium) botulibranchium; A-E: Female; A: frons; B: fronto-ocular triangle; C: basal portion of cibarium; D: cercus and paraproct; E: genital fork; F-I: Male; F: scutum and metanotum; G: abdomen, lateral view; H: ventral plate; I: gonostylus. J-S: S. (I.) souzalopesi; J-Q: Female; J: frons; K: basal portion of cibarium; L-N: fore, mid and hind leg, respectively; O: claw; P: eighth sternite and gonapophyses; Q: cercus and paraproct; R-S: Male; R: gonocoxite and gonostylus; S: ventral plate. T: S. (I.) petropoliense, eighth sternite and gonapophyses.

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illumination, silvery spot of variable size on antero-lateral area (Fig. 69 F); abdomen blackish with light silvery spots as shown in figure 69 G; ventral plate about as long as wide, slightly acuminate distally (Fig. 69 H).Gonostylus elongated about as long as gonocoxite, with external lateral curvature and apical spur (Fig. 69 I). Pupa: Cocoon of thin weave, sparse threads, with straight and reinforced edge (Figs. 70 A, B); length at base 2.6-3.0 mm, on dorsum 2.1-2.6 mm, gill 1.3-1.5 mm; gills very peculiar, each of 2 thick branches: one antero-lateral and one ventral; the first has 2 short basal branches and 4 on distal half, the ventral has several curvatures at different levels and no secondary branches (Fig. 70 C); gill integument with small spiculae; frontoclypeus and thorax with scarce or no tubercles; cephalic trichomes bifid, thoracic trichomes with 2-6 branches; abdominal chaetotaxy as Simulium inaequale. Larva: Maximum length 6.0-6.5 mm; color grayish-brown with greenish-gray dorsolaterally on abdominal segments, aspect as in figure 70 D; antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal antennal segments = 1:0.8-0.9:1.2-1.6; lateromandibular process sinuous and surpassing inferior dental edge; cephalic apotome scarcely ornamented, with negative spots; cephalic fan with 40-46 rays; hypostomium with 5-7 lateral setae (Fig. 70 E); postgenal cleft subtriangular, postgenal bridge about as long as hypostomium length (Fig. 70 F); anal sclerite with single hairs; anal ring with 80-92 rows of 14-15 hooks each; rectal papillae with 2-9 diverticula on each lobe. Distribution: Argentina: Misiones; Brazil: Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Simulium botulibranchium breeds in the fast flow of small, clear creeks with aquatic vegetation. Feeding habits of females are unknown. Discussion: Adults of Simulium (Inaequalium) are very similar externally. Small differences in the shape of the paraproct, gonostylus and ventral plate help differentiate them. The peculiar pupal gill is different from that of any other known Neotropical simuliid. Simulium (Inaequalium) souzalopesi COSCARÓN (Figs. 69 J-S; 70 G-O.) Simulium souzalopesi COSCARÓN, 1980: 296-298. Simulium (Inaequalium) souzalopesi COSCARÓN, 1987: 23; 1991: 171-172; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium souzalopesi PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 81; STRIEDER & PY- DANIEL, 1999: 56; 2000: 58; 2002: 66.

Female: Wing length 2.3-2.5 mm; coloration and morphology similar to that of Simulium botulibranchium; frons a little wider than in Simulium botulibranchium (Fig. 69 J); Sc with about 10 hairs; cibarium with abundant spines, with small median depression at apex (Fig. 69 K); aspect of fore mid and hind legs as in figures 69 L, M, N; claw with welldeveloped subbasal tooth (Fig. 69 O); eighth sternite with about 60 setae; gonapophyses divergent, slightly sclerotized on internal margin (Fig. 69 P); paraprocts subtriangular, about as high as wide and with small protuberance at base (Fig. 69 Q). Male: Wing length 2.2 mm; coloration and morphology, including genitalia, similar to that of Simulium botulibranchium; gonocoxite subtrapezoidal with same length as

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C

A

B

I

G H R J P

Q

M S

E K O D

L

N

F

Figure 70. A-F: S. (Inaequalium) botulibranchium; A-C: Pupa; A: lateral view; B: dorsal view; C: gill; DF: Larva; D: lateral view; E: hypostomium; F: hypostomium and postgenal area. G-O: S. (I.) souzalopesi; G-J: Pupa; G: lateral view; H: dorsal view; I: gill; J: frontoclypeus; K-O: Larva; K: antenna; L: teeth of mandible; M: head, dorsal view; N: hypostomium and postgenal area; O: anal sclerite. P-S: S. (I.) petropoliensye; P-S: Pupa; P: lateral view; Q: dorsal view; R: gill; S: frontoclypeus.

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gonostylus; gonostylus elongated, with ventral subapical curvature externo-laterally positioned (Fig. 69 R); ventral plate relatively narrow basally (Fig. 69 S). Pupa: Cocoon with thin weave and evident threads, slipper shaped with dorsal projection (Figs. 70 G-H); length on dorsum 3.0-3.4 mm; on base 2.8-3.0 mm; gills 2.0-2.1 mm; gills asymmetrical, relatively thickened, each with 6 terminal branches; two primary branches emerging from short basal trunk; dorsal primary branch bifurcated at short distance dorsolaterally; another dorsal secondary branch with strong curvature and bifurcated (Fig. 70 I); gill integument process with small-spaced tubercles; integument of head and exposed portion of thorax with scarce tubercles (Fig. 70 J); frontoclypeal trichomes simple, thoracic trichomes single or bifid; abdominal chaetotaxy as in Simulium subnigrum. Larva: Maximum length 5.5-6.2 mm; coloration light grayish-brown with greenishbrown on thorax dorsally and abdomen dorso-laterally; head light brown, cephalic apotome with diffuse to positive spots, darkened on median and basal area (Fig. 70 M); cervical sclerites well developed; hypostomium with 5-6 setae on each side; postgenal cleft not deep, with length about half of hypostomium (Fig. 70 N); ratio of proximal, medial and distal antennal segments = 1:0.8:1.3 (Fig, 70 K); teeth of mandible as figure 70 L; cephalic fan with about 35 rays; anal ring with about 75 rows with 13 hooks each; anal sclerite with abundant simple hairs, without scales (Fig. 70 O). Distribution: Brazil: Rio de Janeiro. Bionomics: Simulium souzalopesi was collected on aquatic vegetation and stones in small currents about 0.30-0.50 m wide and 0.05-0.10 m deep, with crystalline water, living with S. botulibranchium and S. petropoliense. Feeding habits of the females are unknown. Discussion: Simulium souzalopesi adults are similar to those of S. botulibranchium, showing small differences such as a narrow frons, the fronto-ocular triangle more acute and the cibarium elevated medially. Differences are obvious in the pupal gill, which has a unique morphology not seen in other Neotropical species. Simulium (Inaequalium) petropoliense COSCARÓN (Figs. 69 T; 70 P-S.) Simulium petropoliense COSCARÓN, 1980: 298-301. Simulium (Inaequalium) petropoliense COSCARÓN, 1987: 23; 1991: 172-173; COSCARÓN & COSCARÓN ARIAS, 1997: 111. Inaequalium petropoliense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 1998: 72; STRIEDER & PY- DANIEL, 1999: 55; 2000: 52; 2002: 66.

Female: Wing length 2.6 mm; coloration and external morphology as in Simulium botulibranchium; Sc with about 16 hairs; eighth sternite with about 25 setae on each side; gonapophyses with internal borders separated and parallel (Fig. 69 T); paraprocts as in Simulium souzalopesi. Male: Unknown. Pupa: Cocoon with compact weave and very close threads, with reinforced anterior edge and without dorsal anterior projection (Figs.70 P, Q); length on dorsum 2.7 mm; on base 3.1 mm; gills 1.1 mm; gill asymmetrical with 2 wide primary branches; dorsal branch with 4 short thin terminal branches and ventral branch with several curves, undivided distally (Fig. 70 R); gill integument with abundant thin microscopical projections like hair;

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head and exposed portion of thorax with abundant tubercles (Fig. 70 S); frontoclypeal trichomes with 2-4 branches, thoracic trichomes with 4-6 branches; thorax with dark brown tones not uniformly distributed (Fig. 70 R). Larva: Unknown. Distribution: Brazil: Rio de Janeiro. Bionomics: Simulium petropoliense is similar to S. souzalopesi, which was collected in the same place. Discussion: The peculiar pupal gill allows S. petropoliense to be differentiated from the other Neotropical species. Species inquirendae Simulium parimaensis RAMÍREZ PÉREZ, YARZÁBAL, TAKAOKA, TADA & RAMÍREZ, 1986: 61. Simulium (Inaequalium) parimaensis COSCARÓN, 1991: 155. Inaequalium parimaensis STRIEDER, 1998: 71; STRIEDER & PY-DANIEL, 1999: 46; 2000: 51. Simulium (Psaroniocompsa) parimaensis CROSSKEY & HOWARD, 1997: 58; 2004: 52.

The female description of S. parimaensis corresponds to any Simulium (Inaequalium) species. (As explained before, it is difficult to establish the species identity in this subgenus with only adult material). The pupal gill has 18-20 branches, and is different from known Simulium (Inaequalium) species and closer to Simulium (Ectemnaspis) species. Based on the pupae, it is similar to S. parimaensis from Roraima (Brazil), which is close to the type locality in the Federal Territory of Amazonia (Venezuela). STRIEDER (1998) and STRIEDER & PY-DANIEL (2000) verify that females obtained by dissection correspond to the genus Ectemnaspis. The authors believe a mistake was made when the holotype was mounted on the slide. Based on this assertion, and pending new material reared from the type locality, we maintain S. parimaensis as species inquirenda. Distribution: Venezuela: Amazonas. Simulium luribayae SMART Simulium luribayae SMART, 1944: 132 n. nom; for Trichodagmia angustifrons ENDERLEIN, 1934b: 288. Simulium (Inaequalium) luribayae COSCARÓN & WYGODZINSKY, 1984: 22; 1991: 173.

The type deposited at the Berlin Museum is externally similar to S. subnigrum but with more greenish scutum pilosity. Without knowledge of the pupa, it is impossible to define this species. Simulium pseudoexiguum NUNES DE MELLO & DE ALMEIDA Simulium pseudoexiguum NUNES DE MELLO & DE ALMEIDA, 1974: 65-68. Simulium pseudoexiguum RAMIREZ PÉREZ et al., 1986: 63. Simulium (Inaequalium) pseudoexiguum COSCARÓN, 1987: 22.

As this species was described only from a female, it is impossible to separate it from other species of the subgenus. We consider S. pseudoexiguum as species inquirenda.

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Simulium (Psaroniocompsa) ENDERLEIN (Figs. 71-75; 142.) Type species: Psaroniocompsa opalinifrons ENDERLEIN, 1934a: 192 (= S. incrustatum LUTZ, 1910: 243). Syn. Pliodasina ENDERLEIN, 1936: 124 (Type species Pliodasina guttata ENDERLEIN, 1936: 124).

Female: Medium-sized to small species. General coloration black; frons and clypeus grayish silvery pollinose or with nacreous sheen with greenish-blue tones. Scutum velvety black with silvery lateral and posterior margins, and generally with 1+1 silvery to blackish, according to direction of light, short cuneiform anterior submedian spots that can be continuous posteriorly as thin silvery stripe, sometimes reaching posterior border. Scutum with homogeneous pilosity, gray to bronze; sometimes hairs grouped, simulating scales with golden to violaceous greenish sheen. Metanotum black with velvety gray pollinosity. Abdomen black with tergites I-V with velvety tones, sometimes showing 1+1 small sublateral silvery spots on tergite II; tergites VI-IX with cereous sheen. Frons relatively wide; fronto-ocular triangle scarcely developed. Basal portion of cibarium with sclerotized rim and teeth arranged in several rows reaching cornuae base at middle, glabrous and depressed. Basal portion of R generally without hairs; Sc with or without hairs. Legs with single, spatulate hair. Calcipala well developed. Claws generally with small subbasal tooth. Fore tarsus relatively wide. Genital fork with elongated stem; gonapophyses with subparallel internal border, scarcely sclerotized; eighth sternite with about 40 setae; paraproct short with rounded border. Male: Scutum velvety black with hairs as in female. Pronotal areas grayish dark with silvery lateral and posterior margins; with anterior light source, 1+1 silvery submedian small cuneiform spots. Abdomen black with 1+1 silvery spots on tergites II, VI and VII and another small spot on lateral border of VIII. Gonocoxite subtrapezoidal, about as long as wide; gonostylus shorter than gonocoxite, subquadrate or subtrapezoidal to subtriangular, with great submedian to apical spur. Ventral plate subtriangular, wider than long. Pupa: Cocoon with thick or thin weave, with anterior border from straight to dorsally projected. Gill generally with 6 filaments, exceptionally with 4 or 2 thicker ones. Frontoclypeus and exposed portion of thorax with platelets from abundant to reduced or absent. Head trichomes with one frontal pair and 2 facial pairs; thorax with 5 pairs, generally bifid but can have 5-12 branches in some species. Abdomen with spine combs on distal tergites. Larva: Body without hairs or scales or these very short to well developed on abdomen. Cephalic apotome generally with positive spots, discontinuous, arranged in median and 1+1 sublateral groups. Cephalic capsule with short trichomes, generally bifid. Postgenal cleft deep, with postgenal bridge shorter than half of hypostomial length. Antenna surpassing apex of cephalic fan stem, with median segment generally shorter than fore and hind segments. Mandible with 2 marginal teeth and generally with 1-2 latero-mandibular teeth surpassing level of tooth apex. Hypostomium median tooth about as high as lateral teeth. Cephalic fan with about 35 rays. Anal sclerite generally with scarce simple hairs. Anal ring with about 65 rows with about 12 hooks each. Ninth segment with 1 pair of ventral tubercles. Rectal papillae with 3-8 diverticula on each lobe. Distribution: From Argentina to Panama (Fig. 142).

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The subgenus S. (Psaroniocompsa) includes 18 species that can be arranged in 2 species groups: • INCRUSTATUM: with S. anamariae, S. angrense, S. auripellitum, S. auristriatum, S. bonaerense, S. brevifurcatum, S. fuliginis, S. incrustatum, S. jujuyense, S. limbatum, S. minuanum, S. schmidtmummi and S. stellatum. • SIOLII: with S. damascene, S. guaporense, S. lourencoi, S. siolii, S. tergospinosum. The first group has species with the scutum scarcely silvery ornamented, from absent or reduced to 1+1 anterior submedian spots that can be continued posteriorly by thin stripes not joining with the posterior grayish area; and the male gonostylus smooth without large hairs or scales. The second group is differentiated by having a scutum with wider 1+1 submedian silvery stripes; male gonostylus subtriangular with a dorsal longitudinal carina, and larvae with dorsal protuberances on the abdominal tergites and abundant and large hairs and scales on the integument. Species arrangement in the text was done considering those with more generalized morphology first, showing scutal hairs homogeneously distributed, scarce silvery spots or longitudinal stripes and pollinose frons without shiny iridescent pigmentation. CROSSKEY & HOWARD (1997) include in this subgenus, the species in the AMAZONICUM, AURISTRIATUM, INCRUSTATUM, QUADRIFIDUM, and SIOLII groups. The INCRUSTATUM, AURISTRIATUM and SIOLII groups treated here correspond to the subgenus S. (Psaroniocompsa), AMAZONICUM to S. (Cerqueirellum) PY-DANIEL and QUADRIFIDUM to S. (Coscaroniellum) PY-DANIEL. KEYS

TO

SIMULIUM (PSARONIOCOMPSA)

SPECIES

Key to larvae not included. Females and males 1 – 2(1) – 3(2) –

Scutum hairs grouped, scale like (Fig. 71 K) frequently arranged in longitudinal rows, at least on median and submedian rows (Figs. 71 L, O, Q); frons and clypeus silvery with greenish-blue to violaceous sheen ............................................................ 2 Scutum hairs homogeneously distributed, not grouped or scale like; frons and clypeus with grayish pollinosity, with velvety sheen ................................................... 7 Small species, wing length about 1.8 mm; scutum without silvery submedian cuneiform spots or longitudinal bands ............................................... Simulium fuliginis Medium-sized species, wing length >2.0 mm; scutum with or without silvery spots, if wing length <2.0 mm, scutum with 1+1 submedian longitudinal bands ................................................................................................................................................ 3 Scutum without silvery anterior submedian cuneiform spots; hairs grouped like isolated scales arranged in 1 median and 2 submedian rows (Fig. 71 Q) .............. 4 Scutum with 1+1 silvery anterior submedian cuneiform spots; sometimes prolonged posteriorly, reaching grayish posterior area (Figs. 71 L, O); hairs grouped, appearing like scales sometimes not isolated nor arranged in welldefined rows ........................................................................................................................ 6

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4(3) – 5(4)

Scutum hair bright golden ................................ Simulium anamariae, Simulium stellatum Scutum hair shiny greenish violet or golden-violet .................................................... 5 Female: frons wider than opposite option with parallel sides; fronto-ocular triangle scarcely deep (Fig. 71 R). Male: gonostylus spur very close to inner angle (Fig. 72 H) .......................................................................................... Simulium auristriatum – Female: frons relatively narrow with convergent sides (Fig. 71 S); fronto-ocular triangle relatively deep (Fig. 71 T). Male: gonostylus spur almost halfway to edge ............................................................................................................ Simulium brevifurcatum 6(3) Scutum with 1+1 silvery anterior cuneiform spots not prolonged posteriorly as stripes (Fig. 71 L), if prolonged, they do not surpass anterior 1/4 of scutum; scutum hairs golden-bronze or golden-green ............. Simulium incrustatum, Simulium schmidtmummi – Scutum with 1+1 silvery cuneiform spots prolonged posteriorly as grayish pollinose stripe that reaches posterior grayish area (Fig. 71 O); scutum hairs pale greenish gray ..........................................................................................Simulium limbatum 7(1) Scutum without 1+1 silvery submedian longitudinal stripes; pilosity very abundant with silky appearance. Male gonostylus subtrapezoidal (Figs. 72 A, F) ................................................................................................................................................ 9 – Scutum with 1+1 wide silvery submedian longitudinal bands that extend to posterior grayish area in females or can be shorter and not surpass half length of scutum in males (Figs. 72 J, K); pilosity scarce, without silky appearance; male gonostylus subtriangular with longitudinal carina (Figs. 72 L, M). Female ................................ Simulium siolii, Simulium tergospinosum, Simulium damascenoi Male ....................................................................................................................................... 8 8(7) Scutum with 1+1 silvery submedian, cuneiform spots (Fig. 72 K) .......................... ..................................................................................Simulium siolii, Simulium tergospinosum – Scutum with 1+1 silvery submedian, subtrapezoidal spots (Fig. 72 J) ..................... ................................................................................................................ Simulium damascenoi 9(8) Large species, wing length: female 2.5-2.8 mm, male 2.1-2.6 mm; scutum pilosity very abundant hiding anterior silvery cuneiform spots; scutum hairs from pale greenish yellow to bronze .................................................................... Simulium angrense – Medium-sized species, wing length: female 1.7-2.5 mm, male 1.7-2.4 mm ......... 10 10(9) Claws without subbasal tooth (Fig. 71 I) ....................................... Simulium minuanum – Claws with subbasal tooth .............................................................................................. 11 11(10) Scutum pilosity predominantly bronze to yellowish gray-green; wing length: female 1.7-2.1 mm, male 1.6-2.0 mm ........................................... Simulium auripellitum – Scutum pilosity predominant grayish; wing length: female 2.1-2.5 mm, male 1.92.4 mm ...................................................................Simulium jujuyense, Simulium bonaerense Pupae 1 – 2(1) – 3(2)

Gill with 2 thick subcylindrical branches (Fig. 73 J) ............... Simulium auristriatum Gill with 4-6 thin filamentous branches ........................................................................ 2 Gill with 4 branches (Figs. 73 K, 74 H) ........................................................................ 3 Gill with 6 branches (Figs. 74 B-G) ............................................................................... 8 Cocoon with well-developed dorsal anterior projection (Fig. 73 K) ...................... 4

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– 4(3) – 5(3) – 6(5) – 7(5) – 8(2) – 9(8) – 10(8) – 11(10) – 12(11) – 13(12) – 14(12) – 15(14) –

Cocoon without dorsal anterior projection (Fig. 73 N) or very short (Figs. 73 L, M) ................................................................................................................................................ 5 Cephalic and thoracic trichomes with 2 branches ..........................Simulium stellatum Cephalic trichomes with 1-4 branches, thoracic with 4-12 branches ....................... .......................................................................................................... Simulium schmidtmummi Gill shorter than cocoon length (Figs. 73 M, N); primary branches bifurcating close to base (Fig. 74 I), frontoclypeus without tubercles ........................................ 6 Gill longer than cocoon length (Fig. 73 L); primary branches bifurcating relatively far from base, frontoclypeus with tubercles ................................................................. 7 Cocoon without dorsal anterior projection and with very short ventral projection (Fig. 73 M) ........................................................................................... Simulium damascenoi Cocoon with short dorsal and relatively long ventral projection (Fig. 73 N) ......... ............................................................................................................ Simulium tergospinosum With sclerotized areas on abdominal membrane .................................. Simulium siolii Without sclerotized areas on abdominal membrane ...................... Simulium lourencoi Cocoon with dorsal anterior projection (Figs. 73 H, J) ............................................. 9 Cocoon without dorsal anterior projection (Figs. 73 A-G) .................................... 10 Cocoon and gills expanded laterally; anterior dorsal projection relatively short (Fig. 73 I) ......................................................................................... Simulium brevifurcatum Cocoon and gills not expanded laterally; anterior dorsal projection very elongated and bifurcated, protecting gills laterally (Fig. 73 H) .................... Simulium anamariae Cocoon with open weave, with threads very evident; frontoclypeal and thoracic trichomes with 3-4 branches ............................................................. Simulium guaporense Cocoon with compact weave, with threads not very evident; frontoclypeal and thoracic trichomes with 2 branches (Fig. 74 J) .......................................................... 11 Gill bifurcation of primary ventral and secondary branches very close to base (Fig. 74 C) ............................................................................................. Simulium bonaerense Gill bifurcation of primary ventral and secondary branches not very close to base (Figs. 74 B, D, E-G) ......................................................................................................... 12 Pupa large; gill length 3.1-4.5 mm; frontoclypeus and thorax with abundant (Fig. 73 P) to moderately abundant tubercles (Fig. 73 Q) ................................................ 13 Pupa medium sized; gill length 2.1-4.0 mm, frontoclypeus and thorax with sparse tubercles (Fig. 73 O) ........................................................................................................ 14 Frontoclypeus and thorax with abundant tubercles and trichomes moderately long (Fig. 73 P) ................................................................................................ Simulium angrense Frontoclypeus and thorax with relatively abundant tubercles and trichomes proportionately longer (Fig. 73 Q) ................................................ Simulium incrustatum Primary ventral and secondary gill branches bifurcated relatively close to base (Fig. 74 F) ...............................................................................................Simulium limbatum Primary ventral and secondary gill branches bifurcated farther from base (near two times than the opposite option ) (Figs. 74 D, E, G) ......................................... 15 Gill in profile approximately straight (Fig. 73 C) ............................ Simulium jujuyense Gill in profile with slight median ventral curvature (Figs. 73 A, B) ...................... 16

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16(15) Secondary gill branches with bifurcation surpassing 1/3 of gill length (Fig. 73 A) ...............................................................................................................Simulium auripellitum – Secondary gill branches with bifurcation surpassing 1/4 of gill length (Fig. 73 B) ................................................................................................................. Simulium minuanum CLAVES

DE ESPECIES DE

SIMULIUM (PSARONIOCOMPSA)

Clave para larvas no incluida. Hembras y machos 1

– 2(1) – 3(2) –

4(3) – 5(4) – 6(3)

–

Pelos del escudo agrupados con aspecto de escamas (Fig. 71 K) frecuentemente ordenados en hileras longitudinales, por lo menos en las hileras medianas y submedianas (Figs. 71 L, O, Q); frente y clípeo plateado con brillo verdoso azulado a violáceo .............................................................................................................. 2 Pelos del escudo distribuidos homogeneamente no agrupados con aspecto de escamas, frente y clípeo con polinosidad grisácea, con brillo aterciopelado ........ 7 Especies pequeñas, longitud ala 1,8 mm; escudo sin manchas cuneiformes o bandas longitudinales plateadas ........................................................... Simulium fuliginis Especies medianamente grandes; longitud del ala >2,0 mm; escudo con o sin manchas plateadas, si la longitud de la ala <2,0 mm, muestran 1+1 bandas medianas longitudinales .................................................................................................... 3 Escudo sin las manchas plateadas anteriores submedianas cuneiformes; pelos agrupados como escamas aisladas dispuestas en una hilera mediana y dos submedianas (Fig. 71 Q) ................................................................................................... 4 Escudo con 1+1 manchas plateadas cuneiformes anteriores submedianas, a veces prolongadas posteriormente y llegando al área grisácea posterior (Figs. 71 L, O); pelos agrupados aparentando escamas a veces no bien aisladas y no bien ordenadas en hileras .......................................................................................................... 6 Pelos del escudo dorado brillante ................... Simulium anamariae, Simulium stellatum Pelos del escudo con brillo verde violáceo o dorado violáceo ................................ 5 Hembra: frente de lados paralelos; triángulo frontoocular escasamente profundo (Fig. 71 R). Macho: espolón del gonostilo muy próximo al ángulo interno (Fig. 72 H) ......................................................................................................... Simulium auristriatum Hembra: frente de lados convergentes (Fig. 71 S); triángulo frontoocular relativamente profundo (Fig. 71 T). Macho: espolón del gonostilo próximo a la distancia mediana del borde ........................................................ Simulium brevifurcatum Escudo con 1+1 manchas plateadas cuneiformes no prolongadas posteiormente (Fig. 71 L), si estuvieran prolongadas no sobrepasan el 1/4 anterior del escudo; pelos del escudo con coloración dorado bronceado a dorado verdoso ................... .................................................................... Simulium incrustatum, Simulium schmidtmummi Escudo con 1+1 manchas plateadas prolongadas posteriormente por una banda grisácea polinosa que llega al área grisácea posterior (Fig. 71 O); pelos del escudo de coloración gris verdoso pálido ......................................................Simulium limbatum

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7(1)

Escudo sin 1+1 bandas plateadas longitudinales submedianas; pilosidad muy abundante con apariencia sedosa. Macho: gonostilo subtrapezoidal (Figs. 72 A, F) ................................................................................................................................................ 9 – Escudo con 1+1 bandas plateadas longitudinales submedianas anchas que llegan al borde posterior grisáceo en hembras o pueden ser mas cortas y no sobrepasar la mitad del largo del escudo en machos (Figs. 72 J, K); pilosidad escasa y sin apariencia sedosa. Macho: con gonostilo subtriangular, con una carena longitudinal (Figs. 72 L, M). Hembra .............................. Simulium siolii, Simulium tergospinosum, Simulium damascenoi Macho ................................................................................................................................... 8 8(7) Escudo con 1+1 manchas plateadas submedianas cuneiformes (Fig. 72 K) .......... ..................................................................................Simulium siolii, Simulium tergospinosum – Escudo con 1+1 manchas plateadas submedianas subtrapezoidales (Fig. 72 J) .... ................................................................................................................ Simulium damascenoi 9(7) Especies grandes, longitud ala: hembra 2,5-2,8 mm; macho 2,1-2,6 mm; pilosidad del escudo muy abundante, impidiendo ver las manchas plateadas cuneiformes anteriores; pelos del escudo desde amarillo verdoso pálido a bronceado ............... .................................................................................................................... Simulium angrense – Especies medianas, longitud ala: hembra 1,7-2,5 mm; macho: 1,7-2,4 mm ........ 10 10(9) Uñas sin diente subbasal (Fig. 71 I) ................................................ Simulium minuanum – Uñas con diente subbasal ............................................................................................... 11 11(10) Pilosidad del escudo predominantemente bronceado a amarillento gris-verdoso; longitud ala: hembra 1,7-2,1 mm. Macho 1,6-2,0 mm .............. Simulium auripellitum – Pilosidad del escudo predominantemente grisácea; longitud ala: hembra 2,1-2,5 mm; macho 1,9-2,4 mm ..................................... Simulium jujuyense, Simulium bonaerense Pupas 1 – 2(1) – 3(2) – 4(3) – 5(3) – 6(5)

Branquia con 2 ramas gruesas, subcilíndricas (Fig. 73 J) ......... Simulium auristriatum Branquia con 4-6 ramas finas, filamentosas ................................................................. 2 Branquias con 4 ramas (Figs. 73 K, 74 H) ................................................................... 3 Branquias con 6 ramas (Figs. 74 B-G) ........................................................................... 8 Capullo con bien desarrollada prolongación dorsal anterior (Fig. 73 K) .............. 4 Capullo sin prolongación dorsal anterior (Fig. 73 N) o muy corta (Figs. 73 L, M) ................................................................................................................................................ 5 Tricomas cefálicos y torácicos con 2 ramas .....................................Simulium stellatum Tricomas cefálicos con 1-4 ramas, torácicos con 4-12 tricomas ..................................... .......................................................................................................... Simulium schmidtmummi Branquias más cortas que la longitud del capullo (Figs. 73 M, N); bifurcación de ramas primarias próxima a la base (Fig. 74 I); frontoclípeo sin tubérculos .......... 6 Branquia más larga que el largo del capullo (Fig. 73 L); bifurcación de ramas primarias relativamente lejos de la base, frontoclípeo con tubérculos ................... 7 Capullo sin prolongación dorsal anterior y con muy corta prolongación ventral (Fig. 73 M) ........................................................................................... Simulium damascenoi

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Capullo con corta prolongación dorsal anterior y relativamente larga la prolongación ventral (Fig. 73) ..................................................... Simulium tergospinosum Con áreas esclerotizadas sobre las membranas del abdomen ............ Simulium siolii Sin áreas esclerotizadas sobre las membranas del abdomen ........ Simulium lourencoi Capullo con prolongación anterior dorsal (Figs. 73 H, J) ......................................... 9 Capullo sin prolongación anterior dorsal (Figs. 73 A-G) ........................................ 10 Capullo y branquias expandidas lateralmente; prolongación anterior dorsal relativamente corta (Fig. 73 I) ..................................................... Simulium brevifurcatum Capullo y branquias no expandidas lateralmente; prolongación anterior dorsal muy alargada y bifurcada protegiendo las branquias lateralmente (Fig. 73 H) ....... ................................................................................................................. Simulium anamariae Capullo con tejido abierto, con las fibras bien evidentes; tricomas del frontoclípeo y tórax con 3-4 ramas ......................................................................... Simulium guaporense Capullo con tejido compacto con fibras no muy evidentes; tricomas del frontoclípeo y tórax con 2 ramas (Fig. 74 J) .............................................................. 11 Bifurcación de la rama primaria ventral y ramas secundarias de la branquia muy próximas a la base (Fig. 74 C) .......................................................... Simulium bonaerense Bifurcación de la rama primaria ventral y ramas secundarias de la branquia no muy próximas a la branquia (Figs. 74 B, D, E, G) ............................................................. 12 Pupa grande; longitud branquia 3,1-4,5 mm; frontoclípeo y tórax con abundante (Fig. 73 P) a medianamente abundantes tubérculos (Fig. 73 Q) ............................ 13 Pupa medianamente grande; longitud branquia 2,1-4,0 mm; frontoclípeo y tórax con escasos tubérculos (Fig. 73 O) .............................................................................. 14 Frontoclípeo y tórax con abundantes tubérculos y tricomas moderadamente largos (Fig. 73 P) .................................................................................... Simulium angrense Frontoclípeo y tórax con abundantes tubérculos y tricomas proporcionalmente más largos (Fig. 73 Q) ...................................................................... Simulium incrustatum Rama primaria ventral y secundarias de la branquia bifurcadas relativamente próximas a la base (Fig. 74 F) .............................................................Simulium limbatum Rama primaria ventral y secundarias de las branquias bifurcadas aproximadamente al doble de distancia de la base que en la opción opuesta (Figs. 74 D, E, G) ....... 15 Branquia vista de perfil aproximadamente recta (Fig. 73 C) ........ Simulium jujuyense Branquia vista de perfil con una suave curvatura mediana (Figs. 73 A-B) .......... 16 Bifurcación de ramas secundarias sobrepasando 1/3 de la longitud de la branquia (Fig. 73 A) ...........................................................................................Simulium auripellitum Bifurcación de ramas secundarias sobrepasando 1/4 de la longitud de la branquia (Fig. 73 B) ..............................................................................................Simulium minianum

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INCRUSTATUM species group Simulium (Psaroniocompsa) auripellitum ENDERLEIN (Figs. 71 A-C; 72 A; 73 A, O; 74 J.) Simulium auripellitum ENDERLEIN, 1934b: 286; VULCANO, 1967: 7. Simulium (Psaroniocompsa) auripellitum COSCARÓN & WYGODZINSKY, 1984: 54-58; COSCARÓN, 1987: 23; 1991: 180-181; 2000: 6; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51; SHELLEY et al., 2000: 178-180. Psaroniocompsa auripellita: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122-123.

Female: Wing length 2.1 mm. General coloration as described for subgenus. Frons and clypeus grayish with nacreous sheen and reflecting bluish silvery to velvety appearance. Scutum with homogeneous and abundant pilosity, bronze to yellowish graygreen coloration with silky appearance, allowing 1+1 gray pollinose spots to be seen. Leg coloration as in S. bonaerense. Abdomen blackish with 1+1 gray pollinose spots on tergite II. Frons relatively wide (Fig. 71 A); fronto-ocular triangle reduced. Basal portion of cibarium as in S. limbatum. Sensory vesicle of maxillary palp shorter than half length of palpomere. Sc generally without hair. Claws with short subbasal tooth. Paraprocts relatively short and curved distally (Fig. 71 B); genital fork with lateral branches scarcely sclerotized (Fig. 71 C); eighth sternite with 12-15 setae and gonapophyses with internal border subparallel and slightly sclerotized. Male: Wing length 1.6-2.0 mm. Area around antenna insertion silvery. Scutum velvety black with hair as in female, male basitarsus ratio 6.1-7.5. Abdomen as described for subgenus. Genitalia as in S. bonaerense. Gonostylus subquadrate with strong spur positioned near midsection of distal border (Fig. 72 A). Pupa: Cocoon with small dorsal projection, and reinforced anterior border (Fig. 73 A); threads evident. Cocoon base length 2.5-3.0 mm; on dorsum 2.0-2.7 mm; gills: 1.9-2.7 mm. Gill filaments with 6 terminal relatively joined branches, with short ventral curvature at half total length; secondary bifurcations at relatively long distance from base; bifurcation of dorsal secondaries about 1/3 or more of total gill length (Fig. 73 A). Platelets relatively scarce (Fig. 73 O) and trichomes of frontoclypeus and exposed portion of thorax bifid (Fig. 74 J). Larvae: Color, size and morphology very close to S. incrustatum. Ratio of proximal, medial and distal antennal segments = 1:1.0-1.1:1.0-1.2. Anal sclerite with scarce hairs. Anal ring with 62-65 rows of 11-13 hooks each. Rectal papillae with 6-11 diverticula on each lobe. Distribution: Argentina: Corrientes, Misiones, Santa Fe; Brazil: Goias, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo, Tocantins; Paraguay: Alto Paraná, Asunción, La Cordillera; Uruguay. Bionomics: This species is very abundant in small creeks with moderate flow and clean temperate water. Larval stages live on submerged aquatic vegetation or on grasses at the border of the streams. Females will bite humans and cattle. Discussion: This species is difficult to separate from the other species with generalized characters. Specimens of S. jujuyense in austral and western populations are of smaller size, with gill branches bifurcating nearer to the base and without a ventral concavity, but these differences are not evident in the eastern populations, living in the

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T L

O

Q

R

U

K

P

S

A

M

G

F D

E

N

J C

H

B

I Figure 71. Female. A-C: S. (Psaroniocompsa) auripellitum; A: frons; B: cercus and paraproct; C: genital fork. D-H: S. (P.) bonaerense; D-F: fore, mid and hind leg, respectively; G: genitalia in ventral view showing eighth sternite, gonapophyses, genital fork and spermatheca; H: cercus and paraproct. I-J: S. (P.) minuanum; I: claw; J: cercus and paraproct. K-N: S. (P.) incrustatum; K: grouped hair of scutum; L: scutum; M: anterior veins of wing; N: claw. O-P: S. (P.) limbatum; O: scutum; P: basal portion of cibarium. Q-R: S. (P.) auristriatum; Q: scutum; R: fronto-ocular triangle. S-T: S. (P.) brevifurcatum; S: frons; T: fronto-ocular triangle. U: S. (P.) damascenoi, head, dorsal view and scutum (from HAMADA et al., 2003).

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J

C E K A

D

B F G

L H

I

M

Figure 72. Male. A: S. (Psaroniocompsa) auripellitum, gonocoxite and gonostylus. B-D: S. (P.) bonaerense; B: hairs of legs; C: distal portion of hind basitarsus with calcipala and portion of second tarsite; D: genitalia in ventral view, showing gonocoxite, gonostylus, ventral plate and endoparameres. E-F: S. (P.) limbatum; E: scutum; F: gonostylus. G: S. (P.) anamariae, ventral plate. H: S. (P.) auristriatum, gonocoxite and gonostylus. I: S. (P.) schmidtmummi, gonocoxite and gonostylus (from WYGODZINSKI, 1973). J: S. (P.) damascenoi, scutum and portion of head (from HAMADA et al., 2003). K-M: S. (P.) tergospinosum (from HAMADA, 2000); K: scutum; L: gonocoxite and gonostylus; M: gonostylus dorsal view showing longitudinal carina.

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same areas with S. auripellitum. Simulium minuanum is differentiated by having scutum hairs of silvery with greenish to bronze tones, and claws without a subbasal tooth. Simulium angrense shows hairs on the Sc, and the more abundant scutum hairs do not permit the anterior 1+1 subtriangular silvery spots to be seen. The pupal gill filaments are more expanded and the frontoclypeal and thoracic tubercles are more abundant. Simulium bonaerense has scutum hairs more grayish in coloration and the pupal gill bifurcation is closer to the base. Simulium incrustatum is similar, but the grouped scutum hairs, more abundant platelets on the pupal frontoclypeus and thorax, and gill bifurcation of the secondary branches at a greater distance from the base, allow it to be separated. Simulium (Psaroniocompsa) minuanum STRIEDER & COSCARÓN (Figs. 71 I, J.) Simulium (Psaroniocompsa) minuanum STRIEDER & COSCARÓN, 2000: 105-111; COSCARÓN, 2003: 5; CROSSKEY & HOWARD, 2004: 51. Psaroniocompsa minuana: STRIEDER, 2004: 124.

Female: Very close to some others species of subgenus; it is characterized by wing length 2.0-2.2 mm, scutum hair homogeneously distributed; frons relatively narrow and frontal angle lower than in closest species; sensory vesicle of maxillary palp surpassing half length of palpomere; claws without subbasal tooth (Fig. 71 I); paraproct longer than high and with rounded border (Fig. 71 J); gonapophysis with internal border slightly sclerotized and distal border with nearly straight margin. Male: Wing length 1.8-2.0 mm, no special differences from other species of the group. Pupa: Cocoon without anterior projection. Primary ventral branch elongated, with bifurcation surpassing 3 times primary dorsal branch bifurcation length (Fig. 73 B). Frontoclypeus and exposed portion of thorax with scarce platelets, fewer than in S. auripellitum; trichomes of thorax with 1-3 branches. Larva: Maximum length 3.4-3.6 mm. Body integument with abundant short hairs. Cephalic apotome slightly positive. Antenna surpassing apex of cephalic fan stem; ratio of antennal segments: proximal, medial and distal = 1:0.6-0.7:1.1-1.2. Hypostomium with 4 setae on each side. Mandible with one latero-mandibular process. Postgenal cleft deep with rounded margin. Anal ring with 60-66 rows with 8-13 hooks each. Rectal papillae with 8-10 diverticula on each lobe. Distribution: Argentina: Corrientes, Entre Ríos, Misiones; Brazil: Rio Grande do Sul. Bionomics: Larval stages live in creeks of fast, clear water, with soft milky tones, flowing over aquatic vegetation or submerged tree branches. Female feeding habits are unknown. Discussion: As it is difficult to differentiate S. minuanum from other closely related species, we have added a table with differentiating characteristics: Simulium Simulium Simulium angrense auripellitum minuanum ‡ wing length (mm)

2.5-3.0 ¯x = 2.6 n=3

2.1-3.0 ¯x = 2.3 n=6

2.0-2.2 ¯x = 2.1

Simulium Simulium jujuyense bonaerense 2.1 -2.4

2.2-2.5

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Simulium Simulium Simulium angrense auripellitum minuanum † wing length (mm)

Simulium Simulium jujuyense bonaerense

2.1-2.6 ¯x = 2.3 n=3

1.8-2.0 ¯x = 1.9

1.8-2.0 ¯x = 1.9

1.9-2.2

2.0-2.4

Scutum hair coloration

Tin to yellowish green

Tin to golden; male silvery-gray

Silvery with bright green to tin

Silverygreen to tin

Tin to grayish

Frontal angle

103°

110-120°

100°

115-120°

110°

Ratio of length of third palpomere to sensory vesicle

2.9-3.0 ¯x = 2.9 n=2

2.2-2.5 ¯x = 2.3 n=4

2.3-2.5 ¯x = 2.4 n=3

2.0-2.5 ¯x = 2.2

2.5-2.8 ¯x = 2.6

Ratio of length/width of hind basitarsus, female

6.2-8.9 ¯x = 7.4 n=3

7.0-9.0 ¯x = 7.8 n=5

7.3-8.2 ¯x = 7.7

6.4-7.7 ¯x = 7.1

6.0-7.0 ¯x = 7.1

Ratio of length/width of hind basitarsus, male

6.4-6.5 = 6.4

6.1-7.5 = 6.6 n=4

7.5-7.9 = 7.7

7.7-8.0 = 7.9

6.0-7.3 = 6.5

Claw with subbasal tooth

+

+

-

+ reduced

+ reduced

Pupa with tubercles on frontoclypeus and thorax

Very numerous

Relatively scarce

More scarce

Cocoon ventral length (mm)

3.0-3.7

2.5-3.0

2.5-2.9

2.8-3.0

3.8-4.0

Cocoon dorsal length (mm)

2.7-3.2

2.0-2.7

1.7-2.0

2.5-2.7

3.2-3.5

Gill length

3.1-4.5

1.9-2.7

1.7-2.3

3.4-3.5

2.5-3.0

Larva antennal ratio of segments: proximal, medial and distal

1:1.0:1.0

1:1.0-1.1: 1.0-1.2

1:0.66-0.71: 1.15-1.27

1:1.0.0.9

1:1.0:0.9-1.0

Medium to From scarce scarce to numerous

Simulium (Psaroniocompsa) jujuyense PATERSON & SHANNON (Figs. 73 C; 74 A, B.) Simulium jujuyense PATERSON & SHANNON, 1927: 739; WYGODZINSKY, 1951a: 206-214; COSCARÓN & WYGODZINSKY, 1960: 1124-1125; VULCANO, 1967: 14. Simulium (Psaroniocompsa) jujuyense: COSCARÓN & WYGODZINSKY, 1984: 67-69; COSCARÓN, 1987: 23; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51; STRIEDER & COSCARÓN, 2000: 111; COSCARÓN, 2003: 5; COSCARÓN ARIAS, 2003: 198, 204, 208, 215. Psaroniocompsa mendozana ENDERLEIN, 1936: 123. Simulium spinifer KNAB, 1914b: 81. Simulium hoffmanni VARGAS, 1943a: 138 (n. nom. Trichodagmia angustitarse ENDERLEIN, 1934b: 289). Simulium sicuani SMART, 1944: 132 (n. nom. Ectemnaspis limbata ENDERLEIN, 1934b: 282). Simulium incaicum VARGAS, 1945a: 148 (unjustified repl. name for E. limbata).

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Female: Wing length 2.1-2.4 mm. Coloration general blackish. Frons and clypeus velvety grayish. Scutum with silvery to bronze, silky pilosity, homogeneously distributed. Legs dark brown to black on darkened areas. Abdomen with 1+1 lateral silvery spots on tergite II. Wings without hair on Sc. Genitalia as in S. bonaerense. Male: Wing length 1.9-2.2 mm; general coloration as in female. Genitalia as in S. bonaerense. Pupa: Cocoon yellowish whitish-gray, with straight border and without anterior reinforcement (Fig. 73 C); weave superficially smooth, with thin threads. Length at base 2.8-3.0 mm; on dorsum 2.5 mm, gill 3.4 mm. Gill bifurcation of secondary branches more proximal than in S. auripellitum, but more distal than in S. bonaerense. Primary ventral branch bifurcation more proximal than secondary ventral (Fig. 73 C) to about same level (Fig. 74 B). Frontoclypeal and thoracic tubercles relatively scarce, fewer than in S. incrustatum. Abdominal chaetotaxy as in figure 73 A. Larva: Maximum length 4.5-5.0 mm. General coloration light yellowish gray, with greenish violaceous on abdominal segment bands. Ratio of antennal segments: proximal, medial and distal = 1:1.0:0.9. Mandible, hypostomium and postgenal cleft as in S. incrustatum. Anal gill with 8-10 diverticula on each lobe. Distribution: Argentina: Buenos Aires, Chaco, Córdoba, Corrientes, Entre Ríos, Formosa, Jujuy, La Pampa, La Rioja, Mendoza, Río Negro, Salta, Tucumán; Bolivia; Colombia; Ecuador: Cotopaxi, Esmeraldas; Peru: Calca, Cuzco. Bionomics: The preimaginal stages live on aquatic vegetation and submerged trees branches, especially Salix humboldtiana in moderate to fast-flowing creeks and rivers. Females attack humans, cows and horses. Discussion: The closest species are S. auripellitum and S. bonaerense, which are sympatric in some places, and can be differentiated by the yellowish green scutum pilosity and the pupal gill branching, as explained in the description. Simulium bonaerense can be distinguished by the big size and especially by the lower gill branching. This similarity produced misidentification by COSCARÓN & WYGODZINSKY (1960: 1124) and COSCARÓN (1968: 66) and in part by COSCARÓN & WYGODZINSKY (1984: 57) in relation to material from Chaco, Corrientes, Entre Ríos and Formosa, which was corrected by STRIEDER & COSCARÓN (2000: 111). Simulium (Psaroniocompsa) bonaerense COSCARÓN & WYGODZINSKY (Figs. 71 D-H; 72 B-D; 73 D; 74 C; 75 A, B.) Simulium (Psaroniocompsa) bonaerense COSCARÓN & WYGODZINSKY, 1984: 59-62; COSCARÓN, 1987: 23; 1991: 183-184; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51. Simulium jujuyense PATERSON & SHANNON, 1927; COSCARÓN & WYGODZINSKY, 1960: 1124; COSCARÓN, 1968: 66 (misidentification).

Female: Wing length 2.2-2.5 mm. General coloration blackish. Frons and clypeus gray pollinose, frons with velvety aspect. Scutum with hair bronze to grayish color, homogeneously arranged, permitting subtriangular anterior silver spots to be seen. Leg coloration in accordance with figures 71 D-F. Claw subbasal tooth present but small.

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A D

B

I

C

E

M F G

H P J

K

L N

O Q

Figure 73. Pupa. A-N: General view; A: S. (Psaroniocompsa) auripellitum; B: S. (P.) minuanum; C: S. (P.) jujuyense; D: S. (P.) bonaerense; E: S. (P.) angrense; F: S. (P.) incrustatum; G: S. (P.) limbatum; H: S. (P.) anamariae; I: S. (P.) brevifurcatum; J: S. (P.) auristriatum; K: S. (P.) schmidtmummi (from WYGODZINSKY, 1973); L: S. (P.) siolii; M: S. (P.) damascenoi (L-M from PY-DANIEL, 1988); N: S. (P.) tergospinosum (from HAMDA, 2000). O-Q Frontoclypeus; O: S. (P.) auripellitum; P: S. (P.) angrense; Q: S. (P.) incrustatum.

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I C B H G K

J

A

D F

E

Figure 74. Pupa. A: S. (Psaroniocompsa) jujuyense, abdominal chaetotaxy. B-I: Gills; B: S. (P.) jujuyense; C: S. (P.) bonaerense; D: S. (P.) angrense; E: S. (P.) incrustatum; F: S. (P.) limbatum; G: S. (P.) anamariae; H: S. (P.) schmidtmummi (H from WYGODZINSKY, 1973); I: S. (P.) damascenoi (I from PY-DANIEL, 1988). JK: Trichomes; J: S. (P.) auripellitum, trichomes and tubercles of thorax; K: S. (P.) schmidtmummi, multibranched trichomes (from WYGOODZINSKY, 1973).

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Abdomen with 1+1 small silvery spots on tergite II. Frons relatively wide but narrower than in S. auripellitum; fronto-ocular triangle very reduced. Wings with hair on Sc. Genitalia as in figure 71 G. Eighth sternite with about 40 setae and gonapophyses with abundant microtrichiae, internal border subparallel and scarcely sclerotized. Paraproct short (Fig. 71 H) and genital fork with stem well sclerotized. Male: Wing 2.0-2.4 mm. General coloration as in female and with same pattern as in other males of subgenus. Legs as in female with filiform and spatuliform hair (Fig. 72 B). Genitalia as in figure 72 D. Pupa: Cocoon light brown, straight and slowly reinforced on anterior border (Fig. 73 D); weave superficially smooth with evident threads; length at base 3.8-4.0 mm, at dorsum 3.2-3.5 mm, gills 2.5-3.0 mm. Gill branching very close to base (Fig. 74 C) and relatively open. Frontoclypeus and exposed portion of thorax with platelets medially, abundant as in S. incrustatum. Abdominal chaetotaxy as in S. jujuyense. Larva: General coloration from greenish to yellowish brown. Morphology as in S. incrustatum, only differentiated by scarce gill diverticula that only reach 4 on each lobe in S. bonaerense (Fig. 75 B). Antenna as in figure 75 A, with ratio of proximal, medial and distal segments = 1:1:0.9-1.0. Rectal papillae with 76-80 rows with 15-16 hooks each. Distribution: Argentina: Buenos Aires (center and south west area), Río Negro (Alto Valle). Bionomics: This species lives in environments similar to those of S. jujuyense. It is present all year, but the highest peak is in the spring (COSCARÓN et al., 2000). Females bite humans and cattle, and are a problem for inhabitants. Discussion: The closest species are S. auripellitum, S. jujuyense and S. minuanum. The larger size and pupal gill branching close to the base allow it to be differentiated from them. Also, S. auripellitum and S. minuanum show differences in scutum hair coloration, which is bronze to golden with silvery greenish tones, and the latter species does not have a subbasal tooth on the claws. Simulium (Psaroniocompsa) angrense PINTO (Figs. 73 E, P; 74 D.) Simulium angrensis PINTO, 1931: 683. Simulium angrense: VARGAS, 1945a: 112; VULCANO, 1967: 6. Simulium (Psaroniocompsa) angrense COSCARÓN & WYGODZINSKY, 1984: 52-54; COSCARÓN, 1987: 23; 1991: 179; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51. Psaroniocompsa angrense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122.

The type of this species was not found at the Instituto Oswaldo Cruz by the first author in 1979 and the description of COSCARÓN & WYGODZINSKY (1984) that is used as a guide here, is based on material collected close to the type locality (Angra dos Reis, Rio de Janeiro), and in accordance with the original description. Female: Wing length 2.5-2.8 mm. Frons and clypeus from velvety grayish pollinose to bronze to greenish yellow; scutum hair tin to yellowish green, arranged homogeneously, difficult to see the 1+1 anterior subtriangular silvery spots. Sc with 10-12 hairs. Claw with small subbasal tooth. Genitalia as in S. bonaerense.

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Male: Wing length 2.1-2.6 mm. General coloration as in female. Area around antenna insertion velvety grayish. Scutum black, velvety with hair arranged as in female. Sc without hairs. Genitalia similar to those of S. bonaerense. Pupa: Cocoon with small anterior projection and reinforced on anterior border (Fig. 73 E), with resistant weave and visible threads. Length on base 3.0-3.7 mm, on dorsum 2.7-3.2 mm, gills 3.1-4.5 mm. Gill branching near base (Fig. 73 E), secondary branches bifurcating frequently at same level (Fig. 74 D); frontoclypeus with abundant tubercles (Fig. 73 P); trichomes bifid and a little shorter than in S. incrustatum. Larva: Maximum length 4.5-5.5 mm. General coloration yellowish green. General morphology close to that of S. incrustatum but mandible with thinner marginal teeth and rectal papillae diverticula numbering 10-12 on each lobe. Distribution: Brazil: Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: Unknown. Discussion: The closest species is S. auripellitum, which can be differentiated by its smaller size, scutum pilosity more silky yellowish and pupal gill with secondary branches bifurcating at a greater distance from the base. Simulium incrustatum, which is present in the same area, can be differentiated by the scutum hair arranged in groups simulating scales, and the pupal primary ventral and secondary branches bifurcating at greater distance from the base. Simulium (Psaroniocompsa) incrustatum LUTZ (Figs. 71 K-N; 73 F; 74 E, Q; 75 C-J.) Simulium incrustatum LUTZ, 1910: 243-245; PINTO, 1931: 703; LANE & PORTO, 1940: 194; VARGAS, 1945a: 148; VULCANO, 1967: 13; COSCARÓN & WYGODZINSKY, 1973a: 170. Simulium (Psaroniocompsa) incrustatum: COSCARÓN & WYGODZINSKY, 1984: 62-67; COSCARÓN, 1987: 23; 1991: 184-186; 2003: 5; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 51; SHELLEY et al., 1997: 24; COSCARÓN ARIAS, 2003: 198, 204, 208-215. Psaroniocompsa incrustata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 133-135; 2004: 123-124. Psaroniocompsa opalinifrons ENDERLEIN, 1934a: 192; 1936: 112. Simulium opalinifrons: VARGAS, 1945a: 100; WYGODZINSKY, 1951a: 217; VARGAS & DIAZ NÁJERA, 1953b: 137. Simulium (Psaroniocompsa) opalinifrons: COSCARÓN & WYGODZINSKY, 1973a: 161-172. Simulium yarzabali RAMÍREZ PÉREZ, 1980: 66-68; 1983: 9; SHELLEY et al., 1987a: 463 (revalidated).

Female: Wing length 2.0-2.8 mm. General coloration blackish. Frons and clypeus with silvery pruinosity, bluish-green sheen iridescent. Scutum velvety black with golden to bronze hair grouped, simulating scales (Fig. 71 K) and arranged in longitudinal rows specially at sides of anterior half (Fig. 71 L). Scutellum and metanotum black, the latter with gray pollinosity. Legs brown with darkened areas blackish showing, with some illumination, whitish fore tibia internally and 3/4 hind basitarsus externally. Abdomen blackish with 1+1 silvery pollinose spots on tergite II. Fronto-ocular triangle very short; Sc with 4-12 hairs, chaetotaxy of anterior wing veins as in figure 71 M. Claws with short basal tooth (Fig. 71 N). Genitalia as in S. bonaerense; genital fork with well-sclerotized stem.

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K C

D L

O

P F A

Q H

E M

G N

J

I

B

Figure 75. Larva. A-B: S. (Psaroniocompsa) bonaerense; A: antenna; B: rectal papillae. C-J: S. (P.) incrustatum; C: general aspect; D: head, dorsal view; E: antenna; F: mandible; G: trichomes of head, dorsum; H: postgenal cleft and hypostomium; I: anal sclerite; J: distal portion of abdomen with rectal papillae. K-N: S. (P.) auristriatum; K: general aspect; L: head, dorsal view; M: antenna; N: hypostomium. O-Q: S. (P.) siolii ( from PY-DANIEL, 1988); O: general aspect; P: abdomen with more magnification showing dorsal prominences and trichomes; Q: teeth of mandible with lateromandibular process (O-Q from PY-DANIEL, 1988).

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Male: Wing length 1.9-2.8 mm. General coloration as in female. Area around antennae insertion silvery with iridescent sheen. Scutum velvety black with golden hair grouped as in female, but not arranged in longitudinal rows. Sc with 0-2 hairs. Abdominal pattern as in subgenus. Genitalia as in S. bonaerense. Pupa: Cocoon without dorsal anterior projection, slowly reinforced on rim, weave not compacted, threads visible (Fig. 73 F). Length at dorsum 2.5-2.9 mm; at base 3.0-3.2 mm; gills 3.1-4.5 mm. Generally gill bifurcation of primary ventral and secondary branches relatively far from base, variable distance among specimens, with secondary branches bifurcating about 1/2-1/3 of total gill length, and primary ventral bifurcation at half of anterior distance (Fig. 74 E). Cephalothoracic trichomes mostly bifid and relatively elongated; platelets of frontoclypeus and exposed portion of thorax moderately abundant, some grouped, leaving clear areas especially on thorax (Fig. 74 Q). Larva: Maximum length 4.6-5.0 mm. General coloration light greenish-yellow. General aspect as in figure 75 C. Body integument with microscopic hairs, like flattened multibranched scales as seen with high magnification. Cephalic apotome with small, discontinuous, median and sublateral positive spots (Fig. 75 D); Antennae surpassing stem length of cephalic fan; ratio of proximal, medial and distal segments = 1:1.0-1.1:1.2 (Fig. 75 E). Mandible with thick first marginal tooth (Fig. 75 F). Head tegument with bifid hairs (Fig. 75 G) Cephalic fan with about 35-40 rays. Hypostomium with 4 setae on each side; postgenal cleft deep and rather rounded (Fig. 75 H). Anal ring with about 62-70 rows with 12-15 hooks each; anal sclerite with hairs single or with several branches (Fig. 75 I). Rectal papillae with 5-8 diverticula on each lobe (Fig. 75 J). Distribution: Argentina: Corrientes, Misiones; Brazil: Bahia, Federal District, Mato Grosso do Sul, Minas Gerais, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Roraima, Santa Catarina, São Paulo; Ecuador: Cotopaxi, El Oro, Esmeraldas, Guayas, Los Rios, Manabi; Paraguay: Alto Paraná, La Cordillera; Venezuela: Amazonas, Bolivar. Bionomics: This species breeds in small creeks with clean water, on submerged aquatic vegetation. Females are haematophagous and the bite produces serious annoyance in Brazil. Discussion: The presence of hair grouped and forming scales on the scutum can confuse the identification with S. limbatum, S. anamariae, S. auristriatum, S. brevifurcatum, S. cuasiexiguum, S. exiguum and S. paraguayense. The three last-mentioned species correspond to S. (Notolepria), with different genitalia and pupal gill and trichomes. From limbatum, it can be differentiated by the presence of 1+1 silvery submedian stripes going from anterior subtrianglular spots to the posterior border of the scutum, and grayish pilosity. From S. anamariae, S. auristriatum and S. brevifurcatum it differs by the presence of larger hair groups and greenish violaceous coloration, plus the cocoon with a dorsal projection and different gill morphology. Pliodasina guttata (ENDERLEIN) from Paraguay has some similarity with the female type of S. incrustatum, but since it is not well preserved, it is not possible to define its identity; no other stages are known and we prefer to maintain it as species inquirendae. Simulium yarzabali was synonymized by RAMÍREZ PÉREZ (1983: 9), and rejected by SHELLEY (1987: 464). We consider it a geographic variant and accept the RAMÍREZ PÉREZ criteria.

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Simulium (Psaroniocompsa) limbatum KNAB (Figs. 71 O, P; 72 E, F; 73 G; 74 F.) Simulium limbatum KNAB, 1915c: 259; RAMÍREZ PÉREZ et al., 1982: 45; RAMÍREZ PÉREZ, 1983: 43; SHELLEY et al., 1984: 143; SHELLEY et al., 1997: 27. Simulium (Psaroniocompsa) limbatum: COSCARÓN, 1991: 188; CROSSKEY & HOWARD, 1997: 57; 2004 : 51. Simulium (Psaroniocompsa) aequifurcatum LUTZ, 1910: 259, 263; COSCARÓN & WYGODZINSKY, 1984: 49; COSCARÓN, 1987: 23; 1991: 186. Simulium machadoi RAMÍREZ PÉREZ, 1971: 341. Simulium machadoallisoni VULCANO, 1981: 276 (n. nom. to Simulium machadoi RAMÍREZ PÉREZ). Simulium meruoca NUNES DE MELLO et al., 1973: 59.

Female: Wing length 1.8-2.2 mm. Color generally blackish. Frons and clypeus with silvery pruinosity and blue sheen. Scutum black with one median and 1+1 submedian silvery pollinose thin bands reaching posterior grayish area (Fig. 71 O), (evident with some special illumination). Scutum hair pale gray-greenish, arranged in groups but fewer in number than in S. incrustatum. Sc without hair, cibarium with evident spines arranged in several rows (Fig. 71 P). Eighth sternite with about 20-25 setae on each side, and gonapophysis with internal rim slightly divergent. Male: Wing length 1.8-2.0 mm. General coloration as in female but gray pollinosity of scutum bands shorter and thinner posteriorly (Fig. 72 E). Genitalia as in S. bonaerense but gonostylus with wrinkled border (Fig. 72 F). Pupa: Cocoon with slight dorsal anterior projection and reinforced border (Fig. 73 G). Length on base 2.6-3.0 mm; on dorsum 2.4-2.6 mm. Gill with 6 branches emerging close to base; primary ventral branch bifurcated about half of secondary dorsal and ventral bifurcation level, (as shown in figure 74 F), or secondary dorsal branch bifurcating about two times distance of secondary ventral bifurcation. Frontoclypeus and exposed portion of thorax with numerous platelets, similar to condition in S. incrustatum, but with variation in number and not arranged in groups. Trichomes bifid and shorter than in S. incrustatum. Larva: Maximum length 4.0 mm. Body integument with scarce and short filiform, generally single hairs. Cephalic apotome similar to that of S. incrustatum. Postgenal bridge shorter than half of hypostomium length. Ratio of antennal segments: proximal, median and distal = 1:0.8-0.9:1.1. Mandible with 2 latero-mandibular processes. Lateral sclerite of thoracic proleg with 14-19 teeth arranged in 4-6 groups. Anal ring with 65-70 rows with 10-12 hooks each. Rectal papillae with 3 diverticula on each lobe. Distribution: Brazil: Amapá, Bahia, Espírito Santo, Mato Grosso, Paraiba, Pará, Pernambuco, Rio de Janeiro, Roraima, São Paulo; Guyana: Rupununi; Venezuela: Amazonas, Anzoategui, Aragua, Barinas, Bolivar, Carabobo, Cojedes, Guarico, Lara, Miranda, Monagas, Portuguesa, Táchira, Trujillo, Yaracuy. Bionomics: This species breeds in small creeks, about 1-5 m wide and not more 1 m deep, not torrential, flowing over aquatic vegetation. Females were collected biting humans and horses. Discussion: Simulium limbatum is very close to S. incrustatum. They can be differentiated by the presence in S. incrustatum of 1+1 short silvery bands on the scutum that do not continue from the anterior cuneiform spots nor surpass 1/4 of the scutum length, and golden scutum hair arranged in groups a little bigger than in S. limbatum. Also, the pupal

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gill branches more distal from the base. The latter character shows great variation in both species and can produce confusion, especially when the material is not reared. Gill branching similar to that in S. limbatum can be found in the range of variation for S. incrustatum (SHELLEY et al., 1997). COSCARÓN & WYGODZINSKY (1984) and COSCARÓN (1987, 1991) used S. aequifurcatum LUTZ to describe this species, based on material deposited at Institute Oswaldo Cruz. It corresponds to a pupa, without exact provenance. No reared material from the LUTZ collection was found. The best characters to separate S. limbatum from S. incrustatum are in the adults. It is best to use S. limbatum to name this taxon, which was described from a female, and to accept the synonymy of S. aequifurcatum with S. incrustatum proposed by SHELLEY et al. (1984). Material indicated as aequifurcatum by COSCARÓN & WYGODZINSKY (1984), COSCARÓN (1987, 1991) & COSCARÓN et al. (1992) corresponds to S. limbatum. Simulium (Psaroniocompsa) fuliginis FIELD Simulium fuliginis FIELD, 1969b: 281. Simulium (Psaroniocompsa) fuliginis: COSCARÓN & WYGODZINSKY, 1984: 77; COSCARÓN, 1987: 24; 1991: 188; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 52.

This species was described based on females from Panama. The description, with good illustrations, is similar to that for S. limbatum KNAB. It is differentiated only by its small size (wing length 1.8 mm) and absence of 1+1 submedian thin silvery stripes on the scutum, evident with some illumination. We could not see the types which are not in the U.S.N.M. as indicated by the author. With the identity in doubt, we prefer not to synonymize this species with S. limbatum, pending study of type material of all stages. Simulium (Psaroniocompsa) anamariae VULCANO (Figs. 72 G; 73 H; 74 G.) Simulium anamariae VULCANO, 1962: 246; 1967: 6. Simulium (Psaroniocompsa) anamariae: COSCARÓN & WYGODZINSKY, 1984: 69-71; PY-DANIEL et al., 1985: 97-98; COSCARÓN, 1987: 23; 1991: 189-190; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 52. Psaroniocompsa anamariae: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 122.

Female: Wing length 2.5-2.8 mm. Frons and clypeus grayish with bluish silvery iridescent sheen. Scutum black without anterior spots, with abundant hair groups like scales arranged in rows, one median and several laterals as in S. incrustatum. Humeral callus light brown. Scutellum, metanotum and pleurae black; abdomen with tergite I brown with nacreous sheen pollinosity at sides; tergite II with 1+1 lateral gray pollinose spots; following tergites brownish with cereous sheen. Fronto-ocular triangle reduced, wider than long. Legs with spatuliform hairs; claws with small subbasal tooth. Eighth sternite, paraproct, cercus and gonapophysis as in S. bonaerense. Genital fork with enlarged stem similar to that of S. auripellitum. Male: Wing length 2.0-2.5 mm. General coloration blackish, ornamented as typical for subgenus. Gonocoxite subtrapezoidal, about as long as wide; gonostylus half length of gonocoxite, subtrapezoidal with spur subapical. Ventral plate as in figure 72 G.

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Pupa: Cocoon relatively flattened with two anterior, dorsal enlarged projections (Fig. 73 H); weave not compressed, with evident threads. Length on base 3.2-3.6 mm; on dorsum without projections, 2.8-3.2 mm; gills 3.8-4.5 mm. Primary ventral and secondary dorsal and median branch bifurcated far from base; the two latter about at half of total length (Fig. 74 G). Frontoclypeus with 2 pairs of frontal trichomes elongated and with 2-3 branches and 1 pair facials simple. Tubercles of cephalothorax abundant, but relatively small. Larva (based on PY-DANIEL et al., 1985): Maximum length 5.1-5.3 mm. Body integument with spiculae of 3-4 branches. Ratio of antennal segments: proximal, medial and distal = 1:0.6:1.2. Cephalic fan with 45-46 rays. Mandibles with 2 sinuous lateromandiblular processes surpassing teeth apical border. Anal ring with about 66 rows with 912 hooks each. Rectal papillae with 8-10 diverticula on each lobe. Distribution: Brazil: Rio de Janeiro, Rio Grande do Sul, São Paulo. Bionomics: Unknown. Discussion: The most similar species are S. brevifurcatum and S. auristriatum, which have similar scutum pilosity with grouped hairs arranged in longitudinal rows, and without 1+1 anterior silvery cuneiform spots. It is easy to separate them by the peculiar two projections of the S. anamariae cocoon. Also, the gills have an open arrangement with very basal branching in S. brevifurcatum, and the gill has two thick branches in S. auristriatum. Simulium (Psaroniocompsa) auristriatum LUTZ (Figs. 71 Q, R; 72 H; 73 J; 75 K-N.) Simulium auristriatum LUTZ, 1910: 245-246, 266-267; LUTZ & NUNEZ TOVAR, 1928: 131; PINTO, 1931: 684; D’ANDRETTA & D’ANDRETTA, 1949: 55-56; VULCANO, 1967: 7. Simulium (Psaroniocompsa) auristriatum: COSCARÓN & WYGODZINSKY, 1984: 71-74; COSCARÓN, 1987: 23; 1991: 190-191; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 56; 2004: 50. Simulium infuscatum LUTZ, 1910: 236; PINTO, 1931: 704; PORTO, 1939: 369. Psaroniocompsa auristriata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

Female: Wing length 2.5-2.6 mm. General coloration blackish. Scutum with hair grouped as big scales greenish blue to violet and arranged in longitudinal rows (Fig. 71 Q). Scutellum wide and fronto-ocular triangle reduced (Fig. 71 R). Sc without hairs. Genitalia as in S. bonaerense; genital fork with stem well sclerotized and elongated, about 1/3 longer than each posterior branch. Male: Wing length 2.3-2.5 mm. Coloration and disposition of scutum hair as in female. Gonocoxite similar to that of S. anamariae; gonostylus subquadrate with spur near internal corner (Fig. 72 H). Pupa: Cocoon projected dorsally (Fig. 73 J); weave soft with threads obvious. Length at base 3.0-3.5 mm, at dorsum 2.8-3.0 mm, gills 4.3-5.4 mm. Gill with 2 thick branches, emerging from short basal trunk; gills with some constrictions similar to pseudoannulations (Fig. 73 J), surface of gills with small spiculae. Frontoclypeus and thorax with abundant platelets. Trichomes bifid and some single. Larva: Maximum length 5.5 mm; general coloration yellowish brown with transverse band grayish green on abdominal segments, shape as in figure 75 K. Cephalic apotome scarcely pigmented (Fig. 75 L). Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.7-0.8:1.1-1.2 (Fig. 75 M). Cephalic fan with 45

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rays. Mandible as in S. incrustatum. Hypostomium as figure 75 N. Postgenal bridge about as long as hypostomium. Anal sclerite with hairs single or bifid, longer than in S. incrustatum. Anal ring with about 65 rows of 12 hooks each. Rectal papillae unknown. Distribution: Brazil: Mato Grosso do Sul, Minas Gerais, Rio de Janeiro, São Paulo. Bionomics: This species breeds in small creeks, with moderate-flowing, clear water. Female feeding habits are unknown. Discussion: The adults are very close to those of S. (Notolepria) species, S. incrustatum, S. anamariae and S. brevifurcatum. Simulium (Notolepria) species are smaller but have a more acuminate paraproct. The S. incrustatum scutum is similar in appearance, but the hair groups are not aligned in rows. Simulium anamariae has golden hair groups on the scutum and S. brevifurcatum a grayish spot on the pronotal corner. The pupal gill shape permits it to be differentiated from other Neotropical black fly species. Simulium (Psaroniocompsa) brevifurcatum LUTZ (Figs. 71 S, T; 73 I.) Simulium brevifurcatum LUTZ, 1910: 259, 263 , 264; PINTO, 1931: 694; VULCANO, 1962: 255; 1967: 8. Simulium (Psaroniocompsa) brevifurcatum: COSCARÓN & WYGODZINSKY, 1984: 75; PY-DANIEL et al., 1985: 96-97; COSCARÓN, 1987: 23; 1991: 191-193; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 56; 2004: 50. Psaroniocompsa brevifurcata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 123.

Female: Wing length 2.5-2.6 mm. Scutum, scutellum and metanotum velvety black; scutum without anterior cuneiform silvery spots, and hair groups appearing like scales with metallic golden violaceous coloration, ordered in longitudinal rows. Pronotum corners with grayish spots. Abdomen black with I-IV segments velvety; subsequent segments with cereous sheen; II-V with silvery large dorso-lateral spot on each side; Sc with 2-3 setae near base. Legs yellowish with some portion darkened and abundant spatuliform hairs. Frons relatively narrow and convergent (Fig. 71 S). Fronto-ocular triangle wider than high but deeper than in remaining species in subgenus (Fig. 71 T). Genitalia as in S. bonaerense, but paraproct shorter, slowly surpassing cercus ventral border; genital fork as in S. auristriatum. Male: Wing length 2.3 mm. Coloration as in female; disposition of hair groups less orderly than in female. Legs with abundant brownish spatuliform hairs. Gonostylus subquadrate, with small median depression on distal rim and robust subapical spur positioned as in S. limbatum. Pupa: Cocoon laterally expanded and with dorsal projection (Fig. 73 I); weave thick with visible threads; length at base 3.5-4.0 mm; at dorsum 3.8 mm; gills 3.4-4.0 mm. Gill with 6 branches with dorsal secondary branch bifurcated very close to base (Fig. 73 I); all filaments markedly expanded laterally. Frontoclypeus and exposed portion of thorax with small and abundant platelets; frontoclypeus and trichomes bifid. Larva (based on PY-DANIEL et al., 1985): Maximum length 5.7 mm. Body integument with spiculae of 4-6 branches. Ratio of antennal segments: proximal, medial and distal = 1:0.9:0.9-1.0; cephalic fan with 52-53 rays. Mandible with latero-mandibular process straight, surpassing apex. Rectal papillae with 68 rows and 10-12 hooks each. Distribution: Argentina: Misiones; Brazil: Paraná, Rio Grande do Sul, Rio de Janeiro, São Paulo.

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Bionomics: This species breeds in clear creeks of moderate to slow flow. Female feeding habits are unknown. Discussion: The adults closest to this species are those of S. anamariae and S. auristriatum; differences were explained in those species. The cocoon and pupal gill branching pattern permit differentiation of this species. Simulium (Psaroniocompsa) schmidtmummi WYGODZINSKY (Figs. 72 I; 73 K; 74 H, K.) Simulium (Psaroniocompsa) schmidtmummi WYGODZINSKY, 1973: 1-11; COSCARÓN & WYGODZINSKY, 1984: 77; COSCARÓN, 1987: 23; 1991: 193; COSCARÓN & COSCARÓN ARIAS, 1997: 115; CROSSKEY & HOWARD, 1997: 57; 2004: 50.

Female (based on WYGODZINSKY, 1973): Wing length 3.0-3.3 mm. Frons and clypeus from silver gray to nacreous. Scutum velvety black with 1+1 anterior silvery subtriangular spots. Hair appressed, isolated or in small groups somewhat longitudinally arranged, but not forming distinct lines. Pleurae silver gray to nacreous; scutellum velvety black. Abdomen with tergite I from piceous to black; II-V blackish with III-V velvety black at center. Tergite II with 1+1 large silver whitish spots; III-V with hind border silvery; VI-IX piceous, polished. Frons relatively wide. Sc glabrous. Claws with minute subbasal tooth; leg hairs spatuliform and single. Eighth sternite with 10 setae on each side; gonapophyses with internal margin subparallel and abundant microtrichiae; paraproct very short; genital fork with slender and well-sclerotized stem. Male: Wing length 3.0 mm. General coloration as in female. Abdomen velvety black. Segments II, IV-IX with 1+1 silver white lateral spots, segments IV and V not distinct. Ratio of length/width of hind basitarsus = 5.8-6.6; spatuliform hair more abundant than in female, especially on femora and tibiae. Gonostylus subquadrate (Fig. 72 I), remaining genitalia parts similar to those of S. bonaerense. Pupa: Cocoon with anterodorsal projection with blunt apex (Fig. 73 K). Cocoon translucent with threads evident. Length at dorsum 3.5-4.0 mm; at base 4.0-4.2 mm; gills 4.3 mm. Gill with 4 filaments, branching from base, with very short primary branches (Fig. 74 H). Frontoclypeus and thorax with moderately abundant small irregularly arranged platelets; head trichomes with simple to four branches; thoracic trichomes with 4-12 branches, some of them clavate at apex (Fig. 74 K). Larva: Maximum length 6.5 mm. General coloration yellowish white slightly reddish posteriorly. Body integument glabrous. Ninth segment with 1+1 ventral tubercles. Ratio of antennal segments: proximal, medial and distal = 1:1.1:1.2. Mandible marginal teeth 2 in number, relatively thin, especially second, sometimes there is another very small third tooth. Cephalic fan with 50-55 rays. Postgenal cleft deeper than postgenal bridge length. Anal ring with about 66 rows of 14-15 hooks each. Anal sclerite with single hairs. Rectal papillae with 12-16 diverticula on each lobe. Distribution: Colombia: Boyaca, Cundinamarca, in the Páramo area between 3050-3450 m. Bionomics: The larvae live on trailing grasses and stones in clear, small streams about 1 m wide and 0.15 m deep, with temperatures of about 9-12 °C. Female feeding habits are unknown.

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Discussion: Several species with hairs grouped and simulating scales on the scutum can confuse identification. The golden to bronze hair coloration helps to separate this species from S. incrustatum, the closest species. Simulium auristriatum and S. brevifurcatum don’t show 1+1 anterior cuneiform spots. The pupa gill with 4 branches allows it to be separated from S. (Psaroniocompsa) and S. (Notolepria) species. Simulium (Psaroniocompsa) stellatum GIL ACEVEDO, FIGUEIRÓ & MAIA HERZOG Simulium (Psaroniocompsa) stellatum GIL ACEVEDO et al., 2005 (922): 1-12.

(Based on original description). Female: Wing length 2.5 mm. Scutum totally black with lateral and posterior margin silver pruinose; pilosity golden, grouped hair arranged in longitudinal rows. Abdomen silvery pruinose with segments I-IV velvety, remaining black shiny. Claw with small subbasal tooth. Genitalia as in S. anamariae. Male: Wing length 1.9 mm. Color and disposition of scutum hairs as in female. Gonostyle subquadrate with tiny sub apical spur; ventral plate with smooth pointed apex. Pupa: Cocoon with dorsal projection, surpassing cocoon base, weave compact and thin, length at base 3.5-4.2 mm; gill length 3.4-4.1 mm. Gill with 4 branches emerging from short trunk. Larvae: Maximum length 6.0-7.2 mm. Cephalic apotome with one median longitudinal 1+1 anterolateral and 1+1 sublateral positive spots. Antenna with distal segment relatively long, ratio of proximal, medial and distal segments = 1:1:1.2: postgenal cleft rounded apically. Ninth segment with 1+1 ventral tubercles; anal ring with 67 rows of 913 hooks each; rectal papillae with 8-10 diverticula on each lobe. Distribution: Brazil: Minas Gerais, Rio de Janeiro. Bionomics: This species was collected in springs at about 2000 m in altitude, attached to aquatic vegetation. No information about female feeding habits is available. Discussion: Adults are similar to those of S. anamariae, which has 6 branches in the pupa gill. The presence of 4 pupal gill branches makes S. stellatum similar to S. schmidtmummi and S. (SIOLII) group species; these can be differentiated by the absence of an anterodorsal process on the pupal cocoon and some other characters such as scutum coloration, gonostyle shape and larval morphology. The closest species is S. schmidtmummi, with few differences such as the scutum with 1+1 anterior silvery subtriangular spots, the trichomes not arranged in rows and the pointed apex of the larval postgenal cleft. SIOLII species group (Figs. 75 O-Q.) The S. SIOLII species group has similarities with S. (Psaroniocompsa) s. str. (COSCARÓN, 1987), S. (Cerqueirellum) and S. (Coscaroniellum) and this is the reason that CROSSKEY & HOWARD (1997, 2004) put all the species groups in S. (Psaroniocompsa) s. l. The following diagnosis includes the characters that help to define the SIOLII species group.

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Female: Small species (maximum wing length 1.7 mm). Scutum blackish with 1+1 silvery submedian stripes that join with posterior grayish area. Basal portion of cibarium with sclerotized border and acute teeth with median area scarcely depressed, with small plates. Fronto-ocular triangle relatively deep. Claws without subbasal tooth. Genitalia as in S. (Psaroniocompsa) s. str. Male: Scutum blackish with 1+1 submedian anterior silvery cuneiform spots from narrow to wide joining with lateral grayish area. Gonostylus subtriangular, with one apical strong and blunt spur, and a dorso-longitudinal carina. Pupa: Cocoon slipper shaped. Frontoclypeus and thorax with trichomes single or bifid and tubercles from rounded to acuminate, absent, on anterior and central area of thorax; gill with 4-6 filiform branches. Larva: Cephalic apotome with positive spots. First abdominal segment darkened like band around body. Body segments II-VIII with subconical to rounded dorsal protuberances and integument with abundant and large setae. Distal antennal segment longer than other two. Mandible with 1-2 latero-mandibular processes; without supramarginal setae. Five species are included: S. siolii, S. lourencoi, S. damascenoi, S. tergospinosum and S. guaporense. Simulium (Psaroniocompsa) siolii PY-DANIEL (Figs. 73 L; 75 O-Q.) Simulium (Psaroniocompsa) siolii PY-DANIEL, 1988: 294; CROSSKEY & HOWARD, 1997: 57; 2004: 51. Simulium siolii: HAMADA et al., 2003: 353. Psaroniocompsa siolii: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

(Based on PY-DANIEL, 1988). Female: General coloration black. Scutum blackish bordered laterally and posteriorly by gray pollinose band; frontal illumination shows 1+1 silvery pollinose longitudinal stripes from anterior to posterior border; pilosity composed of recumbent yellowishgreen hair. Pleura blackish with gray pollinosity. Frons relatively wide; fronto-ocular triangle obsolescent. Sc and basal section of R without hair. Legs with hairs single and spatuliform. Claw without subbasal tooth. Eighth sternite with 8-10 setae on each side; gonapophyses with abundant microtrichiae and internal margin parallel; paraproct short. Genital fork as in S. auripellitum. Male: Scutum black with 1+1 anterior cuneiform grayish pruinose spot, that with some illumination joins grayish posterior area; pilosity as in female. Abdomen brown. Ratio of length/width of hind basitarsus = 6.3-7.0. Gonocoxite longer than gonostylus; gonostylus subtrianglular with longitudinal carina and strong apical spur. Pupa: Cocoon slightly projected ventrally, and with small dorsal projection (Fig. 73 L); anterior border reinforced. Length at base 2.5-3.3 mm; at dorsum 2.0-2.5 mm; gills 2.32.5 mm. Gills with 4 terminal branches, with relatively short primary branch; bifurcation of secondary branches at short distance from base. Frontoclypeus and exposed portion of thorax with relatively low number of small tubercles, irregularly arranged in groups; head trichomes single or bifid, thoracic trichomes single to trifid.

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Larva: Maximum length 3.4-4.1 mm. Cephalic apotome with positive spots diffuse, and abundant single and bifid trichomes. Body with acuminate dorsal protuberances 2+2 on segments II-V and 1+1 on VI; ninth segment with 1+1 ventral tubercles (Fig. 75 O). Body integument with abundant hair and scales with several acute projections (Fig. 75 P). Antennae surpassing apex of cephalic fan stem; ratio of proximal, median and distal segments = 1:0.9-1.2:0.8-1.1. Mandible with 2-3 very thin marginal teeth and one curved latero-mandibular tooth surpassing teeth apical border (Fig. 75 Q). Hypostomium with 47 setae on each side; postgenal cleft deep, with postgenal bridge about 1/4 hypostomium length. Anal sclerite with hair single or bifid not obvious. Anal ring with about 63-66 rows with 9-13 hooks each. Rectal papillae with 11-16 diverticula on each lobe. Distribution: Brazil: Amapá, Mato Grosso, Rondônia. Bionomics: This species breeds in small creeks flowing over deciduous leaves. Discussion: Simulium tergospinosum is the closest species, but it has gill branches bifurcated very close to the base and the gill is shorter than the cocoon. Simulium schmidtmummi is also close, but can be differentiated by the gill branching very close to the base, absence on the larval abdomen of dorsal prominences and abundant and long trichomes, postgenal bridge longer than the hypostomium and a trapezoidal male gonostylus. Simulium (Psaroniocompsa) lourencoi PY-DANIEL Simulium (Psaroniocompsa) lourencoi PY-DANIEL, 1988: 300 ; CROSSKEY & HOWARD, 1997: 57; 2004: 51. Psasroniocompsa lourencoi: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

(Based on PY-DANIEL, 1988). Female and Male: A short abstract of a Congress communication (HAMADA & AZEVEDO, 2002) is not sufficient to make a diagnosis. Pupa: Cocoon elongated ventrally a little more than in S. siolii and with small projection dorsally, anterior border scarcely reinforced; length at dorsum 1.7-1.9 mm, at base 2.0-2.1 mm, gills 1.9-2.4 mm. Cephalic and thoracic trichomes long, single or bifid. Tubercles of frontoclypeus and thorax rounded, somewhat abundant. Gills with 4 filaments, with branching similar to that of S. siolii. Larva: Maximum length 3.2-3.7 mm. Head and body with hair and on abdomen on acuminate dorsal projections as in S. siolii, except on prominences of segments VII or VIII. Cephalic apotome with positive spots. Antennal length, proportion of segments, mandible teeth, postgenal cleft and anal sclerite with abundant hairs sclerotized like scales are very similar to those of S. siolii. Anal ring with 61-62 rows of 12-13 hooks each. Rectal papillae with 6-8 diverticula on each lobe. Ninth sternite with 1+1 subtrianglular tubercles. Distribution: Brazil: Goiás. Bionomics: Unknown. Discussion: This species is very similar to S. siolii. Small differences in the degree of abdominal groove sclerotization in the pupa and the greater number of and shorter rectal papillae diverticula, plus weakly defined cephalic apotome ornamentation in the larva, are indicated by the species author as differences between the species.

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Simulium (Psaroniocompsa) damascenoi PY-DANIEL (Figs. 71 U; 72 J, 73 M; 74 I.) Simulium (Psaroniocompsa) damascenoi PY-DANIEL, 1988: 304; 1989: 506-507; CROSSKEY & HOWARD, 1997: 57; 2004: 51; HAMADA et al., 2003: 353.

(Based on PY-DANIEL, 1988 and HAMADA et al., 2003). Female: Wing length 1.6 mm. Scutum black with 1+1 longitudinal wide silvery stripes (Fig. 71 U). Abdominal tergites I-II brown with silvery pollinosity, III-V brown with tergal plates bright; VI-IX with peculiarly pale greenish sheen. Fronto-ocular triangle absent. Claws without subbasal tooth. Eighth sternite setae 9 or 10 in number on each side; gonapophysis, paraproct and genital fork as in S. siolii. Male: Wing length 1.4 mm. Scutum black with 1+1 antero-lateral subquadrate silvery spots (Fig. 72 J). Pleurae brown with silver pollinosity, scutellum and metanotum brown with grayish pollinosity. Abdomen brown. Gonostylus subtriangular with apical spur. Pupa: Cocoon with reinforced border and very small dorsal anterior projection (Fig. 73 M), threads weakly evident. Gill with 4 branches originating from short primary branches (Fig. 74 I). Head without platelets; exposed portion of thorax with only small and acuminate tubercles on posterior half. Trichomes of head and thorax single or bifid. Larva: Maximum length 3.7-3.9 mm. Abdomen with acuminate dorsal protuberances and trichomes as in S. siolii. Protuberances on segment VIII rounded apically. Ninth segment without ventral tubercles. Cephalic apotome with positive spots and scarce, single trichomes. Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.5-1.7:0.9-1.0. Postgenal cleft deep and postgenal bridge shorter than half of hypostomium length. Mandible similar to that of S. siolii. Anal ring with 70-76 rows with 14-17 hooks each. Rectal papillae with 4-6 diverticula on each lobe. Distribution: Brazil: Amapá. Bionomics: This species breeds in rivers of about 25-27 °C and 80-100 m in width in forests and savannas, with rocky bottoms, attaching to Podostomaceae, deciduous leaves and streamside vegetation. Female feeding habits are undescribed. Discussion: The species is similar to S. siolii, S. lourencoi, S. guaporense, S. schmidtmummi and S. stellatum. The first three species show platelets on the pupal frontoclypeus and the exposed portion of the thorax. Simulium schmidtmummi and S. stellatum also have a dorsal anterior projection on the cocoon (but it is larger), absence of dorsal protuberances in the larva, and a trapezoidal gonostylus in the male. In S. siolii and S. lourencoi, the medial antennal segment is proportionally shorter and bifurcation of the gill secondary branches are at a greater distance from the base. Simulium guaporense has 6 pupal gill branches. Simulium (Psaroniocompsa) tergospinosum HAMADA (Figs. 72 K-M; 73 N.) Simulium (Psaroniocompsa) tergospinosum HAMADA, 2000: 819; CROSSKEY & HOWARD, 2004: 51.

(Based on HAMADA, 2000). Female: Wing length 1.7 mm. General color black. Frons and clypeus with silvery blue pollinosity. Scutum black with 1+1 thick light silvery stripes, similar to that of S. damascenoi, with thin golden reddish hair unevenly distributed. Pleurae dark brown with silver pollinosity. Scutellum and metanotum brownish. Sc bare. Legs yellowish brown with

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tarsi black on fore leg; hind leg with femur, tibial apex and terminal tarsomeres dark brown. Abdomen brown with greenish membranous area and basal fringe highlighted with gold. Tergite II with silvery blue pollinosity; VI-VIII slightly bright. Fronto-ocular triangle obsolescent; claw without subbasal tooth. Eighth sternite with about 14 setae on each side; gonapophysis, paraproct and genital fork similar to those of S. siolii. Male: Wing length 1.7 mm. General coloration as in female. Antenna, hair and insertion area with silvery-blue pollinosity. Scutum black with golden reddish hair and 1+1 silvery, anterior submedian cuneiform spots, reaching nearly middle of scutum length (Fig. 72 K). Scutellum and metanotum brownish with silver pollinosity. Leg coloration as in female. Abdomen dark brown with basal fringe gold; tergites II, V-VIII with 1+1 lateral silvery spots. Gonostylus longer than half of gonocoxite length (Fig. 72 L), elongate, subtriangular with dorsal internal longitudinal ridge and strong apical spur (Fig. 72 M). Pupa: Cocoon with basal anterior enlargement and without dorsal projection (Fig. 73 N), with reinforced anterior border, and weave with evident threads. Length at base 2.8 mm, at dorsum 2.4 mm, gills 2.0 mm. Gills with 4 filaments bifurcated close to base with primary branches very short, similar to those of S. damascenoi. Frontoclypeus and anterior portion of thorax without tubercles; posterior exposed portion of thorax with rounded or pointed tubercles. Head trichomes simple to trifid, thoracic trichomes from simple to with 4 branches. Larva: Maximum length 4.3-4.6 mm. Cephalic apotome with light positive spots and small simple trichomes. Antennae surpassing cephalic fan stem apex; ratio of proximal, medial and distal segments = 1:0.8-1.2:1.0-1.1. Mandible with thin marginal teeth and 1-2 latero-mandibular processes, simple or bifid, reaching mandible teeth inferior margin. Body with abundant scales multiply branched. Abdomen with 1+1 acuminate prominences dorsally and laterally on segments II-VI and only dorsally on segment VII. Ninth segment with 1+1 ventral tubercles. Anal sclerite area with short branched setae. Anal ring with 62-69 rows with 12-15 hooks each. Rectal papillae with 8-11 diverticula on each lobe. Distribution: Brazil: Amazonas. Bionomics: This species was collected in large black water river waterfalls and creeks in the Amazonian region, on Podostemaceae and fallen leaves. Females bite humans. Discussion: The closest species is S. damascenoi but it can be differentiated by the presence in the larva of abdominal dorsal prominences on segment VIII and absence of ventral tubercles on the ninth segment. Simulium siolii and S. lourencoi have the pupal gill bifurcating far from the base, and gills as long as or longer than the cocoon. Simulium schmidtmummi has a large dorsal projection on the cocoon and male subquadrate gonostylus. Simulium (Psaroniocompsa) guaporense PY-DANIEL Simulium (Psaroniocompsa) guaporense PY-DANIEL, 1989b: 502; CROSSKEY & HOWARD, 2004: 51. Simulium (Psaroniocompsa) siolii PY-DANIEL, 1988: (in part). Psaroniocompsa guaporense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

(Based on PY-DANIEL, 1989). Female and Male: Unknown. Pupa: Cocoon with reinforced border and small antero-dorsal projection, leaving free 2/3 portion of thorax; weave with evident threads. Gill with 6 terminal

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branches; primary dorsal branch bifurcated close to base. Primary ventral and secondary branches bifurcated near half of total length. Frontoclypeus and exposed portion of thorax with moderate number of small tubercles rounded or acuminate, arranged in irregular groups; tubercles of thorax acuminate on anterior half; trichomes with 3-4 branches. Larva: Maximum length 4.3 mm. Abdomen dorsally with hair and protuberances as in S. siolii, but with 1+1 protuberances on tergite I, presence of well-sclerotized scales, small ventral expansion of each abdominal segment and absence of protuberances on segment VIII. Cephalic apotome scarcely ornamented but with positive spots and abundant single elongated hairs. Postgenal cleft deep with postgenal bridge about 1/5 of hypostomium length. Antennal apex reaching apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9:1.3. Mandible and anal sclerite as in S. siolii. Anal ring with 79 rows with 10-15 hooks each. Rectal papillae diverticula number unknown. Distribution: Brazil: Rondônia. Bionomics: Unknown. Discussion: The SIOLII group species are very similar. The presence of six gill branches allows the pupa to be readily separated from those of the other species. In the larva, the other species have fewer integument scales that are less sclerotized on the terminal portion of the abdomen, and also have dorsal protuberances on segment VIII. Species inquirendae Simulium catarinense PINTO Simulium catarinense PINTO, 1931: 695; COSCARÓN, 1991: 193. Simulium (Psaroniocompsa) catarinense: COSCARÓN & WYGODZINSKY, 1984: 77.

The description is based on a male, with illustrations that allow it to be placed in this subgenus, but which makes it impossible to determine the species level. Distribution: Brazil: Santa Catarina. Simulium guttatum (ENDERLEIN) Pliodasina guttata ENDERLEIN, 1936: 124. Simulium guttatum: VARGAS & DIAZ NÁJERA, 1953b: 137; STONE, 1963: 14; COSCARÓN & WYGODZINSKY, 1984: 77; COSCARÓN, 1987: 24; 1991: 193.

The type material is from Paraguay and deposited at the Berlin Museum. VARGAS & DIAZ NAJERA stated that it is close to S. incrustatum, and STONE considered it as S. incrustatum. We examined the female but it is not well preserved, the coloration is deficient and does not permit us to determine to which species it corresponds. We prefer to maintain it as species inquirendae.

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Simulium (Cerqueirellum) PY-DANIEL (Figs. 76-79; 143.) Type-Species: Simulium amazonicum GOELDI, 1905: 138.

Female: Relatively small, maximum wing length 2.8 mm. General coloration black. Frons and clypeus gray nacreous. Scutum black bordered laterally and posteriorly by silvery pollinose area, and with 1+1 thick wide submedian silvery bands reaching posterior grayish area. These silvery longitudinal bands generally have some light and anterior, internal elongated darker, shadow-like spots. Abdomen with tergites I-IV velvety black, with lateral and posterior borders light grayish, subsequent tergites black with cereous sheen; tergite II with metallic greenish blue sheen at sides. Frons generally wider than high; fronto-ocular triangle absent or very reduced. Female antennae as long as male antennae. Infra-frontal sutures not developed. Sc and basal sector of R without hair; legs with hairs single and spatuliform; calcipala well developed and claws generally without subbasal tooth. Basal portion of cibarium well sclerotized, with abundant acute teeth in several rows, which reach laterally to cornuae and leave (depressed) median area free. Gonapophyses with abundant microtrichiae, straight distal border, and internal rim slightly subparallel; paraproct shorter than high, cercus with subcircular border. Male: Scutum ornamentation as in female or totally black with only 1+1 anterior cuneiform silvery spots. Abdomen black with 1+1 gray nacreous spots on segments II, VI, VII and very small on VIII. Gonocoxite subquadrate wider than long; gonostylus about as long as gonocoxite, subtriangular or subtrapezoidal elongated with 1-2 (exceptionality to 5-7) subconical spurs on or near apex; endoparameres with abundant hooks. Pupa: Cocoon generally slipper shaped, thick weave and frequently anteriorly reinforced. Head with 2 pairs of frontal and 1 pair facial simple trichomes; 5-6 thoracic trichomes simple, bifid, or up to 4 branches. Frontoclypeus and thorax with smooth tubercles, not acute; gill with 4-10 branches. Larva: Maximum length 5.5 mm. Body generally with light coloration showing dark band around first abdominal segment. Body integument without scales or spines, but with some hair. Cephalic apotome, generally scarcely ornamented and without positive spots; with obvious hairs. Gular cleft deep with rounded margin, postgenal bridge about 1/4 or more shorter than hypostomium. Hypostomium median tooth generally below lateral teeth. Antenna as long as or longer than cephalic fan stem, medial article generally longer or as long as distal article. Mandible with only one row of internal teeth; with or without latero-mandibular process and with one row of setae on supramarginal inferior border. Ninth segment with or without pair of ventral tubercle. Anal sclerite with relatively short ventral branches, with enlarged union with dorsal branches. Anal ring with 50-70 rows of 8-14 hooks each. Rectal papillae with 3-9 diverticula. Distribution: Found from central Argentina to Mexico, with greater distribution in the intertropical region (Fig. 143). Discussion: This subgenus includes 11 species. CROSSKEY & HOWARD (1997, 2004) include these species in the AMAZONICUM species group of S. (Psaroniocompsa). This subgenus is differentiated from Psaroniocompsa by the presence on the female scutum of 1+1 intervittae submedian silvery bands with cuneiform elongated interior spots slightly darkened like a shadow; in the male by the subtriangular or subtrapezoidal elongated

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gonostylus with 1-7 spurs; in the pupa by thoracic trichomes with 1-4 branches; and in the larva by a supramarginal setal row on the mandibular inferior border. PY-DANIEL (1988) gave a detailed description of the subgenus and of several species of this taxon, including a species key for all life stages except the egg. SHELLEY et al. (1997, 2000, 2002) also treated some of the species. Here we include a diagnosis of the species, with a key for the different stages. We also include S. delponteianum, the southernmost species, noting several differences from the common species, such as a more grayish colored scutum, greater number of spurs on the gonostylus and the absence of supramarginal setae on the mandible; and S. argentiscutum with different scutum ornamentation, and a shoe-shaped cocoon with nonfilamentous gill branches. In both cases, this is the closest subgenus in which they can be placed. Also, can be objected the inclusion of S. minusculum, differentiated by a subquadrate gonostylus and absence of supramarginal setae on the larval mandible. Another difficult species is S. quadrivittatum, which has been placed here in S. (Coscaroniellum). KEYS

TO

SIMULIUM (CERQUEIRELLUM)

SPECIES

Females 1

–

2(1)

– 3(2)

– 4(3)

Scutum with black, narrow median stripe reduced to thin line on anterior 2/3 and 1+1 black short sublateral stripes; with frontal light showing 1+1 cuneiform anterior submedian intervittae blackish stripes reaching to half length of scutum (Fig. 76 U); with posterior light, the black median and posterior sublateral stripes shorter and intervittae cuneiform spots lighter and shorter, with nacreous sheen (Fig. 76 V) ........................................................................................ Simulium argentiscutum Scutum with black, thick median stripe, and 1+1 black sublateral stripes covering 2/3 of scutum, anterior intervittae spots present or absent, if present, they can be short (Fig. 76 F) or large surpassing half of scutum length (Figs. 76 C, H), sometimes varying according to direction of light ..................................................... 2 Scutum with 1+1 black sublateral stripes short, divergent, covering about 1/2 to 1/3 of scutum, depending on direction of light; with anterior light, the 1+1 short blackish diffuse intervittae spots (Fig. 76 F) evident; with posterior light, intervittae generally not evident ................................................... Simulium amazonicum Scutum with 1+1 black sublateral stripes longer, surpassing 2/3 of scutum; with or without anterior intervittae spots of different length and shape according to light direction ...................................................................................................................... 3 Frons pollinose without sheen; scutum with abundant pilosity, sometimes not allowing limit of stripes to be seen, giving them lead gray tonality (Fig. 76 Y); claws generally with subbasal tooth (Fig. 76 Z); fronto-ocular triangle evident (Fig. 76 W) ............................................................................................... Simulium delponteianum Frons with nacreous or metallic sheen; scutum with sparse pilosity permitting full extension of stripes to be seen; claws generally without subbasal tooth; frontoocular triangle absent (Fig. 76 G) ................................................................................... 4 Anterior cuneiform intervittae spots of scutum shadowed anteriorly, short, not surpassing anterior 1/3 of scutum, varying according to direction of light (Figs. 76 A, B) ...................... Simulium minusculum, Simulium roraimense, Simulium venezuelense

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Anterior cuneiform intervittae spots of scutum shadowed anteriorly, elongate, surpassing anterior 1/2 of scutum, varying according to direction of light (Figs. 76 C, E, H, I, R) ................................................................................................................. 5 Wing length about 2.5 mm ................................................................ Simulium chaquense Wing length about 1.3-2.1 mm ....................................................................................... 6 Wing length about 1.7-2.1 mm ........ Simulium oyapockense, S. cuneatum, S. ganalesense Wing length about 1.3-1.5 mm ...................................................... Simulium sanguineum

Males 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(5) – 7(5) – 8(7) – 9(8) –

Scutum with 1+1 anterior silvery spots joining grayish posterior area (Fig. 77 F) ................................................................................................................................................ 2 Scutum with 1+1 anterior silvery spots, not joining grayish posterior area (Figs. 77 A, C, D, L, R) ...................................................................................................................... 4 Wing length 2.2-2.3 mm; gonostylus generally with 2 apical spurs (Fig. 77 J) ....... .................................................................................................................. Simulium chaquense Wing length 1.2-1.6 mm; gonostylus generally with 1 apical spur as in Figure 77 O ................................................................................................................................................ 3 Scutum with 1+1 silvery submedian stripes relatively wide (Fig. 77 F) ................... ................................................................................................................ Simulium ganalesense Scutum with 1+1 silvery submedian stripes relatively thin, especially on posterior 1/2 (Fig. 77 E) ..................................................................................... Simulium roraimense Gonostylus with 5-10 spurs (Fig. 77 T); scutum with silvery submedian stripes thin anteriorly (Fig. 77 R) ............................................................ Simulium delponteianum Gonostylus with only 1 spur; scutum with silvery submedian stripes wide anteriorly, subtriangular ..................................................................................................... 5 Scutum with 1+1 silvery submedian stripes not reaching 1/2 of scutum length (Fig. 77 A) ............................................................................................................................ 6 Scutum with 1+1 silvery submedian stripes surpassing 1/2 of scutum length (Figs. 77 C, D, K-M) .......................................................................................................... 7 Scutum with 1+1 anterior silvery spots subtrianglular; gonostylus subtriangular with apical spur .................................................................................. Simulium venezuelense Scutum with 1+1 anterior silvery spots comma shaped (Fig. 77 A); gonostylus subtrapezoidal with subapical spur (Fig. 77 B) ........................... Simulium minusculum Scutum with 1+1 silvery submedian stripes not reaching 1/2 of scutum length .............................................................................................................. Simulium amazonicum Scutum with 1+1 silvery submedian stripes reaching nearly to grayish posterior area of scutum (Figs. 77 C, D, K, Q) ............................................................................ 8 Wing length 1.7-2.0 mm ...................................................................... Simulium cuneatum Wing length 1.4-1.5 mm ................................................................................................... 9 Scutum with 1+1 black sublateral stripes wide anteriorly, reaching nearly to anterior border (Figs. 77 C, D) ...................................................... Simulium oyapockense Scutum with 1+1 black sublateral stripes narrow anteriorly, not reaching (Fig. 77 Q), or only laterally reaching (Fig. 77 K) nearly to anterior border ...................... 10

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10(9) –

Scutum with 1+1 silvery submedian stripes posteriorly reaching or very close to posterior grayish area (Fig. 77 K) .................................................. Simulium sanguineum Scutum with 1+1 silvery submedian stripes posteriorly not reaching posterior grayish area (Fig. 77 Q) ................................................................. Simulium argentiscutum

Pupae 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(6) – 8(3) – 9(8) –

Gill with 10 branches (Figs. 78 J, L) ................................................. Simulium cuneatum Gill with 4-8 branches ....................................................................................................... 2 Gill with 4 branches ......................................................................... Simulium venezuelense Gill with 6-8 branches ....................................................................................................... 3 Gill with 6 branches (Figs. 78 B, E, N, O) ................................................................... 4 Gill with 8 branches (Fig. 78 H) ..................................................................................... 8 Cocoon shoe shaped (Figs. 78 M); weave structure spongy, without evident threads; gill short with wrinkled border (Fig. 78 N) ............... Simulium argentiscutum Cocoon slipper shaped (Figs. 78 A, C, G, J), weave structure compact, with evident threads; gill elongated with smooth border (Figs. 78 B, E, O) .................. 5 Species relatively large; cocoon length basally 3.6-4.6 mm; gill length 1.8-2.5 mm; thoracic trichomes simple (Fig. 78 O) ...................................... Simulium delponteianum Species relatively small; cocoon length basally 2.0-2.2 mm; gill length 0.7-1.8 mm; thoracic trichomes with 1-4 branches (Figs. 78 B, E) ................................................ 6 Frontoclypeus and exposed portion of thorax with few tubercles ..... Simulium minusculum Frontoclypeus and exposed portion of thorax with abundant tubercles (Fig. 78 D) ....... 7 Frontoclypeus and exposed portion of thorax with rounded tubercles .................. ........................................................................... Simulium ganalesense, Simulium oyapockense Frontoclypeus and exposed portion of thorax with some rounded but acuminate tubercles on apex of frontoclypeus and dorsomedian area of thorax ................. Simulium roraimense Species relatively large; cocoon length basally 3.0-3.5 mm, 2.5-2.7 mm dorsally; gills 1.8-2.0 mm .................................................................................... Simulium chaquense Species relatively small, cocoon length basally 2.0-2.7 mm, dorsally 1.6-2.3 mm, gills 1.0-1.8 mm .................................................................................................................. 9 Dorsal secondary branch single, ventral branch bifid (Fig. 78 I) ....... Simulium sanguineum Dorsal secondary branch bifid, ventral branch single .............. Simulium amazonicum

Larvae Unknown: Simulium venezuelense. 1 – 2(1) –

Antenna not surpassing apex of cephalic fan stem; postgenal bridge about 1/5-1/6 of hypostomium length; ninth segment without ventral tubercles ..... Simulium argentiscutum Antenna surpassing apex of cephalic fan stem; postgenal bridge about 1/2-1/6 of hypostomium length; ninth segment generally with ventral tubercles .................... 2 Anal ring with 70-78 rows with 14-18 hooks each; mandible without row of supramarginal setae ....................................................................... Simulium delponteianum Anal ring with 40-66 rows with 8-13 hooks each; mandible generally with row of supramarginal setae ............................................................................................................ 3

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Mandible without row of supramarginal setae ........................... Simulium minusculum Mandible with row of supramarginal setae .................................................................. 4 Cephalic fan with 17-22 rays ........................................................................................... 5 Cephalic fan with 37-45 rays ........................................................................................... 8 Mandible without latero-mandibular process .............................................................. 6 Mandible with 1-2 latero-mandibular processes ......................... Simulium sanguineum Maximum length 4.5-5.0 mm; cephalic fan with 34-40 rays ....... Simulium chaquense Maximum length 3.6-4.3 mm; cephalic fan with 16-22 rays ..................................... 7 Postgenal bridge about 1/3 or shorter than hypostomium length; anal ring with 52-58 rows of hooks ....................................................................... Simulium amazonicum Postgenal bridge about 1/2 hypostomium length; anal ring with 60-70 rows of hooks ........................................................................................................ Simulium cuneatum Mandible with 2 latero-mandibular processes; cephalic fan with 27-38 rays .......... ............................................................................................................... Simulium oyapockense Mandible with 1 latero-mandibular process; cephalic fan with 37-42 rays ............. .............................................................................. Simulium roraimene, Simulium ganalesense

As the species of this subgenus are especially differentiated by the adult scutal ornamentation and pupal gills, we will reduce the diagnosis to the principal characters. CLAVES

DE ESPECIES DE

SIMULIUM (CERQUEIRELLUM)

Hembras 1

–

2(1)

– 3(2)

Escudo con una banda negra mediana angosta, reducida a una fina línea en 2/3 anteriores y 1+1 banda negra corta sublateral; con luz frontal presenta 1+1 bandas negruzcas cuneiformes anteriores submedianas, llegando hasta la mitad del largo del escudo (Fig. 76 U); con luz posterior las bandas mediana y posterior sublaterales, se presentan más cortas y las manchas submedianas anteriores cuneiformes también cortas y más claras, con brillo nacarado (Fig. 76 V) ............. ............................................................................................................ Simulium argentiscutum Escudo con banda negra mediana gruesa y 1+1 bandas negras sublaterales cubriendo los 2/3 del largo del escudo; manchas anteriores entre bandas presentes o ausentes, si están presentes pueden ser cortas (Fig. 76 F) o largas sobrepasando la mitad del largo del escudo (Figs. 76 C, H), a veces variando según la dirección de la luz .......................................................................................................... 2 Escudo con 1+1 bandas negras sublaterales cortas, divergentes, cubriendo alrededor de la mitad o un tercio del largo del escudo de acuerdo a la iluminación; con luz anterior presenta 1+1 manchas difusas cortas entre bandas (Fig. 76 F); con luz posterior generalmente estas manchas no son evidentes ............. Simulium amazonicum Escudo con 1+1 bandas negras sublaterales sobrepasando 2/3 del largo del escudo; con o sin manchas anteriores entre bandas presentando diferente longitud y forma según la posición de la luz ................................................................................ 3 Frente polinosa sin brillo; escudo con abundante pilosidad, a veces impidiendo ver el límite de las bandas dando un aspecto general gris plomizo (Fig. 76 Y); uñas

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generalmente con diente subbasal (Fig. 76 Z); triángulo frontoocular evidente (Fig. 76 W) ...................................................................................... Simulium delponteianum Frente con brillo nacarado o metálico; escudo con escasa pilosidad permitiendo ver la extensión de las bandas; uñas generalmente sin diente subbasal; triángulo frontoocular ausente (Fig. 76 G) .................................................................................... 4 Manchas del escudo sombreadas, anteriores, cuneiformes entre bandas de acuerdo a la posición de la luz son cortas, no sobrepasando el 1/3 anterior (Figs. 76 A, B) ...................... Simulium minusculum, Simulium roraimense, Simulium venezuelense Manchas del escudo sombreadas, anteriores, cuneiformes entre bandas, de acuerdo a la posición de la luz son alargadas sobrepasando la mitad del largo del escudo (Figs. 76 C, E, H, I, R) ........................................................................................ 5 Largo del ala alrededor de 2,5 mm .................................................. Simulium chaquense Largo del ala alrededor de 1,3-2,1 mm ......................................................................... 6 Largo del ala alrededor de 1,7-2,1 mm ........................................................................... ........................................... Simulium cuneatum, Simulium ganalense, Simulium oyapockense Largo del ala alrededor de 1,3-1,5 mm ........................................ Simulium sanguineum

Machos 1 – 2(1) – 3(2) – 4(2) – 5(4) – 6(5) – 7(5)

Escudo con 1+1 manchas anteriores plateadas uniéndose al área posterior grisácea (Fig. 77 F) ............................................................................................................. 2 Escudo con 1+1 manchas anteriores plateadas no uniéndose al área posterior grisácea (Fig. 77 A, C, D, L, R) ....................................................................................... 4 Longitud ala 2,2-2,3 mm; gonostilo generalmente con dos espolones apicales (Fig. 77 J) ................................................................................................ Simulium chaquense Longitud ala 1,2-1,6 mm; gonostilo generalmente con un espolón apical similar a (Fig. 77 O) ........................................................................................................................... 3 Escudo con 1+1 bandas plateadas submedianas relativamente anchas (Fig. 77 F) ................................................................................................................ Simulium ganalesense Escudo con 1+1 bandas plateadas submedianas relativamente finas, especialmente sobre la 1/2 posterior (Fig. 77 E) ..................................................... Simulium roraimense Gonostilo con 5-10 espolones (Fig. 77 T); escudo con bandas plateadas submedianas finas anteriormente (Fig. 77 R) .......................... Simulium delponteianum Gonostilo con solamente 1 espolón; escudo con bandas plateadas submedianas anchas anteriormente en forma triangular ................................................................... 5 Escudo con 1+1 bandas plateadas submedianas no llegando a la mitad del largo del escudo (Fig. 77 A) ....................................................................................................... 6 Escudo con 1+1 banda plateada submediana sobrepasando la mitad del largo del escudo (Figs. 77 C, D, K-M) ............................................................................................ 7 Escudo con 1+1 bandas plateadas submedianas subtriangulares; gonostilo de forma subtriangular con espolón apical ....................................... Simulium venezuelense Escudo con 1+1 bandas plateadas en forma de coma (Fig. 77 A); gonostilo subtrapezoidal con espolón subapical (Fig. 77 B) ...................... Simulium minusculum Escudo con 1+1 bandas plateadas submedianas no alcanzando a la mitad del largo del escudo ................................................................................ Simulium amazonicum

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Escudo con 1+1 banda plateada submediana llegando cerca del área posterior grisácea (Figs. 77 C, D, K, Q) .......................................................................................... 8 Longitud ala 1,7-2,0 mm ..................................................................... Simulium cuneatum Longitud ala 1,4-1,5 mm .................................................................................................. 9 Escudo con 1+1 bandas negras sublaterales anchas anteriormente y llegando cerca del borde anterior (Figs. 77 C, D) ....................................... Simulium oyapockense Escudo con 1+1 bandas negras sublaterales angostas anteriormente no llegando (Fig. 77 Q), o sólo lateralmente (Fig. 77 K) cerca del borde anterior .................. 10 Escudo con 1+1 bandas plateadas submedianas llegando posteriormente o muy cerca del área grisácea posterior (Fig. 77 K) ............................... Simulium sanguineum Escudo con 1+1 bandas plateadas submedianas no llegando posteriormente al área posterior grisácea (Fig. 77 Q) ............................................. Simulium argentiscutum

Pupas 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(6) – 8(3) – 9(8) –

Branquias con 10 ramas (Figs. 78 J, L) ............................................. Simulium cuneatum Branquia con 4-8 ramas .................................................................................................... 2 Branquias con 4 ramas ..................................................................... Simulium venezuelense Branquias con 6-8 ramas .................................................................................................. 3 Branquia con 6 ramas (Figs. 68 B, E, N, O) ................................................................. 4 Branquia con 8 ramas (Fig. 78 H) .................................................................................. 8 Capullo en forma de zapato (Fig. 78 M); estructura del tejido esponjoso, sin fibras evidentes; branquia corta y de borde rugoso (Fig. 78 N) ..... Simulium argentiscutum Capullo en forma de chinela (Figs. 78 A, C, G, J) estructura del tejido compacta, con fibras evidentes; branquia alargada con borde liso (Figs. 78 B, E, O) ............ 5 Especies relativamente grandes, largo del capullo en la base 3,6-4,6 mm; largo de la branquia 1,8-2,5 mm; tricomas torácicos simples (Fig. 78 O) ....................... ........................................................................................................... Simulium delponteianum Especies relativamente pequeñas; longitud del capullo en la base 2,0-2,2m, largo branquia 0,7-1,8 mm; tricomas torácicos con 1-4 ramas (Figs. 78 B, E) ............... 6 Frontoclípeo y porción expuesta del tórax con escasos tubérculos .......................... ............................................................................................................... Simulium minusculum Frontoclípeo y porción expuesta del tórax con abundantes tubérculos (Fig. 78 D) ................................................................................................................................................ 7 Frontoclípeo y porción expuesta del tórax con tubérculos redondeados ................ ........................................................................... Simulium ganalesense, Simulium oyapockense Frontoclípeo y porción expuesta del tórax con tubérculos algunos redondeados pero aguzados en el ápice en frontoclípeo y área dorso-mediana del tórax ........... ................................................................................................................. Simulium roraimense Especies relativamente grandes; largo del capullo en la base 3,0-3,5 mm; en el dorso 2,5-2,7 mm; branquias 1,8-2,0 mm ....................................... Simulium chaquense Especies relativamente pequeñas; largo capullo en la base 2,0-2,7 mm; en el dorso 1,6-2,3 mm; branquias 1,0-1,8 mm ................................................................................. 9 Rama secundaria dorsal simple, ventral bífida (Fig. 68 I) ........ Simulium sanguineum Rama secundaria dorsal bífida, ventral simple ........................... Simulium amazonicum

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Larvas Desconocidos: Simulium venezuelense. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(6) – 8(4) –

Antena no sobrepasando el ápice del tallo del abanico cefálico; puente postgenal aproximadamente 1/5-1/6 del largo del hipostomio; noveno segmento sin tubérculos ventrales ....................................................................... Simulium argentiscutum Antena sobrepasando el ápice del tallo del abanico cefálico; puente postgenal aproximadamente 1/2-1/6 del largo del hipostomio; noveno segmento generalmente con divertículos ventrales ....................................................................... 2 Anillo anal con 70-78 hileras con 14-18 ganchos cada una; mandíbulas sin hileras de cerdas supramarginales ........................................................... Simulium delponteianum Anillo anal con 40-66 hileras con 8-13 ganchos cada una; mandíbula generalmente con una hilera de cerdas supramarginales ..................................................................... 3 Mandíbula sin hilera de setas supramarginales ........................... Simulium minusculum Mandíbula con hilera de setas supramarginales ........................................................... 4 Abanico cefálico con 17-22 rayos ................................................................................... 5 Abanico cefálico con 37-45 rayos ................................................................................... 8 Mandíbula sin proceso latero-mandibular .................................................................... 6 Mandíbula con 1-2 procesos latero-mandibular ......................... Simulium sanguineum Longitud máxima 4,5-5,0 mm; abanico cefálico con 34-40 rayos ..... Simulium chaquense Longitud máxima 3,6-4,3 mm; abanico cefálico con 16-22 rayos ........................... 7 Puente postgenal aproximadamente 1/3 más corto que el largo del hipostomio; anillo anal con 52-58 hileras de ganchos .................................... Simulium amazonicum Puente postgenal aproximadamente 1/2 del largo del hipostomio; anillo anal con 60-70 hileras de ganchos ..................................................................... Simulium cuneatum Con 2 procesos latero-mandibulares; abanico cefálico con 27-38 rayos .................. ............................................................................................................... Simulium oyapockense Con 1 proceso látero-mandibular; abanico cefálico con 37-42 rayos ....................... ............................................................................. Simulium roraimense, Simulium ganalesense

Siendo que las especies de este subgénero son diferenciados por la ornamentación del escudo de los adultos y por las branquias de las pupas, reduciremos los diagnósticos a los caracteres principales. Simulium (Cerqueirellum) minusculum LUTZ (Figs. 76 A, B; 77 A, B; 78 A, B; 79 D.) Simulium minusculum LUTZ, 1910: 253; NATHAN et al., 1982: 221-222; SHELLEY et al., 1982: 4-8, 22-25; COSCARÓN, 1983: 90. Psaroniocompsa minuscula ENDERLEIN, 1936: 123; PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 124. Simulium (Psaroniocompsa) minusculum PY-DANIEL, 1983: 181; CROSSKEY & HOWARD, 1997: 56; 2004: 50; SHELLEY et al., 2000: 181-182. Simulium (Cerqueirellum) minusculum: COSCARÓN, 1987: 24; 1991: 199-200.

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A

B

D

C

333

F

P K

E

O

I

H

R L

Q

M G

U

Z

V Y

S W

X

N

T

J

Figure 76. Female. A-B: S. (Cerqueirellum) minusculum, scutum with different illumination. C-D: S. (C.) oyapockense, scutum with different illumination. E: S. (C.) ganalesense, scutum. F: S. (C.) amazonicum, scutum (redesigned from SHELLEY et al., 1982). G-K: S. (C.) chaquense; G: fronto-ocular triangle; H-I: scutum with different illumination; J: hair of legs; K: abdomen, lateral view. L-M: S. (C.) sanguineum, scutum with different illumination (from TIDWELL et al., 1981). N-T: S. (C.) cuneatum; N: basal portion of cibarium; O-R: scutum with different illumination (O-P: lateral view, Q-R dorsal view); S: eighth sternite, gonapophyses, genital fork and spermatheca; T: cercus and paraproct (left mounted in slide). U-V: S. (C.) argentiscutum, scutum with different illumination. W-Z: S. (C.) delponteianum; X: basal portion of cibarium; W: fronto-ocular triangle; Y: scutum; Z: claws.

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A

C

F

D

G

E

H

K

I

R L

M

Q

B

S

N

P J

O T

Figure 77. Male. A-B: S. (Cerqueirellum) minusculum; A: scutum; B: gonostylus. C-D: S. (C.) oyapockense, scutum , with different illumination. E: S. (C.) roraimense, scutum. F: S. (C.) ganalesense, scutum. G-J: S. (C.) chaquense; G: scutum; H-I: abdomen in dorsal and lateral position; J: gonocoxite and gonostylus. K: S. (C.) sanguineum, scutum. L-P: S. (C.) cuneatum; L-M: scutum with different illumination; N: genitalia in ventral view showing gonocoxite, gonostylus, ventral plate and endoparameres; O: gonostylus dorsal view; P: ventral plate. Q: S. (C.) argentiscutum, scutum. R-T: S. (C.) delponteianum; R-S: scutum (S: lateral view); T: gonocoxite and gonostylus.

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Female: Wing length 1.8-2.0 mm. Frons slightly nacreous; clypeus with grayish pruinosity. Scutum velvety black with 1+1 submedian silvery stripes reaching grayish posterior area (Figs. 76 A, B), anteriorly on intervittae area 1+1 short cuneiform spots (lighter with posterior illumination and blackish with anterior illumination) that can be more elongated as in figure 76 B. Fronto-ocular triangle small; cibarium and genitalia as in S. cuneatum. Male: Wing length 1.4-1.6 mm. Scutum black with 1+1 anterior cuneiform silvery spots with length 1/5 to 1/2 of scutum length (Fig. 77 A). Gonostylus subtrapezoidal with submedian spur with truncate apex (Fig. 77 B). Pupa: Cocoon weave with threads obvious, slightly reinforced border and small dorsal projection (Fig. 78 A); length at base 2.0-2.3 mm, on dorsum 1.8-2.0 mm, gill 0.91.3 mm. Gills with 6 branches bifurcated close to base (Fig. 78 B). Frontoclypeus without tubercles, thorax with scarce tubercles acuminate on dorsum; cephalic trichomes single or bifid; thoracic trichomes with 2-4 branches. Larva: Maximum length 3.4 mm. Cephalic apotome with some positive spots; few evident and scarce hairs. Antenna surpassing stem apex of cephalic fan as in figure 79 D; ratio of proximal, medial and distal segments = 1:1:1. Postgenal bridge a little shorter than hypostomium length. Mandible without latero-mandiblular process; row of supra-marginal setae absent. Cephalic fan with 27-34 rays. Anal sclerite without hair or scales. Anal ring with 53-58 rows of 14-18 hooks each. Body integument with trichomes single or bifid. Distribution: Argentina: Corrientes, Jujuy, Misiones, Salta; Brazil: Amazonas, Bahia, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Piauí, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, São Paulo, Tocantins; Guyana; Venezuela: Bolivar. Bionomics: This species breeds in rivers in areas of relatively fast-flowing water, on aquatic vegetation. It shows intense anthropophily and is implicated as a vector of Mansonella ozzardi in Guyana (NATHAN et al., 1982) and Onchocerca volvulus in Toototobi, Brazil (SHELLEY et al., 1982: 24). Discussion: Simulium oyapockense, S. roraimense and S. ganalesense have gills with 6 branches and similar scutum ornamentation, but they have a subtriangular gonostylus with a spur on the apex. They also have abundant frontoclypeal and thoracic tubercles. The absence of supramarginal setae on the larval mandible and subtrapezoidal gonostylus suggest placing S. minusculum in the INCRUSTATUM group of S. (Psaroniocompsa). The wide submedian vittae, similar to those of Cerqueirellum species, support its inclusion in S. (Cerqueirellum). Simulium (Cerqueirellum) oyapockense FLOCH & ABONNENC (Figs. 76 C, D; 77 C, D; 78 C-F; 79 A-C.) Simulium oyapockense: FLOCH & ABONNENC, 1946b: 4; RAMÍREZ PÉREZ, 1983: 60; SHELLEY, 1988: 341. Simulium (Cerqueirellum) oyapockense: PY-DANIEL, 1983: 168; COSCARÓN, 1987: 25; COSCARÓN & COSCARÓN ARIAS, 2000a: 179-182; COSCARÓN ARIAS, 2003: 196, 204, 208, 215. Cerqueirellum oyapockense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium (Psaroniocompsa) oyapockense: SHELLEY et al., 1997: 19; CROSSKEY & HOWARD, 1997: 56; 2004: 50 (cytoforms). Simulium pseudosanguineum RAMÍREZ PÉREZ & PETERSON, 1981a: 154 =159. Simulium sanchezi RAMÍREZ PÉREZ, YARZABAL & PETERSON, 1982: 71. Simulium cuasisanguineum RAMÍREZ PÉREZ, YARZABAL & PETERSON, 1982: 36.

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Simulium pseudoamazonicum RAMÍREZ PÉREZ & PETERSON, 1981a: 151-154. Several cytoforms and morphoforms are mentioned by CROSSKEY & HOWARD, 1997: 56.

Female: Wing length 1.6-2.1 mm. General coloration blackish, with silver pruinosity. Frons silver, iridescent, clypeus silver pruinose. Scutum with three black vittae bordered by silver pruinosity, submedian silver stripes showing elongated shadow comma-shaped mark with antero-lateral illumination, or when observed laterally with antero-superior light position (Fig. 76 C); with anterior superior illumination black median vittae narrower and more elongated posteriorly, submedian shadow intervittae elongated anteriorly and commashaped marks shorter (Fig. 76 D); with posterior illumination black vittae slightly shorter and silver area increased, with submedian bands reaching anterior scutal margin and lacking anterior comma-shaped marks. Scutum pilosity golden, metanotum and scutellum black. Abdomen dark brown to black with 1+1 silver spot on tergite II; tergites III-V velvety black with narrow postero-lateral area silver pruinose, tergite VI-VIII shiny and waxy. Frons with subparallel sides slightly convergent ventrally, fronto-ocular triangle absent, basal portion of cibarium with 23-27 acuminate teeth on each side, extending from borders of deep central area to cornua; sensory vesicle of palp 1/2 or nearly 1/3 length of palpomere; claws without subbasal tooth; paraproct short with rounded border; sternite VIII with 14-18 hairs and gonapophysis slowly divergent below with sclerotized borders. Male: Wing length 1.7-1.8 mm. Coloration similar to that of female. Scutum with 1+1 black, submedian, silvery cunae extending for more than half of scutum length, but generally not joining with silvery posterior area (Figs. 77 C-D); length of cunae change with light position. Genitalia in accordance with the morphology of subgenus. Gonocoxite subquadrate, about as long as high; gonostylus subtriangular with strong club-shaped spine; ventral plate with short basal arms and rounded border. Pupa: Cocoon length on dorsum 2.1-2.6 mm (¯x = 2.3 mm) on base 2.5-3.2 mm (¯x = 2.7 mm) with light yellowish brown coloration. Cocoon slipper shaped, slightly protruding dorsally and basally, and covering gill base (Fig. 78 C). Cocoon made of soft and thick silk with closely woven threads obscuring pupa; gill length 1.2-1.4 mm (n = 5), (¯x = 1.3 mm). Frontoclypeus with moderately abundant rounded platelets; facial trichomes bifid with 1+1 or 2+2 single frontal trichomes (Fig. 78 D). Respiratory gill with six filamentous branches subdivided near base, with short basal trunk and three short primary branches (Fig. 78 E). Disc of thorax with numerous rounded platelets, but with some pointed on posterior area and 5+5 trichomes mainly bifid (Fig. 78 F). Larva: Maximum length 4.1-4.7 mm (¯x = 4.3 mm). Color (in alcohol) yellowish white with greenish spots (Fig. 79 A) and circular band around anterior third of abdomen; head yellowish grey; cephalic apotome light brownish at base, with pale median longitudinal stripe and 1+1 submedian spots, covered by abundant long and single to bifid trichomes leaving free only the pigmented area (Fig. 79 C). Deep subcircular postgenal cleft. Ratio of hypostomium/postgenal bridge = 2.0-2.7; hypostomial teeth at same level, with 4+4 to 5+5 lateral setae and 7-8 lateral serrations on each side. Antenna thin, with ratio of segments: proximal, medial and distal = 1:0.9-1.1:1-1.2. Mandible with 5-6 internal teeth, 2 marginal teeth, (the second small and isolated from anterior), fringe of supramarginal setae and presence of 2 (3) latero-mandibular processes as Fig. 79 B. Cephalic fan with 2738 principal rays. Thoracic proleg lateral sclerite with about 40 teeth in 7-8 groups. Body

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K F Figure 78. Pupa. A-B: S. (Cerqueirellum) minusculum; A: general view; B: gill and trichomes with high magnification. C-F: S. (C.) oyapockense; C: general view; D: frontoclypeus and trichomes; E: gill; F: thoracic trichomes with high magnification. G-H: S. (C.) chaquense: G: general view; H: gill and thoracic trichomes. I: S. (C.) sanguineum, portion of thorax with trichomes and base of gill. J-L: S. (C.) cuneatum; J: general view; K: frontoclypeus and trichomes; L: gill and thoracic trichomes. M-N: S. (C.) argentiscutum (from SHELLEY & LUNA-DIAS, 1980); M: general view; N: gills. O: S. (C.) delponteianum, portion of thorax with trichomes and gill.

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Figure 79. Larva. A-C: S. (Cerqueirellum) oyapockense; A: lateral view; B: apical portion of mandible, showing teeth and supra-marginal setae; C: cephalic apotome. D: S. (C.) minusculum, antenna. E-G: S. (C.) chaquense; E: antenna; F-G: rectal papillae in dorsal and lateral views. H-M: S. (C.) cuneatum; H: head, dorsal view; I: antenna; J: postgenal cleft and hypostomium; K: anterior border of hypostomium; L: lateral sclerite of thoracic proleg; M: anal sclerite and hairs of the area.

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integument with abundant palmate trichomes 0.02 mm in length. Anal sclerite with few hairs. Anal ring with 57-63 rows with 12-15 hooks on each. Rectal papillae with 2-3 diverticula on each lobe. Distribution: Argentina: Salta, Santiago del Estero, Tucumán; Brazil: Amapá, Amazonas, Mato Grosso, Minas Gerais, Pará, Paraná, Rondônia, Roraima; Colombia: Vaupes; Ecuador: Manabí; French Guiana; Guyana; Paraguay: Maracayú; Uruguay; Venezuela: Amazonas, Bolivar, Guarico. Bionomics: This species breeds in plains rivers with swift water, on aquatic vegetation. Females bite humans and are considered vectors of Onchocerca volvulus (SHELLEY 1988: 341). Discussion: The female and male, especially with regard to the scutum ornamentation, are very difficult to separate from the other species of the S. (Cerqueirellum) = (amazonicum) group, such as S. minusculum, S. roraimense, S. ganalesense, S. amazonicum, S. chaquense, S. sanguineum, S. venezuelense and S. cuneatum. The pupal gills with 4-8-10 filaments allow S. oyapockense to be separated from five species. The closest species is S. ganalesense, which can be differentiated only by the male scutum with submedian silvery stripes joining with the grayish posterior area, which also occurs in S. roraimense (and exceptionally, too, in some specimens of S. oyapockense). The female is less sclerotized on the dentate border of the cibarium. Simulium minusculum and S. roraimense can be differentiated by the short intervittae shadow stripe, which does not surpass the scutum anterior one third. The male of S. minusculum has a subtrapezoidal gonostylus with a subapical spur; the pupal frontoclypeus and thorax have sparse tubercles; and the larval mandible does not have supramarginal setae. The S. roraimense pupal gills show primary ventral branch bifurcation at greater distances from the base than do the secondary branches. The difficulties of separating S. oyapockense from S. roraimense and S. ganalesense based on morphological characters make it necessary to use other methods to separate them. Simulium (Cerqueirellum) ganalesense VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 76 E; 77 F.) Simulium (Byssodon) ganalesense VARGAS et al., 1946: 114, 165; VARGAS & DÍAZ NÁJERA, 1957b: 163, 253-254; COSCARÓN, 1987: 16. Simulium (Cerqueirellum) ganalesense: COSCARÓN & IBAÑEZ-BERNAL, 1994: 1-7. Simulium (Psaroniocompsa) ganalesense: CROSSKEY & HOWARD, 1997: 56; 2004: 50; SHELLEY et al., 2002: 155-158.

Female: Wing length 1.6-1.9 mm. General coloration (material discolored) dark brown; cephalic appendages grayish-brown, frons iridescent. Scutum dark brown with golden hair; silvery longitudinal band wide anteriorly and 1+1 elongated shadow bands (Fig. 76 E). Abdomen black with median posterior subrectangular tergite II and subquadrate dark brown tergites III-V. Frons slightly convergent; fronto-ocular triangle absent and claw without subbasal tooth. Basal portion of cibarium with blunt teeth on lateral area. Paraproct very short and scarcely surpassing cercus. Eighth sternite with 8-10 setae on each side. Male: Wing length 1.6 mm. Scutum black with 1+1 silvery bands reaching posterior grayish area (Fig. 77 F). Ratio of length/width of hind basitarsus = 6.0-6.5. Gonostylus subtriangular with big apical spur.

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Pupa: Cocoon with slight antero-ventral projection and reinforced border; thin weave with evident threads; length on base 1.6 mm, on dorsum 1.4 mm; gills 1.2 mm. Frontoclypeus and exposed portion of thorax with rounded tubercles; frontal trichomes single, facial trichomes bifid, thoracic trichomes with 1-3 branches. Gill with 6 branches bifurcated relatively close to base. Larva: Maximum length 3.5-4.5 mm. Cephalic apotome scarcely ornamented, with abundant single and bifid trichomes. Ratio of antennal articles: proximal, medial and distal = 1:1.1:1.2-1.3. Mandible with one latero-mandibular process. Cephalic fan with 20-22 principal rays. Postgenal cleft acuminate distally, postgenal bridge half of hypostomium length. Hypostomium median tooth surpassing lateral teeth. Anal sclerite with 1-3 rows of scales and scarce hairs intermixed. Anal ring with 54-68 rows with 9-13 hooks each; rectal papillae with 3-6 diverticula on each lobe. Distribution: Belize: Cayo; Mexico: Morelos, San Luis de Potosí, Tabasco. Bionomics: In Belize, S. guanalense was collected in rivers about 50 m wide to flat creeks 2 m wide, flowing over submerged grasses. In this country, females bite humans and are very voracious. No information is available from Mexico. Discussion: The female thoracic ornamentation is very similar to that of S. chaquense, S. sanguineum and S. cuneatum. Different numbers of gill branches permit S. guanalense to be separated from the species that have a greater number of branches. Simulium oyapockense and S. roraimense, which also have six branches, can be separated in the male by the scutum with 1+1 submedian cuneiform silvery stripes not reaching the posterior grayish area, and in the female by the scutum with 1+1 submedian silvery stripes narrower with shorter shadow spots; in S. roraimense the shadow intervittae spots are shorter. Belize material shows some differences in the male scutum, pupal size and distances of gill bifurcation (SHELLEY et al 2002: 157). The authors believe that it is necessary to have more material and to use cytological and molecular studies to determine if these variations are inter- or intraspecific variation. Simulium (Cerqueirellum) roraimense NUNES DE MELLO (Fig. 77 E.) Simulium roraimense NUNES DE MELLO, 1974: 45; RAMÍREZ PÉREZ et al., 1982: 67; SHELLEY et al. , 1982: 22; RAMÍREZ PÉREZ, 1983: 66; SHELLEY et al., 1987: 22. Simulium (Cerqueirellum) roraimense PY-DANIEL, 1983: 171; COSCARÓN, 1987: 25; 1991: 196, 198, 199. Cerqueirellum roraimense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium (Psaroniocompsa) roraimense: CROSSKEY & HOWARD, 1997: 56; 2004: 50.

Description extracted in part from RAMÍREZ PÉREZ et al., 1982; SHELLEY et al., 1982; PY-DANIEL, 1983. Female: Wing length 1.5-1.6 mm. Color very close to that of S. oyapockense, S. ganalesense and S. minusculum. Scutum with 1+1 silvery submedian stripes showing, with some light, a shadow cuneiform inter-vittae spot, shorter in S. minusculum and a little longer than in S. amazonicum. Claw without subbasal tooth. Genitalia as in S. cuneatum. Male: Wing length 1.2-1.5 mm. Scutum black with 1+1 silvery submedian stripes, reaching grayish posterior area (Fig. 77 E). Scutum hair yellowish. Genitalia as in S. cuneatum.

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Pupa: Cocoon with small dorsal anterior projection, and anterior border reinforced; weave with evident threads. Length at base 2.1-2.2 mm, at dorsum 1.8 mm, gills 1.7-1.8 mm. Gill with 6 branches showing primary ventral bifurcation at greater distance from base than for secondary branches. Frontoclypeus and exposed portion of thorax with tubercles rounded and acuminate. Head and thoracic trichomes simple and bifid. Larva: Maximum length 3.4-3.7 mm. Cephalic apotome with positive spots and short trichomes with 1-3 branches. Antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1-1.5:1-1.4. Mandible with simple lateromandibular process. Cephalic fan with 40-42 rays. Postgenal cleft subcirclular, postgenal bridge half of hypostomium length. Anal ring with about 68 rows with 11-13 hooks each. Rectal papillae with 6-9 diverticula on each lobe. Distribution: Brazil: Roraima; Venezuela: Amazonas. Bionomics: The species was collected in rivers on aquatic vegetation and submerged tree branches. Discussion: Similarities in the female scutum and pupal gill branches with S. oyapockense and S. minusculum can produce misidentifications, but the silvery submedian stripes of the male scutum in the mentioned species do not reach the grayish posterior area, and bifurcation of the pupal primary ventral branch is lower, not drawn away from the bifurcations of the secondary branches. The closest species is S. ganalesense; only small differences in length of the intervittae submedian shadow spots and proportionately shorter pupal gill permit its separation. Here, also cytological or molecular studies will be necessary to assure the differentiation of these species. Simulium (Cerqueirellum) amazonicum GOELDI (Fig. 76 F.) Simulium amazonicum GOELDI, 1905: 138; SHELLEY, 1982: 67; 1988: 203; SHELLEY et al., 1982: 8-21. Simulium (Cerqueirellum) amazonicum: PY-DANIEL, 1983: 167. Simulium (Psaroniocompsa) amazonicum: CROSSKEY & HOWARD, 1997: 56; 2004: 50. Simulium tallaferroae RAMÍREZ PÉREZ, 1971: 339. Cerqueirellum amazonicum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

Description from SHELLEY et al., 1982 and PY-DANIEL, 1983 gives details. Female: Wing length 1.2-1.9 mm. Frons and clypeus black with bluish silver pruinosity. Scutum with anterior illumination showing black with silver gray pruinosity, leaving median black and 1+1 sublateral bands, and 1+1 diffuse shadow-like spots on anterior border (Fig. 76 F); with posterior light, the median band is wider posteriorly and shorter sublaterally; scutum with silver-yellow appressed hair. Abdomen with tergite I brownish black, tergites IIV velvety black, anterior and posterior margins silver pruinose, tergites VI-IX shiny black. Eighth sternite with about 12 setae on each side. Claws without subbasal tooth. Male: Coloration generally as in female. Scutum black bordered with silver pruinosity that extends as 1+1 wide triangular wedge submedially, similar to that of S. argentiscutum, but not reaching half scutum length. Genitalia as in S. cuneatum. Pupa: Cocoon with reinforced border and small dorsal projection. Length at base 2.0-2.7 mm; dorsally 1.6-2.3 mm; gills 1.0-1.5 mm. Gill with 8 filaments, with branching relatively close to base, arrangement similar to that of S. chaquense but with some variations as shown by

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SHELLEY et al. (1982). Frontoclypeus and thorax with small numerous acuminate tubercles; trichomes bifid except frontals that can be single, those on thorax with 3 or 4 branches. Larva: Maximum length 3.6-4.3 mm. Color yellowish to brown with gray mottling. Cephalic apotome scarcely ornamented, with positive spots and with numerous hairs from thin and single to thicker with 2-4 branches. Antenna surpassing stem apex of cephalic fan. Ratio of antennal segments: proximal, medial and distal = 1:0.9-1: 1-1.1. Postgenal cleft large with rounded margin; postgenal cleft 1/3 or shorter than hypostomium length. Hypostomium with 3-4 lateral setae on each side. Mandible with 1 or 2 marginal teeth; latero-mandibular process absent. Cephalic fan with 12-15 rays. Thoracic proleg lateral sclerite with about 15-17 teeth arranged in about 8 process. Body integument with spines. Ventral papillae obsolescent. Rectal papillae with about 8 diverticula on each lobe. Anal ring with 52-58 rows with 9-12 hooks each. Distribution: Brazil: Acre, Amazonas, Roraima; Colombia: Amazonas; Venezuela: Amazonas, Apure, Barinas, Guarico. Bionomics: Immature stages live in large rivers on herbaceous vegetation and submerged bushes at places with fast current up to 15 m deep. This species is a very dangerous pest, attacking humans, horses, cows and sheep in great quantity. It is mentioned as a vector of Mansonella ozzardi and Onchocerca volvulus. Discussion: The closest species are those with 8 filamentous branches in the pupa, such as S. chaquense and sanguineum. Simulium sanguineum can be differentiated by the branching of the gill (Fig. 78 I), and in S. amazonicum, as for S. chaquense, in the dorsal branch (Fig. 78 H). Simulium chaquense can be differentiated in the pupa by the large size and presence of small spines on abdominal tergites III-IV; in the female by the 1+1 anterolateral vittae more elongated and in the male by the 1+1 silvery submedian stripe reaching the posterior grayish area. Based on the original description of S. tallaferroae, the synonymy proposed by SHELLEY et al. (1982: 8) is accepted. Simulium (Cerqueirellum) chaquense COSCARÓN (Figs. 76 G-K; 77 G-J; 78 G, H; 79 E-G.) Simulium chaquense COSCARÓN, 1971: 33-41; SHELLEY et al., 1980: 785; SHELLEY et al., 1982: 8; PYDANIEL, 1983: 170; COSCARÓN, 1983: 81. Simulium (Cerqueirellum) chaquense: PY-DANIEL, 1981: 170; COSCARÓN, 1987: 24; 1991: 201. Simulium (Psaroniocompsa) chaquense: CROSSKEY & HOWARD, 1997: 56; 2004: 50.

As in other species of this group, large differences occur in the scutum ornamentation. Female: Wing length 2.5 mm. Scutum black with 1+1 silvery submedian bands reaching posterior grayish area, with longitudinal shadow elongated nacreous band over silvery bands, showing variation in vittae shape, in accordance with illumination (Figs. 76 H, I). Abdomen blackish with pattern as in figure 76 K. Fronto-ocular triangle absent (Fig. 76 G); legs with single and spatuliform hairs (Fig. 76 J). Genitalia as in S. cuneatum. Male: Wing length 2.2-2.3 mm. Scutum ornamentation very similar to that of female, with 1+1 silvery submedian bands reaching posterior grayish area (Fig. 77 G), but in some specimens not reaching posterior area, as in S. cuneatum. Abdomen coloration as in figures 77 H, I. Gonostylus about as long as gonocoxite, subtriangular in shape with blunt apex and 1-3 apical spurs (Fig. 77 J). Other genital pieces as in S. cuneatum.

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Pupa: Cocoon with very short ventral projection and reinforced anterior border (Fig. 78 G); length at base 3.0-3.5 mm, dorsally 2.5-2.7 mm, gill 1.8-2.0 mm. Gill with 8 branches with branching near base, showing bifurcation in both secondary dorsal branches (Fig. 78 H); this bifurcation can be at greater distances from base. Frontoclypeus and thorax with rounded to acuminate tubercles, especially over median longitudinal area of thorax in moderate to abundant quantity; trichomes from single to three branched. Larva: Maximum length 4.5-5.0 mm. Cephalic apotome yellowish brown, scarcely ornamented, with few evident spots and with hairs of 1-3 branches. Postgenal cleft deep, subcirclular anteriorly; postgenal bridge about half of hypostomium length. Antenna surpassing apex of cephalic fan, ratio of proximal, media and distal segments = 1:1:1.1 (Fig. 79 E). Mandible with marginal tooth and one latero-mandibular process (only visible on the most southern specimen); cephalic fan with 36-40 ray; anal ring with about 62-66 rows of 11-12 hooks each. Rectal papillae with 4-6 diverticula on each lobe (Figs. 79 F, G). Anal sclerite with scarce scales and single hair. Distribution: Argentina: Buenos Aires, Chaco, Corrientes, Formosa, Salta, Santa Fe, Santiago del Estero, Tucumán. Bionomics: This species breed in small creeks with low to regular flow, but also in medium (Salado River) to exceptionally large rivers (Paraná River) over grass or submerged tree branches (also found on stones, plastic objects, etc.). In Paraná River, it was found in small numbers only two times during one year of collections, in a place where S. cuneatum is very abundant. In Salado River in the center of Buenos Aires province it was very abundant during high river levels, which produce overflows; it occurred in small currents at the sides of the Salado River. As it is an anthropophilic species, it is a serious pest in several cities and towns such as Junin along the Salado River where it also attacks cows and horses. Discussion: The closest species are S. amazonicum and S. sanguineum, with 8 branches and basal branching. Simulium sanguineum can be separated by the secondary branches bifurcating on the ventral branch; in S. amazonicum and S. chaquense differentiation is on the dorsal branch. Small differences are found in the pupa of both species: S. amazonicum is small and the few tubercles on the head and thorax are acuminate; in S. chaquense, these are only conical on the median area of the thorax. Simulium. amazonicum has a male scutum with 1+1 cuneiform silvery bands not surpassing the median length of the scutum, and in S. chaquense, these reach the posterior grayish area. Simulium amazonicum females have less expanded shadows in the intervittae spots. The distinct environment where they breed is another difference of both species. Non morphological methods probably can help to resolve this identification problem, very common in Neotropical black flies. Simulium chaquense female coloration is similar to that of S. cuneatum, but the pupal gill with 10 branches allows their separation. Simulium (Cerqueirellum) sanguineum KNAB (Figs. 76 L, M; 77 K; 78 I.) Simulium sanguineum KNAB, 1915c: 279; TIDWELL et al., 1981: 79; SHELLEY et al., 1982: 26; SHELLEY, 1988: 207. Simulium (Cerqueirellum) sanguineum: PY-DANIEL, 1983: 168; COSCARÓN, 1991: 196. Simulium (Psaroniocompsa) sanguineum: CROSSKEY & HOWARD, 1997: 56; 2004: 50.

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This species was described in detail by TIDWELL et al. (1981), and PY-DANIEL (1983) added more information on larval stages. Diagnosis was extracted from these authors. Female: Wing length 1.3.0-1.5 mm. Scutum with 1+1 submedian bluish-silver to golden-silver iridescent stripes reaching posterior grayish border and varying in shape in accordance with light position, intervittae shadow spots elongated (Figs. 76 L, M). Claws without subbasal tooth. Genitalia as in S. cuneatum. Male: Wing length 1.4-1.5 mm. Scutum black with 1+1 silvery cuneiform anterior spots projected by thin stripe reaching nearly posterior grayish area and sometimes meeting it (Fig. 77 K). Genitalia as in S. cuneatum. Gonostylus subtriangular with single spur as in S. amazonicum. Pupa: Cocoon with small dorsal anterior projection. Cocoon length 1.7-2.5 mm; gills 1.4-1.8 mm. Gills with 8 branches; branching close to base; secondary ventral branches bifurcated (Fig. 78 I). Frontoclypeus and thorax with rounded tubercles in regular quantity and showing irregular pattern; frontal trichomes single, facial trichomes bifid, thoracic trichomes with 2 to 4 branches. Larva: Maximum length 3.8-4.5 mm. Antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1:1. Mandible with 2 long, sinuous or straight latero-mandibular processes. Anal ring with 52-55 rows of 8-12 hooks each. Rectal papillae with 3-5 diverticula on each lobe. Distribution: Colombia: Cauca, Choco; Panama: Darien. Bionomics: Immature stages were collected in rivers with turbid water on submerged branches about 0.5-3.0 m deep. The region is a tropical rain forest, below 200 m in altitude, but the immatures have also been found at 1000 m in clear, fast-flowing streams. This species is anthropophilic, and is implicated as a vector of Mansonella ozzardi. Discussion: The closest species are S. amazonicum and S. chaquense. The bifurcation of the secondary dorsal branches in the latter two species permits them to be separated from S. sanguineum. Simulium (Cerqueirellum) cuneatum (ENDERLEIN) (Figs. 76 N-T; 77 L-P; 78 J-L; 79 H-M.) Psaroniocompsa cuneata ENDERLEIN, 1936: 124. Simulium cuneatum VARGAS, 1945a: 129; VULCANO, 1967: 9; COSCARÓN, 1983: 90-95. Simulium (Cerqueirellum) cuneatum: PY-DANIEL, 1983: 181; COSCARÓN, 1987: 24; 1991: 202; COSCARÓN & COSCARÓN ARIAS, 2000: 25. Simulium (Psaroniocompsa) cuneatum CROSSKEY & HOWARD, 1997: 57; 2004: 52.

Female: Wing length 1.7-2.1 mm. Coloration similar to that of S. chaquense, complemented with small variations in relation to the illumination direction (Figs. 76 OR). Abdomen as in S. chaquense but lateral silver gray spots on segment III-V are less evident. Frons relatively wide. Basal portion of cibarium with 2-3 rows of acute teeth (Fig. 76 N); claws without subbasal tooth. Eighth sternite with about 40 setae, gonapophysis with abundant microtrichiae; genital fork with posterior branches relatively wide, spermatheca subovoidal (Fig. 76 S); paraprocts relatively short (Fig. 76 T). Male: Wing length 1.7-2.0 mm. Scutum black with 1+1 silvery submedian cuneiform spots reaching or surpassing half of scutum length, but not joining with posterior grayish

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area (Figs. 77 L, M). Scutum hair golden to silvery, homogeneously distributed. Ratio of length/width of hind basitarsus = 5.7. Genitalia as in figure 77 N; gonostylus subtriangular with one apical spur (Fig. 77 O); ventral plate with slightly curved terminal border (Fig. 77 P); endoparameres with strong hooks. Pupa: Cocoon with small anterior dorsal projection (Fig. 78 J), weave compact with evident threads. Length on base 2.0-2.3 mm, at dorsum 2.5-2.7 mm; gills 1.2-1.8 mm. Frontoclypeus and exposed portion of thorax with scarce and apically acute tubercles (Fig. 78 K). Head trichomes single, thoracic trichomes mostly single but can be bifid. Gill with 10 branches, with branching close to base, showing primary dorsal and median 4 branches each (Fig. 78 L). Larva: Maximum length 3.5-4.0 mm. Cephalic apotome with scarce ornamentation (Fig. 79 H) and numerous short single hairs. Antennae surpassing apex of head fan stem; shape as in figure 79 I; ratio of proximal, medial and distal segments = 1:1.1:1.2 (Fig. 79 I). Mandible without latero-mandibular process. Cephalic fan with 16-22 rays. Postgenal cleft subcircular (Fig. 79 J); postgenal bridge about half of hypostomium length. Hypostomium with central tooth lower than lateral teeth (Fig. 79 K). Lateral sclerite of thoracic proleg with teeth in 3-5 groups with 2-5 teeth each (Fig. 79 L). Anal sclerite with wide and short ventral branches with scarce single hairs (Fig. 79 M). Anal ring with 60-70 rows with 10-13 hooks each. Body integument with numerous single short hairs. Rectal papillae with about 6 elongate diverticula on each lobe. Distribution: Argentina: Chaco, Corrientes, Entre Ríos, Formosa, Misiones, Santa Fe; Paraguay: Cordillera. Bionomics: The species breeds in large rivers such as Paraná, and Paraguay, in fastflowing places, over leaves of tree branches, particularly of Salix humboldtiana. It is very abundant during the dry season (August-October; COSCARÓN & COSCARÓN ARIAS, 2000). Simulium cuneatum is an anthropophilic species and very dangerous near the river areas as in Corrientes in the spring time. Discussion: Simulium cuneatum is very close to S. chaquense and S. minusculum, which sometimes are sympatric and also the other species of this group or subgenus. It can be separated by the number of pupal gill branches, which in other species are of 4-8 branches. Simulium (Cerqueirellum) venezuelense RAMÍREZ PÉREZ & PETERSON Simulium venezuelense RAMÍREZ PÉREZ & PETERSON, 1981: 157; PY-DANIEL, 1983: 180; RAMÍREZ PÉREZ, 1983: 123. Simulium (Psaroniocompsa) venezuelense: CROSSKEY & HOWARD, l997: 56; 2004: 50.

(Diagnosis based on RAMÍREZ PÉREZ & PETERSON, 1981). Female: Wing length 1.6-1.7 mm. Blackish species with scutum and abdomen ornamentation as in S. minusculum. Cibarium, fronto-ocular triangle, legs, claws and paraprocts similar to those of S. cuneatum. Male: Wing length 1.5-1.6 mm. Scutum black with 1+1 silvery short and wide anterior cuneiform spots not reaching to half of scutum length (probably with another illumination direction, they get longer). Genitalia as in S. cuneatum.

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Pupa: Cocoon slipper shaped with anterior reinforcement; gill length 1.7 mm. Gill with 4 branches, bifurcated close to base, with primary ventral branch about 3 times longer than dorsal branch. Frontoclypeus and thorax without platelets. Cephalic trichomes single, thoracic trichomes with 2-3 branches. Larva: Unknown. Distribution: Venezuela: Bolivar; Brazil: Amapá, Tocantins. Bionomics: Unknown. Discussion: This is a peculiar species, with adults similar to the known species of this group, but with a pupal gill of only 4 branches, which permits it to be readily differentiated. Simulium (Cerqueirellum) argentiscutum SHELLEY & LUNA DIAS (Figs. 76 U, V; 77 Q; 78 M.) Simulium argentiscutum SHELLEY & LUNA-DIAS, 1980: 105; HAMADA & ADLER, 2001: 119. Simulium (Cerqueirellum) argentiscutum: PY-DANIEL, 1982: 174. Simulium (Psaroniocompsa) argentiscutum: CROSSKEY & HOWARD, 1997: 57; 2004: 52. Cerqueriellum argentiscutum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118.

The species was well described by the authors of the species name, and PY-DANIEL (1982) added more information. The information for this diagnosis mostly was extracted from the above-mentioned authors. Female: Wing length 1.8-2.2 mm. Scutum black with intense silver-gray pruinosity, leaving black narrow median and 1+1 sublateral bands, and 1+1 intervittae darkened cuneiform shadow spots, elongated with frontal light and shorter with posterior illumination (Figs. 76 U-V). Abdomen very similar to that of S. amazonicum. Claw without subbasal tooth. Genitalia as in S. cuneatum. Male: Wing length 1.9-2.0 mm. Scutum color and ornamentation very similar to that of S. amazonicum, but 1+1 silvery submedian cuneiform bands surpassing half of scutum length (Fig. 77 Q). Scutum hair yellowish. Genitalia as in S. cuneatum. Pupa: Cocoon shoe shaped with gelatinous consistency, without evident threads (Fig. 78 M). Length at base 2.8-3.4 mm, on dorsum 1.9-2.2 mm, gills 0.8-0.9 mm. Gill with 6 branches bifurcated close to base, but ventral primary branch bifurcated at greater distance, about 4-5 times its length from base (Fig. 78 N); filaments with winkled margins. Larva: Maximum length 4.4-5.3 mm. Cephalic apotome with positive spots and single or bifid trichomes. Antenna reaching apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1-1.3:1.3-1.5. Mandible without latero-mandibular tooth; well-developed marginal teeth and presence of supramarginal setae. Postgenal cleft very deep, reaching near hypostomium base, interrupting postgenal cleft centrally. Cephalic fan with 25-28 rays. Ninth segment without ventral tubercles. Anal ring with 52-65 rows of 816 hooks each. Rectal papillae with 5-8 diverticula on each lobe. Distribution: Brazil: Acre, Amazonas, Bahia, Rondônia; Colombia: Amazonas; Peru. Bionomics: The species breeds in large, fast-flowing rivers. It bites humans and is considered a vector of Mansonella ozzardi (SHELLEY, 1980). Discussion: Simulium argentiscutum differs from similar species in the female by the extensive silvery gray pruinosity of the scutum, the narrow median and 1+1 sublateral

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black vittae, and very enlarged intervittae shadow spots, with reddish brown to black tones in some specimens when illumination changes. The cocoon is shoe shaped and has a gelatinous consistency. The larva has a deeper postgenal cleft, presence of multibranched scales on the body integument and absence of ventral papillae. Simulium (Cerqueirellum) delponteianum WYGODZINSKY (Figs. 76 W-Z; 77 R-T; 78 O.) Simulium delponteianum WYGODZINSKY, 1961: 30; COSCARÓN, 1983: 95; PY-DANIEL, 1983: 181; SHELLEY et al., 1980: 785; SHELLEY et al., 1982: 4. Simulium (Cerqueirellum) delponteianum: COSCARÓN, 1987: 24; 1991: 203. Simulium (Psaroniocompsa) delponteianum: CROSSKEY & HOWARD, 1997: 57; 2004: 52. Cerqueirellum delponteianum: STRIEDER, 2004: 125.

Female: Wing length 2.5-2.8 mm. Frons blackish with gray pruinosity. Mesonotum blackish gray-pruinose with hair grayish to tin colored; scutum with silvery light anterolateral angles and 1+1 submedian, subovoidal spots projected posteriorly as gray-matte stripes each joining posterior grayish area (Fig. 76 Y). Scutellum, metanotum and pleurae black, the latter two with silver pruinosity. Legs yellowish gray, with darker portion blackish. Abdomen black with tergites I-V velvety and VI-IX with cereous sheen, segment II with 1+1 silvery spots. Fronto-ocular triangle rather deep (Fig. 76 W). Basal portion of cibarium with teeth shorter than in other species of this group, and median thicker invaginated area (Fig. 76 X). Claw with or without small subbasal tooth (Fig. 76 Z). Genitalia as in S. cuneatum. Male: Wing length 2.2-2.7 mm. Scutum silvery black with 1+1 anterior submedian cuneiform silvery spots, which with changing light direction, are continued as thin grayish stripes surpassing half of scutum length (Figs. 77 R-S). Abdomen blackish with 1+1 silvery spots on segments II-VIII. Gonostylus slightly longer than half of gonocoxite length, triangular with blunt apex and 5-10 spurs on dorsal distal area (Fig. 77 T). Ventral plate as in S. cuneatum. Pupa: Cocoon with straight and reinforced border, with weave thin and homogeneous and evident threads; length at base 3.6-4.6 mm; on dorsum 3.0-4.0 mm; gills 1.8-2.5 mm. Frontoclypeus and exposed portion of thorax with medially abundant rounded tubercles; cephalic and thoracic trichomes single and well developed. Gills with 6 branches with branching near base (Fig. 78 O). Larva: Maximum length 5.0 mm. Cephalic apotome with few, weakly evident positive spots, and slightly darkened at base, hair of head dorsally with 1-3 branches. Antenna as long as or scarcely surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1:1.3. Mandible with one latero-mandibular tooth; supramarginal setae absent. Cephalic fan with 28-32 rays. Postgenal cleft subcircular, postgenal bridge about half of hypostomium length. Anal ring with 70-78 rows, with 14-18 hooks each; hair of anal sclerite area scarce and single. Rectal papillae with 3-5 diverticula on each lobe. Body integument with scarce and short single hairs. Distribution: Argentina: Buenos Aires, Corrientes, Entre Ríos; Brazil: Rio Grande do Sul; Uruguay: Artigas.

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Bionomics: The species breed in creeks of plains, with little current in places with slight gradient over submerged aquatic vegetation. Females can bite humans but they are not aggressive; they also attack cattle. Discussion: Simulium delponteianum has similarities with the S. amazonicum group, but can be separated by the grayish thorax, small size of the cibarium teeth, male gonostylus with large number of spurs, and larval mandible without supramarginal setae. These differences make it difficult to include it in this supraspecific taxon. Simulium (Cerqueirellum) argentiscutum has scutum ornamentation and cocoon shape and texture different from other species. Simulium minusculum has the gonostylus shape and absence of mandibular supramarginal setae not concordant with the other species of the group. Simulium chaquense has 2 or 3 spurs on the gonostylus. All of these species show variations from the typical subgeneric characters. Species inquirendae Simulium quadristrigatum ENDERLEIN Simulium quadristrigatum ENDERLEIN, 1934b: 285-286; VARGAS, 1945a: 189; VULCANO, 1967: 19; COSCARÓN, 1983: 96; 1987: 25. Psaroniocompsa quadristrigata: ENDERLEIN, 1936: 123. Simulium (Psaroniocompsa) quadristrigatum: CROSSKEY & HOWARD, 1997: 58; 2004: 52; HERNÁNDEZ & SHELLEY, 2005: 9-10.

This species is not easy to identify. The type is a female in the Berlin Museum. It has a type label and “Paraná”. It is not well preserved; it is pinned through the scutum, and is attacked by fungus and has lost its claws. The original description does not help much. Apparently, it is close to S. delponteianum in the gray scutum tones and leg coloration and wing length, but the cibarium (HERNANDEZ & SHELLEY, 2005; Fig. 42) is like that of S. chaquense and S. cuneatum. It would be very useful to know the shape of the claws, as well as the other life stages, especially the larva to best define the species identity. The type locality (Paraná) does not identify clearly the place it was collected, which can refer to the Brazilian state or to the Paraná River that crosses several states in southern Brazil, Paraguay and Argentina. ENDERLEIN (1937: 13) mentions another two females, from Paraguay but without adequately specifying the locality. We prefer to maintain S. quadristrigatum as species inquirendae. Simulium lutzi KNAB Simulium lutzi KNAB, 1913: 155 (n. nom. to Simulium minutum SURCOUF & GONZÁLEZ RINCONES); SHELLEY et al., 1982: 27; COSCARÓN, 1987: 25. Simulium exiguum: LUTZ, 1909: 141 (preoc.). Simulium minutum SURCOUF & GONZÁLEZ RINCONES, 1911: 290 (repl. name preoc. for exiguum).

SHELLEY et al. (1982) explained, in detail, the history of S. lutzi and the material studied by LUTZ (determined as S. exiguum) from Rio Grande at the limit of São Paulo and Minas Gerais (Brazil). As this material is not well preserved and does not permit proper determination, we prefer to maintain S. lutzi as species inquirendae.

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Subgenus Simulium (Coscaroniellum) PY-DANIEL (Figs. 80-83; 144.) Type-species: Simulium quadrifidum LUTZ, 1917: 66.

Female: Small species, maximum wing length 2.5 mm. Coloration generally blackish to reddish dark brown. Scutum with 1+1 silvery submedian bands that can reach posterior grayish area or only on anterior half as cuneiform spots. Abdomen similar to that of S. (Cerqueirellum). Frons wider than high, with infrafrontal sutures well developed. Legs with filiform and petaloid hair; claw with small subbasal tooth. Basal portion of cibarium with median invaginated area well sclerotized, with small blunt projections at sides and teeth mostly apically blunt. Paraprocts short; genital fork stem distally wider. Gonapophysis distally surpassing eighth sternite; internal border slightly concave and well sclerotized; surface with abundant spiculae. Spermatheca relatively large. Male: Coloration blackish to reddish dark brown with 1+1 anterior submedian cuneiform silvery spots. Abdomen similar to that of S. (Cerqueirellum). Gonostylus about as long as gonocoxite, subtrapezoidal, longer than wide, with large spur distally and slightly flattened; ventral plate and endoparameres as in S. (Cerqueirellum). Pupa: Cocoon compact with threads showing in some species, poorly defined weave with spongy aspect. Gill with 4-8 branches. Frontoclypeus and thorax mostly with apically acute tubercles, and trichomes with 2-3 branches. Larva: Maximum length 4.0 mm. Body generally with dark band on first abdominal segment; integument with elongate multibranched trichomes. Cephalic apotome with median and lateral spots generally weak. Antenna surpassing apex of cephalic fan stem, with distal article proportionately long. Postgenal cleft deep, with postgenal bridge shorter than hypostomium length. Mandibles with latero-mandibular process and without supramarginal setae. Rectal papillae with diverticula. Ninth abdominal segment ventrally with 1 pair of tubercles. Distribution: This taxon is a heterogeneous group of exclusively Amazonian or periamazonian species, composed of S. quadrifidum, S. cerradense, S. cauchense and, with some restrictions, S. goeldii, S. ulyssesi, S. daltanhani and S. quadrivittatum (Fig. 144). KEYS

TO

SIMULIUM (COSCARONIELLUM)

SPECIES

Females 1 – 2(1) – 3(2)

Scutum black with 1+1 silvery submedian stripes reaching or not reaching silvery posterior area (Figs. 80 E, Q, R, W) .............................................................................. 2 Scutum reddish brown to blackish without 1+1 silvery submedian stripes or only 1+1 anterior rectangular thin silvery spots ................................................................... 5 Scutum with 1+1 anterior submedian comma-shaped silvery stripes not reaching half scutum length ................................................................................ Simulium cauchense Scutum with 1+1 submedian silvery stripes reaching silvery posterior area ......... 3 Scutum with anteriorly wide silvery submedian stripes, and posteriorly wide dropshaped median black stripe (Fig. 80 E); 1+1 cibarium submedian protuberances blunt (Fig. 80 D) ............................................................................... Simulium quadrifidum

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Scutum with submedian silvery stripes not as wide anteriorly and median black stripe not as wide posteriorly, 1+1 cibarium submedian protuberances acute (Figs. 80 P-Y) ................................................................................................................................. 4 Black median scutum stripes wider posteromedially, and with median thin silvery stripe (Fig. 80 Q) .................................................................................. Simulium cerradense Black median scutum stripes not wider posteromedially, and without median silvery stripe (Fig. 80 W) ............................................................ Simulium quadrivtittatum Scutum black with 1+1 anterior submedian thin rectangular silver spots connected with silver lateral stripe; wing length 2.0 mm; ratio of palpomere length/sensory vesicle = 1.4 .............................................................Simulium daltanhani Scutum reddish without anterior silver spots (Fig. 80 S); wing length 1.8; ratio of palpomere length/sensory vesicle = 2.3-3.1 .......... Simulium goeldii, Simulium ulyssesi

Males 1 – 2(1) – 3(2) – 4(1) – 5(4) –

Scutum with 1+1 anterior submedian stripes (Fig. 81 A) ......................................... 2 Scutum without anterior submedian stripes or only small and thin rectangular anterior spot (Fig. 81 D) ................................................................................................... 4 Scutum with 1+1 submedian silvery stripes joining with posterior grayish area (Fig. 81 K); gonostylus about 1.5 longer than wide at base (Fig. 81 L) ................... ......................................................................................................... Simulium quadrivtittatum Scutum with 1+1 submedian silvery stripes not joining with posterior grayish area (Fig. 81 A); gonostylus about 2 times longer than wide at base (Fig. 81 C) ......... 3 Scutum with 1+1 anterior submedian thin stripes reaching about 1/2 scutum length (Fig. 81 A) ............................................................................. Simulium quadrifidum Scutum with 1+1 anterior submedian cuneiform stripes reduced to anterior 1/4 of scutum length ................................................................................... Simulium cauchense With some illumination, silvery lateral band of scutum joining anteriorly to 1+1 submedian silvery spots (Fig. 81 D); gonostylus with 1 spur; gonostylus length/ width ratio at base = 2.5 ..................................................................... Simulium cerradense With no illumination, silvery lateral band of scutum joining 1+1 submedian silvery spots; gonostylus with 1-3 spurs; gonostylus length/width ratio at base = 1.8-2.1 ................................................................................................................................... 5 Gonostylus with 2 or 3 spurs (Fig. 81 J) ........................................ Simulium daltanhani Gonostylus with 1 spur (Fig. 81 H) ......................... Simulium goeldii, Simulium ulyssesi

Pupae 1 – 2(1) –

Gill with 4 branches .......................................................................................................... 2 Gill with 6-8 branches ....................................................................................................... 3 Gill branches longer than cocoon base (Fig. 82 A); length 3.3-3.4 mm and 2.8 mm respectively; secondary gill branches emerging close to primary gill branches (Fig. 82 C); abdominal tergite V with denticles on anterior border .......... Simulium quadrifidum Gill branches and cocoon base length about same size (length 2.2-2.6 and 2.0-2.6 respectively); secondary gill branches emerging distal to 1/2 gill length; abdominal tergite V without denticles on anterior border ............................... Simulium cauchense

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Gill with 6 branches .......................................................................................................... 4 Gill with 8 branches .......................................................................................................... 6 Basal trunk of gill elongated and bifurcation of secondary branches relatively far from base (Fig. 82 K) .............................................................................. Simulium ulyssesi Basal trunk of gill short and bifurcation of secondary branches near base (Figs. 82 F, L) ....................................................................................................................................... 5 Cocoon weave fibrous, with obvious threads; cocoon with anterodorsal projection (Fig. 82 L); thoracic tubercles rounded apically; abdominal tergite V with denticles on anterior border ...............................................................................Simulium daltanhani Cocoon weave spongy, without obvious threads; cocoon without anterodorsal projection (Fig. 82 E); thoracic tubercles acuminate apically; abdominal tergite V without denticles on anterior border ............................................... Simulium cerradense Cocoon expanded laterally and with open loops (Fig. 82 M); bifurcation of tertiary branches relatively close to base ................................. Simulium quadrivittatum Cocoon not expanded laterally and with compact weave (Fig. 82 G); bifurcation of tertiary branches relatively far from base ........................................ Simulium goeldii

Larvae 1 – 2(1) – 3(2) – 4(3) – 5(4) –

Subesophageal ganglion pigmented; body length 4.8-5.5 mm; postgenal cleft acuminate apically ...............................................................................Simulium daltanhani Subesophageal ganglion not pigmented; body length 3.1-4.7 mm; postgenal cleft apically rounded or not (Figs. 83 D, S) .......................................................................... 2 Anal ring with 80-84 rows of hooks; ratio of postgenal bridge length/ hypostomium height = 3.9; ratio of proximal, medial and distal antennal articles = 1:1.0:1.0 (Fig. 83 N) ............................................................................. Simulium cerradense Anal ring with 57-66 rows of hooks; ratio of postgenal bridge length/ hypostomium height = 1.4-2.9; ratio of proximal, medial and distal antennal articles = 1:0.3-0.9:1.1-1.8 ................................................................................................ 3 Rectal papillae with 10-13 diverticula on each lobe; latero-mandibular process curved and thin (Fig. 83 C); ratio of hypostomium length/postgenal bridge length = 2.0-2.9 ................................................... Simulium quadrifidum, Simulium quadrivittatum Rectal papillae with 4-7 diverticula on each lobe; latero-mandibular process straight or sinuous, relatively thick; ratio of hypostomium length/postgenal bridge length = 1.4-2.1 ..................................................................................................... 4 Latero-mandibular process sinuous, thin and simple apically; proportion of length of proximal, medial and distal antennal articles = 1:1.5:1.6; ratio of hypostomium length/postgenal bridge length = 1.9-2.1 ........................................ Simulium cauchense Latero-mandibular process straight, thick and apically bifid or trifid; proportion of length of proximal, medial and distal antennal articles = 1:1.1:1.4; ratio of hypostomium length/postgenal bridge length = 1.4-1.6 ........................................... 5 Latero-mandibular process apically bifid; cephalic fan rays 38-47; body length = 3.6-3.8 mm ................................................................................................... Simulium goeldii Latero-mandibular process apically trifid; cephalic fan rays 30-36; body length = 4.1-4.2 mm ................................................................................................. Simulium ulyssesi

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CLAVES

DE ESPECIES DE

SIMULIUM (COSCARONIELLUM)

Hembras 1 – 2(1) – 3(2) – 4(3) – 5(1) –

Escudo negro con 1+1 bandas plateadas submedianas llegando o no al área plateada posterior (Figs. 80 E, Q, R, W) ....................................................................... 2 Escudo castaño rojizo a negruzco sin 1+1 bandas submedianas plateadas o solamente reducidas a 1+1 manchas plateadas anteriores rectangulares finas ............................... 5 Escudo con 1+1 bandas plateadas submedianas en forma de coma no llegando a la mitad del largo del escudo ................................................................... Simulium cauchense Escudo con 1+1 bandas plateadas submedianas llegando al área plateada posterior ....... 3 Escudo con bandas submedianas plateadas más anchas anteriormente, y banda mediana negra más ancha posteriormente en forma de gota (Fig. 80 E); cibario con 1+1 protuberancia submedianas romas (Fig. 80 D) ....................... Simulium quadifidum Escudo con bandas submedianas plateadas no tan anchas anteriormente y banda mediana negra no tan ancha posteriormente; cibario con 1+1 protuberancias submedianas aguzadas (Figs. 80 P, Y) ............................................................................ 4 Banda negra mediana del escudo más ancha medio-posteriormente y con una banda mediana plateada fina (Fig. 80 Q) ........................................ Simulium cerradense Banda negra mediana del escudo no más ancha medio-posteriormente y sin una banda mediana plateada (Fig. 80 W) ........................................ Simulium quadrivittatum Escudo negro con 1+1 mancha plateada anterior submediana rectangular conectada con banda lateral plateada; longitud ala 2,0 mm; relación del largo del palpómero/ vesícula sensorial = 1,4 ...............................................Simulium daltanhani Escudo castaño rojizo sin manchas anteriores plateadas (Fig. 80 S); longitud ala 1,8 mm; relación del largo del palpómero / vesícula sensorial = 2,3-3,1 ................ .......................................................................................... Simulium goeldii, Simulium ulyssesi

Machos 1 – 2(1) – 3(2) – 4(1)

Escudo con 1+1 banda plateada submediana (Fig. 81 A) ......................................... 2 Escudo sin 1+1 banda plateada submediana o solamente una mancha anterior rectangular fina (Fig. 81 D) .............................................................................................. 4 Escudo con 1+1 banda plateada submediana uniéndose con el área grisácea posterior (Fig. 81 K); gonostilo aproximadamente 1,5 más largo que ancho en la base (Fig. 81 L) ............................................................................. Simulium quadrivittatum Escudo con 1+1 banda plateada submediana no uniéndose con el área grisácea posterior (Fig. 81 A); gonostilo aproximadamente 2 veces más largo que ancho en la base (Fig. 81 C) ............................................................................................................... 3 Escudo con 1+1 banda fina anterior submediana, llegando cerca de la mitad del largo del escudo (Fig. 81 A) ........................................................... Simulium quadrifidum Escudo con 1+1 banda cuneiforme anterior submediana reducida al 1/4 anterior del largo del escudo .............................................................................. Simulium cauchense Con cierta iluminación las bandas plateadas del escudo se unen con 1+1 manchas plateadas anteriores (Fig. 81 D); gonostilo con 1 espolón; relación largo/ancho en la base del gonostilo = 2,5 ................................................................. Simulium cerradense

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Con ninguna iluminación las bandas plateadas laterales del escudo se unen con 1+1 manchas plateadas submedianas; gonostilo con 1- 3 espolones; relación del largo/ancho en la base del gonostilo = 1,8-2,1 ........................................................... 5 Gonostilo con 2 o 3 espolones (Fig. 81 J) ..................................... Simulium daltanhani Gonostilo con 1 espolón (Fig. 81 H) ....................... Simulium goeldii, Simulium ulyssesi

Pupas 1 – 2(1)

–

3(1) – 4(3) – 5(4) – 6(3) –

Branquia con 4 ramas ........................................................................................................ 2 Branquias con 6-8 ramas .................................................................................................. 3 Branquias más largas que el capullo en la base (Fig. 82 A); largo 3,3-3,4 mm y 2,8 mm respectivamente; ramas secundarias de la branquia emergen cerca de las ramas primarias (Fig. 82 C); V tergito abdominal con dentículos sobre el borde anterior ............................................................................................... Simulium quadrifidum Largo de las branquias y de la base del capullo aproximadamente del mismo tamaño (largo 2,2-2,6 mm y 2,0-2,6 mm respectivamente; ramas secundarias emergiendo lejos de la mitad del largo de la branquia. V tergito abdominal sin dentículos sobre el borde anterior ..................................................... Simulium cauchense Branquia con 6 ramas ........................................................................................................ 4 Branquia con 8 ramas ........................................................................................................ 6 Tronco basal de la branquia alargada y bifurcación de las ramas secundarias relativamente lejos de la base (Fig. 82 K) ........................................... Simulium ulyssesi Tronco basal de la branquia corto y bifurcación de las ramas secundarias próxima a la base (Figs. 82 F, L) ..................................................................................................... 5 Tejido del capullo fibroso, con fibras evidentes; capullo con prolongación dorsal anterior (Fig. 82 L); tubérculos del tórax redondeados apicalmente; tergito abdominal V con dentículos sobre el borde anterior .................. Simulium daltanhani Tejido del capullo esponjoso, sin fibras evidentes; capullo sin prolongación anterior dorsal (Fig. 82 E); tubérculos del tórax aguzados apicalmente; tergito abdominal V sin dentículos sobre el borde anterior .................... Simulium cerradense Capullo extendido lateralmente y con agujeros (Fig. 82 M); bifurcación de las ramas terciarias relativamente cercanas a la base .................. Simulium quadrivittatum Capullo no expandido lateralmente y con tejido compacto (Fig. 82 G); bifurcación de las ramas terciarias relativamente lejos de la base ......................... Simulium goeldii

Larvas 1 – 2(1)

Ganglio subesofágico pigmentado; longitud del cuerpo 4,8-5,5 mm; hendidura postgenal aguzada apicalmente ........................................................ Simulium daltanhani Ganglio subesofágico no pigmentado; longitud del cuerpo 3,1-4,7 mm; hendidura postgenal redondeada o no apicalmente (Figs. 83 D, S) ............................................ 2 Anillo anal con 80-84 hileras de ganchos; relación puente postgenal / altura del hipostomio = 3,9; relación de los artejos antenales proximal, mediano y distal = 1:1:1 (Fig. 83 N) .................................................................................... Simulium cerradense

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Anillo anal con 57-66 hileras de ganchos; relación puente postgenal / altura del hipostomio = 1,4-2,9; relación de los artejos antenales proximal, mediano y distal = 1:0,3-0,9:1,1-1,8 .............................................................................................................. 3 Papilas rectales con 10-13 divertículos en cada lóbulo; proceso látero-mandibular curvo (Fig. 83 C); relación largo del hipostomio/ puente postgenal = 2,0-2,9 ...... .................................................................... Simulium quadrifidum, Simulium quadrivittatum Papilas rectales con 4-7 divertículos en cada lóbulo; proceso látero-mandibular recto o sinuoso; relación largo hipostomio/ puente postgenal = 1,4-2,1 ............. 4 Proceso látero-mandibular sinuoso, fino y apicalmente simple; relación de los artejos antenales proximal, mediano y distal = 1:1,5:1,6 relación largo hipostomio/ puente postgenal = 1,9-2,1 ......................................... Simulium cauchense Proceso látero-mandibular recto, grueso y apicalmente bífido o trífido; relación de los artejos antenales proximal, mediano y distal = 1:1,1:1,4; relación largo hipostomio/ puente postgenal = 1,4-1,6 ...................................................................... 5 Proceso látero-mandibular apicalmente bífido; rayos del abanico cefálico en número de 38-47; longitud del cuerpo = 3,6-3,8 mm ....................... Simulium goeldii Proceso látero-mandibular apicalmente trífido; rayos del abanico cefálico en número de 30-36; longitud del cuerpo = 4,1-4,2 mm ..................... Simulium ulyssesi

QUADRIFIDUM species group Simulium (Coscaroniellum) quadrifidum LUTZ (Figs. 80 A-M; 81 A-C; 82 A-D; 83 A-K.) Simulium quadrifidum LUTZ, 1917: 66; PINTO, 1931: 709; VULCANO, 1967: 19; TIDWELL et al., 1982: 62; SHELLEY et al., 1984: 152; HAMADA & GRILLET, 2001: 34; HAMADA & FOUQUE, 2001: 955; HAMADA & ADLER, 2001: 119; ALVAN-AGUILAR et al., 2005: 250-256. Simulium (Coscaroniellum) quadrifidum: PY-DANIEL, 1983: 184; COSCARÓN, 1987: 25; 1991: 207. Coscaroniellum quadrifidum PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium (Psaroniocompsa) quadrifidum: SHELLEY et al., 1997: 15; CROSSKEY & HOWARD, 1997: 57, 2004: 51; COSCARÓN ARIAS, 2003: 196, 205, 208, 215. ?Simulium rasii RAMÍREZ PÉREZ, 1980: 60. Simulium torrealbai RAMÍREZ PÉREZ, 1980: 64.

Female: Wing length 1.7-2.0 mm. Frons and clypeus with bluish-green nacreous pruinosity. Scutum black with 1+1 silvery submedian stripes becoming thinner posteriorly, reaching grayish posterior area; median black stripe wide posteriorly (Fig. 80 E); with changing light position, silvery submedian stripe becomes thinner and does not reach grayish posterior area. Scutum silvery. Pleurae dark grayish with silvery sheen. Abdomen velvety black on tergites I-V, VI-IX with cereous sheen; tergite II with 1+1 large lateral silvery spots; III-V with thin grayish pruinose band on posterior border (Fig. 80 J). Frons relatively wide (Fig. 80 A); fronto-ocular triangle relatively short (Fig. 80 B). Sensory vesicle of maxillary palp about half of palpomere and with numerous tubercles (Fig. 80 C). Basal portion of cibarium well sclerotized apically with large rounded submedian process and short teeth (Fig. 80 D). Leg shape and ornamentation as in figures 80 F, G, H. Claws with short subbasal tooth (Fig. 80 I). Genitalia with

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Y

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Figure 80. Female. A-M: S. (Coscaroniellum) quadrifidum; A: frons; B: fronto-ocular triangle; C: maxillary palp ; D: basal portion of cibarium; E: scutum; F-H: fore, mid and hind leg, respectively; I: claw; J: abdomen, K: eighth sternite, gonapophyses and genital fork; L: spermatheca; M: cercus and paraproct. N-R: S. (C.) cerradense; N: frons; O: fronto-ocular triangle; P: basal portion of cibarium; Q-R: scutum with different position of light. S-U: S. (C.) goeldii; S: scutum; T: basal portion of cibarium; U: genital fork. V: S. (C.) cauchense, frons. W-Z: S. (C.) quadrivittatum; W: scutum; X; abdomen; Y: basal portion of cibarium; Z: cercus and paraproct.

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general aspect as in figure 80 K. Eighth sternite with about 20 setae; spermatheca subspherical (Fig. 80 L); paraproct short (Fig. 80 M). Male: Wing length 1.6-1-7 mm. Scutum black with 1+1 silvery thin anterior submedian cuneiform spots that can reach distally to half of scutum length (Fig. 81 A); (this submedian spot can be absent in some specimens (SHELLEY et al., 1997)). Genitalia as in figure 81 B; gonostylus subtriangular in shape with large spur (Fig. 81 C); ventral plate slowly acuminated apically. Pupa: Cocoon with reinforced border and slightly projected distally (Fig. 82 A), with compact weave and evident threads; length at base 2.8-2.9 mm, on dorsum 2.0-2.5 mm, gills 3.3-3.8 mm with 4 filiform branches, bifurcating near base (Fig. 82 C). Frontoclypeus and exposed portion of thorax with regular quantity of acuminate tubercles (Fig. 82 B); cephalic and thoracic trichomes bifid (Fig. 82 D). Abdominal tergite V with denticles on anterior border. Larva: Maximum length 4.0 mm. Cephalic apotome light brown with median and lateral spots not evident (Fig. 83 A). Antenna with distal article elongated (Fig. 83 B), ratio of proximal, medial and distal articles = 1:0.7-0.9:1.2-1.3. Cephalic fan with 35-40 rays. Mandible as in figure 83 C. Postgenal cleft wide and deep, postgenal bridge about 1/3 of hypostomium length (Fig. 83 D). Hypostomium with lateral teeth acuminate and longer than median tooth (Fig. 83 E). Body integument with abundant multibranched hairs as shown in figures 83 G-K. Ninth sternite with 1+1 ventral papillae. Anal ring with 55-60 rows with 10-12 hooks each. Anal sclerite with abundant hair with 1-4 branches and irregular size (Fig. 83 F). Rectal papillae with 1013 diverticula on each lobe. Distribution: Bolivia: Cochabamba; Brazil: Amapá, Amazonas, Pará, Rondônia, Roraima; Colombia; Ecuador: Esmeraldas, Napo, Pastaza; French Guiana; Guyana; Suriname; Venezuela: Amazonas, Bolívar. Bionomics: This species breeds in small creeks with clear water, frequently in shaded places. Females have zoophilic habits (SHELLEY et al., 1997). Discussion: This species is similar to S. cauchense and S. cerradense. The first species can be distinguished by the silvery submedian stripes of females and males that are shorter and do not reach beyond half the scutum length. In the pupa, the gills are proportionally shorter, and they have denticles on abdominal tergite V. The larva has a shorter distal antennal article and longer medial article. Simulium cerradense can be differentiated from S. quadrifidum in the female by the presence of intervittae stripes on the scutum and the acuminate submedian process of the cibarium, in the male by the anterior subtrapezoidal spots, in the pupa by the spongy cocoon weave and the gills with 6 branches, and in the larva by the cephalic apotome with evident positive spots and the medial antennal article longer than the proximal and the distal. Simulium rasii was synonymized with some doubt, because material from Venezuela, Monagas and San Antonio, determined by RAMÍREZ PÉREZ, has pupal gills with thicker and shorter primary branches and more tubercles on the frontoclypeus and thorax.

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Figure 81. Male. A-C: S. (Coscaroniellum) quadrifidum; A: scutum; B: genitalia, ventral view showing gonocoxite, gonostylus, ventral plate and endoparameres; C: gonostylus, dorsal view (mounted on slide). D-G: S. (C.) cerradense; D: scutum; E: ventral plate, median sclerite and endoparameres; F: gonocoxite and gonostylus, ventral view; G: gonostylus (mounted on slide). H-I: S. (C.) goeldii, H: gonocoxite and gonostylus on dorsal view; I: ventral plate. J: S. (C.) daltanhani, gonostylus (from HAMADA & ADLER, 1998). K-L: S. (C.) quadrivittatum; K: scutum, L: gonostylus.

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Figure 82. Pupa. A-D: S. (Coscaroniellum) quadrifidum; A: lateral view; B: frontoclypeus with trichomes and tubercles; C: portion of thorax with trichomes and gill; D: thoracic trichomes and tubercles. E-F: S. (C.) cerradense; E: dorsolateral view; F: portion of thorax with trichomes and gill. G-J: S. (C.) goeldii; G: dorsal view; H: frontoclypeus trichomes and sparse tubercles; I: portion of thorax with trichomes and gill; J: thoracic tubercles with high magnification. K: S. (C.) ulyssesi, basal portion of gill. L: S. (C.) daltanhani, lateral view (from HAMADA & ADLER, 1998). M: S. (C.) quadrivittatum, dorsal view.

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Figure 83. Larva. A-K: S. (Coscaroniellum) quadrifidum; A: cephalic apotome; B: antenna; C: teeth of mandible; D: postgenal cleft and hypostomium; E: hypostomium; F: anal sclerite; G-K: different body trichomes. L-P: S. (C.) cerradense; L: lateral view; M: head, dorsal view; N: antenna; O: teeth of mandible; P: lateral sclerite of thoracic proleg. Q-S: S. (C.) goeldii; Q: antenna; R: teeth of mandible; S: postgenal cleft and hypostomium. T: S. (C.) ulyssesi, latero-mandibular process.

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Simulium (Coscaroniellum) cerradense COSCARÓN, CERQUEIRA, SATO & LA SALVIA (Figs. 80 N-R; 81 D-G; 82 E, F; 83 L-P.) Simulium (Coscaroniellum) cerradense COSCARÓN et al., 1992: 111-113. Simulium (Coscaroniellum) sp. A COSCARÓN, 1991: 206. Simulium (Psaroniocompsa) cerradense: CROSSKEY & HOWARD, 1997: 57; 2004: 51. Coscaroniellum cerradense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium cerradense: HAMADA et al., 2004: 1-8.

Female: Wing length 1.8 mm. Scutum black with 1+1 silvery submedian stripes that join with grayish posterior area, variation in width of stripes in accordance lighting (Figs. 80 Q, R); hair golden to silvery. Abdomen with tergites I-IV velvety black, V-IX grayish with cereous sheen. Frons with weakly developed infra-frontal sutures (Fig. 80 N). Fronto-ocular triangle wider than high (Fig. 80 O). Basal portion of cibarium with 1+1 submedian process distally acuminate and teeth of invaginated area very small (Fig. 80 P); claws with subbasal tooth; leg hair filiform and petaloid. Genitalia with general aspect as in S. quadrifidum; eighth sternite with about 20 setae on each side. Male: Wing length 1.7 mm. Scutum velvety black with 1+1 anterior submedian silvery spots, subtrapezoidal (Fig. 81 D); changing light position shows narrow band joining to grayish lateral area. Ventral plate, median sclerite and endoparameres as in figure 81 E. Gonostylus subtrianglular, twice as long as wide (Figs. 81 F, G). Pupa: Cocoon weave spongy, thick not transparent, with reinforcement on anterior border (Fig. 82 E). Length at base 2.6-2.8 mm, at dorsum 2.1-2.4 mm, gills 3.04.0 mm. Frontoclypeus and exposed portion of thorax with abundant acuminate spots; trichomes bifid and large (Fig. 82 F). Gill with 6 branches bifurcated near base. Abdominal tergite V without denticles on anterior border. Abdominal terminal spines short and with slightly acute apex. Larva: Maximum length 4.3-4.7 mm. General aspect as in figure 83 L. Cephalic apotome with positive spots evident (Fig. 83 M). Hairs of dorsum of head single. Antenna surpassing apex of cephalic fan stem. Ratio of proximal, medial and distal articles = 1:1:1.1 (Fig. 83 N). Mandible with 2 marginal teeth and strong latero-mandibular process (Fig. 83 O). Postgenal bridge longer than half of hypostomium length. Cephalic fan with 40-46 rays. Body integument with short hairs with 2.5 branches. Lateral sclerite of thoracic proleg with 22-24 teeth arranged in 7-9 groups (Fig. 83 P). Ninth sternite with 1+1 ventral tubercles, slightly acuminate apically. Anal sclerite area with abundant unbranched hairs. Anal ring with 80-84 rows with 12-13 hooks each. Rectal papillae with 5-6 diverticula on each lobe. Distribution: Brazil: Bahia. Bionomics: This species was collected in creeks 5-10 m wide and 0.5-2.0 m deep, in crystalline waters with little to moderate flow over aquatic vegetation. Female habits are unknown. The distributional area corresponds to the Cerrado region of northeastern Brazil. Discussion: The ornamentation of the female scutum is similar to that of S. quadrifidum and S. (Cerqueirellum) species, S. (Notolepria) and some species of S. (Psilopelmia) and S. (Aspathia). Differentiation from S. quadrifidum was explained under that species. It can be distinguished from the Cerqueirellum species group by the shape of the male gonostylus,

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from S. (Notolepria) and S. (Psilopelmia) by the shape of the female paraprocts, and from S. (Aspathia) by the shapes of the male gonostylus and ventral plate. The spongy cocoon weave of S. cerradense permits it to be differentiated from these similar species. Only S. argentiscutum has a similar cocoon, but the pupal gill and cocoon shape permit separation. Simulium (Coscaroniellum) cauchense FLOCH & ABONNENC (Fig. 80 V.) Simulium cauchense FLOCH & ABONNENC, 1946a: 1; SHELLEY et al., 1984: 144-145; HAMADA & GRILLET, 2001: 34; HAMADA & ADLER, 2001: 119; ALVAN-AGUILAR et al., 2005: 250-256. Simulium (Coscaroniellum) cauchense: PY-DANIEL, 1983: 188; COSCARÓN, 1987: 25; 1991: 206. Coscaroniellum cauchense PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium (Psaroniocompsa) cauchense: SHELLEY et al., 1997: 17; CROSSKEY & HOWARD, 1997: 57; 2004: 51. Simulium sextobecium NUNES DE MELLO, 1974: 15. Simulium rangeli RAMÍREZ PÉREZ, RASSI & RAMÍREZ, 1977: 163.

(Based on description by PY-DANIEL, 1983). Female: Wing length 1.5-1.8 mm. Frons and clypeus with pruinosity sheen bluish green. Scutum black with 1+1 silvery anterior submedian cuneiform spot not reaching beyond half length of scutum with anterior or posterior illumination, abdomen coloration very close to quadrifidum. Frons with pronounced infra-frontal sutures (Fig. 80 V); basal portion of cibarium, claws and genitalia shape as in S. quadrifidum. Male: Wing length 1.4-1.7 mm. Scutum black with 1+1 small cuneiform anterior submedian silvery spots. Genitalia as in S. quadrifidum. Pupa: Cocoon similar to that of S. quadrifidum; length along base 1.6-2.5 mm, on dorsum 1.7-2.4 mm, gills 2.2-3.1 mm. Gills with 4 branches bifurcated near the base, dorsal primary branch slightly longer than ventral; dorsal bifurcation can be near midpoint of gill. Frontoclypeus and thorax with tubercles and trichomes as in S. quadrifidum. Abdominal tergite V without denticles on anterior border. Larva: Maximum length 3.0-3.8 mm. Color and morphology similar to S. quadrifidum. Cephalic apotome with obvious positive spots. Ratio of proximal, medial and distal antennal articles = 1:0.4-0.5:1.5-1.6. Mandible teeth as in S. cerradense; cephalic fan with 36-38 rays. Anal sclerite as in S. quadrifidum but with hair more evident. Anal ring with 6072 rows with about 13 hooks each. Rectal papillae with 3-8 diverticula each. Body integument with single to multibranched trichomes. Distribution: Brazil: Amapá, Amazonas, Pará, Roraima; French Guiana; Guyana; Venezuela: Amazonas, Bolivar. Bionomics: Females are zoophilic (SHELLEY et al., 1984: 144). Discussion: The most similar species are S. quadrifidum and S. cerradense, but the more elongated submedian silvery stripes that join with the posterior grayish area on the scutum of females separate them. The male of S. quadrifidum has the silvery submedian stripes thicker and more elongated, reaching half the length of the scutum, and in S. cerradense the silvery submedian stripes are reduced to anterior subtrapezoidal spots. Simulium cerradense can be separated also by the pupal gill with 6 branches. Simulium goeldii and S. ulyssesi have some similarities, but their scutum is reddish brown and their pupal gills have 8 and 6 branches, respectively.

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Simulium (Coscaroniellum) goeldii CERQUEIRA & NUNES DE MELLO (Figs. 80 S-U; 81 H, I; 82 G-J; 83 Q-S.) Simulium goeldii CERQUEIRA & NUNES DE MELLO, 1967: 125; PY-DANIEL, 1981: 176; RAMÍREZ PÉREZ, 1983: 47; CROSSKEY & HOWARD, 1997: 81; HAMADA & ADLER, 2001: 119; RÍOS-VELÁSQUEZ et al., 2002: 114-117. Simulium (Coscaroniellum) goeldii: COSCARÓN, 1987: 25. Simulium (Psaroniocompsa) goeldii: CROSSKEY & HOWARD, 2004: 51. Simulium scorzai RAMÍREZ PÉREZ, 1980b: 59.

Female: Wing length 1.7 mm. Frons and clypeus with silvery pruinosity and nacreous sheen; scutum dark and scutellum dark reddish brown; scutum bordered laterally and posteriorly by silvery pruinose area, with 1+1 sublateral, transverse silvery spots that with some lighting join lateral grayish area (Fig. 80 S); with posterior lighting, spots reduced to small oblique bands; pleurae and metanotum blackish with nacreous pruinosity; scutum hair golden to tin colored. Abdomen blackish with 1+1 lateral gray nacreous area on segment II-V. Basal portion of cibarium with median invagination and 1+1 submedian projections (Fig. 80 T); all this area with small acuminate teeth. Frontoocular triangle wider than high. Claws with subbasal tooth. Genitalia as in quadrifidum; eighth sternite with about 12 setae on each side; paraproct short, spermatheca enlarged; genital fork with well sclerotized stem (Fig. 80 U). Male: Wing length 1.7 mm. General coloration as in female; scutum without obvious 1+1 anterolateral silvery spots. Gonostylus surpassing gonocoxite by half length, subtrapezoidal elongate shape and with 1 subapical spur (Fig. 81 H). Ventral plate slightly acuminate apically (Fig. 81 I). Pupa: Cocoon with thin weave and slight anterodorsal projection (Fig. 82 G); threads weakly evident. Length along base 2.2-2.3 mm, on dorsum 1.7-2.0 mm, gills 2.3-2.5 mm. Frontoclypeus and exposed portion of thorax with sparse (Fig. 82 H) and elongate tubercles with blunt apices (Fig. 82 J). Trichomes long, frontal bifid, facial with 3-7 branches, thoracic with 2-5 branches. Gills with 8 filamentous branches, relatively long basal trunk and 2 short primary branches; primary dorsal branch bifurcated at short distance; dorsal secondary branches bifurcated at short distance, and both dorsal tertiary branches bifurcated near half of gill length (Fig. 82 I); abdominal tergite V with barely evident denticles. Larva: Maximum length 3.5 mm. Color light reddish brown, with darker ring around first abdominal segment. Cephalic apotome not ornamented with positive spots. Cephalic integument without hair. Antenna thin with short median article (Fig. 83 Q); ratio of proximal, medial and distal articles = 1:0.3-0.4:1.4-1.8. Mandible with robust latero-mandibular process (Fig. 83 R). Cephalic fan with 33-38 rays. Postgenal cleft dome shaped; postgenal bridge about 3/4 of hypostomium height (Fig. 83 S); teeth of hypostomium evident. Lateral sclerite of thoracic proleg elongated, with about 17 teeth arranged in 5-6 groups. Anal ring with 54-56 rows with about 9 hooks each. Anal sclerite with sparse single hairs on posterior area. Distribution: Brazil: Amazonas, Pará, Rondônia; Venezuela: Amazonas. Bionomics: This species breeds in small creeks, generally on leaves in fast flow, in clear or dark acidic waters, with temperatures of about 25 °C, which are typical in the Amazonian region. Females were collected biting humans.

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Discussion: This peculiar species is easy to distinguish by the arrangement of the pupal gill, tubercle shape of the frontoclypeus and thorax, coloration and ornamentation of the scutum, and gonostylus shape, all of which make it difficult to include it in the S. quadrifidum species group. The closest species are S. ulyssesi, which can be distinguished by the pupal gill with 6 branches and the latero-mandibular process that is trifid distally, and S. daltanhani, which has 6 branches in the pupal gill and a male gonostylus with 3 spurs. Simulium (Coscaroniellum) ulyssesi PY-DANIEL & COSCARÓN (Figs. 82 K; 83 T.) Coscaroniellum ulyssesi PY-DANIEL & COSCARÓN, 2001: 52-57. Simulium 6 B1 HAMADA & ADLER, 1999: 44. Simulium ulyssesi: RÍOS-VELÁSQUEZ et al., 2002: 114-117.

Female and male coloration and morphology are similar to that of S. goeldii. Female: Fronto-ocular triangle reduced, basal portion of cibarium well sclerotized with teeth bordering median invagination short and blunt, and laterally more developed and apically acute. Male: Gonostylus subtrapezoidal with flattened spur subapically positioned. Pupa: Cocoon shape as in S. goeldii, length along base 1.8-2.7 mm; on dorsum 2.1-2.9 mm; gills 4.5-5.2 mm. Thoracic tubercles subconical; trichomes elongated, facial with 3 branches, frontal bifid, thoracic with 3-4 branches. Gill with 6 filamentous branches and long basal trunk, primary branches bifurcated at short distance, and dorsal secondary branches bifurcated at twice length of primaries (Fig. 82 K) Abdominal tergite V with small teeth on anterior border. Larva: Maximum length 4.1-4.2 mm. Coloration as for S. goeldii. Cephalic apotome with small single hairs. Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal articles = 1:0.4-0.5:1.1-1.2. Mandible with 1 strong latero-mandibular process straight and apically with 3 branches (Fig. 83 T). Cephalic fan with 30-36 rays. Postgenal cleft shape, ratio of postgenal bridge to hypostomium, anal sclerite and number of anal ring hook rows similar to S. goeldii. Distribution: Brazil: Amazonas. Bionomics: Simulium ulyssesi lives in the same environment as S. goeldii. Discussion: This species is very similar to S. goeldii with which it is sympatric. It is possible to differentiate it because S. goeldii has a pupa with a shorter gill of 8 branches, a shorter cocoon, and a shorter larva with a relatively shorter medial antennal article and the apex of the latero-mandibular process with 2 branches. Simulium (Coscaroniellum) daltanhani HAMADA & ADLER (Figs. 81 J; 82 L.) Simulium (Coscaroniellum) daltanhani HAMADA & ADLER, 1998a: 317; 2001: 119. Simulium (Psaroniocompsa) daltahani: CROSSKEY & HOWARD, 2004: 51.

Female: Wing length 2.0 mm. General coloration similar to that of S. goeldii; scutum blackish to reddish brown with 1+1 rectangular silvery anterolateral spots joining with lateral silvery area, 4 short submedian pruinose bands faintly visible; scutum hair golden; scutellum and metanotum blackish, the latter with silvery pruinosity. Abdomen similar to

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that of S. goeldii. Fronto-ocular triangle evident, rather shallow, infra-frontal sutures short; basal portion of cibarium well sclerotized and with submedian projections and teeth as in S. goeldii; sensory vesicle of maxillary palp elongated, about 2/3 of palpomere length; claws with subbasal tooth; genitalia similar to those of S. goeldii. Male: Wing length 1.9 mm. Scutum blackish, with 1+1 narrow anterolateral silver spots connected with silver lateral band, and 1+1 faintly visible silvery submedian stripes; hair reddish gold. Gonostylus about half of gonocoxite length, subtrapezoidal with 1-3 subapical spurs (Fig. 81 J). Ventral plate and endoparameres as in S. goeldii. (Pupa and larva based on HAMADA & ADLER, 1998a). Pupa: Cocoon similar to that of S. goeldii. Length along base 2.7-3.4 mm, on dorsum 3.0-3.4 mm, gills 3.9-4.0 mm. Frontoclypeal and thoracic tubercles rounded distally. Head and dorsum of thorax with bifid trichomes. Gills with 6 filamentous branches, with trunk short and subdivided at short distance from base, with secondary dorsal branches very short (Fig. 82 L). Larva: Maximum length 4.8-5.5 mm. Cephalic apotome slightly darkened on median and basal area. Postgenal cleft dome shaped. Postgenal bridge 0.5-0.8 times hypostomium length. Subesophageal ganglion heavily pigmented. Antenna about same length as cephalic fan stem; ratio of proximal, medial and distal articles = 1:0.7-0.9:1.2-1.5. Cephalic fan with 49-51 rays; latero-mandibular process single. Anal sclerite with single to trifid hairs. Anal ring with 60-66 rows of 12-13 hooks each. Rectal papillae with 9-10 diverticula on each lobe. Distribution: Brazil: Amazonas. Bionomics: Simulium daltanhani breeds in small, shallow, acidic streams with sandy bottoms and warm temperatures (26-29 °C) flowing over fallen leaves and grasses. Female feeding habits are unknown. Discussion: The similar scutal ornamentation and cocoon with an anterior projection can confuse S. daltanhani with S. goeldii and S. ulyssesi, but the latter two don’t have the submedian faintly visible silvery stripes. Simulium goeldii also can be differentiated by the 8 pupal gill branches; S. ulyssesi has 6 branches but they are directed forward, not laterally expanded, and the larval latero-mandibular process has 3 branches. QUADRIVITTATUM species group The S. quadrivittatum group has particular characters, such as narrow straight silvery submedian stripes joined posteriorly by a transverse straight, narrow band before joining the posterior grayish area on the scutum, and a cibarium with 1+1 submedian elongated thin processes with acute teeth. The following characters indicate the similarity of this group to Simulium (Psaroniocompsa), S. (Cerqueirellum) and S. (Coscaroniellum): female scutum with 1+1 submedian stripes and, in some illumination, with shadow-like intervittae spots, short rounded paraproct, subconical gonostylus, wing with basal sector of R without hair; long bifid cephalic and thoracic trichomes on the pupa, a single dark ring on the first abdominal segment of the larva, and the cephalic apotome with positive median, lateral and posterior spots, Simulium (Coscaroniellum) is the closest group, as evidenced in the female by 1+1 submedian processes on the cibarium, relatively deep fronto-ocular

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triangle, and infra-frontal sutures; in the male by the subconical gonostylus; and in the larva by the hypostomium with intermediate teeth about 1/3-1/2 the height of the median and lateral teeth, and the distal antennal article nearly two times longer than the medial article. This species group consists of only one species. Simulium (Coscaroniellum) quadrivittatum LOEW (Figs. 80 W-Z; 81 K, L; 82 M.) Simulium quadrivittatum LOEW, 1862: 186; WYGODZINSKY, 1953b: 334-336; SHELLEY & ARZUBE, 1985: 285; COSCARÓN, 1987: 22. Simulium (Psilopelmia) quadrivittatum. SHELLEY et al., 1989: 336; CROSSKEY & HOWARD, 1997: 60; 2004: 54; SHELLEY et al., 2002: 167-169. Simulium (Psaroniocompsa) quadrivittatum COSCARÓN ARIAS, 2003: 196, 204, 209, 215. Psilopelmia quadrivittatum: RUBTSOV & GARCIA AVILA, 1972: 19-23. Wilhelmia mallochi ENDERLEIN, 1925: 208. Simulium fairchildi VARGAS, 1942b: 458. Simulium (Byssodon) quadrivittatum: VARGAS & DIAZ-NÁJERA, 1948b: 349; 1957b: 162, 262. Simulium metallicum MALLOCH, 1914: 25 part (nec BELLARDI, 1859). ?Simulium (Byssodon) benjamini DALMAT, 1952: 339-344; 1955: 121-126 (n. syn.).

Female: Wing length 1.6-2.0 mm. Frons and clypeus blackish with bluish pruinosity. Scutum black with 1+1 silvery pruinose submedian stripes (Fig. 80 W), joining with dark grayish posterior area; vittae when posteriorly illuminated show anterior small shadow-like intervittae spots. Fronto-ocular triangle deep, wider than high. Abdomen blackish with posterior border grayish, wide on segments III-V (Fig. 80 X). Cibarium basal portion with sclerotized rim, slightly depressed medially and with 1+1 thin and sclerotized submedian process and acute teeth (Fig. 80 Y). Basal portion of R without hair, claws with small subbasal tooth. Gonapophysis with straight distal edge; paraproct short with rounded edge (Fig. 80 Z). Male: Wing length 1.8 mm. Scutum black with 1+1 silvery thin submedian stripes that surpass half of scutum length (Fig. 81 K); (in specimens without hair, submedian silvery stripes are wider with less silvery color). Gonostylus subconical with blunt apex and elongated subapical spur (Fig. 81 L). Pupa: Cocoon slipper shaped with basal and dorsal protuberances, lateral sides expanded with open loops and softly reinforced on anterior edge (Fig. 82 M), length along base 2.4-2.6 mm, on dorsum 1.6-1.8 mm, gills with 2.8-3.2 mm. Gill with 8 branches with secondary and tertiary branches bifurcated closer to base than in S. goeldii. Frontoclypeus and thorax with sparse rounded tubercles; cephalic and thoracic trichomes bifid and relatively long. Larva: Cephalic apotome with positive median, mediolateral and posterolateral spots. Antenna longer than cephalic fan stem; median article shorter than other two articles; cephalic fan with 40-42 rays. Ratio of proximal, medial and distal article = 1:0.60.7:1.4-1.5. Hypostomium with intermedian teeth lower by at least one third median and lateral teeth. Postgenal bridge about as long as hypostomium height. Mandible with 1 latero-mandibular process; supramarginal setae absent. Body with dark band around first abdominal segment; integument with sparse hair dorsally. Ninth abdominal segment with ventral tubercles. Anal ring with 62-65 rows with 11-13 hooks each.

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Distribution: Belize: Belize, Cayo, Stan Creek, Toledo; Colombia: Choco; Costa Rica: Cartago, Limón; Cuba; Dominican Republic; Ecuador: Cotopaxi, El Oro, Esmeraldas, Manabi, Pichincha; Guatemala; Mexico: Campeche, Chiapas, Oaxaca, Puebla, Veracruz; Nicaragua; Panama; Puerto Rico; Venezuela. Bionomics: This species breeds in small creeks with clear water and slow current, on leaves of aquatic vegetation in forested areas of lowlands (Ecuador). Females bite cattle and humans and are considered vectors of Onchocerca volvulus in Ecuador. Discussion: Simulium quadrivittatum can be confused with species of S. (Cerqueirellum), S. quadrifidum, S. metallicum and S. haematopotum by its scutum, but the peculiar cibarium shape allows it to be separated easily. Simulium (Ectemnaspis) ENDERLEIN (Figs. 84-90; 145-148.) Simulium (Ectemnaspis) ENDERLEIN, 1934b: 281. Type-species: Ectemnaspis macca ENDERLEIN, 1934b = Simulium bicoloratum MALLOCH, 1912.

Female: Medium to large size, wing length 1.5-3.0 mm. General coloration yellowish to blackish brown. Scutum yellowish-orange without silvery longitudinal bands, or if present, not reaching posterior border, frequently with median and sublateral areas brown to blackish, bordered by yellow extended in 1+1 areas over humeral plates and 1 median posterior area. Scutellum yellow to yellowish brown. Metanotum generally blackish. Abdomen generally yellow to brownish on tergites I-IV, blackish on posterior area. Pleurae from yellow to black pollinose. Legs from yellow to blackish with some portions black. Fronto-ocular triangle generally wider than high. Basal portion of cibarium with sclerotized rim, from untoothed to covered by abundant teeth with median area slightly covered by small tubercles or flanked by 1+1 submedian large process covered by small teeth. Basal sector of R generally haired. Claws generally with stout subbasal tooth. Paraproct subtrianglular, apically elongated and frequently with rounded basal lobe. Gonapophysis divergent below, lightly sclerotized internally, without hair or spiculae. Genital fork with stem as long as length of posterior branches joined. Male: Wing length 1.8-3.0 mm. Pigmentation as in female. Scutum unbanded, rarely with 1+1 silvery anterior wedges. Posterior basitarsus ratio length/width 3.5-7.5. Gonostylus about same length as gonocoxite, subcylindrical, with slight distal curvature and 1 terminal spur. Ventral plate in ventral view generally about as long as wide with slight lateral concavity; endoparameres with strong hooks. Pupa: Cocoon slipper shaped sometimes with dorsal anterior projection and longitudinal dorsal crest; weave from resistant with evident threads to soft like felt or spongy and without obvious threads. Gill generally with 6-8 thin filaments, but can be from 2-250, and in some cases with thick and irregular filaments. Two pairs of trichomes on frontoclypeus and 5 on thorax, well developed with 3-45 branches. Abdomen with terminal spur short. Larva: Maximum length 4.8-7.0 mm. Tegument with or without trichomes. Cephalic apotome generally with pigmentation on basal or subbasal area with light median subbasal window. Postgenal cleft deep, nearly reaching hypostomium, with postgenal cleft shorter than hypostomium height. Antenna with medial and distal segment about same length and

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both longer than proximal. Cephalic fan with 35-50 rays. Anal sclerite without scales. Anal ring with 68-113 rows of hooks. Diverticula of rectal papillae generally 3-13 but can be 1-20. Simulium (Ectemnaspis) can be subdivided into 4 species groups: BICOLORATUM, ROMANAI, PERFLAVUM and DINELLII. SIMULIUM (ECTEMNASPIS) species groups BICOLORATUM species group: yellowish scutum and scutellum, scutum frequently slightly darkened to blackish median and laterally, but bordered by yellow, metanotum blackish; abdomen with segments yellowish to brown. Cibarium smooth with thin sclerotized rim between cornuae. Paraproct subtriangular, nearly as long as wide at base, blunt, rounded apically. Gonostylus subcylindrical, slightly curving distally; ventral plate with slight lateral constriction. Cocoon frequently with thick spongy or felt-like weave, reinforced anterior border and sometimes with dorsal ridge; frontoclypeus and thorax sometimes with big trichomes with about 50 branches; gill with 2-10 branches; larval cephalic apotome generally darkened on posterior half, with negative median, lateral and basal spots; rectal papillae diverticula of normal size. Fifteen species in the group: S. anaimense, S. antonii, S. arcabucense, S. bicoloratum, S. bicornutum, S. cormonsi, S. furcillatum, S. ignescens, S. jaimeramirezi, S. pautense, S. pifanoi, S. roquemayu, S. rubiginosum, S. tolimaense, S. tunja. ROMANAI species group: scutum with variable coloration from yellowish to totally black but bordered by yellow, scutellum yellowish, metanotum black; abdomen with segment I (II-III) yellowish, the following ones dark-brown; cibarium with median area slightly depressed and covered with tubercles or plaques laterally to cornuae; base with thin teeth-like spines; paraproct subtriangular, nearly as long as wide at base, with blunt apex. Male coloration mostly blackish, gonostylus subcylindrical and slightly curved distally; ventral plate with marked lateral constriction. Pupal cocoon from compact to thick with spongy or felt-like weave; gill with 6-8 branches. Larval ornamentation of cephalic apotome and general morphology as in S. bicoloratum species group. Six species in the group: S. albanense, S. gabaldoni, S. lutzianum, S. mayuchuspi, S. romanai, S. wolffhuegeli. PERFLAVUM species group: yellowish scutum, scutellum and metanotum; abdomen with segments I (II-III) yellowish and following ones dark brown; cibarium with median portion elevated, sometimes scarcely invaginated at center and with space between cornuae, covered by acute teeth-like spines; paraproct subtriangular nearly 2 times longer than wide at base, acute distally and with abundant robust spiculae posteriorly. Male coloration similar to that of female; gonostylus subcylindrical, about as long as or longer than gonocoxite. Ventral plate as long as wide, with lateral constriction. Pupal cocoon frequently with irregular weave, leaving open areas anteriorly, and with 8-20 branches; larval cephalic apotome without peculiar ornamentation and rectal papillae with 10-15 diverticula elongated. Six species in the group: S. kabanayense, S. perflavum, S. rorotaense, S. suarezi, S. trombetense, S. maroniense. DINELLII group: scutum, scutellum and metanotum yellowish, sometimes with 1+1 submedian silvery nacreous stripes on anterior half; abdomen with segments yellowish, V-IX yellowish brown; fronto-ocular triangle wider than high; cibarium with 1+1 submedian relatively thick process covered with small teeth leaving median curved smooth or wrinkled

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concavity; paraproct slightly longer than wide at base, and rounded distally. Male with same coloration as female; gonostylus subcylindrical with slight curvature, slightly shorter than gonocoxite; ventral plate nearly wider than long with or without very slight lateral concavity. Pupal cocoon with compact weave, threads evident; frontoclypeal and thoracic trichomes with 2-8 branches; gill with 8 branches. Larval cephalic apotome mostly uniformly colored without particular ornamentation; postgenal cleft deep ,reaching base of hypostomium. Five species in the group: S. antillarum, S. dinellii, S. ochraceum, S. nuneztovari, S. shewellianum. KEYS

TO

SIMULIUM (ECTEMNASPIS)

SPECIES

Females Simulium flavipictum not included. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(4) – 7(6) – 8(3) – 9(8) –

Scutum yellowish with median and 1+1 sublateral areas orange to blackish brown (Figs. 84 A, B); metanotum predominately blackish brown or totally blackish gray ................................................................................................................................................ 2 Scutum totally yellowish; metanotum predominately yellowish ............................. 16 Cibarium untoothed and with thin sclerotized rim (Figs. 84 I, J) ............................. ....................................................................................... BICOLORATUM sp. group......3 Cibarium toothed with border of median area depressed and covered with tubercles, and with acute lateral teeth (Figs. 84 K-P) ................................................... ................................................................................................ ROMANAI sp. group......11 Basal sector of R without hair (Fig. 85 A) ................................................................... 4 Basal sector of R with hair .............................................................................................. 8 Scutum yellowish orange, without brown color (Fig. 84 A) ..................................... 5 Scutum yellowish to brown dark on disc ...................................................................... 6 Wing length 3.0 mm; abdominal tergites I-IV yellowish; paraproct length about 1/ 4 longer than cerci width at base ........................................................... Simulium antonii Wing length 2.4-2.6 mm; abdominal tergites I-III yellowish; paraproct as long as cerci width at base (Figs. 85 G, H) ........................ Simulium pautense, Simulium pifanoi Wing length 2.8-2.9 mm; paraproct longer than basal width (Fig. 85 I) .................. ......................................................................................................................... Simulium tunja Wing length 2.4-2.7 mm; paraproct about as long as basal width (Fig. 85 E) ...... 7 Abdominal tergites I-IV totally grayish yellow .............................................................. ......................................... Simulium furcillatum, Simulium bicornutum, Simulium tolimaense Abdominal tergites I-IV yellow with tergal plates III-IV brownish on posterior border ...................................................................................................Simulium arcabucense Scutellum and abdominal tergites I-IV brownish yellow .......................................... 9 Scutellum yellowish or greenish orange; abdominal tergites II-IV yellow to yellowish brown ................................................................................................................ 10 Maxillary palp sensory vesicle length less than 1/3 palpomere length (Fig. 84 Y) .................................................................................................................... Simulium ignescens Maxillary palp sensory vesicle length nearly 1/2 palpomere length (Fig. 84 X) .... ...............................................................................................................Simulium rubiginosum

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Scutum totally yellow; abdominal tergites I-III yellowish, IV brown on tergal plate; length of paraproct about as long as basal width (Fig. 85 F) .......................... .................................................................................................................Simulium roquemayu Scutum generally yellow with extensive black (Fig. 84 B), but central area from brown to orange; abdominal tergites I-II yellowish orange, III-IV blackish brown; length of paraproct about 1/3 longer than basal width .............................................. ...................................... Simulium bicoloratum, Simulium cormonsi, Simulium jaimeramirezi Basal portion of R glabrous (Fig. 85 A); basal portion of cibarium with rounded tubercles medially and sparse and diminutive spiculae laterally (Fig. 84 K) ........... ................................................................................................................... Simulium albanense Basal portion of R with hair; basal portion of cibarium with tubercles or small plates of different shape medially and generally with abundant small teeth-like spines laterally (Figs. 84 L-P) ......................................................................................... 12 Scutum, scutellum, metanotum and abdomen blackish (Fig. 84 C); claw without subbasal tooth (Fig. 85 B) ............................................................... Simulium mayuchuspi Scutum, scutellum, metanotum and abdomen yellowish in variable proportion; claw with subbasal tooth (Fig. 85 C) ............................................................................ 13 Scutum orange; median area of cibarium with subovoidal tubercles (Fig. 84 L) ...................................................................................................................Simulium gabaldoni Scutum yellowish orange but frequently with blackish brown spots on median and 1+1 sublateral areas; cibarium with subquadrate to subspheroidal small plates on median area (Figs. 84 M-P) ............................................................................................ 14 Abdominal tergites I-II (III) yellowish ........................................... Simulium lutzianum Abdominal tergites I yellowish ...................................................................................... 15 Abdominal tergites II-IV yellowish to yellowish brown with median tergal plate on III-V dark brown (Fig. 84 G); Sc with 0-3 hairs ................... Simulium wolffhuegeli Abdominal tergites II-IV blackish brown to black; Sc with 6-9 hairs ...................... .................................................................................................................... Simulium romanai Abdominal tergites IV-VIII generally blackish brown with tergal plate yellowish; cibarium median area with 1+1 submedian wide and short elevations covered by strong teeth followed by minute teeth (Figs. 84 Q, R); paraproct apically acuminate (Figs. 85 M, N) to moderately blunt (Fig. 85 K) ....................................... ......................................................................................... PERFLAVUM sp. group......17 Abdominal tergites I-IV yellowish (Fig. 84 H); tergites V-VII from yellow to yellowish brown or yellowish to light gray; cibarium median area with 1+1 submedian elongated processes flanking median depressed well-sclerotized area, superficially from smooth to wrinkled (Figs. 84 T-W); paraproct blunt apically (Figs. 85 O-Q) ...................................................................... DINELLII sp. group......21 Legs and abdominal tergites all yellowish; fronto-ocular triangle higher than wide; paraproct apically blunt (Fig. 85 K); wing length 2.5-2.7 mm ......... Simulium perflavum Legs and abdominal tergites, at least IV-VII, blackish brown, paraproct generally acuminate apically (Figs. 85 M, N); wing length 2.2-2.3 mm ................................. 18 Metanotum yellow to yellowish brown ....................................................................... 19 Metanotum blackish ........................................................................................................ 20

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19(18) Sensory vesicle of maxillary palp about 1/3-1/4 length of palpomere (Fig. 84 b) ........................................................................... Simulium maroniense, Simulium trombetense – Sensory vesicle of maxillary palp about 1/2 length of palpomere (Fig. 84 c) ....... .................................................................................................................. Simulium rorotaense 20(18) Abdominal tergite III yellowish; ratio of paraproct length/width at base = 1.3 (Fig. 85 L) .......................................................................................... Simulium kabanayense – Abdominal tergite III blackish; ratio of paraproct length/width at base = 1.7 (Fig. 85 N) ........................................................................................................... Simulium suarezi 21(16) Scutum homogeneously pigmented without silvery submedian longitudinal bands .............................................................................................................................................. 22 – Scutum with 1+1 short silvery submedian longitudinal bands (Fig. 84 E) ......... 23 22(21) Sensory vesicle of maxillary palp shorter than 1/2 of palpomere length, (Fig. 84 a); cibarium with 1+1 acuminate submedian processes (Fig. 84 U); wing length 1.9 mm ..................................................................................................... Simulium shewellianum – Sensory vesicle of maxillary palp longer than 1/2 of palpomere length; cibarium with 1+1 submedian processes blunt (Fig. 84 T); wing length about 2.1 mm ....... .................................................................................. Simulium dinellii, Simulium nuneztovari 23(21) Wing 2.2-2.3 mm; scutum matte yellow with 1+1 wedges connected with silvery anterolateral area (Fig. 84 E); cibarium with median depressed area wrinkled or with small spines (Fig. 84 V); metanotum dark brown ................ Simulium ochraceum – Wing 1.9 mm; scutum reddish with 1+1 wedges not well connected with anterolateral silvery area (Fig. 84 D); cibarium with median depressed area smooth, without spines or wrinkling and bordered basally by thick rim (Fig. 84 W); metanotum yellowish .................................................................. Simulium antillarum Males Unknown: Simulium anaimense, S. antonii, S. roquemayu, S. tolimaens. 1 – 2(1) – 3(2) –

Scutum, scutellum, metanotum and abdomen dark grayish; scutum with posterior 1/4 black (Fig. 86 C); ventral plate with strong lateral constrictions (Fig. 87 G) ............................................................................................................... Simulium mayuchuspi Scutum and abdomen all yellowish, or having this pigment reduced to border of scutum and abdominal tergite I (Figs. 86 A, D; ventral plate with or without lateral constrictions (Figs. 87 B, H) ............................................................................................ 2 Scutum yellowish orange, with black in different proportions from small dark brown to blackish spots on median and sublateral areas of disc (Fig. 86 B); metanotum blackish ........................................................................................................... 3 Scutum yellowish; metanotum brownish yellow to dark brown ............................ 14 Gonostylus about 1/2 to 2/3 of gonocoxite length, subquadrate with distal border truncate (Figs. 86 M, O); ventral plate wide with length/width ratio = 3.04.0 (Figs. 87 E, I) ................................................................................................................ 4 Gonostylus about 3/4 or more of gonocoxite length, subcylindrical with acuminate distal border (Figs. 86 K, N); ventral plate not as wide as in above, with length/width ratio = 1.1-1.9 (Figs. 87 D, H) ............................................................... 5

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Scutum black, with yellow area restricted to pronotal angles, laterally and narrow posterior border; scutellum dark brown and abdomen blackish; gonostylus with 810 terminal spurs (Fig. 86 O) .......................................................... Simulium wolffhuegeli – Scutum yellowish with blackish pigmentation on disc as in Simulium romanai (Fig. 86 D), in some specimens; abdominal tergites I-III yellowish; gonostylus with 1 apical spur (Fig. 86 M) ......................................................................... Simulium albanense 5(3) Lateral areas of ventral plate strongly concave (Fig. 87 H) ...................................... 6 – Lateral areas of ventral plate slightly concave (Figs. 87 A, C, D) ........................... 7 6(5) Length/width ratio of hind basitarsus = 4.8-5.1; scutum frequently mostly black as in figure 86 D ..................................................................................... Simulium romanai – Length/width ratio of hind basitarsus = 6.5-7.6; scutum yellowish orange ........... ................................................................................................................. Simulium lutzianum 7(5) Basal sector of R haired ................................................................................................... 8 – Basal sector of R glabrous ............................................................................................... 9 8(7) Abdominal tergites I-IV yellowish brown, the following tergites black ................... .............................................................................. Simulium rubiginosum, Simulium ignescens – All abdominal tergites black ................... Simulium bicoloratum, Simulium jaimeramirezi 9(7) Wing length 2.9-3.0 mm; abdominal tergites I-III or IV yellowish brown to yellow (Fig. 86 F); ventral plate with slight lateral concavity ............................................... 10 – Wing length 2.5 mm; abdominal tergites I-II or I-III yellowish, the following tergites blackish; ventral plate laterally convex or with slight concavity .............. 11 10(9) Abdominal tergites I-IV greenish yellow with small dark brown area on tergite IV (Fig. 86 F); length/width ratio of posterior basitarsus = 3.5 ................. Simulium bicornutum – Abdominal tergites I-III yellowish brown with large brown dark area on tergite IV (Fig. 86 G); length/width ratio of posterior basitarsus = 4.0 ......... Simulium furcillatum 11(9) Ventral plate laterally convex (Fig. 87 B); gonostylus shorter than gonocoxite (Figs. 86 I-J) ....................................................................................................................... 12 – Ventral plate not convex laterally (Figs. 87 A, D); gonostylus about same length as gonocoxite (Figs. 86 H, L) ............................................................................................. 13 12(11) Scutum with 1+1 elongated lateral blackish stripe and median T shaped one on anterior 1/3 (Fig. 86 B) ......................................................................... Simulium pautense – Scutum with 1+1 short lateral brownish stripes ................................ Simulium pifanoi 13(11) Length/width ratio of posterior basitarsus = 4.3; wing length 2.7 mm; abdominal tergite III yellowish with tergal plate brownish to black .................... Simulium tunja – Length/width ratio of posterior basitarsus = 7.5; wing length 2.5 mm; abdominal tergite III yellow with tergal plate brownish ................................ Simulium arcabucense 14(2) Ventral plate with strong lateral concavity (Fig. 87 F); abdominal tergites black ...................................................................................................................Simulium gabaldoni – Ventral plate generally without strong lateral concavity (Fig. 87 O); abdominal tergites all yellow or reddish-yellowish brown ........................................................... 15 15(14) Scutum with short silvery anterior submedian bands (Fig. 86 E); ventral plate laterally convex (Fig. 87 O) ............................................................................................ 16 – Scutum homogeneously ornamented, without bands; ventral plate generally with small lateral concavity (Figs. 87 J-N) ........................................................................... 17

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16(15) Scutum yellowish orange, with 1+1 silvery stripes on anterior third ....................... .................................................................................................................. Simulium ochraceum – Scutum yellowish-reddish brown, with 1+1 silvery stripes on anterior half or surpassing it .......................................................................................... Simulium antillarum 17( 15) Ventral plate distal portion narrowing slowly (Fig. 87 N); gonostylus shorter than gonocoxite (Fig. 86 S) ..................................................................................................... 18 – Ventral plate distal portion narrowing abruptly after 1/2 its length (Figs. 87 K-M); gonostylus not shorter than gonocoxite (Figs. 86 P-R) ............................................... ......................................................................................... PERFLAVUM sp. group......20 18(17) Abdomen blackish brown; wing length 1.8 mm ....................... Simulium shewellianum – Abdomen all reddish yellow to greenish yellow; wing length 2.0-2.5 mm .......... 19 19(18) Abdomen reddish yellow; wing length 2.5 mm .................................. Simulium dinellii – Abdomen greenish yellow; wing length 2.0 mm ........................ Simulium nuneztovari 20(17) Metanotum and abdomen yellowish; gonostylus longer than gonocoxite; wing length 2.6-2.7 mm ................................................................................ Simulium perflavum – Metanotum and abdomen brownish; gonostylus not longer than gonocoxite; wing length 1.8-2.6 mm ............................................................................................................ 21 21(20) All abdominal tergites brown to blackish brown; wing length 2.0-2.1 mm ............. ................................................................................................................ Simulium trombetense – Abdominal tergites I or II yellowish to yellowish brown, remaining tergites blackish brown; wing length 1.8-2.6 mm .................................................................... 22 22(21) Abdominal tergite I brown or yellowish brown, remaining tergites dark brown ............................................................................... Simulium suarezi, Simulium kabanayense – Abdominal tergites I-II yellowish brown, remaining tergites dark brown ............... .............................................................................. Simulium maroniense, Simulium rorotaense Pupae Unknown: Simulium antonii. 1 – 2(1) – 3(2) – 4(1) – 5(4)

Gill composed of 2-5 branches; cocoon with prominent anterodorsal projection (Figs. 88 C, D, F) ................................................................................................................ 2 Gill composed of 6-250 filaments; cocoon without prominent anterodorsal projection ............................................................................................................................. 4 Gill branches relatively thin (Fig. 88 F) ........................................... Simulium tolimaense Gill composed of 2-4 very thick branches ................................................................... 3 Gill with 2 similar branches and small internal protuberance on each (Fig. 88 C) ................................................................................................................ Simulium bicornutum Gill with 4 branches, 1 thick inferior branch and 3 thin ones positioned overhead (Fig. 88 D) .............................................................................................Simulium furcillatum Cocoon with anterior open loops (Fig. 88 L); gill with 17-250 relatively thick branches (Figs. 89 N, O) ................................................................................................... 5 Cocoon without anterior open loops (Fig. 88 G); gill with 6-12 thin branches .......... 8 Gill with 7 thick and short basal branches with 100 to 250 thin, short filaments (Fig. 89 O) ........................................................................................... Simulium trombetense

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– 8(4) – 9(8) – 10(9) – 11(8) – 12(11) – 13(12) – 14(13) – 15(12) – 16(15) – 17(11) –

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Gill with 14-23 non arborescent branches, without thin filaments ......................... 6 Gill with 20 relatively thick branches emerging from enlarged short common trunk (Fig. 89 N) ...................................................................................... Simulium suarezi Gill with 10-23 thin branches basally emerging from primary, secondary, tertiary or more subdivided branches .......................................................................................... 7 Gill branch length changing gradually from top to bottom, dorsal branches shorter than ventral branches (Fig. 89 L); in frontal view, filaments form rosette basally; frontoclypeus and thorax with abundant tubercles and trichomes with robust basal portion (Fig. 88 V) ...................................................... Simulium maroniense Gill branches all about same length (Fig. 89 M); in frontal view not forming rosette basally; frontoclypeus and thorax with sparse tubercles and trichomes with basal portion not robust ..................................................................... Simulium rorotaense Gill with 10-12 filaments .................................................................................................. 9 Gill with 6-8 filaments .................................................................................................... 11 Frontoclypeus without tubercles except for small patch on base; frontoclypeal trichomes, with 3-6 branches; gills with 10 filaments .................... Simulium ignescens Frontoclypeus with regularly spaced tubercles; frontoclypeal trichomes with 1-2 branches (Fig. 88 Q); gills with 10-12 filaments (Fig. 89 K) ................................... 10 Frontoclypeus and thorax with few tubercles; gill about as long as cocoon length; gill with 10-12 filaments (Fig. 89 K) ............................................ Simulium kabanayense Frontoclypeus and thorax with abundant tubercles (Fig. 88 Q); gill shorter than cocoon length (Fig. 88 G); gill with 10 filaments ................................. Simulium tunja Gill with 6 branches ........................................................................................................ 12 Gill with 8 branches ........................................................................................................ 17 Cocoon whitish with enlarged anterior margin and with dorsal longitudinal ridge (Fig. 88 H) .......................................................................................................................... 13 Cocoon not light colored, without enlarged anterior margin, without dorsal ridge .............................................................................................................................................. 15 Cephalic and thoracic trichomes curled at base (Fig. 88 Y); gill branches very open (Fig. 89 F) ..............................................................................................Simulium anaimense Cephalic and thoracic trichomes straight at base (similar to Fig. 88 P); gill branches not very open .................................................................................................. 14 Cocoon anterior edge strongly oblique ..................................... Simulium jaimeramirezi Cocoon anterior edge slightly oblique ............................................... Simulium cormonsi Gill about same length as cocoon length (Figs. 88 A, E) ........................................ 16 Gill shorter than cocoon length ......................................................... Simulium gabaldoni Cocoon without anteroventral projection (Fig. 88 A); bifurcation of tertiary branches near base (Fig. 89 A) ........................................................ Simulium arcabucense Cocoon with anteroventral projection (Fig. 88 E), bifurcation of median and ventral tertiary branches not near base (Fig. 89 B) ......................... Simulium pautense Gill terminal branches emerging very close to base, very short primary and secondary branches (Fig. 89 J) .......................................................... Simulium perflavum Gill terminal branches not emerging close to base, longer primary and secondary branches (Figs. 89 C, D) ................................................................................................. 18

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18(17) Frontoclypeal and thoracic tubercles longer than wide and acuminate distally (Fig. 88 Z) ........................................................................................................ Simulium albanense – Frontoclypeal and thoracic tubercles shorter than wide and blunt distally ......... 19 19(18) Cocoon usually soft and thick; threads not very evident, with felt-like or spongy texture, generally with thickened anterior border and dorsal longitudinal ridge (Figs. 88 B, I, J) ................................................................................................................. 20 – Cocoon compact; threads thin, evident, anterior border can be reinforced but not thickened and without dorsal ridge (Fig. 88 K) ......................................................... 25 20(21) Frontoclypeus and thorax with 35-40 branches (Fig. 88 M); gill with minute tubercles giving wrinkled appearance ........................................... Simulium bicoloratum – Frontoclypeus and thorax with 3-8 branches; gill branches without minute tubercles, having smooth appearance .......................................................................... 21 21(20) Gill branches relatively open at base (Fig. 89 H) ...................................................... 22 – Gill branches relatively closed at base (Fig. 89 G) .................................................... 23 22(21) Frontoclypeus and thorax with sparse tubercles ....................................................... ............................................................................................ (in part)......Simulium lutzianum – Frontoclypeus and thorax with abundant tubercles (Fig. 88 T) ........................... .................................................................................................................... Simulium romanai 23(21) Frontoclypeus and thorax with sparse tubercles (Fig. 88 R) ..... Simulium lutzianum – Frontoclypeus and thorax with moderate (Fig. 88 O) to abundant tubercles (Fig. 88 S) .................................................................................................................................... 24 24(23) Frontoclypeus and thorax with sparse tubercles (Fig. 88 O); trichomes with 3-7 branches ...................................................................................................... Simulium pifanoi – Frontoclypeus and thorax with abundant tubercles (Fig. 88 S); trichomes with 712 branches ......................................................................................... Simulium mayuchuspi 25(19) Tertiary branches emerging beyond midlength of gill (Fig. 89 D) ........................... .................................................................................................................Simulium roquemayu – Tertiary branches emerging at basal 1/3 of gill (Figs. 89 E, I) .............................. 26 26(25) Frontoclypeal and thoracic trichomes with 2-4 branches relatively elongated (Fig. 88 X) .................................................................................................. Simulium shewellianum – Frontoclypeal and thoracic trichomes with variable number of not so elongated branches (Fig. 88 W) ........................................................................................................ 27 27(26) Frontoclypeus and thorax with sparse tubercles (Figs. 88 P, W) ........................... 28 – Frontoclypeus and thorax with abundant tubercles (Fig. 88 U) ............................ 29 28(27) Cocoon prolonged dorsally and basally; gill length 2.7-3.5 mm; tubercles of frontoclypeus absent or sparse (Fig. 88 P) ..................................Simulium rubiginosum – Cocoon without anterior projections; gill length 2.0-2.2 mm; tubercles more abundant than described above (Fig. 88 W) ........................................ Simulium dinellii 29(27) Cocoon with open weave ................................................................ Simulium nuneztovari – Cocoon compact, without open weave (Fig. 88 K) .................................................. 30 30(29) Thoracic trichomes with 4-9 branches; arrangement of primary and secondary branches not close to base (Fig. 89 I) ............................................ Simulium wolffhuegeli – Thoracic trichomes with 2-6 branches; arrangement of primary and secondary branches close to base ..................................... Simulium ochraceum, Simulium antillarum

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Larvae Unknown: Simulium antonii, S. cormonsi, S. gabaldoni, S. jaimeramirezi, S. nuneztovari, S. roquemayu, S. tolimaense. 1 – 2(1) – 3(2) – 4(3) – 5(1) – 6(5) – 7(6) – 8(6) – 9(8) – 10(9) –

Medial antennal segment similar or smaller than proximal one (Fig. 90 O); anal ring with 84-100 rows of hooks; cephalic apotome darkened on basal border and with median longitudinal band (Fig. 90 H) .................................................................. 2 Medial antennal segment usually longer than proximal one (Fig. 90 N); anal ring, usually with fewer than 84 rows of hooks; cephalic apotome with different ornamentation ..................................................................................................................... 5 Ratio of length of postgenal bridge/hypostomium = 0.1-0.2 (Fig. 90 K) .............. .............................................................................. Simulium maroniense, Simulium rorotaense Ratio of length of postgenal bridge/hypostomium = 0.3-0.4 ................................. 3 Anal ring with about 84 rows of hooks; ratio of length of postgenal bridge/ hypostomium = 0.4 ......................................................................... Simulium kabanayense Anal ring with 96-110 rows of hooks; ratio of length of postgenal bridge/ hypostomium = 0.3 ........................................................................................................... 4 Ratio of antennal segments: proximal, medial and distal = 1:0.9-1.0:1.1-1.3; cephalic fan with 45-50 rays ............................................................ Simulium trombetense Ratio of antennal segments: proximal, medial and distal = 1:0.8:1.2; cephalic fan with 40 rays ............................................................................................... Simulium suarezi Anal ring with over 80 rows of hooks; body length usually over 6.0 mm ............ 6 Anal ring with less than 80 rows of hooks; body length usually less than 6.0 mm .............................................................................................................................................. 11 Ratio of length of postgenal bridge/hypostomium = 0.3-0.4 (Fig. 90 J); anal ring with 83-85 rows of hooks ................................................................................................ 7 Ratio of length of postgenal bridge/hypostomium = 0.2; anal ring with variable number of rows .................................................................................................................. 8 Body length 5.7 mm; cephalic fan with 35 rays ........................... Simulium furcillatum Body length 6.0-6.7 mm; cephalic fan with 45-50 rays ................................................ ......................................................................... Simulium rubiginosum, Simulium bicoloratum Anal ring with 86-88 rows with 17-20 hooks each ......................... Simulium ignescens Anal ring with 77-85 rows and 12-16 hooks each ...................................................... 9 Cephalic fan with about 30-32 rays; rectal papillae with 8-11 diverticula on each lobe; anal sclerite without hairs .............................................................. Simulium pifanoi Cephalic fan with about 37-40 rays; rectal papillae with 10-16 diverticula on each lobe (Fig. 90 T); anal sclerite with hairs ...................................................................... 10 Medial segment of antenna longer than distal one (Fig. 90 N); ratio of antennal segments: proximal, medial and distal =1:1.7:1.4-1.6; body length 6.5 mm ........... .................................................................................................................. Simulium perflavum Medial segment of antenna shorter than distal one (Fig. 90 M); ratio of antennal segments: proximal, medial and distal = 1:1.7:2.5; body length 5.8 mm ................. ................................................................................................................ Simulium bicornutum

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11(5) – 12(11) – 13(12) – 14(13) – 15(14) – 16(14) – 17(12) – 18(17) – 19(18) – 20(19) –

CLAVES

Ratio of length of postgenal bridge/hypostomium 0.2-0.3; cephalic apotome darkened subbasally with lighter submedian area (Fig. 90 F) ........ Simulium wolffhuegeli Ratio of length of postgenal bridge/hypostomium less than 0.10-0.25; cephalic apotome darkened mediobasally with lighter subbasal window (Figs. 90 B, D, E) .............................................................................................................................................. 12 Anal ring with 70-79 rows of hooks ............................................................................ 13 Anal ring with fewer than 69 rows of hooks ............................................................. 17 Cephalic fan with 42-48 rays ............................................................... Simulium pautense Cephalic fan with 27-39 rays ......................................................................................... 14 Cephalic fan with 27-35 rays ......................................................................................... 15 Cephalic fan with 36-39 rays ......................................................................................... 16 Body length 4.8-5.2 mm; postgenal cleft subtrianglular (Fig. 90 I); cephalic apotome without darkened areas and with very few hairs (Fig. 90 C) .......... Simulium arcabucense Body length 6.0 mm; postgenal cleft dome shape (Fig. 90 J); cephalic apotome slightly darkened on basal border and with abundant hairs (Fig. 90 D) ....... Simulium albanense Body length about 6.0 mm ..................................................................... Simulium dinellii Body length about 4.6 mm ............................ Simulium ochraceum, Simulium antillarum Cephalic fan with about 47 rays; anal ring with 52 rows of hooks ...................... 18 Cephalic fan with about 35 rays; anal ring with 55-74 rows of hooks .................... ................................................................................................................. Simulium lutzianum Anal sclerite with trichomes and scales (Fig. 90 R) ............................. Simulium tunja Anal sclerite with only trichomes, without scales ..................................................... 19 Body length about 6.0 mm .................................................................. Simulium romanai Body length 4.3-4.7 mm ................................................................................................. 20 Body integument with foliaceus trichomes (Fig. 90 P); ratio of length of postgenal bridge/hypostomium = 0.2 ...............................................................Simulium anaimense Body integument without foliaceus trichomes; ratio of length of postgenal bridge/hypostomium = 0.1 ..................... Simulium mayuchuspi, Simulium shewellianum DE ESPECIES DE

SIMULIUM (ECTEMNASPIS)

Hembras 1 – 2(1) – 3(2) – 4(3) –

Escudo amarillento con área mediana y 1+1 áreas sublaterales anaranjadas hasta a veces castaño negruzco (Figs. 84 A, B); metanoto predominantemente castaño negruzco o totalmente gris negruzco ............................................................................ 2 Escudo totalmente amarillento; metanoto predominantemente amarillento ...... 16 Cibario sin dientes y con borde fino esclerotizado (Figs. 84 I, J) ............................. ....................................................................................... grupo sp. BICOLORATUM......3 Cibario dentado con área mediana de borde deprimido, cubierto con tubérculos y lateralmente con dientes aguzados (Figs. 84 K-P) ......... grupo sp. ROMANAI......11 Sector basal de R sin pelos (Fig. 85 A) .......................................................................... 4 Sector basal de R con pelos ............................................................................................. 8 Escudo naranja amarillento, sin color castaño (Fig. 84 A) ........................................ 5 Escudo amarillento a castaño oscuro sobre el disco .................................................. 6

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Largo ala 3,0 mm; tergitos abdominales I-IV amarillentos; largo de paraprocto aproximadamente 1/4 del ancho del cerco en la base ...................... Simulium antonii – Largo ala 2,4-2,6 mm; tergitos abdominales I-III amarillentos; paraprocto tan largo como el ancho del circo en la base (Figs. 85 G, H) ......Simulium pautense, Simulium pifanoi 6(4) Largo ala 2,8-2,9 mm; paraprocto más largo que ancho en la base (Fig. 85 I) ...... ......................................................................................................................... Simulium tunja – Largo ala 2,4-2,7 mm; paraprocto aproximadamente similar al ancho en la base (Fig. 85 E) ............................................................................................................................ 7 7(6) Tergitos abdominales I-IV totalmente amarillo grisáceo ............................................. ......................................... Simulium bicornutum, Simulium furcillatum, Simulium tolimaense – Tergitos abdominales I-IV amarillo con placas tergales III-IV castaño sobre el borde posterior ................................................................................... Simulium arcabucense 8(7) Escutelo y tergitos abdominales I-IV castaño amarillento ........................................ 9 – Escutelo naranja amarillento o naranja verdoso; tergitos abdominales II-IV amarillo a castaño amarillento ....................................................................................... 10 9(8) Longitud vesícula sensorial del palpo maxilar menos de 1/3 de la longitud del palpómero (Fig. 84 Y) ........................................................................... Simulium ignescens – Longitud vesícula sensorial del palpo maxilar menos de 1/2 de la longitud del palpómero (Fig. 84 X) ......................................................................Simulium rubiginosum 10(8) Escudo totalmente amarillo, tergitos abdominales I-III amarillentos, IV castaño sobre placa tergal; longitud de paraprocto aproximadamente tan largo como ancho en la base (Fig. 85 F) ...............................................................Simuliun roquemayu – Escudo generalmente amarillo con abundante negro (Fig. 84 B), con área central desde castaño a anaranjado; tergitos abdominales I-II amarillo-anaranjado, III- IV castaño negruzco; longitud del paraprocto aproximadamente 1/3 mas largo que el ancho basal tan largo como ancho en la basal ............................................................... ....................................... Simuliun bicoloratum, Simulium cormonsi, Simulium jaimiramirezi 11(2) Porción basal de R glabro (Fig. 85 A); porción basal del cibario con tubérculos redondeados medianamente, y escasos y diminutas espículas lateralmente (Fig. 84 K) .............................................................................................................. Simulium albanense – Porción basal de R con pelos; porción basal del cibario con tubérculos y pequeñas placas de diferente forma medianamente, y generalmente con dientes abundantes y pequeños como espinas lateralmente (Figs. 84 L-P) ............................................. 12 12(11) Escudo, escudete, metanoto y abdomen negruzcos (Fig. 84 C); uña sin diente subbasal (Fig. 85 B) ........................................................................... Simulium mayuchuspi – Escudo, escudete, metanoto y abdomen amarillentos en proporción variable; uña con diente subbasal (Fig. 85 C) ..................................................................................... 13 13(12) Escudo naranja; área mediana del cibario con tubérculos subovoidales (Fig. 84 L) ...................................................................................................................Simulium gabaldoni – Escudo naranja amarillento pero frecuentemente con manchas castaño negruzcas medialmente y 1+1 áreas sublaterales; cibario con pequeñas placas subcuadradas a subesferoidales sobre el área mediana (Figs. 84 M-P) ............................................. 14 14(13) Tergitos abdominales I-II (III) amarillentos ................................. Simulium lutzianum – Tergitos abdominales I amarillento .............................................................................. 15

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15(14) Tergitos abdominales II-IV amarillentos a castaño amarillentos, con la porción mediana de las placas tergales del III-V castaño oscuro (Fig. 84 G); Sc con 0-3 pelos ...................................................................................................... Simulium wolffhuegeli – Tergitos abdominales II-IV castaño negruzco a negro; Sc con 6-9 pelos ............... .................................................................................................................... Simulium romanai 16(1) Tergitos abdominales IV-VIII generalmente castaño negruzco con placa tergal amarillenta; área mediana del cibario con 1+1 elevación submediana ancha, corta y cubiertas por fuertes dientes seguidos por pequeños dientes (Figs. 84 Q, R); paraprocto apicalmente desde aguzado (Figs. 85 M, N) a moderadamente romo (Fig. 85 K) ..................................................................... grupo sp. PERFLAVUM......17 – Tergitos abdominales I-IV amarillentos (Fig. 84 H); tergitos V-VII desde amarillo a castaño amarillento o amarillo a gris claro; área mediana del cibario con 1+1 prolongaciones submedianas alargadas, flanqueando un área mediana deprimida bien esclerotizada, superficialmente desde lisa a rugosa (Figs. 84 T-W); paraprocto romo apicalmente (Figs. 85 O-Q) .................................. grupo sp. DINELLII......21 17(16) Patas y tergitos abdominales totalmente amarillentos; triángulo fronto-ocular más alto que ancho; paraprocto apicalmente romo (Fig. 85 K); largo ala 2,5-2,7 mm .................................................................................................................. Simulium perflavum – Tergitos abdominales presentando por lo menos IV-VII castaño negruzco; paraprocto apicalmente generalmente aguzado (Figs. 85 M, N); largo ala 2,2-2,3 mm ............... 18 18(17) Metanoto amarillo a castaño amarillento .................................................................... 19 – Metanoto negruzco .......................................................................................................... 20 19(18) Vesícula sensorial del palpo maxilar alrededor de 1/3-1/4 del largo del palpómero (Fig. 84 b) ........................................................ Simulium maroniense, Simulium trombetense – Vesícula sensorial del palpo maxilar alrededor de 1/2 del largo del palpómero (Fig. 84 c) ............................................................................................... Simulium rorotaense 20(18) Tergito abdominal III amarillento; relación del largo/ancho en la base del paraprocto = 1,3 (Fig. 85 L) .......................................................... Simulium kabanayense – Tergito abdominal III negruzco; relación del largo/ancho en la base del paraprocto = 1,7 (Fig. 85 N) ................................................................. Simulium suarezi 21(16) Escudo homogeneamente pigmentado sin bandas plateadas submedianas ........ 22 – Escudo con 1+1 bandas plateadas submedianas cortas (Fig. 84 E) ..................... 23 22(21) Vesícula sensorial del palpo maxilar más corta que lo mitad del largo del palpómero (Fig. 84 A); cibario con 1+1 prolongación submediana aguzada (Fig. 84 U); largo ala 1,9 mm ................................................................. Simulium shewellianum – Vesícula sensorial del palpo maxilar más larga que la mitad del largo del palpómero; cibario con 1+1 prolongación submediana roma (Fig. 84 T); largo ala 2,1 mm ....... .................................................................................. Simulium dinellii, Simulium nuneztovari 23(21) Largo del ala 2,2-2,3 mm; escudo amarillo opaco con 1+1 cuñas plateadas conectadas con el área plateada anterolateral (Fig. 84 E); área mediana deprimida del cibario rugosa o con pequeñas espinas (Fig. 84 V); metanoto castaño oscuro ......... Simulium ochraceum – Largo del ala 1,9 mm; escudo amarillo rojizo con 1+1 cuñas no bien conectadas con el área plateada anterolateral (Fig. 84 D); área mediana deprimida del cibario lisa, sin espinas, no rugosa y bordeada basalmente por un grueso borde (Fig. 84 W); metanoto amarillento .................................................................. Simulium antillarum

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Machos Desconocidos: Simulium anaimense, S. antonii, S. roquemayu, S. tolimaense. 1 – 2(1) – 3(2) – 4(3) – 5(3) – 6(5) – 7(5) – 8(7) – 9(7) – 10(9)

Escudo, escudete, metanoto y abdomen gris oscuro; escudo con el área del cuarto posterior negra (Fig. 86 C); placa ventral con una fuerte contracción lateral (Fig. 87 G) .................................................................................................... Simulium mayuchuspi Escudo y abdomen desde todo amarillo o reducido este pigmento al borde del escudo y al tergito abdominal I (Figs. 86 A, D); placa ventral con o sin contracción lateral (Fig. 87 B, H) .......................................................................................................... 2 Escudo naranja amarillento pasando al negro en diferente proporción desde pequeñas manchas castaño oscuro a negruzco sobre las áreas medianas y sublaterales del disco (Fig. 86 B); metanoto negruzco .............................................. 3 Escudo amarillento; metanoto amarillo castaño a castaño oscuro ........................ 14 Gonostilo aproximadamente 1/2 a 2/3 del largo del gonocoxito, subcuadrado con borde distal truncado (Figs. 86 M, O); placa ventral ancha con una relación de largo /ancho = 3.0-4.0 (Figs. 87 E, I) ........................................................................... 4 Gonostilo aproximadamente 3/4 o más del gonocoxito, subcilíndrico con borde distal aguzado (Figs. 86 K, N); placa ventral no tan ancha como en el dilema anterior con un índice de largo/ancho = 1,1-1,9 (Figs. 87 D, H) ........................... 5 Escudo negro, reducida el área amarilla lateralmente a los ángulos pronotales y al borde posterior angosto; escudete castaño oscuro y abdomen negruzco; gonostilo con 6-8 espolones terminales (Fig. 86 O) ..................................... Simulium wolffhuegeli Escudo amarillento con pigmentación negra sobre el disco (como en Simulium romanai Fig. 86 D), en algunos especimenes; tergito abdominal I-III amarillento, gonostilo con un espolón apical (Fig. 86 M) ................................... Simulium albanense Costados laterales de la placa ventral con una fuerte concavidad (Fig. 87 H) ..... 6 Costados laterales de la placa ventral con una suave concavidad (Figs. 87 A, C, D) ..... 7 Relación largo/ ancho del basitarso posterior = 4,8-5,1; escudo con frecuencia mayormente negro como Figura 86 D .............................................. Simulium romanai Relación largo/ancho del basitarso posterior = 6,5-7,6; escudo naranja amarillento ............................................................................................ Simulium lutzianum Sector basal de R con pelos ............................................................................................. 8 Sector basal de R glabro ................................................................................................... 9 Tergitos abdominales I-IV castaño amarillento, los siguientes tergitos negro ........ .............................................................................. Simulium rubiginosum, Simulium ignescens Tergitos abdominales todos negros .................................................................................. .................................................. Simulium bicoloratum, S, cormonsi, Simulium jaimeramirezi Largo ala 2,9-3,0 mm; tergitos abdominales I-III o IV castaño amarillento a amarillo (Fig. 86 F); placa ventral con escasa concavidad lateral .......................... 10 Largo ala 2,5 mm; tergitos abdominales I-II ó I-III amarillentos, los siguientes tergitos negruzcos, placa ventral lateralmente convexa o con suave concavidad ................... 11 Tergitos abdominales I-IV amarillo verdoso, con pequeña área castaño oscuro sobre el tergito IV (Fig. 86 F); relación largo/ancho del basitarso posterior = 3,5 ................................................................................................................ Simulium bicornutum

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Tergitos abdominales I-III amarillento, con una mancha grande sobre tergito IV castaño oscuro (Fig. 86 G); relación largo/ancho del basitarso posterior = 4,0 ... .................................................................................................................Simulium furcillatum Placa ventral convexa lateralmente (Fig. 87 B); gonostilo más corto que el gonocoxito (Figs. 86 I - J) .............................................................................................. 12 Placa ventral no convexa lateralmente (Figs. 87 A, D); gonostilo aproximadamente del mismo largo que el gonocoxito (Figs. 86 H, L) .................................................. 13 Escudo con 1+1 banda negruzca lateral alongada y una mediana en forma de T en el tercio anterior (Fig. 86 B) ................................................................. Simulium pautense Escudo con 1+1 banda castaña corta lateral ...................................... Simulium pifanoi Relación largo/ancho del basitarso posterior = 4,3; largo del ala 2,7 mm; tergito abdominal III amarillento con la placa tergal castaño a negro ......... Simulium tunja Relación largo/ancho del basitarso posterior = 7,5; largo del ala 2,5 mm; tergito abdominal III todo amarillo con placa tergal castaño ............... Simulium arcabucense Placa ventral con fuerte concavidad lateral (Fig. 87 F); tergitos abdominales negros ......................................................................................................Simulium gabaldoni Placa ventral generalmente sin fuerte concavidad lateral (Fig. 87 O); tergitos abdominales amarillos o castaño rojizo amarillentos ............................................... 15 Escudo con cortas bandas plateadas submedianas anteriores (Fig. 86 E); placa ventral lateralmente convexa (Fig. 87 O) .................................................................... 16 Escudo con ornamentación homogénea sin bandas; placa ventral generalmente con una pequeña concavidad lateral (Figs. 87 J-N) .................................................. 17 Escudo naranja amarillento, con 1+1 bandas plateadas submedianas en el tercio anterior ................................................................................................... Simulium ochraceum Escudo castaño rojizo amarillento con 1+1 bandas plateadas en la mitad anterior o sobrepasandola ................................................................................. Simulium antillarum Placa ventral con porción distal haciéndose angosta suavemente (Fig. 87N); gonostilo más corto que el gonocoxito (Fig. 86 S) ................................................... 18 Placa ventral con porción distal volviéndose abruptamente angosta después de la mitad de su largo (Figs. 87 K-M); gonostilo aproximadamente del mismo largo o más largo que el gonocoxito (Figs. 86 P-R) .......... grupo sp. PERFLAVUM......20 Abdomen castaño negruzco; largo ala 1,8 mm ........................ Simulium shewellianum Abdomen amarillo rojizo a amarillo verdoso; largo del ala 2,0-2,5 mm .............. 19 Abdomen amarillo rojizo; largo ala 2,5 mm ........................................ Simulium dinellii Abdomen amarillo verdoso; largo ala 2,0 mm ............................ Simulium nuneztovari Metanoto y abdomen amarillentos; gonostilo más largo que el gonocoxito; longitud ala 2,6-2,7 mm ...................................................................... Simulium perflavum Metanoto y abdomen amarillentos; gonostilo no más largo que el gonocoxito; longitud ala 1,8-2,6 mm .................................................................................................. 21 Todos los tergitos abdominales castaño a castaño negruzco; largo ala 2,0-2,1 mm ................................................................................................................ Simulium trombetense Tergitos abdominales I o II amarillo o castaño amarillento, restantes tergitos castaño negruzco; largo ala 1,8-2,6 mm ...................................................................... 22 Tergito abdominal I castaño o castaño amarillento, restantes castaño oscuro ....... ............................................................................... Simulium suarezi, Simulium kabanayense

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Tergito abdominal I-II castaño amarillento, restantes castaño oscuro ..................... .............................................................................. Simulium maroniense, Simulium rorotaense

Pupas Desconocida: Simulium antonii. 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(5) – 7(6)

– 8(4) – 9(8) – 10(9) –

Branquia compuesta por 2-5 ramas; capullo con prominente prolongación antero dorsal (Figs. 88 C, D, F) .................................................................................................... 2 Branquia compuesta por 6-250 filamentos; capullo sin prominente prolongación anterior-dorsal ..................................................................................................................... 4 Ramas relativamente finas (Fig. 88 F) .............................................. Simulium tolimaense Branquia compuesta por 2-4 ramas muy gruesas ....................................................... 3 Branquia compuesta por 2 ramas similares con una pequeña protuberancia sobre cada una (Fig. 88 C) ........................................................................... Simulium bicornutum Branquia con 4 ramas, 1 gruesa inferior y 3 finas sobrepuestas a la cabeza (Fig. 88 D) ............................................................................................................Simulium furcillatum Capullo con aberturas anteriores (Fig. 88 L); branquias relativamente gruesas en número de 17-250 (Figs. 89 N, O ) ................................................................................ 5 Capullo sin aberturas anteriores (Fig. 88 G); ramas de la branquia finas y en número de 6-12 .................................................................................................................. 8 Branquias con 7 ramas basales gruesas, y finos y cortos en número superior a 100 y hasta 250 (Fig. 89 O) ...................................................................... Simulium trombetense Branquias con 14-23 ramas no arborescentes, sin filamentos finos ....................... 6 Ramas de las branquias relativamente gruesas en número de 20, emergiendo de un corto y grueso tallo común (Fig. 89 N) .............................................. Simulium suarezi Ramas de las branquias no gruesas basalmente en número de 10-23 emergiendo desde ramas primarias, secundarias y terciarias o más ramas subdivididas ........... 7 Ramas de las branquias cambiando gradualmente de tamaño desde arriba hacia abajo, siendo las ramas dorsales más cortas que las ventrales (Fig. 89 L); en vista frontal los filamentos forman una roseta basalmente; frontoclípeo y tórax con abundantes tubérculos, y tricomas con porción basal robusta (Fig. 88 V) .......... Simulium maroniense Ramas branquiales todas del mismo largo (Fig. 89 M); en vista frontal los filamentosa no forman una roseta, frontoclípeo y tórax con moderado número de tubérculos, y tricomas con la porción basal no robusta .............. Simulium rorotaense Branquias con 10-12 filamentos ..................................................................................... 9 Branquias con 6-8 filamentos ........................................................................................ 11 Frontoclípeo sin tubérculos, excepto un pequeño grupo sobre la base de la frente; tricomas del frontoclípeo con 3-6 ramas; branquias con 10 filamentos ............... Simulium ignescens Frontoclípeo con tubérculos regularmente distribuidos; tricomas del frontoclípeo con 1-2 ramas (Fig. 88 Q); branquias con 10-12 filamentos (Fig. 89 K) ............. 10 Frontoclípeo y tórax con escasos tubérculos; branquias aproximadamente tan largas como el largo del capullo; branquias con 10-12 filamentos (Fig. 89 K) ...... .............................................................................................................. Simulium kabanayense Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 Q); branquias más cortas que el largo del capullo (Fig. 88 G); branquias con 10 filamentos ................ Simulium tunja

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Branquia con 6 ramas ...................................................................................................... 12 Branquia con 8 ramas ...................................................................................................... 17 Capullo blanquecino con el del capullo (Fig. 88 H) ................................................. 13 Capullo no blanquecino, sin margen anterior engrosado ni carena dorsal .......... 15 Tricomas cefálicos y torácicos con base en forma de rulo (Fig. 88 Y); ramas de las branquias bien abiertas (Fig. 89 F) ...................................................Simulium anaimense Tricomas cefálicos y torácicos con base recta (similar a Fig. 88 P); ramas de las branquias no muy abiertas ............................................................................................. 14 Borde anterior del capullo fuertemente oblicuo ..................... Simulium jaimeramirezi Borde anterior del capullo escasamente oblicuo ............................. Simulium cormonsi Largo de branquia aproximadamente igual que el del capullo (Figs. 88 A, E) .......... 16 Largo de branquia más corto que el del capullo ............................ Simulium gabaldoni Capullo sin prolongación anterior ventral (Fig. 88 A); bifurcación de las ramas terciarias próximas a la base (Fig. 89 A) ........................................ Simulium arcabucense Capullo con prolongación anterior ventral (Fig. 88 E); bifurcación de las ramas terciarias mediana y ventral no próximas a la base (Fig. 89 B) .... Simulium pautense Ramas terminales de las branquias emergiendo muy próximas a la base, mostrando ramas primarias y secundarias muy cortas (Fig. 89 J) ............................. .................................................................................................................. Simulium perflavum Ramas terminales de las branquias emergiendo no próximo a la base, mostrando ramas primarias y secundarias largas (Figs. 89 C, D) ............................................... 18 Tubérculos del frontoclípeo y tórax más largos que anchos y aguzados distalmente (Fig. 88 Z) ............................................................................................... Simulium albanense Tubérculos del frontoclípeo y tórax más cortos que anchos y romos distalmente .............................................................................................................................................. 19 Capullo generalmente blando y grueso; fibras del tejido no bien evidentes con aspecto de fieltro o esponjoso, generalmente con borde anterior engrosado y con una carena dorsal longitudinal (Figs. 88 B, I, J) ......................................................... 20 Capullo compacto, fino, fibras del tejido evidentes, borde anterior puede estar reforzado pero no engrosado y sin una carena dorsal (Fig. 88 K) ........................ 25 Tricomas del frontoclípeo y tórax con 35-40 ramas (Fig. 88 M); superficie de las ramas de las branquias con pequeños tubérculos dando apariencia rugosa ........... ............................................................................................................... Simulium bicoloratum Tricomas del frontoclípeo y tórax con 3-8 ramas; ramas de la branquia sin tubérculos y con apariencia lisa ..................................................................................... 21 Ramas de las branquias relativamente abiertas en la base (Fig. 89 H) ................. 22 Ramas de las branquias relativamente cerradas en la base (Fig. 89 G) ................ 23 Frontoclípeo y tórax con escasos tubérculos ........ (en parte)......Simulium lutzianum Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 T) ....... Simulium romanai Frontoclípeo y tórax con escasos tubérculos (Fig. 88 R) ........... Simulium lutzianum Frontoclípeo y tórax con medianamente (Fig. 88 O) a abundantes tubérculos (Fig. 88 S) .................................................................................................................................... 24 Frontoclípeo y tórax con escasos tubérculos (Fig. 88 O); tricomas con 3-7 ramas ....................................................................................................................... Simulium pifanoi

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Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 S); tricomas con 7-12 ramas .................................................................................................... Simulium mayuchuspi Ramas terciarias emergen sobrepasando la mitad del largo de la branquia (Fig. 89 D) .................................................................................................................Simulium roquemayu Ramas terciarias emergen debajo de 1/3 de la distancia a la base (Figs. 89 E, I) .............................................................................................................................................. 26 Tricomas del frontoclípeo y tórax con 2-4 ramas relativamente largos (Fig. 88 X) ............................................................................................................. Simulium shewellianum Tricomas del frontoclípeo y tórax con variable número de ramas y no tan alargadas (Fig. 88 W) ....................................................................................................... 27 Frontoclípeo y tórax con escasos tubérculos (Figs. 88 P, W) ................................. 28 Frontoclípeo y tórax con abundantes tubérculos (Fig. 88 U) ................................. 29 Capullo prolongado dorsal y basalmente; largo branquia 2,7-3,5 mm; tubérculos del frontoclípeo ausentes o escasos (Fig. 88 P) ..........................Simulium rubiginosum Capullo sin prolongaciones anteriores; largo branquia 2,0-2,2 mm; tubérculos más abundantes que lo mostrado en el dilema opuesto (Fig. 88 W) ...... Simulium dinellii Capullo con fibras abiertas .............................................................. Simulium nuneztovari Capullo compacto sin fibras abiertas (Fig. 88 K) ...................................................... 30 Tricomas torácicos con 4-9 ramas; ramificación de las ramas primarias y secundarias no próximas a la base (Fig. 89 I) .............................. Simulium wolffhuegeli Tricomas torácicos con 2-6 ramas; ramificación de las ramas primarias y secundarias próximas a la base ...................... Simulium ochraceum, Simulium antillarum

Larvas Desconocidas: Simulium antonii, S. cormonsi, S. gabaldoni, S. jaimeramirezi, S. nuneztovari, S. roquemayu, S. tolimaense. 1 – 2(1) – 3(2) – 4(3) –

Artejo mediano de la antena similar o menor que el proximal (Fig. 90 O); anillo anal con 84-100 hileras de ganchos; apotoma cefálico oscurecido en el borde basal y una banda mediana longitudinal (Fig. 90 H) ............................................................. 2 Artejo mediano de la antena generalmente más largo que el proximal (Fig. 90 N); anillo anal generalmente con un número de hileras por debajo de 84; apotoma cefálico con diferente ornamentación ........................................................................... 5 Relación del largo puente postgenal/hipostomio = 0,1-0,2 (Fig. 90 K) .................. .............................................................................. Simulium maroniense, Simulium rorotaense Relación del largo puente postgenal/hipostomio = 0,3-0,4 ..................................... 3 Anillo anal con alrededor de 84 hileras de ganchos; relación del largo puente postgenal/hipostomio = 0,4 .......................................................... Simulium kabanayense Anillo anal con 96-110 hileras de ganchos; relación del largo puente postgenal/ hipostomio = 0,3 ................................................................................................................ 4 Relación de los artejos antenales proximal, mediano y distal = 1:0,9-1,0:1,1-1,3; abanico cefálico con 45-50 rayos .................................................... Simulium trombetense Relación de los artejos antenales proximal, mediano y distal = 1:0,8:1,2; abanico cefálico con 40 rayos ............................................................................... Simulium suarezi

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Anillo anal sobrepasando 80 hileras de ganchos; longitud del cuerpo sobre 6,0 mm ........................................................................................................................................ 6 Anillo anal por debajo de 80 hileras de ganchos; longitud del cuerpo generalmente por debajo de 6,0 mm ..................................................................................................... 11 Relación del largo puente postgenal/hipostomio =0,3-0,4 (Fig. 90 J); anillo anal con 83-85 hileras de ganchos .......................................................................................... 7 Relación del largo puente postgenal/hipostomio = 0,2; anillo anal con número variable de hileras de ganchos ......................................................................................... 8 Longitud del cuerpo 5,7 mm; abanico cefálico con 35 rayos ............................. .................................................................................................................Simulium furcillatum Longitud del cuerpo 6,0-6,7 mm; abanico cefálico con 45-50 rayos ........................ ......................................................................... Simulium rubiginosum, Simulium bicoloratum Anillo anal con 86-88 hileras, con 17-20 ganchos cada una ......... Simulium ignescens Anillo anal con 77-85 hileras, con 12-16 ganchos cada una ..................................... 9 Abanico cefálico con alrededor de 30-32 rayos; papilas rectales con 8-11 divertículos en cada lóbulo; esclerito anal sin pelos .......................... Simulium pifanoi Abanico cefálico con alrededor de 37-40 rayos; papilas rectales con 10-16 divertículos en cada lóbulo (Fig. 90 T); esclerito anal con pelos ........................... 10 Artejo mediano de la antena más largo que el distal (Fig. 90 N); relación de los artejos antenales: proximal, mediano y distal = 1:1,7:1,4-1,6; longitud cuerpo 6,5 mm .......................................................................................................... Simulium perflavum Artejo mediano de la antena más corto que el distal (Fig. 90 M); relación de los artejos antenales: proximal, mediano y distal = 1:1,7:2,5; longitud del cuerpo 5,8 mm ........................................................................................................ Simulium bicornutum Relación del largo puente postgenal/hipostomio 0,2-0,3; apotoma cefálico oscurecido subbasalmente con área submediana más clara (Fig. 90 F) ................... ................................................................................................................ Simulium wolffhuegeli Relación del largo puente postgenal/hipostomio 0,10-0,25; apotoma cefálico oscurecido medio basalmente con una ventana subbasal más clara (Figs. 90 B, D, E) .............................................................................................................................................. 12 Anillo anal con 70-79 hileras de ganchos ................................................................... 13 Anillo anal con por debajo de 69 hileras de ganchos .............................................. 17 Abanico cefálico con 42-48 rayos ....................................................... Simulium pautense Abanico cefálico con 27-39 rayos ................................................................................. 14 Abanico cefálico con 27-35 rayos ................................................................................. 15 Abanico cefálico con 36-39 rayos ................................................................................. 16 Longitud cuerpo 4,8-5,2 mm; hendidura postgenal subtriangular (Fig. 90 I); apotoma cefálico sin áreas oscurecidas y escasos pelos (Fig. 90 C) .......................... ................................................................................................................Simulium arcabucense Longitud cuerpo 6,0 mm; hendidura postgenal en forma de domo (Fig. 90 J); apotoma cefálico levemente oscurecido sobre el borde basal y con abundantes pelos (Fig. 90 D) .................................................................................... Simulium albanense Longitud cuerpo alrededor de 6,0 mm ................................................ Simulium dinellii Longitud cuerpo alrededor de 4,6 mm ....... Simulium ochraceum, Simulium antillarum

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17(12) Abanico cefálico con alrededor de 47 rayos; anillo anal con 52 hileras de ganchos .............................................................................................................................................. 18 – Abanico cefálico con alrededor de 35 rayos; anillo anal con 55-74 hileras de ganchos .................................................................................................. Simulium lutzianum 18(17) Esclerito anal con pelos y escamas (Fig. 90 R) ..................................... Simulium tunja – Esclerito anal con solamente pelos, sin escamas ....................................................... 19 19(18) Largo del cuerpo 6,0 mm ..................................................................... Simulium romanai – Largo del cuerpo 4,3-4,7 mm ........................................................................................ 20 20(19) Tegumento del cuerpo con tricomas foliáceos (Fig. 90 P); relación largo puente postgenal/hipostomio 0,2 ..................................................................Simulium anaimense – Tegumento del cuerpo sin tricomas foliáceos; relación largo puente postgenal/ hipostomio 0,1 ............................................ Simulium mayuchuspi, Simulium shewellianum BICOLORATUM species group Simulium (Ectemnaspis) rubiginosum (ENDERLEIN) (Figs. 84 I, X; 86 K; 87 C; 88 P; 89 E; 90 A.) Chirostilbia rubiginosa ENDERLEIN, 1934b: 287. Simulium rubiginosum: WYGODZINSKY, 1971: 19. Simulium (Ectemnaspis) rubiginosum: COSCARÓN, 1984: 294-295; 1987: 28; 1990: 121; 1991: 248-249; CROSSKEY & HOWARD, 1997: 30; 2004: 21. Simulium ignescens: WYGODZINSKY, 1951b: 223 (part.); 1953: 325; COSCARÓN & WYGODZINSKY, 1960: 1124.

Female: Wing length 2.4-2.9 mm. Scutum from orange to mostly black, but always bordered anterolaterally and posteromedially by yellowish brown or grayish brown band as in S. romanai; scutellum yellowish, metanotum from yellowish brown to velvety black, pleurae yellowish orange with episternum and sternites black pruinose. Abdomen with tergites I-IV yellow to yellowish-brown, V-IX dark brown. Legs yellowish with dark portion from light brown to totally black. Sensory vesicle of maxillary palp about half palpomere length (Fig. 84 X). Basal portion of cibarium with sclerotized border and without teeth (Fig. 84 I). Wings with hair on basal sector of R. Paraproct 2/3 longer than wide. Male: Wing length 2.4-2.7 mm. Thorax and abdomen with similar coloration as in female but darker. Abdomen with posterior border of tergite I black, V-VIII velvety black. Hind basitarsus with sides approximately parallel, ratio of length/width = 4.0. Gonostylus slightly shorter than gonocoxite, subcylindrical, with parallel sides on basal 2/3, becoming thin and curved on distal one third, with apical spur (Fig. 86 K). Ventral plate about as long as wide, without or with slight lateral invagination in frontal view (Fig. 87 C). Pupa: Cocoon slipper shaped, moderately compact weave with threads evident, slightly projected dorsally and basally; length along base 3.6-4.3 mm, on dorsum 3.1-3.5 mm, gills 2.7-3.5 mm. Frontoclypeus and thorax with few tubercles arranged homogeneously and enlarged trichomes with 2-6 branches (Fig. 88 P). Gill with 8 filaments, grayish dark, not bifurcated at base, open at several levels, with tendency to meet at apex (Fig. 89 E). Larva: Maximum length 6.0 mm. Cephalic apotome pale to brownish, scarcely darkened on basal half (Fig. 90 A). Antennae shorter than cephalic fan stem; ratio of

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A

B

C

D

E

I J K M

F

G H

O

N

L

Q R P U

b

W a

S

Y

T

V

X

Z

c

Figure 84. Female. A-H: Scutum and scutellum; A: S. (Ectemnaspis) pautense; B: S. (E.) bicoloratum; C: S. (E.) mayuchuspi; D: S. (E.) antillarum; E: S. (E.) ochraceum, abdomen; F: S. (E.) bicoloratum; G: S. (E.) wolffhuegeli; H: S. (E.) shewellianum. I-W Cibarium; I: S. (E.) rubiginosum; J: S. (E.) arcabucense; K: S. (E.) albanense; L: S. (E.) gabaldoni; M-N: S. (E.) lutzianum, showing different disposition of tubercles and spines ; O: S. (E.) romanai; P: S. (E.) wolffhuegeli; Q: S. (E.) kabanayense; R: S. (E.) perflavum; S: S. (E.) suarezi; T: S. (E.) dinellii; U: S. (E.) shewellianum; V: S. (E.) ochraceum; W: S. (E.) antillarum. X-Y, a-c: Sensory vesicle of palpus; X: S. (E.) rubiginosum; Y: S. (E.) ignescens; Z: S. (E.) dinellii; a: S. (E.) shewellianum; b: S. (E.) maroniense; c: S. (E.) rorotaense.

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proximal, medial and distal segments = 1:1.3-1.4:1.5-1.6. Cephalic fan with about 45 rays. Mandible without latero-mandibular process; with 1-3 marginal teeth, the first about 2 times larger than following teeth. Anal ring with 85 rows with about 15 hooks each. Rectal papillae generally with 4-9 diverticula each, but some specimens with only 0-2 diverticula. Distribution: Argentina: Buenos Aires, Jujuy, Mendoza, Salta, San Juan, Tucumán; Bolivia: Cochabamba; Peru: Cuzco. Bionomics: This species breeds in creeks of crystalline water, with fast flow (not torrential) in mountain areas. Larvae and pupae occur on rocks and leaves. Female feeding habits are unknown. Discussion: The closest species is S. ignescens. Differences are explained under that species. Simulium (Ectemnaspis) ignescens ROUBAUD (Fig. 84 Y.) Simulium (Ectemnaspis) ignescens ROUBAUD, 1906b: 107; WYGODZINSKY, 1951b: 223-229; 1971: 14-19; LEON & WYGODZINSKY, 1953: 31; RAMÍREZ PÉREZ, 1971 : 315; 1983: 87. Eusimulium ignescens: SOURCOUF & GONZALEZ RINCONES, 1911: 281; PINTO, 1931: 732. Simulium (Ectemnaspis) ignescens COSCARÓN, 1984: 295-296; 1990: 121; CROSSKEY & HOWARD, 1997: 29; 2004: 21; COSCARÓN ARIAS, 2003: 195, 200, 205, 208, 215. S. ignescens is similar to S. rubiginosum, and we include here only the differences.

Female: Sensory vesicle of maxillary palp shorter than half of palpomere length, and with more abundant tubercles than in S. rubiginosum (Fig. 84 Y). Male: Hind basitarsus wider at middle. Pupa: Gill with 10 filamentous branches. Frontoclypeus with sparse tubercles arranged on both sides near facial trichomes. Larva: Distal antennal segment not longer than medial segment. Anal ring with 8688 rows with 17-20 hooks each. Rectal papillae with 15-20 diverticula on each. Distribution: Bolivia: Chapare; Colombia: Cauca, Cundinamarca, Valle; Ecuador: Chimborazo, Cotopaxi, Imbabura, Napo, Pastaza, Pichincha, Tungurahua; Peru: Cuzco; Venezuela: Merida. Bionomics: Simulium ignescens breeds in fast-flowing, small creeks of mountain areas (1100-3050 m), with clear water, on submerged tree branches, aquatic vegetation and rocks. Females bite humans in Ecuador and mules in Bolivia (LEÓN & WYGODZINSKY, 1953). Discussion: The species is similar to S. rubiginosum, which can be differentiated by the 8 pupal gill branches. Females can be differentiated by the length of the palpal sensory vesicle, which is longer in S. rubiginosum. Simulium (Ectemnaspis) roquemayu COSCARÓN (Figs. 85 F; 89 D.) Simulium (Ectemnaspis) roquemayu COSCARÓN, 1984: 296-298; 1990: 122; 1991: 251; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female: Wing length 2.5 mm. Scutum and scutellum yellowish, metanotum black, legs blackish; abdomen with tergites I-III yellowish, IV blackish-brown medially, V-IX greenish-gray. Fronto-ocular triangle about as wide as high. Sensory vesicle of maxillary

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palp about half palpomere length. Basal portion of cibarium scarcely sclerotized, more intensely sclerotized laterally. Wings with hair on basal sector of R; Sc with about 22 setae. Claws with subbasal tooth. eighth sternite well sclerotized medially, laterally with about 25 setae on each side. Paraproct subtriangular in shape, about as long as wide with rounded distal border, abundant microtrichiae and without basal lobe (Fig. 85 F). Male: Unknown. Pupa: Cocoon slipper shaped with thick spongy weave, slightly elevated dorsally as longitudinal carina. Length along base 2.6 mm, on dorsum 3.0 mm, gills 3.3 mm. Gills, head, thorax and abdomen light brown. Frontoclypeus and thorax with numerous smooth tubercles; large trichomes with 6-7 branches and curved base. Gills with 8 filamentous branches, with tertiary branches bifurcated at more than half total gill length (Fig. 89 D). Larva: Unknown. Distribution: Bolivia: Cochabamba. Bionomics: Unknown. Discussion: Simulium antonii has coloration similar to that of S. roquemayu but can be differentiated by the absence of hair on the basal sector of R. Also, S. bicornutum and S. furcillataum have similar females, but the pupal gills with 2 thick branches readily distinguish them. Simulium pifanoi shows similarities in scutum ornamentation, but abdominal tergite III has a dark brown tergal plate, tergite V is velvety black and VI-IX are blackish. The pupal gill branches are tightly arranged, forming a bundle. Simulium (Ectemnaspis) pifanoi RAMÍREZ PÉREZ (Figs. 85 G; 86 J; 88 O; 89 C; 90 T.) Simulium pifanoi RAMÍREZ PÉREZ, 1971: 346; 1983: 62. Simulium (Ectemnaspis) pifanoi: COSCARÓN, 1984: 301; 1987: 28; 1990: 122-123; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female: Wing length 2.4-2.6 mm. Scutum orange, with yellowish pronotal area and scutellum; metanotum black. Abdomen with tergites I-II yellow; III-IV yellowish with dark brown tergal plate; V velvety black; VI-IX shiny blackish. Legs black. Frons convergent. Fronto-ocular triangle about as high as wide. Basal sector of R without hair. Gonapophysis with internal border well sclerotized. Eighth sternite with about 30 setae on each side; paraproct slightly longer than wide at base (Fig. 85 G). Male: Wing length 2.5-2.6 mm. Coloration as in female but abdominal segments IIIIV dark-brown and scutum with dark stripes laterally. Gonostylus 3/4 of gonocoxite length (Fig. 86 J); ventral plate without lateral constriction. Pupa: Cocoon with basal anterior projection; weave with thin threads; length along base 3.0-3.3 mm, on dorsum 2.6-2.8 mm, gills 3.2-3.5 mm. Frontoclypeus and thorax with sparse tubercles and large trichomes with 3-7 branches (Fig. 88 O). Gill with 8 branches with arrangement similar to that of S. roquemayu but arranged more open and with lower bifurcation on tertiary branches (Fig. 89 C). Larva: Maximum length 5.0-5.5 mm. Cephalic apotome slightly darkened on basal half. Ratio of antennal segments: proximal, medial and distal = 1:1.3-1.4:1.6-1.7. Mandible with 1 latero-mandibular process. Postgenal bridge about 1/8 of hypostomium height. Cephalic fan with 30-32 rays. Lateral sclerite of proleg with 26-30 teeth arranged in 7-8 groups. Anal

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P Figure 85. Female. A: Basal section of wing of S. (Ectemnaspis) albanense (C = costa, Sc = subcosta, R = radius, bR = basal sector of R). B-C: Claw; B: S. (E.) mayuchuspi; C: lutzianum. D-Q: Paraproct; D: S. (E.) bicoloratum; E: S. (E.) furcillatum; F: S. (E.) roquemayu; G: S. (E.) pifanoi; H: S. (E.) pautense; I: S. (E.) tunja; J: S. (E.) wolffhuegeli; K: S. (E.) perflavum; L: S. (E.) kabanayense; M: S. (E.) rorotaense; N: S. (E.) suarezi; O: S. (E.) dinellii; P: S. (E.) shewellianum; Q: S. (E.) ochraceum.

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sclerite without hair. Anal ring with 77-84 rows with 12-14 hooks each. Rectal papillae with 8-11 diverticula on each lobe (Fig. 90 T). Body integument without hair. Distribution: Colombia: Cauca, Cundinamarca; Venezuela: Mérida. Bionomics: Unknown. Discussion: Simulium roquemayu is very similar and the differences were described under that species. Simulium arcabucense is without hair on the basal sector of R and has a similarly shaped paraproct but can be differentiated by the 6 branches of the pupal gill and in some specimens by the dark brown area on the thoracic disc. Simulium (Ectemnaspis) arcabucense COSCARÓN (Figs. 86 H; 87 A; 88 A; 89 A; 90 C.) Simulium (Ectemnaspis) arcabucense COSCARÓN, 1990: 123-125; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female: Wing length 2.7 mm. Scutum and scutellum yellowish; sometimes the first with sublateral stripes and median anterior area dark brown. Metanotum black, pleurae with silver pruinosity. Legs black with lighter portions on tibiae. Abdomen with tergites III light yellow, darkened laterally; III-IV yellow with tergal plate brownish on posterior border; V-IX blackish. Basal sector of R without hair. Eighth sternite with about 25 setae; paraproct about as long as wide. Male: Wing length 2.5 mm. Coloration as in female, sometimes with 1+1 lateral and median frontal black stripes. Ratio of length/width of hind basitarsus 7.5. Gonostylus about 3/4 of gonocoxite length (Fig. 86 H). Ventral plate with slight lateral concavity (Fig. 87 A). Pupa: Cocoon with thicker anterior rim, spongy weave and thin threads (Fig. 88 A); length along base 2.9-3.0 mm, on dorsum 2.5-2.7 mm, gills 3.0-3.6 mm. Frontoclypeus and thorax with numerous large tubercles arranged in irregular rows and wrinkled apically. Cephalic and thoracic trichomes large, with thick base and 6-13 branches. Gill with 6 branches, emerging near base (Fig. 89 A). Larva: Maximum length 4.8-5.2 mm. Cephalic apotome light, without special ornamentation (Fig. 90 C). Ratio of antennal segments: proximal, medial and distal = 1:1.0-1.1:1.8-2.0. Postgenal bridge about 1/3-1/4 of hypostomium height. Mandible with 1 latero-mandibular process. Cephalic fan with about 27 rays. Lateral sclerite of thoracic proleg with about 25 teeth in 8-9 groups. Anal sclerite without hair. Anal ring with about 70 rows with 12-13 hooks each. Body integument without hairs. Distribution: Colombia: Boyacá. Bionomics: Unknown. Discussion: The most similar species is S. pautense, which can be differentiated by the shorter gonostylus, thinner pupal cocoon, frontoclypeal and thoracic tubercles uniformly arranged and apically smooth, gill branches bifurcating at a greater distance from the base, darker cephalic apotome with negative spots and greater number of rays. Simulium gabaldoni also has similar adult coloration, but the cibarium has the median area well sclerotized with tubercles and lateral spines, the pupal tubercles are less elevated, and the larval cephalic apotome has distinct spots. Simulium pifanoi, with similar adult ornamentation, can be differentiated by the pupal gill with 8 branches. Simulium species A and B LEWIS & LEE-PORTER (1964: 97-98) can be differentiated by the medially elevated cibarium (sp. A) and the abdomen not darkened laterally and the longer paraproct (sp. B).

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Figure 86. Male. A-E: Scutum and scutellum; A: S. (Ectemnaspis) furcillatum; B: S. (E.) pautense; C: S. (E.) mayuchuspi; D: S. (E.) romanai; E: S. (E.) ochraceum. F-G: Abdomen; F: S. (E.) bicornutum; G: S. (E.) furcillatum. H-S: Gonocoxite and gonostylus; H: S. (E.) arcabucense; I: S. (E.) pautense; J: S. (E.) pifanoi; K: S. (E.) rubiginosum; L: S. (E.) tunja; M: S. (E.) albanense; N: S. (E.) romanai; O: S. (E.) wolffhuegeli; P: S. (E.) kabanayense; Q: S. (E.) perflavum; R: S. (E.) rorotaense; S: S. (E.) shewellianum.

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Simulium (Ectemnaspis) pautense COSCARÓN & TAKAOKA (Figs. 84 A; 85 H; 86 B, I; 87 B; 88 E; 89 B; 90 B.) Simulium (Ectemnaspis) pautense COSCARÓN & TAKAOKA, 1989: 191; COSCARÓN, 1990: 125, CROSSKEY & HOWARD, 1997: 29; 2004: 21; COSCARÓN ARIAS, 2003: 195, 200, 206, 208, 215, 219.

Female: Wing length 2.4-2.6 mm. Scutum orange, greenish orange laterally; scutellum greenish yellow (Fig. 84 A); metanotum velvety black with 1+1 small grayish patches. Pleura mostly pale yellow. Legs dark brown. Abdominal tergites I-IV yellow, with tergal plates III and IV pale brown, tergal plates V-IX blackish brown; waxy, bright except plate V velvety, bright. Basal portion of cibarium with reinforced border. Basal sector of R bare. Eighth sternite with darker middle area and about 27 setae on each side. Paraproct about as long as wide at base (Fig. 85 H). Male: Wing length 2.3-2.5 mm. Scutum pale but yellow orange on top and with Tshaped black stripe on anterior third and pair of black, broad longitudinal stripes sublaterally (Fig. 86 B). Ratio of length/width of hind basitarsus = 5. Gonostylus about 4/5 of gonocoxite length (Fig. 86 I), ratio of length/wide = 2.0. Ventral plate without lateral concavity (Fig. 87 B). Pupa: Cocoon with small basal anterior projection (Fig. 88 E); length along base 3.03.3 mm, on dorsum 2.4-2.8 mm, gills 2.7-3.3 mm. Frontoclypeus and thorax with rounded and smooth tubercles; trichomes curved basally and with 3-6 branches. Gill with 6 branches (Fig. 89 B). Larva: Maximum length 5.5-5.8 mm. Posterior half of cephalic apotome darkened, with negative spots (Fig. 90 B). Postgenal cleft deep and narrow; postgenal bridge about 1/6 of hypostomium height. Antennae shorter than stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1.0-1.2:1.6-2.0. Cephalic fan with 42-48 rays. Mandible with 1 latero-mandibular process. Lateral sclerite of thoracic proleg with 23-25 teeth. Anal sclerite with hairs. Anal ring with 73-76 rows with 14-15 hooks each. Rectal papillae with 12 diverticula on each lobe. Distribution: Ecuador: Azuay, Cuenca, Pichincha. Bionomics: Unknown. Discussion: Simulium arcabucense is similar but has a longer gonostylus (ratio of length/ width = 2.7), the trichomes of the pupal frontoclypeus and thorax with a straight and wide base, the tubercle borders wrinkled and the tertiary gill branches bifurcated nearer the base. Simulium pifanoi can be differentiated by the 8 gill branches and longer gonostylus. Simulium gabaldoni has fewer tubercles on the pupal frontoclypeus and thorax and the basal portion of the cibarium has tubercles and spines, and the legs are entirely black. Simulium (Ectemnaspis) antonii WYGODZINSKY Simulium antonii WYGODZINSKY, 1953b: 325. Simulium (Ectemnaspis) antonii: COSCARÓN, 1984: 302; 1987: 29; 1991: 249; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female: Wing length 3.0 mm. Scutum pale orange, pale yellow border laterally and posteriorly, sometimes with dark brown sublateral stripes. Scutellum orange; metanotum

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Figure 87. Male. Ventral plate. A: S. (Ectemnaspis) arcabucense. B: S. (E.) pautense. C: S. (E.) rubiginosum. D: S. (E.) tunja. E: S. (E.) albanense. F: S. (E.) gabaldoni. G: S. (E.) mayuchuspi. H: S. (E.) romanai. I: S. (E.) wolffhuegeli. J: S. (E.) perflavum. K: S. (E.) kabanayense. L: S. (E.) rorotaense. M: S. (E.) dinellii. N: S. (E.) shewellianum. O: S. (E.) ochraceum. P: S. (E.) antillarum.

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black; pleurae orange with episternum on ventral half blackish with silver pruinosity. Legs dark brown to black. Abdomen with tergites I-IV yellowish, sometimes IV darkened, V-IX dark brown to blackish; I-V matte, the following tergites shiny. Basal sector of R bare. Paraproct 1/3 longer than wide. Male, Pupa and Larva: Unknown. Distribution: Bolivia: Cochabamba. Bionomics: Unknown. Discussion: Without knowledge of the pupa and male, it is difficult to separate S. antonii from other species of the subgenus Without hairs on the basal sector of R and with similar scutum and abdomen coloration, the closest species are S. pautense, S. pifanoi, S. arcabucense and S. furcillatum, which are smaller (wing length 2.4-2.7 mm). In S. pautense, the paraproct is as long as wide, and in S. furcillatum the scutum is brownish and the metanotum grayish brown. Simulium tunja has the paraproct about half as wide as long and no dark brown sublateral stripes on the scutum. Simulium (Ectemnaspis) tunja COSCARÓN (Figs. 85 I; 86 L; 87 D; 88 G, Q; 90 R.) Simulium (Ectemnaspis) tunja COSCARÓN, 1990: 126-128; CROSSKEY & HOWARD, 1997: 30; 2004: 21.

Female: Wing length 2.8-2.9 mm. Scutum yellowish with median and sublateral area from brownish orange to black, scutum rim and scutellum yellow, and metanotum black. Abdomen with tergites I-II yellowish, III (IV) yellow with tergal plates brownish to dark brown IV-V generally velvety black, VI blackish, waxy bright. Basal sector of R bare. Eighth sternite with about 40 hairs on each side; paraproct about 2/3 as wide as long, with thick apex (Fig. 85 I). Male: Wing length 2.7 mm. Scutum blackish; abdomen as in female but more darkened, with segment III tergal plate largest and darker, VI-VII with 1+1 silvery spots. Ratio of length/width of hind basitarsus = 4.3. Gonostylus slightly shorter than gonocoxite (Fig. 86 L). Ventral plate with slight lateral depression (Fig. 87 D). Pupa: Cocoon with anterior basal projection (Fig. 88 G), weave spongy; length along base 3.2-3.7 mm, on dorsum 2.9-3.1 mm, gills 2.1-2.4 mm. Frontoclypeus and thorax with numerous smooth tubercles (Fig. 88 Q); trichomes single or bifid. Gills with 10 branches, primary dorsal and median branches bifurcated near base. Larva: Maximum length 5.0-5.2 mm. Cephalic apotome scarcely pigmented, without positive spots. Postgenal bridge about 1/6 of hypostomium length. Antennae about same length as cephalic stem apex; ratio of proximal, medial and distal segments = 1:1.2-1.5:1.21.5. Cephalic fan with 29-32 rays. Anal sclerite with hairs and scales (Fig: 90 R). Anal ring with 67-70 rows with 12-13 hooks each. Rectal papillae with 16-18 diverticula on each lobe. Distribution: Colombia: Boyacá, Cundinamarca. Bionomics: Unknown. Discussion: Simulium kabanayense has similar coloration but can be differentiated by the presence of hair on the basal sector of R, small teeth on the cibarium, smaller tubercles on the frontoclypeus and thoracic trichomes with 3-6 branches. Simulium ignescens, with 10 branches, has a thinner cocoon, the frontoclypeus and thorax without tubercles and hair on the basal sector of R. Simulium bicoloratum, also with 10 branches, has

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Figure 88. Pupa. A-L: General aspect; A: S. (Ectemnaspis) arcabucense; B: S. (E.) bicoloratum; C: S. (E.) bicornutum; D: S. (E.) furcillatum; E: S. (E.) pautense; F: S. (E.) tolimaense; G: S. (E.) tunja; H: S. (E.) anaimense; I: S. (E.) lutzianum; J: S. (E.) romanai; K: S. (E.) wolffhuegeli; L: S. (E.) suarezi. M-X: Frontoclypeus; M: S. (E.) bicoloratum; N: S. (E.) pautense; O: S. (E.) pifanoi; P: S. (E.) rubiginosum; Q: S. (E.) tunja; R: S. (E.) lutzianum; S: S. (E.) mayuchuspi; T: S. (E.) romanai; U: S. (E.) wolffhuegeli; V: S. (E.) maroniense; W: S. (E.) dinellii; X: S. (E.) shewellianum. Y: Trichome of S. (E.) anaimense. Z: Tubercles of S. (E.) albanense.

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Figure 89. Pupa. Gill. A: S. (Ectemnaspis) arcabucense. B: S. (E.) pautense. C: S. (E.) pifanoi. D: S. (E.) roquemayu. E: S. (E.) rubiginosum. F: S. (E.) anaimense. G: S. (E.) lutzianum. H: S. (E.) romanai. I: S. (E.) wolffhuegeli. J: S. (E.) perflavum. K: S. (E.) kabanayense. L: S. (E.) maroniense. M: S. (E.) rorotaense. N: S. (E.) suarezi. O: S. (E.) trombetense (based on HAMADA & ADLER, 1988). P: S. (E.) shewellianum.

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the frontoclypeal and thoracic trichomes with more than 20 branches and hair on the basal sector of R. Simulium antonii has similar coloration but without the darker median and sublateral stripes sometimes present in S. tunja, but the unknown male, pupa and larva of S. antonii are inconvenient in evaluating the possible synonymy of these species. Simulium (Ectemnaspis) tolimaense COSCARÓN (Fig. 88 F.) Simulium (Ectemnaspis) tolimaense COSCARÓN, 1984: 301 n. nom. to Simulium (Ectemnaspis) simplex WYGODZINSKY & COSCARÓN, 1982: 11 (preoc. by Simulium simplex GIBBINS, 1936); COSCARÓN, 1990: 128; CROSSKEY & HOWARD, 1997: 30; 2004: 21.

Female (Pharate): Scutum very similar to that of males of S. bicornutum and S. furcillatum; fronto-ocular triangle about as long as wide. Cibarium as in S. furcillatum. Claw with subbasal tooth well developed Male (Pharate): Pigment pattern not fully developed, mesonotum yellowish brown. Genitalia not discernible. Pupa: Cocoon with anterior subrectangular flap-like projection dorsally and also projected basally (Fig. 88 F); walls of cocoon thick, individual threads not perceptible; length along base 3.4-3.5 mm, on dorsum 3.2-3.5 mm, gills 1.8-2.0 mm. Frontoclypeus and thorax with relatively abundant tubercles; trichomes with 5-8 branches, with curved base. Gills with 5 branches, surface minutely wrinkled. First branch perpendicular to stem of gill. Secondary branch bifurcated; lower branch subdivided subapically, with resulting branch about as long as common stem. Upper branch also bifurcated but the two branches longer than common stem. Larva: Unknown. Distribution: Colombia: Tolima. Bionomics: The species has been collected in the Central Cordillera at 3100 m, on the slope of the hillside. No more information is available. Discussion: The scutum ornamentation is typical of the species of the Simulium (Ectemnaspis) subgenus. The unique gill number and branching permit identification. Simulium (Ectemnaspis) bicornutum WYGODZINSKY & COSCARÓN (Figs. 86 F; 88 C; 90 M.) Simulium (Ectemnaspis) bicornutum WYGODZINSKY & COSCARÓN, 1982: 2; COSCARÓN, 1984: 300; 1990: 128; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female (Pharate): Scutum and scutellum yellow; color pattern of scutum similar to that of S. bicoloratum. Abdominal tergites I-IV grayish yellow, V-IX dark grayish brown. Fronto-ocular triangle approximately as deep as wide. Eighth sternite with about 30 setae on each side; gonapophysis subtriangular, glabrous, with few bands of heavier sclerotization; paraproct about as long as wide at base, rounded apically. Male: Wing length 3.0 mm. Scutum orange, sublateral pattern dark brown to black. Metanotum black. Abdomen with segment I brown, II-III pale greenish yellow, IV brown dorsally, V-IX velvety black; VI with 1+1 large silvery spots laterally (Fig: 86 F). Basal sector of R glabrous. Ratio of length/width of hind basitarsus = 3.5. Gonostylus as long as gonocoxite, elongate, subconical, with apical portion somewhat curved. Basal plate subtriangular, its sides almost straight.

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Pupa: Cocoon with a broad median projection dorsally. Walls of cocoon thick; length of cocoon along base 3.7 mm, on dorsum 4.0 mm, gills 1.8 mm. Frontoclypeus and thorax with few or without tubercles; trichomes of head with 4-8 branches, of thorax with 2-6 branches. Gill with 2 thick branches, roughly half-moon shaped in lateral view; arms pointed apically, each with small tooth-like process on inner surface (Fig. 88 C). Larva: Maximum length 5.8 mm. Cephalic apotome rather uniformly pigmented, somewhat darker at base. Ratio of antennal segments: proximal, medial and distal = 1:1.7:2.5 (Fig: 90 M). Cephalic fan with about 40 rays. Postgenal cleft narrow and deep; postgenal bridge about 1/5 of hypostomium length. Ninth segment without ventral tubercles. Anal sclerite with abundant and minute setae. Anal ring with about 85 rows with 16 hooks each. Rectal papillae with about 16 diverticula on each lobe. Distribution: Colombia: Cundinamarca. Bionomics: The species has been collected in the Paramo region at and altitude of 3300-3700 m. The immatures live on trailing blades of grass in torrential, small creeks 13 m wide, with clear water. Discussion: The unique pupal gill allows this species to be differentiated from other Neotropical black flies. Simulium (Ectemnaspis) furcillatum WYGODZINSKY & COSCARÓN (Figs. 85 E; 86 A, G; 88 D.) Simulium (Ectemnaspis) furcillatum WYGODZINSKY & COSCARÓN, 1982: 6; COSCARÓN, 1984: 300; 1990: 128; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female: Wing length 2.4 mm. Scutum orange brown, lateral borders and lateral incision posterior to pronotal incisions, greenish yellow. Scutellum greenish yellow. Metanotum grayish brown with 1+1 anterior grayish spots with certain illumination. Abdominal segments I-IV with terga grayish yellow, fourth tergal plate light brown. Cibarium smooth with slight median depression and large, heavily pigmented cornuae. Wing without hairs on basal sector of R. Claws with small but distinct subbasal tooth. Eighth sternite with inner border conspicuously sclerotized. Paraproct subtriangular, about as long as wide at base (Fig. 85 E). Male: Wing length 2.9 mm. Scutum and scutellum pale greenish yellow; scutum with wide median dark marking covering anterior 3/4 and 1 pair of lateral wedgeshaped spots beginning at anterior sixth of dark central marking (Fig. 86 A). Metanotum dark grayish brown. Abdominal segment I brown, II-III with tergal plates pale grayish yellow; pleuron and sternum grayish white; V-IX dark brown, VIII with 1+1 lateral silvery spots (Fig. 86 G). Ratio of length/width of hind basitarsus about 4.0. Gonocoxite about 2.5 times as long as wide with well-developed apical spine. Ventral plate subtriangular. Pupa: Cocoon translucent, with flap-like projection dorsally at middle. Fabric of cocoon uniform, parchment like, individual threads difficult to discern. Cocoon length ventrally 3.2 mm, on dorsum 3.7 mm, gills 2.1 mm. Frontoclypeus and thorax without tubercles, except group between facial and frontal trichomes; trichomes with 3-6 branches. Gill consisting of 2 rigid arms forming U shape, lower arm of gill pointed, apical half of upper arm fork-like, with 3 tine-like processes (Fig. 88 D).

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Figure 90. Larva. A-H: Cephalic apotome; A: S. (Ectemnaspis) rubiginosum; B: S. (E.) pautense; C: S. (E.) arcabucense; D: S. (E.) albanense; E: S. (E.) romanai; F: S. (E.) wolffhuegeli; G: S. (E.) perflavum; H: S. (E.) rorotaense. I-L: Hypostomium and postgenal cleft; I: S. (E.) arcabucense; J: S. (E.) albanense; K: S. (E.) rorotaense; L: S. (E.) shewellianum. M-O: Antennae; M: S. (E.) bicornutum; N: S. (E.) perflavum; O: S. (E.) rorotaense. P: Trichomes of body integument of S. (E.) anaimense. Q: Apical portion of mandible of S. (E.) mayuchuspi. R: Anal sclerite of S. (E.) tunja. S-T: Diverticula of rectal papillae lobes; S: S. (E.) perflavum; T: S. (E.) pifanoi.

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Larva: Maximum length 5.7 mm. Cephalic apotome rather homogeneously pigmented, somewhat darker at base. Ratio of antennal segments: proximal, medial and distal = 1:1.3:1.8. Cephalic fan with about 35 rays. Postgenal cleft deep, narrowly subtriangular; postgenal bridge about half of hypostomium length. Lateral sclerite of proleg with about 24 teeth. Body integument glabrous. Anal ring with 84 rows with 14-16 hooks each. Rectal papillae with about 16 diverticula each. Distribution: Colombia: Boyacá, Cundinamarca. Bionomics: This species breeds on blades of grasses trailing in rapid, clean streams in the Paramos of the Eastern Cordillera at 3200-3400 m and at water temperatures of about 10 °C. Discussion: The most similar species is S. bicornutum, which has a pupal gill with the two arms undivided, unlike in S. furcillatum, which has the upper arm divided into 3 tinelike appendages. Adult coloration is similar between these two species. Simulium (Ectemnaspis) bicoloratum MALLOCH (Figs. 84 B, F; 85 D; 88 B, M.) Simulium bicoloratum MALLOCH, 1912: 649; VARGAS & DIAZ NAJERA, 1953b: 140; WYGODZINSKY, 1953b: 327-330; 1958: 133. Simulium (Ectemnaspis) bicoloratum WYGODZINSKY, 1971: 19-31, 34; COSCARÓN, 1984: 298-299; 1987: 28; 1990: 128; 1991: 249-251; CROSSKEY & HOWARD, 1997: 29; 2004: 21; COSCARÓN ARIAS, 2003: 195, 200, 205, 209, 215, 218. Odagmia bicolorata: ENDERLEIN, 1925: 208; 1930: 97. Simulium gaudeatum KNAB, 1914b: 81; 1914c: 123. Ectemnaspis macca ENDERLEIN, 1934b: 281 (preoc.); 1936: 122. Simulium molli VARGAS, 1943a: 142; 1945a: 165. Dasypelmoza macca ENDERLEIN, 1934b: 276. (n. syn.).

Female: Wing length 2.8-3.2 mm. Frons and clypeus silver gray. Scutum yellow with extensive area blackish as in figure 84 B; central area can change from brown to orange. Scutellum yellow; metanotum velvety black with 1+1 silvery spots anteromedially. Anepisternum and katepisternum black, remaining pleural area brownish. Abdomen with segments I-II golden yellow, fringe of first segment blackish; dorsum of III-IV velvety black, dorsum VI entirely yellow to partly or entirely dark brown or black; VII-IX brown to black; terga III-V dull, VI-IX polished (Fig. 84 F). Cibarium with large heavily pigmented cornuae, space between cornuae smooth. Basal sector of R with setae. Claws with medium-sized subbasal tooth. Eighth sternite with about 15-20 setae on each side; gonapophysis subtriangular, glabrous, rather conspicuously sclerotized on median half. Paraproct slightly longer than wide at base, subconical (Fig. 85 D). Male: Wing length 2.8-3.2 mm. Color of scutum much as in female, but lightcolored area of center of posterior portion shorter, not as long as posterior black area. Scutellum yellow, darkened at sides and in many cases narrowly along base. Metanotum as in female. Abdomen velvety black with 1+1 large silvery spots on tergite VI. Hind basitarsus 4 times as long. Ventral plate subtriangular, strongly constricted at sides, producing abruptly narrowed apical portion, slightly incised medially. Pupa: Cocoon very thick, spongy, threads poorly visible; anterior margin conspicuously thickened, dorsal surface with 2 or 3 longitudinal ridges along whole

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length of cocoon (Fig. 88 B); length along base 3.5-4.0 mm, on dorsum 2.7-3.0 mm, gills 2.0 mm. Frontoclypeus and thorax with numerous large tubercles with small protuberances, arranged in irregular fashion. Trichomes large, heavily sclerotized, almost black with 30-40 branches (Fig. 88 M). Gill with 8 branches emerging from 3 short primary branches. Surface of branches and filaments with minute tubercles of 2 sizes, giving wrinkled appearance. Larva: Maximum length 6.5-7.0 mm. Cephalic apotome yellowish with positive spots. Antennae as long as stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.5:1.1-1.5. Mandible with 1 latero-mandibular process. Cephalic fan with 40-50 rays. Hypostomium with intermediate teeth scarcely prominent, median and lateral teeth about same height. Postgenal cleft pointed, subtriangular; postgenal bridge slightly shorter than hypostomium. Body integument without hair. Anal sclerite with small setae. Anal ring with 80-85 rows of about 17 hooks each. Rectal papillae with 15-20 elongate diverticula on each lobe. Distribution: Bolivia; Colombia: Cauca, Cundinamarca, Tolima; Ecuador: Pichincha; Peru: Lima; Venezuela: Mérida. Bionomics: This species has been collected on aquatic vegetation, grasses and under stones in mountainous areas at 2000-3600 m in fast-flowing creeks 3-5 m wide and with water temperatures of about 9 °C. It also has been found in irrigation ditches. Female feeding habits are unknown. Discussion: Simulium bicoloratum varies in color. WYGODZINSKY (1971) considered this condition to justify its position in Simulium (Ectemnaspis). The closest species are S. cormonsi, S. jaimeramirezi, S. romanai and S. lutzianum. Differentiation from similar species is explained in the description of each similar species. Dasypelmoza macca, based on the original description, fits into the range of variation of S. bicoloratum. Simulium (Ectemnaspis) cormonsi WYGODZINSKY Simulium cormonsi WYGODZINSKY, 1971: 31-32; RAMÍREZ PÉREZ, 1983: 43. Simulium (Ectemnaspis) cormonsi; COSCARÓN, 1984: 299; 1987: 28; 1990: 129; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female and Male: Not distinguishable by color or morphology from S. bicoloratum. Female: fronto-ocular triangle wider than high; paraproct longer and narrower. Male: gonostylus slightly narrowed and ventral plate more constricted laterally, with apical half narrower. Pupa: Cocoon as in S. bicoloratum. Trichomes as in S. bicoloratum. Gill length 3.0 mm. Gill with 6 branches, emerging from 3 short primary branches. Larva: Unknown. Distribution: Venezuela: Mérida. Bionomics: The immature stages of this species are found on vegetation and stones at altitudes of 3000-3400 m in torrential streams 3-5 m wide, with water temperatures of about 13-15 °C. Discussion: Differences from S. bicoloratum are in the number of gill branches and the slight morphological differences in shape, as explained in the description.

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Simulium (Ectemnaspis) jaimeramirezi WYGODZINSKY Simulium jaimeramirezi WYGODZINSKY 1971: 32-34; RAMÍREZ PÉREZ, 1983: 48. Simulium (Simulium) jaimeramirezi: RAMÍREZ PÉREZ & VULCANO, 1973: 395. Simulium (Ectemnaspis) jaimeramirezi COSCARÓN, 1984: 299; 1987: 28; 1990: 129; CROSSKEY & HOWARD, 1997: 29; 2004: 21.

Female and Male: Color and structure, including genitalia, similar to those of S. bicoloratum. Pupa: Cocoon with structure as in S. bicoloratum, dorsally with only median longitudinal ridge; length at along 4.4 mm, on dorsum 2.9-3.2 mm; gills 1.9-2.8 mm. Frontoclypeus and thorax with tubercles less numerous than in S. bicoloratum. Larva: Unknown. Distribution: Venezuela: Mérida. Bionomics: This species was collected in torrential streams 3-5 m wide, at 30503400 m, on the undersurfaces of stones at 13-15 °C. Discussion: Simulium jaimeramirezi is very similar to S. cormonsi and S. bicoloratum, but can be differentiated by the weak coloration and small frontoclypeal and thoracic trichomes. It also can be separated from S. bicoloratum by the number of gill branches. It would be useful to have more material and conduct cytological or molecular studies comparing this species with S. cormonsi. Simulium (Ectemnaspis) anaimense COSCARÓN & MUÑOZ DE HOYOS (Figs. 88 H, Y; 89 F; 90 P.) Simulium (Ectemnaspis) anaimense COSCARÓN & MUÑOZ DE HOYOS, 1995: 587-592. Simulium (Psilopelmia) anaimense: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female (Pharate): Scutum yellow. Cibarium with tubercles in space between cornuae. Sensory vesicle of maxillary palp about half length of palpomere. Claw subbasally with medium-sized, pointed projection. Male (Pharate): Scutum and scutellum yellowish; metanotum blackish. Pupa: Cocoon whitish, thick, reinforced at the anterior rim and on median dorsal ridge (Fig. 88 H); texture felt like, threads only visible at high magnification. Length along base 3.4 mm, on dorsum 2.6 mm, gill 2.3 mm. Frontoclypeal and thoracic tubercles superficially rugose, moderate in number, but abundant on base of frontoclypeus and insertion of gill; trichomes with curled base and 4-13 branches (Fig. 88 Y). Gill whitish, thick, projected in several planes, dotted homogeneously and with 6 branches (Fig. 89 F). Larva: Maximum length 4.0-5.5 mm. Cephalic apotome brown, dark basally, with pattern similar to that of S. pautense. Antenna surpassing length of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.0-1.1:1.5-1.6. Mandibles with 1 latero-mandibular process. Postgenal cleft deep, dome shaped; postgenal bridge about 1/6 hypostomium length. Cephalic fan with 28-34 rays. Body integument with palmiform and simple trichomes intermixed (Fig. 90 P). Thoracic proleg sclerite with about 16 teeth in 6-7 groups. Ninth segment with 2 ventral papillae. Anal sclerite with abundant unbranched hairs. Anal ring with 64-70 rows with 11-14 hooks each. Rectal papillae with 12-18 diverticula on each lobe.

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Distribution: Colombia: Tolima on the Cordillera Central. Bionomics: This species breeds in medium-sized torrential streams with clear, cold water of 12 °C. Larvae and pupae were collected from trailing roots in swift flow. Discussion: Simulium anaimense is similar in cocoon shape to several species of the S. bicoloratum and S. romanai groups, but the unique, relatively thick and short gill branches distinguish it. Also, the tubercles on the cibarium are similar to those of the S. romanai group. It is also similar to S. albanense in relation to the integumentary setae of the larva. ROMANAI species group Simulium (Ectemnaspis) romanai WYGODZINSKY (Figs. 84 O; 86 D, N; 87 H; 88 T; 89 H; 90 E.) Simulium romanai WYGODZINSKY, 1951b: 229. Simulium (Ectemnaspis) romanai: COSCARÓN, 1984: 302-303; 1987: 29; 1990: 129-131; 1991: 252; COSCARÓN ARIAS, 2003: 195, 200, 205, 209, 215, 219. Simulium (Psilopelmia) romanai: CROSSKEY & HOWARD, 1997: 60; 2004: 54. Simulium adolfolutzi WYGODZINSKY, 1951b: 234.

Female: Wing length 1.9-2.8 mm. Scutum yellowish to brown blackish on central area. Scutellum orange; metanotum black. Abdomen with tergite I yellowish, II-IV brown with center of tergite III dark brown or all tergites black, V-IX shiny dark brownish gray. Legs yellowish brown with distal portion of femur, tibia and tarsomeres dark brown to mostly brown. Cibarium with well-sclerotized rim, depressed at center, medially with rounded protuberances and laterally with abundant small acuminate teeth-like spines (Fig. 84 O). Basal sector of R with hair. Ratio of length/width of posterior basitarsus = 6.6. Eighth sternite with about 25 setae, gonapophyses subtriangular, membranous, with divergent internal border and slightly sclerotized; paraproct 2/3 longer than wide with rounded apex, and genital fork with median stem length about same as 2 lateral branches together. Male: Wing length 2.1-2.4 mm. Coloration as in female but often darker as in figure 86 D. Abdomen velvety black with 1+1 lateral silvery spots on tergites VI-VII. Ratio of length/width of hind basitarsus = 4.8-5.1. Gonostylus subcylindrical with sides subparallel on basal 2/3 and becoming thin toward apex, with 1 apical spur (Fig. 86 N). Ventral plate wider than long, with strong lateral depression, in lateral view appearing as 2 superimposed pieces (Fig. 87 H). Pupa: Cocoon thick, weave spongy, slightly projected basally and frequently with dorsal longitudinal ridge, especially on anterior portion. Frontoclypeus and thorax with abundant tubercles (Fig. 88 T); trichomes with 4-12 branches. Gill with 8 branches relatively open from base (Fig. 89 H). Larva: Maximum length 6 mm. Cephalic apotome with darkened median basal area, head spots negative (Fig. 90 E). Antenna not surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.0-1.4:0.9-1.5. Mandible with 1 lateromandibular process; cephalic fan with about 37 rays. Postgenal cleft very deep, acuminate anteriorly; postgenal bridge very short. Body integument without hairs, anal sclerite without scales or hairs. Lateral thoracic proleg sclerite with about 21 teeth. Anal sclerite with about 68 rows with 12 hooks each. Rectal papillae with 5-9 diverticula on each lobe.

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Distribution: Argentina: Catamarca, Córdoba, Jujuy, La Rioja, Salta, San Juan, Tucumán; Bolivia: Potosí; Colombia: Cauca, Cundinamarca, Valle; Ecuador: Los Rios, Pastaza, Pichincha; Peru: Junin; Venezuela: Mérida. Bionomics: This species breeds in creeks with clear, fast flow over stones and aquatic vegetation. Female feeding habits are unknown. Discussion: The most similar species is S. lutzianum because that species shows variation in coloration, creating confusion. Simulium lutzianum can be differentiated in the female by abdominal tergites II-III, which are frequently yellowish, and the cibarium with several rows of rounded tubercles; in the male by the lack of black on the scutum disc and thinner hind legs; in the pupa by the sparse tubercles on the frontoclypeus and thorax; and in the larva by the lighter cephalic apotome and hair on the body integument. Cytological studies by MORENO RAMÍREZ (Magister thesis, 1997) of S. lutzianum from Colombia and S. romanai from Colombia and Argentina reveal different cytotypes. Simulium bicoloratum is similar in pupal appearance and color of the adult, but the smooth basal portion of the cibarium permits it to be differentiated. Simulium (Ectemnaspis) lutzianum PINTO (Figs. 84 M, N; 85 C; 88 I, R; 89 G.) Simulium lutzianum PINTO, 1931: 748; RAMÍREZ PÉREZ, 1971: 311; SHELLEY & ARZUBE, 1985: 330. Simulium (Ectemnaspis) lutzianum: COSCARÓN, 1984: 303; 1987: 20; 1990: 131-132; 1991: 253-254.; COSCARÓN ARIAS, 2003: 195, 200, 205, 209, 215, 218. Ectemnaspis lutziana PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simulium (Psilopelmia) lutzianum: SHELLEY et al., 1997: 34; CROSSKEY & HOWARD, 1997: 60; 2004: 54. Simulium lewisi RAMÍREZ PÉREZ, 1971: 349. Simulium (Ectemnaspis) lewisi: COSCARÓN, 1984: 304; 1990: 132-133; SHELLEY et al., 1989: 93. Simulium iguazuense COSCARÓN, 1976: 147; 1984: 304; 1987: 29. Simulium (Psilopelmia) alirioi RAMÍREZ PÉREZ & VULCANO, 1973: 383 n. syn.

Female: Wing length 1.8-2.2 mm. Scutum and scutellum yellowish orange, metanotum blackish. Pleurae dark brown, with silver pruinosity. Legs dark brown except basal 2/ 3 of fore tibiae, and mid and hind basitarsus, which are whitish with a reduced dark area at apex of femora and tibiae. Abdomen with tergites I-II (III) yellowish to light brown, (III) IV-IX brownish, dark gray to blackish, III-IV opaque, V-IX shiny. Tergal plates IV-IX evident. Fronto-ocular area wider than high. Cibarium with sclerotized rim; central area with 5-6 rows of rounded tubercles and acute small teeth-like spines sublaterally (Figs. 84 M-N); some specimens similar to S. romanai, wrinkled without definite tubercles. Basal sector of R with hair. Ratio of length/width of hind basitarsus = 8. Claw with subbasal tooth (Fig. 85 C). Genitalia similar to those of S. romanai. Male: Wing length 1.6-2.0 mm. Coloration similar to that of female. Abdomen with segments I-III yellowish or only I brownish, rest of tergites black, VI-VII with 1+1 lateral silver spots. Ratio of length/width of hind basitarsus = 6.5-7.6. Genitalia as in S. romanai. Pupa: Cocoon thick, weave spongy, with very thin fibers; anterior border reinforced and dorsal medium longitudinal ridge (Fig. 88 I); length along base 2.2-3.4 mm, on dorsum 1.8-2.8 mm, gills 1.8-3.3 mm. Frontoclypeal and thoracic tubercles sparse with rounded borders (Fig. 88 R). Cephalic trichomes with 2-3 branches, thoracic with 5-7

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branches. Gill with 8 branches from very open to meeting basally. Bifurcation of secondary and tertiary branches at variable distances from base (Fig. 89 G). Larva: Maximum length 3.7-4.8 mm. Cephalic apotome without special ornamentation, slightly darkened on posterior border, with short single hairs. Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9-1.2:1.31.6. Mandible with 1 latero-mandibular process. Cephalic fan with about 35 branches. Postgenal cleft nearly reaching hypostomium base. Body integument with numerous small hairs. Thoracic proleg lateral sclerite with about 20 teeth. Anal ring with 55-74 rows with 11-13 hooks each. Rectal papillae with 8-12 diverticula on each lobe. Distribution: Argentina: Misiones; Bolivia: Cochabamba; Brazil: Goiás, Paraná, Roraima, Santa Catarina; Colombia: Cauca, Cundinamarca, Valle; Ecuador: Chimborazo, El Oro, Esmeraldas, Guayas, Imbabura, Los Rios, Manabí, Napo, Pastaza, Pichincha, Tungurahua, Zamora-Chinchipe; Panama: Panama; Peru: Cuzco, Huanuco, Junin; Venezuela: Amazonas, Aragua, Barinas, Bolivar, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Miranda, Monagas, Táchira, Trujillo, Yaracuy, Zulia. Bionomics: Simulium lutzianum breeds in torrential creeks of mountainous areas, on aquatic vegetation and stones. Female feeding habits are unknown. Discussion: Simulium lutzianum shows great variation in morphology, including the arrangement of the cibarial tubercles and size of the teeth, extent of yellow on the female and male abdomen and the length of the pupal gill and distance of bifurcation of the secondary branches. SHELLEY et al. (1997: 36) synonymized S. lewisi based on molecular and cytological studies in Ecuador (SAWYER, 1991), considering that the color variation is not linked to cytospecies and that cytoforms A, B and C belong to the S. lutzianum species complex. Additional non morphological studies with material from other regions will be necessary to elucidate the extent of S. lutzianum variation. Similarity with S. romanai is explained in that species description. Simulium bicoloratum is very similar in pupal shape and adult coloration, but has a smooth cibarium between the cornuae. Because the description of S. alirioi is included in the range of variation of S. lutzianum, it is synonymized here. Simulium (Ectemnaspis) gabaldoni RAMÍREZ PÉREZ (Figs. 84 L; 87 F.) Simulium gabaldoni RAMÍREZ PÉREZ, 1971: 352; 1983: 46. Simulium (Ectemnaspis) gabaldoni: COSCARÓN, 1984: 307-308; 1987: 29; 1990: 134. Simulium (Psilopelmia) gabaldoni: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female: Wing length 2.0-2.1 mm. Scutum yellowish-orange, scutellum yellowlemon, metanotum dark brown. Legs brown with darkened areas. Abdomen with tergite I yellow-lemon, II-III yellow with posterior border brown, IV-IX blackish. Cibarium with several rows of tubercles on median area and acute teeth at sides to cornuae (Fig. 84 L). Basal sector of R with 1 row of hair. Claws with subbasal tooth. Paraproct about as long as wide at base, very similar to that of S. arcabucense. Male: Wing length 1.8-1.9 mm. Color as in female but darker; legs with more extensive blackish area. Abdomen all black except 1+1 lateral silvery spots on segments VI-VII. Gonostylus subcylindrical, slightly shorter than gonocoxite. Ventral plate about as long as wide, with lateral constrictions (Fig. 87 F).

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Pupa: Cocoon thick, threads not evident, with slight anterior projection; length along base 2.3-2.6 mm, on dorsum 2.0-2.5 mm, gills 2.6-3.2 mm. Frontoclypeus and thorax with sparse tubercles; trichomes with 3-6 branches. Gills with 6 branches, bifurcated near base. Larva: Unknown. Distribution: Colombia: Valle; Venezuela: Monagas and Sucre. Bionomics: In Colombia, this species was collected at an altitude of 1400 m. No other information is available. Discussion: The coloration and number of gill branches of this species are similar to the condition in S. arcabucense and S. pautense, but these species have a smooth cibarium and no hairs on the basal sector of R. Simulium lutzianum is very similar, but has a pupa with 8 gill branches. Simulium (Ectemnaspis) albanense COSCARÓN (Figs. 84 K; 85 A; 86 M; 87 E; 88 Z; 90 D, J.) Simulium (Ectemnaspis) albanense COSCARÓN, 1990: 135-136. Simulium (Psilopelmia) albanense: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female: Wing length 2.8-3.0 mm. Frons and clypeus brownish with silvery pruinosity. Scutum and scutellum yellowish; scutum in some specimens with median and sublateral areas blackish brown. Pleurae yellowish to light or dark brown. Metanotum black. Legs blackish. Abdomen with segments I-III yellow, III with tergal plate brown, IVV velvety black, VI-IX shiny brown black. Fronto-ocular triangle wider than high. Cibarium with rounded tubercles on median area and with sparse spines laterally (Fig. 84 K). Basal portion of R without hair (Fig. 85 A). Eighth sternite with about 16 setae on each side. Gonapophysis and paraproct very similar to those of S. romanai. Male: Wing length 2.3-2.5 mm. Coloration as in female, with darkened scutum area larger, and frequently abdominal tergite III yellowish. Gonostylus about 1/3 gonocoxite length, subquadrate, with strong apical spur (Fig. 86 M); ventral plate as in figure 87 E. Pupa: Cocoon weave thin with delicate threads, anterior border slightly reinforced; length along base 3.0-3.2 mm, on dorsum 2.7-3.0 mm, gills 2.0-2.5 mm. Frontoclypeus and thorax with numerous elongate tubercles (Fig. 88 Z); trichomes with 4-8 branches. Gill with 8 branches grouped as bundle; bifurcation of tertiary branches at about half total gill length. Larva: Maximum length 6.0 mm. Cephalic apotome homogeneously darkened, with sublateral areas slightly lighter and with numerous multibranched hairs (Fig. 90 D). Postgenal cleft deep, dome shaped (Fig. 90 J); postgenal bridge about 1/8 hypostomium length. Cephalic fan with about 30 rays. Ratio of antennal segments: proximal, medial and distal = 1:1.6:2.3. Mandible with 1 latero-mandibular process. Body integument with hairs single or bifid. Lateral sclerite of thoracic proleg with about 21 teeth. Anal sclerite with abundant hair of 1-4 branches or spatulate shape. Anal ring with 64-77 rows of 12-13 hooks each. Rectal papillae with 3 diverticula on each lobe. Distribution: Colombia: Bogotá, Cundinamarca. Bionomics: Material of this species was collected at altitudes of 2400-3050 m. No more information is available.

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Discussion: The adults are very similar in coloration to other species of this group, but can be differentiated by the short subquadrate gonostylus, elongate tubercles of the pupal frontoclypeus and thorax, and pupal gill filaments arranged in a bundle. Simulium (Ectemnaspis) wolffhuegeli (ENDERLEIN) (Figs. 84 G, P; 86 O; 87 I; 88 K, U; 89 I; 90 F.) Wilhelmia wolffhuegeli ENDERLEIN, 1922: 70. Morops wolffhuegeli ENDERLEIN, 1930: 94. Simulium wolffhuegeli EDWARDS, 1931: 127; WYGODZINSKY, 1950: 88-96; COSCARÓN & WYGODZINSKY, 1960: 1124; COSCARÓN, 1968: 66. Psilopelmia wolffhuegeli ENDERLEIN, 1934b: 284. Tricodagmia haarupiana ENDERLEIN, 1936: 127. Simulium (Ectemnaspis) wolffhuegeli: COSCARÓN, 1984: 308-310; 1987: 29; 1990: 136; 1991: 255-257. Simulium (Psilopelmia) wolffhuegeli: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female: Wing length 1.9-2.1 mm. Scutum yellowish-orange with yellow hair, scutellum grayish yellow; metanotum dark brown; pleurae yellow with grayish pruinosity. Legs grayish yellow with apex of femora, tibiae and tarsomeres dark brown. Abdomen yellowish brown with great variation in color, mostly with tergites I-IV yellowish, tergal plate of tergites III-V darker, VI-IX shiny brownish gray (Fig. 84 G). Cibarium with several rows of rounded tubercles medially and numerous spines laterally (Fig. 84 P). Basal sector of R with hair. Claws with short subbasal tooth. Paraproct slightly longer than wide, gonapophysis subovoidal with internal border sclerotized. Male: Wing length 1.8-2.0 mm. Color generally blackish. Scutum black with pronotal and posterior border yellowish gray and covered by abundant silky, grayish pilosity. Scutellum and metanotum black. Abdomen blackish. Legs black, yellowish gray on clear areas. Gonostylus subrectangular, slightly longer than wide, with 7-15 spurs on apical border (Fig. 86 O). Ventral plate wider than long, with small projection mediodistally, without lateral constriction (Fig. 87 I). Pupa: Cocoon weave uniform, with threads evident and surface wrinkled, anterior border straight (Fig. 88 K). Length along base 2.6-3.0 mm, on dorsum 2.0-2.4 mm, gills 1.5-1.9 mm. Frontoclypeus and thorax with numerous tubercles. Cephalic trichomes with 1-3 branches; thoracic with 4-9 branches (Fig. 88 U). Gills with 8 branches (Fig. 89 I). Larva: Maximum length 5.0 mm. Cephalic apotome with variable ornamentation, typically darkened basally and posteriolaterally, sometimes with negative spots (Fig. 90 F). Antenna not surpassing apex of cephalic fan stem; ratio of antennal segments: proximal, medial and distal = 1:1.1-1.4:1.1. Postgenal cleft deep, dome shaped, nearly reaching hypostomium. Body integument with short hairs. Lateral sclerite of thoracic proleg with about 22 teeth. Cephalic fan with 25-31 rays. Anal ring with 65-70 rows with 14 hooks each. Rectal papillae generally with 0-1 diverticula on each lobe. Anal sclerite without scales or hairs, or the latter very scarce. Distribution: Argentina: Buenos Aires, Catamarca, Chubut, Córdoba, Jujuy, La Pampa, La Rioja, Mendoza, Neuquén, Río Negro, Salta, San Juan, San Luis, Santa Cruz, Tucumán; Bolivia: Cochabamba.

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Bionomics: This species breeds in rivers, slower creeks and irrigation canals with clear, but not necessarily crystalline, water and aquatic vegetation, from sea level to 2000 m. Females are haematophagous and dangerous to horses and cows; they generally don’t bite humans. Discussion: Females are very similar to some other species of the S. romanai group, but in S. wolffhuegeli the males have a black thorax, short gonostylus with 7-15 spurs, and ventral plate much wider than high. Simulium (Ectemnaspis) mayuchuspi COSCARÓN (Figs. 84 C; 85 B; 86 C; 87 G; 88 S; 90 Q.) Simulium (Ectemnaspis) mayuchuspi COSCARÓN, 1990: 136-139; COSCARÓN ARIAS, 2003: 198, 205, 209, 215. Simulium (Psilopelmia) mayuchuspi: CROSSKEY & HOWARD, 1997: 60; 2004: 54.

Female: Wing length 2.5-2.6 mm. General coloration blackish. Scutum blackish gray with 1+1 light grayish subtriangular anterior submedian spots and lateral border light grayish (Fig. 84 C); with certain lighting, 1+1 submedian blackish stripes can be seen joining on posterior 1/3, and aforementioned anterior grayish spots change to black; scutellum brown with posterior border yellowish. Hairs of mesonotum from silvery to golden, depending on illumination. Metanotum brown with dark grayish pruinosity. Pleurae and legs brownish black, with light brown on anterior half of fore tibia, anterior 1/5 of femora, anterior 3/4 of hind basitarsus and anterior half of second hind tarsomere; leg hairs yellowish-gray. Abdomen dark brown to black with tergites II-IV velvety black and V-IX shiny brown; ventrally brownish gray. Fronto-ocular triangle not deep. Cibarium with well-sclerotized rim; rounded tubercles medially and abundant acute teeth laterally, as in S. romanai and S. lutzianum. Basal sector of R with hairs. Claws without subbasal tooth (Fig. 85 B). Eighth sternite with 25-30 setae on each side. Gonapophyses subtrianglular, with internal borders slightly divergent and little sclerotized. Male: Wing length 2.5-2.6 mm. Scutum black; with posterior illumination 1+1 grayish anterior submedian triangular spots can be seen continued anterolaterally by short silvery gray bands, and irregular transverse spots on posterior third (Fig. 86 C); with anterior light, scutum gray opaque with median and posterior 1/3 black. Scutellum, metanotum and pleurae as in female. Abdomen black with 1+1 large lateral silvery spots on segments II, III, VI, VII. Ratio of length/width of hind basitarsus = 5.2. Gonostylus about 3 times as long as wide, curved on half distal and with apical spur similar to that of S. arcabucense. Ventral plate with strong lateral constriction (Fig. 87 G). Pupa: Cocoon thick with spongy weave; threads very thin not obvious, slightly thicker on anterior border, and dorsal ridge not well defined, similar to that of S. romanai. Frontoclypeus and thorax with numerous thin tubercles (Fig. 88 S). Cephalic trichomes with 2-7 branches and thoracic with 7-12 branches. Gill with 8 branches, with branching close to base. Larva: Cephalic apotome light brown, slightly ornamented with basal median area darker. Ratio of antennal segments: proximal, medial and distal = 1:1.3:1.5. Cephalic fan with about 36 rays. Mandible with 2 latero-mandibular processes (Fig. 90 Q). Postgenal cleft about 1/7 hypostomium length. Body integument without hairs. Anal sclerite with or without very sparse hairs but no scales. Anal ring with 65 rows with 13-17 hooks each. Rectal papillae with 5-10 diverticula on each lobe.

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Distribution: Ecuador: Pichincha; Peru: Junín. Bionomics: Simulium mayuchuspi breeds in small creeks with fast flowing, crystalline waters, and aquatic vegetation. Female feeding habits are unknown. Discussion: The presence of black females allows this species to be differentiated from other Simulium (Ectemnaspis) species. PERFLAVUM species group Simulium (Ectemnaspis) perflavum ROUBAUD (Figs. 84 R; 85 K; 86 Q; 87 J; 89 J; 90 G, N, S.) Simulium perflavum ROUBAUD, 1906a: 518; LUTZ, 1910: 230; PINTO, 1931: 478; D’ANDRETTA, 1956: 3338; RAMÍREZ PÉREZ, 1971: 301; 1983: 62; COSCARÓN & WYGODZINSKY, 1972: 223; PY-DANIEL, 1989: 254; HAMADA, 1998: 523; HAMADA & ADLER, 1998b: 207; 1999: 273-288; 2001: 119; HAMADA & MC CREADY, 1999: 71; HAMADA & GRILLET, 2001: 35; ALENCAR et al., 2001: 561. Simulium (Ectemnaspis) perflavum: COSCARÓN, 1984: 310; 1987: 29; 1990: 140-141; 1991: 257; STRIEDER et al., 1992: 56. Simulium (Eusimulium) perflavum: LUTZ, 1909: 131. Simulium (Odagmia) perflavum: RUBTZOV, 1940: 532. Eusimulium perflavum: SURCOUF & GONZALEZ RINCONES, 1911: 283; LANE & PORTO, 1939: 169-172; LANE & VULCANO, 1943: 432. Ectemnaspis perflava: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER , 2004: 129-130. Simulium (Psilopelmia) perflavum: CROSSKEY & HOWARD, 1997: 59; 2004: 54. Eusimulium antunesi LANE & PORTO, 1940: 185; LANE & VULCANO, 1943: 436. Simulium mauense NUNES DE MELLO, 1974: 35. Simulium nilesi RAMBAJAN, 1979: 63-77.

Female: Wing length 2.6-2.8 mm. Scutum and scutellum yellowish orange, metanotum, pleurae, sternites and abdomen yellowish brown. Abdominal pleurae grayish. Legs yellowish with darker portion of cibarium covered by abundant acute teeth-like spines, and projected medially with small median invagination (Fig. 84 R). Basal sector of R with hair. Eighth sternite with about 25 setae; gonapophyses with internal margins divergent, slightly produced distally and internal rim poorly sclerotized. Paraproct about 2 times longer than wide at base, narrower on distal 3/4 (Fig. 85 K). Male: Wing length 2.5-2.7 mm. Coloration similar to that of female but slightly darkened. Ratio of length/width of hind basitarsus = 6.0-6.4. Gonostylus as long as gonocoxite, subcylindrical, with apical spur (Fig. 86 Q). Ventral plate about 2 times longer than width, with small lateral constrictions (Fig. 87 J). Pupa: Cocoon with thick and compact weave, slightly projected basally; length along base = 2.1-2.5 mm, on dorsum 2.6-3.0 mm, gills 2.0-2.5 mm. Frontoclypeus and thorax with numerous tubercles; trichomes with 1 or 2 (3-4) branches. Gill with 8 branches arising very close to base with very short basal trunk and without evident primary branches (Fig. 89 J); in some areas (e.g., Corrientes, Argentina) branches darker and thicker. Larva: Maximum length 6.5 mm. Cephalic apotome slightly darkened medially with 1 pair each of basal and sublateral negative spots (Fig. 90 G). Postgenal bridge shorter than half of hypostomium length. Antennae not surpassing stem apex of cephalic fan; ratio of proximal,

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media1 and distal segments = 1:1.7:1.4-1.6 (Fig. 90 N). Mandible with 1-2 latero-mandibular processes. Anal ring with 78-87 rows with 13-14 hooks each. Anal sclerite with hair relatively numerous, no scales. Rectal papillae with 10-15 diverticula on each lobe (Fig. 90 S). Distribution: Argentina: Corrientes, Formosa, Misiones; Brazil: Acre, Amazonas, Amapá, Bahia, Espírito Santo, Goiás, Mato Grosso, Minas Gerais, Pará, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, São Paulo; Paraguay: Caaguazu, Cordillera; Venezuela: Amazonas, Bolivar, Carabobo. Bionomics: Simulium perflavum is a tropical species breeding in small flows that are not fast flowing, and that are frequently in open areas with high insolation. The water is clear to black or muddy yellowish-red in areas that suffer deforestation, road construction, and other degradation. Females are not anthropophilic. Discussion: The yellowish coloration of the female, the male genitalia and the gill branch arrangement differentiate this species from other Simulium (Ectemnaspis) species. Cytologically analyzed populations from 10 states in Brazil and 1 in Venezuela represent a single species (HAMADA & ADLER, 1999). Simulium (Ectemnaspis) kabanayense RAMÍREZ PÉREZ & VULCANO (Figs. 84 Q; 85 L; 86 P; 87 K; 89 K.) Simulium (Psilopelmia) kabanayense RAMÍREZ PÉREZ & VULCANO, 1973: 390; CROSSKEY & HOWARD, 1997: 59; 2004: 53. Simulium kavanayense (sic.): RAMÍREZ PÉREZ, 1983: 48. Simulium kabanayense: HAMADA & ADLER, 1998b: 221; HAMADA & GRILLET, 2001: 34. Simulium (Ectemnaspis) kabanayense: COSCARÓN, 1984: 312; 1987: 29; 1990: 141.

Female: Wing length 2.1-2.3 mm. Scutum orange with silvery lateral rim; scutellum yellowish; pleurae yellowish brown; metanotum dark brown. Legs yellowish brown with darker portion blackish. Abdomen with tergites I-II yellowish, III-IX blackish; tergal plate III brown, V-IX shiny. Fronto-ocular triangle about as wide as long. Cibarium with area between cornuae protruded, covered by acute teeth-like spines scarce at middle (Fig. 84 Q). Paraproct about as long as wide (Fig. 85 L). Male: Wing length 2.0 mm. Coloration as in female, but abdominal tergite I yellowish brown, the following blackish. Gonostylus relatively robust, about as long as gonocoxite (Fig. 86 P); ventral plate with deep lateral constrictions (Fig. 87 K). Pupa: Cocoon slightly projected anteriorly at base; weave compact with obvious threads. Length along base 2.5 mm, on dorsum 2.3 mm, gills 2.0-2.5 mm. Frontoclypeal and thoracic tubercles sparse with smooth surface; cephalic trichomes bifid, thoracic with 2-6 branches. Gill with 10 (12) branches, with branching close to base (Fig. 89 K). Larva: (based on HAMADA & ADLER, 1998). Cephalic apotome without special ornamentation; hypostomium with median tooth at same level as lateral teeth; postgenal cleft pointed; cephalic fan with 40-49 rays; anal ring with 83-85 rows of 12-18 hooks each. Distribution: Brazil: Roraima; Venezuela: Bolivar. Bionomics: Unknown. Discussion: Females of S. kabanayense are very similar to those of S. maroniense, S. rorotaense and S. suarezi, but can be differentiated by the length of the paraproct. The male

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of S. kabanayense can be distinguished by the yellowish brown abdominal tergites I-II, and the pupa by the gill of 14-23 branches. Simulium (Ectemnaspis) maroniense FLOCH & ABONNENC (Figs. 84 B; 88 V; 89 L.) Simulium maroniense FLOCH & ABONNENC, 1946a: 9; RAMÍREZ PÉREZ, 1983: 53; SHELLEY et al., 1984: 153; HAMADA & ADLER, 1998b: 207; 1999: 277; 2001: 119; HAMADA & GRILLET, 2001: 35. Simulium (Ectemnaspis) maroniense: COSCARÓN, 1984: 313; 1987: 29; 1990: 141. Simulium (Psilopelmia) maroniense: SHELLEY et al., 2004: 455-458; CROSSKEY & HOWARD, 2004: 53.

Female and Male: Morphologically and ornamentally similar to that of S. rorotaense, but female gonapophysis slightly longer, fronto-ocular triangle lower, and sensory vesicle of maxillary palp about 1/3 of palpomere length (Fig. 84 B); male ventral plate with deeper constrictions laterally. Pupa: Cocoon similar to that of S. rorotaense. Frontoclypeal and thoracic tubercles more abundant and larger than in S. rorotaense (Fig. 88 V); trichomes elongate with 3-6 branches. Gill with 18-20 filamentous branches that increase in length from dorsal to ventral position (Fig. 89 L) and also are darker than in S. rorotaense. Larva: (based on HAMADA & ADLER, 1998). Antenna with medial segment shorter than distal. Distribution: Brazil: Amapá, Roraima; French Guiana; Venezuela: Amazonas, Bolivar. Bionomics: Unknown. Discussion: This species was synonymized by SHELLEY et al. (1984: 153) but was revalidated by COSCARÓN (1990: 141) and HAMADA & ADLER (1998: 207), who confirmed the validity of this species. Simulium suarezi is very similar; differences are explained in the discussion of that species. HAMADA & ADLER (1999) found four cytotypes of S. maroniense, showing the complexity of this species. Simulium (Ectemnaspis) rorotaense FLOCH & ABONNENC (Figs. 84 c; 85 M; 86 R; 87 L; 89 M; 90 H, K.) Simulium rorotaense FLOCH & ABONNENC, 1946a: 4; PY-DANIEL, 1981: 175; RAMÍREZ PÉREZ, 1983: 66; HAMADA & ADLER, 1998b: 221; 2001: 119 HAMADA & GRILLET, 2001: 35. Simulium (Ectemnsapis) rorotaense: COSCARÓN, 1984: 313; 1987: 29; 1990: 141-143. Simulium (Psilopelmia) rorotaense: SHELLEY et al., 2004: 455-458; CROSSKEY & HOWARD, 1997: 59; 2004: 54. Simulium fulvinotum CERQUERA & NUNES DE MELLO, 1968: 205. ? Simulium wuayaraka ORTIZ, 1957: 163. ? Simulium (Psilopelmia) ignacioi RAMÍREZ PÉREZ & VULCANO, 1973: 387; RAMÍREZ PÉREZ, 1983: 8; HAMADA & ADLER, 1998b: 207.

Female: Wing length 2.0-2.5 mm. Scutum, scutellum and metanotum yellowishorange, scutum with lateral borders nacreous and hair yellowish. Legs yellowish, darkened mostly on apices of segments. Abdominal segments I, II (III) light brown-yellow, IV-IX blackish. Sensory vesicle of maxillary palp longer than half of palpomere length (Fig. 84 c). Cibarium slightly elevated medially, very similar to that of S. suarezi and S. kabanayense.

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Basal portion of R with hair. Eighth sternite with 19-22 setae on each side; gonapophysis slightly sclerotized on internal rim. Paraproct nearly 2 times longer than wide, acute distally, with large microtrichiae on posterior border (Fig. 85 M). Male: Wing length 2.0-2.2 mm. Coloration similar to that of female. Gonostylus subcylindrical, slightly curved on distal third and about same length as gonocoxite (Fig. 86 R). Ventral plate slightly shorter than width at base, with strong lateral depressions (Fig. 87 L). Pupa: Cocoon weave not compact, with open areas especially near aperture Length along base 2.7 mm. Frontoclypeus and thorax slightly sclerotized, tubercles not abundant, trichomes with 2-3 branches. Gills with 18-20 thin filamentous branches, bifurcated at variable distances from base (Fig. 89 M). Larva: Maximum length 5.0 mm. Cephalic apotome with darkened median basal area, without special ornamentation, and with several negative spots near basal border (Fig. 90 H). Ratio of antennal segments: proximal, media1 and distal = 1:0.7:1.4. Postgenal bridge about 1/5 of hypostomium height (Fig. 90 K). Mandible with 1 latero-mandibular process. Cephalic fan with about 45 rays. Rectal papillae with 13-15 elongate diverticula on each lobe. Body integument glabrous. Distribution: Brazil: Amazonas; Venezuela: Amazonas, Bolivar. Bionomics: This species breeds in clear, acidic creeks of fast flow and about 25 °C. Larvae are found on aquatic vegetation. Females bite humans and cattle. Discussion: This species is very similar to S. maroniense and S. suarezi .It can be differentiated by the pupal gill, which viewed frontally looks like an open rosette; branch bifurcation is closer to the base. Simulium wuayaraka was synonymized by RAMÍREZ PÉREZ (1983) and SHELLEY et al. (1984); the female coloration and paraproct are similar, but the cibarium morphology, according to the original drawings, is closer to that of S. shewellianum and S. ochraceum. It would be useful to confirm the identity of this species with more material from different life stages. Simulium ignacioi was synonymized by RAMÍREZ PÉREZ (1983: 8) and accepted by SHELLEY et al. (1984) and according to the original description, it is very similar to S. rorotaense, especially in the pupal gill, but the description is limited. Cytological studies by HAMADA & ADLER (1998) indicate that it is a good species, although morphological characters were not found to separate it from S. rorotaense. Simulium (Ectemnaspis) suarezi RAMÍREZ PÉREZ, RASSI & RAMÍREZ (Figs. 84 S; 85 N; 88 L; 89 N.) Simulium (Psilopelmia) suarezi RAMÍREZ PÉREZ et al., 1977: 173; RAMÍREZ PÉREZ, 1983: 120; HAMADA & ADLER, 1998b: 221; HAMADA & GRILLET, 2001: 35; CROSSKEY & HOWARD, 1997: 60; 2004: 54. Simulium (Ectemnaspis) suarezi: COSCARÓN, 1984: 314-315; 1987: 29; 1990: 143. Ectemnaspis suarezi PY- DANIEL & MOREIRA SAMPAIO, 1995: 118, 120.

Female: Wing length 2.3 mm. Scutum yellowish with silvery lateral borders. Scutellum yellow. Metanotum dark brown. Pleurae and legs brownish yellow. Abdomen with segment I and tergal plate II-III brownish yellow, IV-IX dark brown, V-IX shiny and waxy. Fronto-ocular triangle wider than high. Cibarium with median elevation bearing spines and central depression (Fig. 84 S). Claws with subbasal tooth well developed. Paraproct nearly 2 times longer than wide at base, subtriangular ,acute apically (Fig. 85 N).

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Male: Limited information is available. RAMÍREZ PÉREZ (1983: 120; Fig. 83) shows gonostylus relatively thin, distally curved and longer than gonocoxite; ventral plate longer than wide basally and with strong lateral constriction. Pupa: Cocoon with non compact weave, with holes on anterior border and extended covering laterally on gills (Fig. 88 L). Frontoclypeus and thorax with numerous wrinkled tubercles; trichomes with several to 10 branches. Gill with 20 thick filamentous branches emerging from basal trunk and, when viewed frontally, very open (Fig. 89 N). Larva: (based on HAMADA & ADLER, 1998): Antenna with medial segment shorter than distal. Anal ring with 98-101 rows with 16 hooks each. Cephalic fan with about 40 primary rays. Distribution: Venezuela: Amazonas. Bionomics: Unknown. Discussion: The most similar species are S. maroniense and S. rorotaense. The pupal gill filaments are thinner, and not open, with longer primary and secondary branches. Simulium (Ectemnaspis) trombetense HAMADA, PY-DANIEL & ADLER (Fig. 89 O.) Simulium (Ectemnaspis) trombetense HAMADA et al., 1999 (in HAMADA & ADLER, 1998b: 210); HAMADA & ADLER, 2001: 119; HAMADA & GRILLET, 2001: 35; CROSSKEY & HOWARD, 2004: 54.

(Based on original description of HAMADA & ADLER, 1998). Female: Wing length 2.2 mm. General coloration orange. Frons and clypeus with silver blue pruinosity. Scutum orange with silvery lateral borders, hair golden-brown. Pleurae brownish orange to brown with silver pruinosity. Scutellum and metanotum orange, the latter with some brownish orange. Legs yellowish, dark brown on tibiae, fore basitarsus and tarsomeres. Abdomen with tergites I-II orange, II with silver pruinosity, III-VI greenish orange to gray, VI-VIII orange, shiny. Fronto-ocular triangle very flat. Cibarium with 1+1 anterior submedian elevation covered by well-developed teeth. Basal sector of R with setae, claws with strong subbasal tooth. Paraproct, longer than wide, apically acute, similar to that of S. suarezi. Male: Wing length 2.0-2.1 mm. Coloration as in female. Gonostylus 0.7 times as long as gonocoxite, with strong curvature apically. Ventral plate with lateral constrictions, shape very similar to that of S. kabanayense. Pupa: Cocoon dark brown; weave parchment like with open loops anteriorly, threads distinct; length along base 3.2 mm, on dorsum 3.4-3.7 mm, frontoclypeus and thorax with relatively abundant rounded tubercles; large trichomes with 6-9 branches. Gills with 160250 thin short filaments arising from 7 thick trunks (Fig. 89 O). Larva: Maximum length 6.0-8.0 mm. Cephalic apotome with brown anteromedian, anterolateral and posterior spots. Postgenal cleft deep, tapered apically, postgenal bridge about 1/4 of hypostomium length. Hypostomium with median tooth as long as lateral teeth. Mandible with 1 latero mandibular process, thin, surpassing inferior margin of mandible. Antenna as long as cephalic fan stem; ratio of proximal, media1 and distal segment = 1:0.9-1.0:1.1-1.3. Cephalic fan with 45-50 rays. Anal ring with 89-113 rows with 14-16 hooks each. Rectal papillae with 11-13 elongate diverticula on each lobe. Distribution: Brazil: Amapá, Amazonas, Pará, Roraima; Venezuela: Bolivar.

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Bionomics: This species breeds in forested areas of the Guiana Shield in fast currents. Larvae are found on bedrock, fallen leaves and aquatic vegetation. Female feeding habits are unknown. Discussion: Adults of the S. perflavum species group are very difficult to separate. Simulium perflavum can be separated by the absence of dark patches on the middle legs. The pupal gill with over 100 branches is a very useful character to separate S. trombetense from other closely related species. DINELLII species group Simulium (Ectemnaspis) dinellii JOAN (Figs. 84 T, Z; 85 O; 87 M; 88 W.) Melusina dinellii JOAN, 1912: 363. Simulium dinellii KNAB, 1913: 155; 1914b: 81; EDWARDS, 1933: 210; WYGODZINSKY, 1950: 77-87; BARRETO, 1969: 31. Simulium (Simulium) dinellii: ORFILA, 1935: 1532. Eusimulium dinellii: PATERSON & SHANNON, 1927: 738; PINTO, 1931: 677; LANE & VULCANO, 1943: 434. Eusimulium ochraceum: LANE & PORTO, 1939: 171 (nec WALKER); LANE & VULCANO, 1943: 434. Dasypelmoza miniata ENDERLEIN, 1934b: 275. Trichodagmia miniata ENDERLEIN, 1934a: 194; 1936: 129. Simulium martinezi VARGAS, 1943a: 137; VARGAS, 1945a: 158; SMART, 1945: 508. Simulium (Ectemnaspis) dinellii: COSCARÓN, 1984: 317-318; 1987: 29; 1990: 143; 1991: 258-260; COSCARÓN ARIAS, 2003: 195, 198, 205, 215. Simulium (Psilopelmia) dinellii: CROSSKEY & HOWARD, 1997: 58; 2004: 52. ? Simulium flavipictum KNAB, 1914b: 83. Ectemnaspis dinellii: STRIEDER, 2004: 129.

Female: Wing length 1.6-2.7 mm. Scutum yellowish orange, a little lighter on anterolateral and posteromedian borders; scutellum light yellow, metanotum orange. Pleurae yellowish orange, mesosternum with bluish pruinosity. Legs yellowish orange, with dark brown areas. Abdomen with segments I-IV yellowish green, V-VII brownish green, VI-VII slightly lighter, VIII-IX brownish yellow. Cibarium with median rounded invagination flanked by 1+1 submedian blunt projections covered by short spines (Fig. 84 T). Fronto-ocular triangle about as long as wide. Sensory vesicle of maxillary palp about 1/4 palpomere length (Fig. 84 Z). Basal sector of R with hair. Eighth sternite with about 27 setae; paraproct 1.5 longer than wide at base (Fig. 85 O). Gonapophysis with internal border lightly sclerotized. Male: Wing length 2.5-2.6 mm. Coloration as in female, with abdomen more orange reddish, and 1+1 silvery spots on segments II, VI, VII and IX. Ratio of length/width of hind basitarsus = 6. Gonostylus subcylindrical with distal portion subconical, curved inwardly, and about 2/3 of gonocoxite length. Ventral plate about as long as wide at base, with slight lateral constrictions (Fig. 87 M). Pupa: Cocoon without anterior projection, thin weave, threads sparse and obvious; length along base 3.1-3.7 mm, on dorsum 2.9-3.2 mm, gills 2.0-2.2 mm. Frontoclypeus and thorax with medially sparse tubercles; trichomes with 3-5 branches (Fig. 88 W). Gills with 8 branches, not bifurcating close to base, curved, resembling basket.

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Larva: Maximum length 6.0 mm. Cephalic apotome slightly darker basally, and prolonged by 1+1 submedian stripes, with 1+1 light spots basally and on subapical portion of submedian stripes. Postgenal cleft nearly reaching hypostomium base. Antenna about same length as cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9:1.0. Cephalic fan with 35-43 rays. Lateral sclerite of proleg with about 21 teeth arranged in 10 groups. Anal ring with 76-82 rows, with 12-16 hooks each. Rectal papillae with 3-4 diverticula on each lobe. Distribution: Argentina: Catamarca, Jujuy, La Rioja, Salta, Santiago del Estero, Tucumán; Bolivia: Cochabamba, La Paz; Brazil: Minas Gerais, Rio Grande do Sul; Colombia; Ecuador: Esmeraldas, Pichincha; Peru: Apurimac; Venezuela: Aragua. Bionomics: Simulium dinellii breeds in small, clear watercourses, sometimes in warm flows with high levels of Fe, Su and Cl. Females are anthropophilic and are considered a dangerous species, biting on the face and arms. Discussion: The most similar species is S. shewellianum, but it has dark brown legs, and the cibarium has a more sclerotized rim with 1+1 submedian acute apical projections without spines. Females of S. lutzianum, S. gabaldoni, S. albanense and most species of the S. romanai group are similar, but they have a more flattened basal portion of the cibarium, with tubercles on the median area and numerous spines between the tubercles and cornuae. The males are mostly blackish. Simulium flavipictum has coloration and external morphology similar to that of S. dinellii, although it is a little larger. Simulium flavipictum is known from a female from Peru (Charape River). We synonymize S. flavipictum with doubt, because of the unknown cibarium and genitalia. Simulium (Ectemnaspis) shewellianum COSCARÓN (Figs. 84 H, U; 85 P; 86 S; 87 N; 88 X; 89 P; 90 L.) Simulium (Ectemnaspis) shewellianum COSCARÓN, 1984: 318-319; 1987: 30; 1990: 144; MUÑOZ & COSCARÓN, 1999: 183; COSCARÓN ARIAS, 2003: 200, 205, 209, 215. Simulium (Psilopelmia) shewellianum CROSSKEY & HOWARD, 1997: 58; 2004: 52. Simulium shewellianum: PEPINELLI et al., 2003: 653.

Female: Wing length 1.9 mm. Scutum yellowish with silvery lateral borders, pleurae brownish yellowish, with silvery pruinosity under certain illumination. Scutellum and metanotum yellowish. Legs mostly dark brown with whitish area on mid and hind basitarsus. Abdomen I-IV yellow, V-IX brown, tergite V velvety and VI-IX shiny. Cibarium with well sclerotized rim, with 1+1 submedian acute projections and median depressed area wrinkled (Fig. 84 U). Fronto-ocular triangle longer than wide. Sensory vesicle of maxillary palp longer than half of palpomere (Fig. 84 H). Basal sector of R with hair. Eighth sternite with about 20 setae on each side; gonapophysis more sclerotized along inner margin. Paraproct slightly longer than wide with rounded apex (Fig. 85 P). Male: Wing length 1.8 mm. Coloration similar to that of female. Ratio of length/ width of hind basitarsus = 6.6 Gonostylus very similar to that of S. dinellii with pronounced curvature apically (Fig. 86 S), slightly shorter than gonocoxite. Ventral plate with slight lateral constrictions (Fig. 87 N). Pupa: Cocoon with weave compact, resistant, threads obvious; length along base and dorsum 2.8 mm, gills 2.3 mm. Frontoclypeus and thorax with moderate quantity of

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tubercles (Fig. 88 X); cephalic trichomes with 2-3 branches, thorax with 3-4 branches. Gills with 8 branches, not bifurcated close to base (Fig. 89 P). Larva: Maximum length 4.3-4.5 mm. Cephalic apotome with few ornamentations, with median and sublateral areas slightly lighter and with simple hair. Postgenal bridge absent or very narrow (Fig. 90 L). Cephalic fan with 30-34 rays. Ratio of antennal segments proximal, medial and distal = 1:1.1:1.3. Mandible with 1 latero-mandibular process. Lateral sclerite of proleg with 21 teeth arranged in 7 groups. Anal ring with 72 rows with 11-13 hooks each. Rectal papillae with 6-7 diverticula on each lobe. Eighth segment with 1 pair of ventral papillae. Integument without hair. Distribution: Brazil: São Paulo; Colombia; Ecuador: Los Rios, Pastaza, Pichincha. Bionomics: This species breeds in small creeks with clear water, on aquatic vegetation, at an altitude of about 400 m. Female feeding habits are unknown. Discussion: Simulium lutzianum has similar adult coloration, pupal gill branches and trichomes, but it is possible to distinguish it by the cibarial morphology. Simulium ochraceum also shows some similarity but has 1+1 anterior oblique submedian silvery-whitish stripes on the scutum. Similarity with S. dinellii was treated in the discussion of that species. Simulium (Ectemnaspis) ochraceum WALKER (Figs. 84 E, V; 87 O.) Simulium ochraceum WALKER, 1861: 332; FAIRCHILD, 1990: 706; VARGAS, 1942a: 238; 1946: 173. Simulium (Simulium) ochraceum: DALMAT, 1955: 225. Simulium (Eusimulium) ochraceum: HOFFMAN, 1931: 207 (in part). Eusimulium (Eusimulium) ochraceum: BEQUAERT, 1934: 208. Eusimulium ochraceum: LANE & VULCANO, 1943: 437. Psilopelmia ochraceum: RUBTZOV & GARCÍA AVILA, 1972: 15. Simulium (Psilopelmia) ochraceum: CROSSKEY & HOWARD, 1997: 58; 2004: 52.; SHELLEY et al., 2002: 164. Simulium (Ectemnaspis) ochraceum: COSCARÓN, 1984: 320; 1987: 30; 1990: 147; COSCARÓN ARIAS, 2003: 195, 200, 205, 215, 218. Simulium bipunctatum MALLOCH, 1912: 650. Simulium (Psilopelmia) bipunctatum: SHELLEY et al., 1989: 130; 1997: 28. Simulium (Ectemnaspis) bipunctatum: COSCARÓN, 1984: 321; 1987: 29; 1990: 144. Simulium wolcotti FOX, 1953: 138. ? Simulium scutellatum LANE & PORTO, 1940: 192; LANE & VULCANO, 1943: 434 n. syn. Simulium (Ectemnaspis) scutellatum: COSCARÓN, 1990: 145.

Female: Wing length 1.9-2.3 mm. Scutum yellow, to yellowish red with 1+1 anterolateral silvery spots continued by short submedian stripes not reaching to half of scutum length (Fig. 84 E) but in accordance with DALMAT material from Guatemala, it reaches the prescutellar border. Scutellum yellow, pleurae yellowish brown, showing silvery tones on basal half under certain illumination. Metanotum dark brown, silvery posteriorly. Legs yellowish, darkened on apex of femora and tibiae of hind and mid leg, tibiae and tarsomeres dark brown. Abdomen with segments I-IV yellowish to brownish yellow as also lateral borders of V; VI-IX dark grayish brown; V tergite with velvety appearance, VI-IX shiny, waxy. Frons slightly divergent; fronto-ocular triangle about as long as wide. Cibarium with basal portion well sclerotized, with 1+1 submedian apically

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acute processes, with rounded median area, and rim between cornuae wrinkled or with small denticle-like spines (Fig. 84 V). Sensory vesicle of maxillary palp surpassing half of palpomere length. Basal sector of R with hair. Claws with well-developed subbasal tooth. Eighth sternite with about 15 setae on each side. Gonapophysis with slight sclerotization on internal rim. Paraproct about as long as wide at base, similar to Simulium shewellianum. Male: Wing length 1.9-2.0 mm. Coloration as in female but darker ochre and silvery submedian stripes shorter. Gonostylus elongated, subcylindrical, curved distally with apical spur and slightly shorter than gonocoxite length. Ventral plate mostly shorter than wide at base (Fig. 87 O) and with small lateral constrictions. Pupa: Cocoon compact, slightly thicker dorsally, threads evident. Length along base 2.7-3.8 mm, on dorsum 2.5-3.3 mm, gills 1.7-2.5 mm. Frontoclypeus and thorax with since numerous to moderate number of tubercles; trichomes with 2-4 branches. Gill with 8 branches arising close to base. Larva: Maximum length 4.4-5.0 mm. Cephalic apotome with median dark spot wider basally. Postgenal cleft reaching hypostomium base, postgenal bridge generally absent. Cephalic fan with 28-36 rays. Antenna about as long as or a little longer than length of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1:1.3. Anal sclerite with or without sparse hairs. Anal ring with 76-78 rows of 14-15 hooks each. Rectal papillae with about 8 elongate diverticula on each lobe. Ninth segment with 2 ventral tubercles. Distribution: Brazil: Amazonas, Roraima; Colombia: Cundinamarca; Costa Rica; Cuba; Dominican Republic; Ecuador: Esmeraldas, Pichincha; Guatemala Amatitlan, Chimaltenango, Escuintla, Suchitepequez; Mexico: Chiapas, Oaxaca, Veracruz; Panama: Chiriqui, Coclé, Colón; Peru; Puerto Rico; Venezuela: Aragua, Carabobo, Monagas. Bionomics: Simulium ochraceum breeds in small non torrential creeks and water courses on rocks, decayed leaves and aquatic vegetation. Females bite humans and cattle. It is a vector of Onchocerca volvulus in Mexico and Guatemala. Discussion: Simulium ochraceum is very similar to other members of the S. dinellii species group. Simulium antillarum (synonymized by SHELLEY et al., 1987), with 1+1 submedian silvery spots can be differentiated by the scutum silvery stripes more elongated reaching or surpassing the half of scutum length, metanotum yellowish, the cibarium with a medially depressed area without spines and a thicker basal rim medially, and male with ventral plate more elongated and thinner gonostylus. Simulium shewellianum can be differentiated by the absence of 1+1 submedian silvery stripes on the scutum, and the cibarium medially wrinkled. Simulium ochraceum shows variation in the ventral plate, such as lack of lateral constrictions, especially in Mesoamerican specimens that also have an absence of or very scarce tubercles on the frontoclypeus and thorax. Additional variation includes 1+1 submedian silvery stripes reaching the silver posterior area, and larval cephalic apotome darkened medially and basally (DALMAT, 1955). Three cytotypes are known (HIRAI & HUEMOTO, 1983) in Guatemala and Mexico. Probably the morphological variation corresponds to specific cytological or molecular differences. In accordance with the great variation, S. ochraceum can be considered a species complex. Simulium scutellatum from Colombia, known only from females, has similar coloration and cibarium morphology. Here it is synonymized with doubts. Simulium wolcotti, known from adults, falls morphologically into the range of variation for S. ochraceum.

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Simulium (Ectemnaspis) antillarum JENNINGS (Figs. 84 D, W; 87 P.) Simulium antillarum JENNINGS, 1915: 200. Simulium (Psilopelmia) antillarum: DÍAZ NÁJERA, 1961: 79; STONE, 1969: 312-313; COSCARÓN, 1984: 319; 1987: 29; 1990: 146. Simulium (Psilopelmia) pseudoantillarum RAMÍREZ PÉREZ & VULCANO, 1973: 379. Simulium pseudoantillarum: RAMÍREZ PÉREZ, 1983: 63. Psilopelmia antillarum: RUBTSOV & GARCIA AVILA, 1972: 11-15.

Female and Male: Scutum yellowish with 1+1 submedian silvery spots, reaching or surpassing the half of scutum length (Fig. 84 D), abdomen with segments I-IV yellowish and the following brownish gray,. Cibarium with smooth median area between submedian processes, without spines or wrinkled sclerotized area, bordered basally by thick rim (Fig. 84 W). Male ventral plate longer than wide, with small lateral constrictions (Fig. 87 P). Pupa: Slight differences from S. ochraceum: frontoclypeus and thorax with moderate number of tubercles, more abundant and arranged in small groups, and trichomes with 26 branches. Gill branching near base (DÍAZ NÁJERA, 1961) or primary and secondary branches bifurcated at further distance, as in S. shewellianum. Larva: Similar to that of S. ochraceum; cephalic apotome can show some lighter lateral areas. Distribution: Colombia: Amazonas, Cundinamarca, Valle; Cuba; Dominica; Jamaica; Mexico: Chiapas; Ecuador: Galápagos; Peru: Cuzco, Huanuco; Puerto Rico; Venezuela: Monagas, Sucre. Bionomics: This species breeds in small fast-flowing creeks. Larvae are found on fallen leaves, roots, aquatic vegetation and stones. Females are anthropophilic. Discussion: The typical cibarium is smooth medially, which separates it from S. ochraceum and S. shewellianum; this cibarium is very close to that of S. dinellii, but it has the 1+1 submedian processes thinner and more elongated, with a narrow basal rim. Also, in S. dinellii and S. shewellianum the scutum has no submedian silvery stripes. Simulium pseudoantillarum is in accordance with material from Peru, Colombia and Venezuela, but material from Mexico (DÍAZ NÁJERA 1961) has a pupal gill with branching close to the base. Material from Peru and Galápagos, which we now are considering S. antillarum, has a few differences: a yellow ochreous scutum with thicker silvery submedian stripes. (Fig. 84 D). Surely, here we have another species complex. Simulium (Ectemnaspis) nuneztovari RAMÍREZ PÉREZ, RASSI & RAMÍREZ Simulium nuneztovari RAMÍREZ PÉREZ et al., 1977: 169; RAMÍREZ PÉREZ, 1983: 59. Simulium (Ectemnaspis) nuneztovari: COSCARÓN, 1984: 316; 1987: 29; 1990: 145. Simulium (Psilopelmia) nuneztovari: CROSSKEY & HOWARD, 1997: 58; 2004: 52.

(Information based on RAMÍREZ PÉREZ et al ., 1977 and RAMÍREZ PÉREZ, 1983). Female: Wing length 2.1 mm. Scutum orange with decumbent hair dark golden; no silvery border. Scutellum yellow-lemon with black hair. Metanotum and pleurae yellowish with nacreous or silvery pruinosity. Legs yellowish with tibiae and tarsomeres dark brown. Abdomen yellowish lemon. Fronto-ocular triangle wider than high. Cibarium with 1+1 short blunt submedian projections, covered with some small teeth, as is rim from

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submedian process to cornuae, similar to that of S. antillarum. Paraproct about 2 times longer than wide at base (as in S. bicoloratum). Male: Wing length 2.0 mm. Coloration as in female, abdomen with inferior border of VII-VIII brownish. Gonostylus subcylindrical, shorter than gonocoxite; ventral plate with small lateral constrictions as in S. dinellii. Pupa: Cocoon with weave relatively open, especially on sides; length along base 3.0 mm, on dorsum 2.3 mm. Frontoclypeus and thorax with numerous tubercles; trichomes with 3-8 branches. Gill with 8 branches, with primary and very short secondary branches Larva: Unknown. Distribution: Venezuela: Amazonas. Bionomics: Unknown. Discussion: The closest species are S. dinellii and S. antillarum. The first can be differentiated by the cibarial submedian process, which is thinner and more elongated, and the abdomen with greenish brown to yellowish brown on segments VI-IX. Simulium antillarum has a similar cibarium but has 1+1 silvery submedian stripes on the scutum. Probably by examining more material, it can be established that this species is a variant of the aforementioned species. Species inquirenda Simulium varians LUTZ Simulium (Eusimulium) varians LUTZ, 1909: 131. Simulium varians: LUTZ, 1910: 234; PINTO, 1931: 672; COSCARÓN, 1984: 322; 1990: 148.

Discussion: Simulium varians was described based on 3 females characterized by being dark brown with orange spots from Brazil: São Paulo, Rio Grande, Franca. This material was not found in Instituto Oswaldo Cruz. During a search in the type area, material with this coloration was not collected. It will be necessary to repeat the search for additional material. At present, S. varians is considered species inquirenda. Simulium (Psilopelmia) ENDERLEIN (Figs. 91-100, 149.) Type-species: Psilopelmia rufidorsum ENDERLEIN, 1934b = Simulium escomeli ROUBAUD, 1909. Lanea VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA, 1946: 107. (Type-species Simulium haematopotum: MALLOCH, 1914).

Female: Medium-sized species, wing length 1.5-3.1 mm; general coloration from yellowish brown to grayish brown; scutum yellowish to brown blackish, generally with 1+1 silvery to gray bluish submedian longitudinal vittae or 1 median brown reddish; scutellum from yellowish to brown yellowish, metanotum from yellowish brown to grayish brown; abdominal dorsum generally with median, 1+1 submedian and frequently 1+1 lateral subquadrate darker rows of spots; fronto-ocular triangle generally higher than wide; sensory palpal vesicle less than half palpomere length. Basal portion of cibarium with 1+1 short blunt submedian elevations, covered by short denticles or spiculae, flanking median depressed area, frequently well sclerotized

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halfmoon shaped; basal section of R generally unhaired. Sc with or without hair; claws generally without or with very short subbasal tooth; paraprocts with basal lobe and distally generally projected as anterior thin process; gonapophysis slightly divergent inner sclerotized margin, without hair or with sparse microtrichiae; genital fork with sclerotized stem and well-developed anterior branches. Male: Wing 1.9-2.7 mm; coloration as in female but slightly darker; scutum frequently with 1+1 silvery cuneiform stripes reduced to anterior half anterior or reaching light posterior area; abdomen with variants yellowish, gray to blackish; posterior basitarsus ratio length/width from 3.8-7.8; gonostylus shorter than gonocoxite, subtriangular to subtrapezoidal, frequently wide distally, projected on internal side, and with apical spur; ventral plate in ventral position wider than high, laterally with or without slight concavity; endoparameres with strong hooks. Pupa: Cocoon resistant, weaves with threads evident; slipper shaped, generally with anterior reinforced border and mild anterior basal projection; gill branches generally in number of 8 but can be 4-6 or 20, all forward directed; frontoclypeus and thoracic trichomes mostly single or bifid; abdomen terminal spur small. Larva: Maximum length 6.0 mm; integument without trichomes, spines or scales; cephalic apotome scarcely ornamented, sometimes slightly darkened as median basal bell shape with 1 median and 1+1 lateral subbasal light windows; antenna generally with medial and distal segments similar to or longer than proximal; postgenal cleft moderately deep; postgenal bridge length about as long as or longer than hypostomium height; anal sclerite with 63-105 rows of 10-17 hooks each; diverticula of rectal papillae generally 0-4 but can reach 9-12. Simulium (Psilopelmia) is a relatively homogeneous taxa composed of 24 species. The Neotropical species and one inquirenda are subdivided into two species groups: ESCOMELI with 21 species and BLANCASI with three species. Simulium panamense, S. callidum, and S. pseudocallidum have some unique characters and resemble S. (Ectemnaspis) species. The subgenus Psilopelmia has different arrangements according to CROSSKEY & HOWARD (1997, 2004), including five species groups: DINELLII, PERFLAVUM, ROMANAI, ESCOMELI and QUADRIVITTATUM. The first three are considered to belong to S. (Ectemnaspis) by COSCARÓN (1987: 19); both taxa have a close relationships that some day might lead to including all of the groups in the same subgenus. We maintain S. (Escomeli) as a separate species group from S. (Psilopelmia), being reinforced by the results of cladistic studies that separate them. To S.(Psilopelmia) the S. (Blancasi) species group is also added. KEYS

TO

SIMULIUM (PSILOPELMIA)

SPECIES

Females 1 – 2(1)

Scutum uniformly dull gray, without stripes .................................................................. ......................................................................................... BLANCASI species group......2 Scutum with 1 median or 1+1 longitudinal stripes ...................................................... ......................................................................................... ESCOMELI species group......4 Abdominal tergites III-V velvety black, their center anteriorly (Fig. 98 A) with or without transverse dumbbell-shaped light-colored spots; setae of scutum sparse, not conspicuous under low magnification ......................................... Simulium blancasi

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Abdominal tergites III-V brown or piceous, always without light-colored spots anteriorly (Fig. 99 A); setae of scutum very abundant, conspicuous under low magnification ....................................................................................................................... 3 3(2) Abdominal tergites III-V entirely dark velvety brown (Fig. 99 A); calcipala comparatively large, broadly rounded apically (Fig. 99 D) ............ Simulium tenuipes – Abdominal tergites III-V dark anteriorly, distinctly lighter posteriorly, entire posterior half light colored on V; calcipala much reduced, subtriangular (Fig. 100 A) ......................................................................................................................... Simulium penai 4(1) Scutum grayish with 1 median longitudinal reddish brown band (Fig. 91 A) or unicolorous; cibarium with well-defined small teeth on basal portion (Fig. 91 J) ...... 5 – Scutum with 1+1 light submedian bands generally from anterior to posterior border (Figs. 91 B-E); cibarium with teeth, spiculate or smooth on basal portion (Figs. 91 L-Q) ...................................................................................................................... 7 5(4) Metanotum blackish; wing length 1.9-2.4 mm ........................ Simulium mediovittatum – Metanotum grayish brown; wing length 1.8-2.2 mm .................................................. 6 6(5) Wing length 2.0-2.2 mm; scutellum grayish brown; abdominal tergite II grayish (Fig. 91 G) .................................................................................................... Simulium dugesi – Wing length 1.8 mm; scutellum yellow to yellowish brown; abdominal tergite II grayish yellow .............................................................................................. Simulium ochoai 7(4) Abdominal tergites without rows of darker subquadrate spots; distal portion of paraproct elongated, narrowing slightly (Figs. 92 F, G); claws with well developed subbasal tooth (Figs. 91 V, W) ......................................................................................... 8 – Abdominal tergites with 1 median, 1+1 submedian and sometimes 1+1 sublateral rows of darker subquadrate spots at least from segments III to VI (Figs. 91 H, I); distal portion of paraproct relatively short (Figs. 92 B-E, J) or elongated, narrowing abruptly and projected as narrow appendix (Figs. 92 H-N); claws without subbasal tooth (Figs. 91 T-U) ......................................................................... 10 8(7) Cibarium with small median depression and abundant denticles submedially (Fig. 91 N); paraproct distally elongated, as long as basal width and acuminate distally (Fig. 92 F) ............................................................................................. Simulium panamense – Cibarium with thick rim, with 1+1 slight sublateral protuberances sometimes covered by thin spiculae (Fig. 91 O); distal process of paraproct absent and blunt apically (Fig. 92 G) ............................................................................................................. 9 9(8) Basal sector of R with hair .................................................................. Simulium callidum – Basal sector of R glabrous ......................................................... Simulium pseudocallidum 10(7) Abdominal tergites with 1 median and 1+1 submedian rows of subquadrate darker spots (Fig. 91 H); paraproct distal process shorter than half of basal width (Figs. 92 B, E, J) ................................................................................................................ 11 – Abdominal tergites with 1 median, 1+1 submedian and 1+1 sublateral rows of subquadrate darker spots (Fig. 91 I); distal process of paraproct length about equal to width at base (Figs. 92 C, D) or very elongated (Figs. 92 H, I, K-N) ................ 13 11(10) Scutellum grayish brown; sensory vesicle of maxillary palp shorter than half length of palpomere (Fig. 91 S); paraproct blunt apically (Fig. 92 E) ...................... ........................................................................................................... Simulium haematopotum

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– 12(11) – 13(10) – 14(13) – 15(13) – 16(15) – 17(16) – 18(17) –

Scutellum yellowish brown; sensory vesicle of maxillary palp generally longer than half of length of palpomere; paraproct slightly acuminate apically (Figs. 92 B-J) ...................................................................................................................................... 12 Wing length 2.1-2.7 mm .................................................................... Simulium bivittatum Wing length 2.4-3.0 mm .................................................................... Simulium dandrettai Paraproct apically blunt and distal process about as long as half of basal width (Fig. 92 D); scutum yellowish orange with very thin 1+1 submedian silvery stripe (Fig. 91 B) .......................................................................................................................... 14 Paraproct apically acuminate and distal process as long as basal width (Figs. 92 H, I, K-N); scutum yellowish gray with brown to blackish medial sublateral stripe (Figs. 91 E, F) (except Simulium downsi) ....................................................................... 15 Wing length 1.5-1.6 mm; sensory vesicle of maxillary palp about as long as half of palpomere (Fig. 91 R) ................................................................ Simulium iracouboense Wing length 1.8-2.1 mm; sensory vesicle of maxillary palp shorter than half of palpomere ..................................................................................................Simulium samboli Basal sector of R with hairs; scutum yellowish with light yellowish brown median and sublateral stripes ................................................................................ Simulium downsi Basal sector of R glabrous; scutum yellowish gray with dark brown to black median and sublateral stripes ........................................................................................ 16 Scutum generally yellowish with light brown yellowish to grayish yellow stripes, or with dark brown to blackish on median and sublateral stripes (Fig. 91 E) ............. ..................................................................................................................... Simulium escomeli Scutum grayish yellow with dark brown to blackish on median and sublateral stripes (Fig. 91 F) ............................................................................................................. 17 Distal process of paraproct shorter than half width of base (similar to Simulium dandretta ) (Fig. 92 J) ..........................................................................Simulium zempoalense Distal process of paraproct length, as wide as width of base (Figs. 92 K-N) .............................................................................................................................................. 18 Scutum with median black vittae wider than submedian silvery vittae; sublateral black vittae not reaching anterior rim (Fig. 91 F) ..................... Simulium mangabeirai Scutum with median black vittae about same width as submedian silvery vittae; sublateral black vittae reaching or very close to anterior rim ..................................... ........................... Simulium longithallum, Simulium gonzalezherrejoni, Simulium trivittatum, Simulium bobpetersoni, Simulium veracruzanum

Males 1 – 2(1)

Scutum uniformly dull gray without silvery wedges .................................................... ......................................................................................... BLANCASI species group......2 Scutum with 1+1 silvery wedges (Figs. 93 A-G) ........................................................... ......................................................................................... ESCOMELI species group......4 Abdominal tergites III-V velvety black, their anterior border with distinct lightcolored spot at center (Fig. 98 P); setae of scutum sparse, not conspicuous under low magnification; genitalia with ventral plate constricted laterally near base (Fig. 98 W) ......................................................................................................... Simulium blancasi

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Abdominal tergites III-V dark, sometimes with lighter regions posteriorly but not on anterior border; setae of scutum very abundant, conspicuous under low magnification; genitalia with ventral plate not constricted laterally near base (Fig. 99 J) ....................................................................................................................................... 3 3(2) Length of hind basitarsus more than 4 times its width; calcipala much reduced, subtriangular (Fig. 100 A) .......................................................................... Simulium penai – Length of hind basitarsus less than 4 times its width; calcipala not as reduced, broadly rounded apically (as in female, Fig. 99 D) .......................... Simulium tenuipes 4(1) Scutum and abdomen yellowish, abdomen can be grayish brown but at least with some yellowish orange portion; scutum with submedian silvery wedges only on anterior area (Fig. 93 B) .................................................................................................... 5 – Scutum and abdomen gray to blackish; scutum with silvery wedges with variable extension (Figs. 93 A, C-G) ........................................................................................... 10 5(4) Body of ventral plate subovoidal (Fig. 94 F) ............................................................... 6 – Body of ventral plate of variable shape but not subovoidal .................................... 7 6(5) Basal sector of R with hair .................................................................. Simulium callidum – Basal sector of R glabrous ......................................................... Simulium pseudocallidum 7(5) Ventral plate with strong lateral constrictions on 1/2 of body, continued by narrow subovoidal mediodistal portion (Fig. 94 E) ..................... Simulium panamense – Ventral plate variable, but without strong lateral constrictions on body (Figs. 94 C, H) ................................................................................................................................................ 8 8(7) Gonostylus subtrapezoidal with strong expansion on subterminal external side (Fig. 93 N); ventral plate of body reduced, subtriangular (Fig. 94 H) and narrower than position of branches ....................................................................... Simulium downsi – Gonostylus without lateral or subterminal expansion (Figs. 93 K, L); ventral plate body more developed, not subtriangular and wider than position of branches (Fig. 94 C) ..................................................................................................................................... 9 9(8) Gonostylus subtriangular (Fig. 93 J); ratio of length/width of posterior basitarsus = 7.8 ........................................................................................................... Simulium samboni – Gonostylus subtrapezoidal (Fig. 93 K); ratio of length/width of posterior basitarsus = 6.6 ................................................................................. Simulium iracouboense 10(4) Scutum with 1+1 short silvery stripes not reaching posterior declivity (Fig. 93 A) .............................................................................................................................................. 11 – Scutum with 1+1 long silvery stripes reaching posterior declivity (Fig. 93 G) ......... 19 11(10) Scutum with 1+1 submedian silvery blue areas triangular to oval shaped, contiguous anteriorly with same-colored lateral areas (Fig. 93 A) ........................ 12 – Scutum with 1+1 silvery blue areas not contiguous with same-colored lateral areas (Fig. 93 D) .......................................................................................................................... 18 12(11) Silvery submedian wedges strongly tapered posteriorly, (Fig. 93 A) surpassing or not half of scutum; gonostylus about as long as width of external straight side (Fig. 93 I); ventral plate subtriangular with rounded apex (Fig. 94 A) ................. 13 – Silvery submedian wedges generally surpassing half of scutum; gonostylus expanded over external distal subterminal side (Figs. 93 P, Q, S); ventral plate variable, from subtriangular with or without lateral constrictions to subovoidal ............................... 15

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13(12) Submedian wedges of scutum not reaching posterior declivity (Fig. 93 A) ....... 14 – Submedian wedges of scutum strongly tapered posteriorly and reaching posterior declivity ........................................................................................... Simulium mediovittatum 14(13) Wing length 1.9 mm .................................................................................. Simulium dugesi – Wing length 1.6 mm .................................................................................. Simulium ochoai 15(12) Gonostylus less than 2 times as long as wide at base, distally expanded, wider than at base (Fig. 93 O); ventral plate without lateral constrictions, with rounded borders (Fig. 94 I); ratio of length/width at base = 1.2 ............ Simulium dandrettai – Gonostylus generally over 2 times longer than width at base, not expanded distally; ventral plate usually without rounded borders; ratio of length/width over 1.2 ........................................................................................................................................ 16 16(15) Gonostylus subrectangular, with subparallel sides (Fig. 93 I); ventral plate relatively short without lateral constrictions (Fig. 93 B) ............. Simulium bivittatum – Gonostylus subtrapezoidal, with strong curvature distally and slightly acuminate distally (Fig. 93 N); ventral plate from short and wide (Fig. 94 J) to more elongated with lateral constrictions (Fig. 94 G) ........................................................................... 17 17(16) Ventral plate relatively short with base projected basally (Fig. 94 J), ratio of length/width = 2.3; wing length 1.9-2.1 mm .............................. Simulium bobpetersoni – Ventral plate relatively long, with base not projected basally and laterally with slight constrictions (Fig. 94 G); ratio of length/width = 1.3-1.4 wing length 2.03.0 mm ....................................................................................................... Simulium escomeli 18(11) Ventral plate subtriangular without lateral constrictions (Fig. 94 D); gonostylus strongly enlarged distally, with long curved spur (Fig. 93 L); wing length 1.8-2.1 mm ................................................................................................... Simulium haematopotum – Ventral plate with strong lateral constrictions and distal portion after constriction narrow, about 1/3 width of body (Fig. 94 K); gonostylus not very enlarged distally and with moderate-sized spur (Fig. 93 P); wing length 2.6-3.0 mm ......................... ............................................................................................................ Simulium veracruzanum 19(10) Gonostylus about 2/3 as wide as long (Fig. 93 R); ventral plate without lateral constrictions; distal border of body subcircular ........................ Simulium zempoalense – Gonostylus about 1/2 as long as wide (Fig. 93 Q); ventral plate with or without constrictions and different than above (Figs. 94 L-N) ............................................. 20 20(19) Hind basitarsus wide, ratio of length/width = 3.8; wing length 2.5-2.8 mm; scutum with 1+1 narrow light wedges not reaching posterior edge (Fig. 93 F) .... .............................................................................................................. Simulium mangabeirai – Hind basitarsus narrower with ratio of length/width >5.0; scutum with 1+1 wide light wedges reaching posterior edge (Fig. 93 G) ...................................................... 21 21(20) Abdominal segments I-II yellowish; wing length 2.5 mm .......... Simulium trivittatum – Abdominal segments I-II not yellowish; wing length 2.0-2.5 mm ........................ 22 22(20) 1+1 blackish lateral stripes of scutum not reaching postpronotal lobes (Fig. 93 G); wing length 2.5 mm .................................................................. Simulium gonzalezherrejoni – 1+1 blackish lateral stripes of scutum reaching postpronotal lobes; wing length 2.0 mm ................................................................................................ Simulium longithallum

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Pupae 1 – 2(1) – 3(2)

Gill with 20 branches (Fig. 96 D) .................................................... Simulium panamense Gill with 4-8 branches ....................................................................................................... 2 Gill with 4 branches (Fig. 96 M) ..................................................................................... 3 Gill with 6-8 branches ....................................................................................................... 6 Cocoon well developed covering most of thorax and head (Fig. 95 M); ventral anterior edge more advanced than dorsal ................................... Simulium mangabeirai – Cocoon scarcely developed, not covering most of thorax and head; dorsal anterior edge more advanced than ventral edge (Fig. 98 X) ...................................... ......................................................................................... BLANCASI species group......4 4(3) Gill with short trunk followed by 2 distinct secondary branches bearing 2 + 2 filaments (Fig. 98 Y) ............................................................................... Simulium blancasi – Gills with all 4 filaments seemingly arising from very short common trunk (Figs. 99 L, 100 C) ......................................................................................................................... 5 5(4) Frontoclypeus and exposed portion of thorax with abundant tubercles; surface of cocoon rough, its color brown when examined in liquid ............... Simulium tenuipes – Frontoclypeus and exposed portion of thorax with or without very sparse tubercles (Figs. 100 D, E); surface of cocoon smooth, its color brown with leaden sheen, parchment-like, when examined in liquid ................................. Simulium penai 6(2) Gill with 6 branches .......................................................................................................... 7 – Gill with 8 branches ........................................................................................................ 10 7(6) Gill with wrinkled surface; blunt apex, brown to blackish pigmentation (Figs. 96 I, J); gill branches shorter than cocoon length (Figs. 95 I, J) ....................................... 8 – Gill with smooth surface, generally acuminate apex; yellowish pigmentation; gill branches as long as or longer than cocoon length (Fig. 95 N) ................................ 9 8(7) Gill secondary dorsal outer and inner branches thicker than other branches (Fig. 96 I); frontoclypeal and thoracic tubercles smooth; cocoon without longitudinal ridges (Fig. 95 I) .................................................................................. Simulium dandrettai – Gill with only dorsal outer branch thicker than other branches (Fig. 96 J); frontoclypeal and thoracic tubercles wrinkled; cocoon with several longitudinal ridges (Fig. 95 J) ................................................................... Simulium sp. near dandrettai 9(7) Gill with branches off of primary and secondary branches very close to base (Fig. 96 Q); dorsal thoracic integument with sparse tubercles but with small rugosities simulating tubercles (Fig. 95 O) .....................................................Simulium zempoalense – Gill with branches off of primary ventral and secondary branches not close to base (Fig. 96 P); dorsal thoracic integument smooth with sparse tubercles ........... ................................................................................................................. Simulium trivittatum 10(6) Gill with basal stem short, with primary and secondary branches emerging very close to base, dorsal primary branch thicker than others (Figs. 96 E, F) ............ 11 – Gill with primary and secondary branches emerging from basal stem more distant from base, and with primary branches about same diameter ................................. 12 11(10) Gill tertiary dorsal branch distinctly thicker and 1/3 longer than other branches (Fig. 96 F) ...................................................................................... Simulium pseudocallidum

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Gill tertiary dorsal branch only slightly thicker and with similar length to other branches (Fig. 96 E) ............................................................................... Simulium callidum Gill dorsal secondary and tertiary branches thicker than other branches (Figs. 96 N, O) ................................................................................................................................... 13 Gill dorsal secondary and tertiary branches with diameter similar to that of other branches ............................................................................................................................. 14 Gill dorsal secondary and tertiary branches arising far from base (Fig. 96 N) ...... .......................................................................................................Simulium gonzalezherrejoni Gill dorsal secondary and tertiary branches arising near base (Fig. 96 O) .............. .............................................................................................................. Simulium longithallum Gill length as long as or longer than cocoon length (Fig. 95 L) ................................ ............................................................................................................ Simulium veracruzanum Gill length shorter than cocoon length (Figs. 95 A-G) ........................................... 15 Cocoon weave sparse, with very open weave, without anterior rim (Fig. 95 F) ..... .............................................................................................................. Simulium iracouboense Cocoon weave compact, with threads very close, and with anterior rim (Figs. 95 A-E, G, H, K) ................................................................................................................... 16 Cephalic and thoracic trichomes longer than half of frontoclypeal basal width (Fig. 95 R); cocoon anterior rim reinforced and with anterodorsal projection (Fig. 95 E) .......................................................................................................... Simulium samboni Cephalic and thoracic trichomes not longer than half of frontoclypeal basal width (Figs. 95 Q, T, U); cocoon anterior rim usually not reinforced and without anterodorsal projection ................................................................................................... 17 Cocoon with small dorsal and ventral projection (Fig. 95 G); frontoclypeus and thorax with sparse tubercles (Fig. 95 T) ................................... Simulium haematopotum Cocoon without dorsal projection (Figs. 95 A, K); frontoclypeus and thorax with abundant to sparse tubercles ......................................................................................... 18 Gill branches relatively closed basally, subparallel (Figs. 96 A, K) ........................ 19 Gill branches generally very open basally, and with tendency to converge distally (Figs. 96 B, C, G, H) ........................................................................................................ 20 Secondary dorsal branch of gill thicker than secondary ventral branch; tertiary dorsal branch bifurcated at long distance from base (Fig. 96 K); frontoclypeus and thorax with moderate number of tubercles (Fig. 95 U) ........... Simulium bobpetersoni Secondary dorsal branch of gill with diameter similar to that of secondary ventral branch; tertiary dorsal branch bifurcated at short distance from base (Fig. 96 A); frontoclypeus and thorax with abundant tubercles (Fig. 95 P) ........ Simulium dugesi Median tertiary gill branch bifurcated at short distance from base (Fig. 96 B); frontoclypeus and thorax with sparse tubercles not elevated (Fig. 95 Q) ............... ........................................................................................................................ Simulium ochoai Median tertiary gill branch bifurcated at long distance from base (Figs. 96 C, G, H); frontoclypeus and thorax with abundant to sparse tubercles, in latter case, tubercles elevated as high as wide at base .................................................................. 21 Dorsal tertiary gill branch bifurcated at same distance as or farther than median tertiary branch (Fig. 96 H) ...................................................................... Simulium downsi

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Dorsal tertiary gill branch bifurcated lower than median tertiary branch (Figs. 96 C, G) .............................................................................................................................................. 22 22(21) Frontoclypeus and thorax with abundant tubercles (Fig. 95 S); gill length 2.2-2.5 mm; cocoon length 3.0-3.5 mm ........................................................... Simulium escomeli – Frontoclypeus and thorax with moderate to few tubercles; gill length 1.5-2.1 mm; cocoon length 2.4-2.7 mm ............................................................................................. 23 23(22) Proportionally short gill branches 1.5-1.6 mm (Fig. 95 C): ratio of gill/cocoon length = 1.7 .................................................................................... Simulium mediovittatum – Proportionally long gill branches 2.0-2.1 mm (Fig. 95 D): ratio of gill/cocoon length = 1.2 .......................................................................................... Simulium bivittatum Larvae 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(2) – 7(6) – 8(7) –

Postgenal cleft not developed (Figs. 99 M, 100 G) ........ Simulium tenuipes, Simulium penai Postgenal cleft developed to different depths ............................................................. 2 Postgenal cleft slightly developed (Figs. 97 L, N); ratio of postgenal bridge/ hypostomium = 1.2-1.7 .................................................................................................... 3 Postgenal cleft more developed (Figs. 97 J, K, M); ratio of postgenal bridge/ hypostomium = 0.5-1.1 .................................................................................................... 6 Ratio of postgenal bridge/hypostomium = 1.7; body length of mature larvae 6.8 mm; cephalic apotome with basal half darkened, with light window medially (Fig. 97 G) ................................................................................................... Simulium mangabeirai Ratio of postgenal bridge/hypostomium = 1.2-1.4; body length of mature larvae 4.7-6.3 mm; cephalic apotome basally scarcely darkened and without light window medially (Fig. 97 H) ........................................................................................................... 4 Ratio of postgenal bridge/hypostomium = 1.4; anal ring with about 78 rows of hooks ....................................................................................................Simulium zempoalense Ratio of postgenal bridge/hypostomium = 1.2; anal ring with 66-70 rows of hooks ..................................................................................................................................... 5 Ratio of proximal, medial and distal antennal segments = 1:1.3:1.5; anal ring with about 66 rows of hooks .................................................................... Simulium dandrettai Ratio of proximal, medial and distal antennal segments = 1:1.5:2.0; anal ring with about 70 rows of hooks ................................................................. Simulium longithallum Hypostomium shorter than postgenal bridge (Fig 98 g); ratio of postgenal bridge/ hypostomium length = 1.3 .................................................................... Simulium blancasi Hypostomium as long as or longer than postgenal bridge; ratio of postgenal bridge/hypostomium length = 0.5-1.1 .......................................................................... 7 Hypostomium longer than postgenal bridge (Figs. 97 J, K); ratio of postgenal bridge/hypostomium 0.5-0.9 ........................................................................................... 8 Hypostomium as long as or shorter than hypostomial bridge (Fig 97 M), ratio of postgenal bridge/hypostomium 1.0-1.1 ...................................................................... 14 Ratio of postgenal bridge/hypostomium = 0.5-0.6; cephalic apotome without ornamentation ................................................................. Simulium dugesi, Simulium ochoai Ratio of postgenal bridge/hypostomium = 0.7-0.9; cephalic apotome with basal half darkened and enclosing elongated median light window (Figs. 97 B, C, E) ................ 9

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9(8) – 10(9) – 11(10) – 12(11) – 13(12) – 14(7) – 15(14) – 16(14) – 17(16) – 18(17) –

CLAVES

Body length 5.5-6.0 mm; antenna about as long as stem of cephalic fan; ratio of postgenal bridge/hypostomium = 0.7 ................................................ Simulium escomeli Body length 4.3-5.4 mm; antenna longer than stem of cephalic fan; ratio of postgenal bridge/hypostomium = 0.8-0.9 .................................................................. 10 Anal ring with 89 rows of hooks; distal antennal segment as long as medial segment (Fig. 97 O) ............................................................................ Simulium panamense Anal ring with 63-77 rows of hooks; distal antennal segment longer than medial segment (Fig. 97 P) .......................................................................................................... 11 Cephalic fan with 52-5 rays; body length 5.0-5.5 mm .......... Simulium mediovittatum Cephalic fan with 31-48 rays; body length 4.1-5.0 mm ........................................... 12 Cephalic fan with 31-35 rays; anal ring with 74-91 rows ........ Simulium haematopotum Cephalic fan with 38-48 rays; anal ring with 63-74 rows ........................................ 13 Anal sclerite with hairs and scales; anal ring with 65-74 rows; rectal papillae with 1 diverticulum on each lobe ....................................................... Simulium gonzalezherrejoni Anal sclerite without hairs and scales (Fig. 97 Q); anal ring with 63 rows of hooks; rectal papillae with 2-3 diverticula on each lobe (Fig. 97 S) ......... Simulium iracouboense Anal ring with 100-115 rows with 14-20 hooks each; anal sclerite with abundant hairs and scales (Fig. 97 R); cephalic apotome without ornamentation (Fig. 97 D) .............................................................................................................................................. 15 Anal ring with 60-90 rows with 11-15 hooks each; anal sclerite usually without scales; cephalic apotome with or without ornamentation ....................................... 16 Cephalic fan with 30-32 rays; lateral sclerite of thoracic proleg with about 50 teeth; maximum length 5.5-6.0 mm ................................................... Simulium callidum Cephalic fan with 40-42 rays; lateral sclerite of thoracic proleg with about 37 teeth; maximum length 4.8-5.0 mm ......................................... Simulium pseudocallidum Anal ring with 60-62 rows with 11 hooks each .......................... Simulium bobpetersoni Anal ring with 68-90 rows with 13-15 hooks each ................................................... 17 Body length 5.5-6.0 mm; anal sclerite with abundant scales ....... Simulium veracruzanum Body length 3.9-5.5 mm; anal sclerite without scales ............................................... 18 Cephalic apotome darkened subbasally with median longitudinal elongated and 1+1 transverse subbasal light windows (Figs. 97 A, F); body length 5.1-5.4 mm ................................................... Simulium downsi, Simulium samboni, Simulium trivittatum Cephalic apotome scarcely pigmented, only slightly darkened basally; body length 3.7-4.5 mm ............................................................................................ Simulium bivittatum DE ESPECIES DE

SIMULIUM (PSILOPELMIA)

Hembras 1 – 2(1)

Escudo uniformemente gris opaco ................................ grupo BLANCASI sp.......2 Escudo con una banda mediana o 1+1 bandas longitudinales .................................. ................................................................................................ grupo ESCOMELI sp.......4 Tergitos abdominales III-V negro aterciopelado con o sin manchas claras centrales anteriores transversas (Fig. 98 A); cerdas del escudo esparcidas, no conspicuas con poco aumento ............................................................. Simulium blancasi

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Tergitos abdominales III-V castaño o piceo, sin manchas claras centrales anteriores transversas (Fig. 99 A); cerdas del escudo muy abundantes, conspicuas con bajo aumento ............................................................................................................... 3 Tergitos abdominales III-V totalmente castaño oscuro aterciopelados (Fig. 99 A); calcipala relativamente grande, redondeada apicalmente (Fig. 99 D) ....................... ..................................................................................................................... Simulium tenuipes Tergitos abdominlaes III-V oscuros anteriormente, notoriamente más claros posteriormente, ocupando la mitad posterior en el V; calcipala muy reducida, subtriangular (Fig. 100 A) .......................................................................... Simulium penai Escudo grisáceo con una banda mediana longitudinal castaño rojizo (Fig. 91 A) o unicolor; cibario con dientes pequeños bien definidos en porción basal (Fig. 91 J) ................................................................................................................................................ 5 Escudo con 1+1 bandas longitudinales submedianas generalmente cruzando desde el borde anterior al posterior (Figs. 91 B-E); cibario con dientes, espículas, o liso en la porción basal (Figs. 91 L-Q) .......................................................................... 7 Metanoto negruzco; longitud ala 1,9-2,4 mm ......................... Simulium mediovittatum Metanoto castaño grisáceo; longitud ala 1,8-2,2 mm ................................................. 6 Longitud alar 2,0-2,2 mm; escutelo castaño grisáceo; tergito abdominal II grisáceo (Fig. 91 G) .................................................................................................... Simulium dugesi Longitud ala 1,8 mm; escutelo amarillo a castaño amarillento; tergito abdominal II amarillo grisáceo ......................................................................................... Simulium ochoai Tergitos abdominales sin hileras de manchas oscuras subcuadradas; porción distal del paraprocto alargado, afindandose suavemente (Figs. 92 F, G); uñas con diente subbasal bien desarrollado (Figs. 91 V, W) ................................................................... 8 Tergitos abdominales con 1 hilera mediana de manchas oscuras subcuadradas, 1+1 submedianas y a veces 1+1 sublateral por lo menos en los segmentos III-VI (Figs. 91 H, I); porción distal del paraprocto relativamente corto (Figs. 92 B, E, J) o alargado, afinandose abruptamente y prolongado en un angosto apéndice (Figs. 92 H-N); uñas sin diente subbasal (Figs. 91 T-U) ..................................................... 10 Cibario con una pequeña depresión mediana y abundantes pequeños dientes submedianamente (Fig. 91 N); paraprocto distalmente alargado, tan largo como el ancho basal y acuminado distalmente (Fig. 92 F) ........................ Simulium panamense Cibario con un grueso reborde, con 1+1 suaves protuberancias a veces cubiertas por finas espículas (Fig. 91 O); prolongación distal del paraprocto ausente y con ápice romo (Fig. 92 G) ...................................................................................................... 9 Sector basal de R con pelos ................................................................. Simulium callidum Sector basal de R glabro ............................................................. Simulium pseudocallidum Tergitos abdominales con 1 hilera mediana y 1+1 submediana de manchas oscuras subcuadradas (Fig. 91 H); prolongación distal del paraprocto más corto que la mitad del ancho en la base (Figs. 92 B, E, J) ................................................. 11 Tergitos abdominales con 1 hilera mediana y 1+1 sublateral de manchas oscuras subcuadradas (Fig. 91 I); prolongación distal del paraprocto con un largo aproximado al ancho en la base (Figs. 92 C, D) o muy alargado (Figs. 92 H, I, K-N) .............................................................................................................................................. 13

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11(10) Escutelo castaño grisáceo; vesícula sensorial del palpo maxilar más corta que el palpómero (Fig. 91 S); paraprocto con ápice romo (Fig. 92 E) ................................. ........................................................................................................... Simulium haematopotum – Escutelo castaño amarillento; vesícula sensorial del palpo maxilar generalmente más larga que la mitad del largo del palpómero; paraprocto suavemente aguzado (Figs. 92 B-J) ..................................................................................................................... 12 12(11) Longitud ala 2,1-2,7 mm ................................................................... Simulium bivittatum – Longitud ala 2,4-3,0 mm ................................................................... Simulium dandrettai 13(10) Paraprocto romo apicalmente y prolongación distal aproximadamente tan larga como la mitad del ancho en la base (Fig. 92 D); escudo amarillento-anaranjado con 1+1 banda submediana plateada muy fina (Fig. 91 B) ..................................... 14 – Paraprocto aguzado apicalmente y prolongación distal tan larga como el ancho en la base (Figs. 92 H, I, K-N); escudo gris amarillento con banda mediana y sublaterales castaño a negro (Figs. 91 E, F) (excepto Simulium downsi) ................ 15 14(13) Longitud ala 1,5-1,6 mm; vesícula sensorial del palpo maxilar aproximadamente tan larga como la mitad de la longitud del palpómero (Fig. 91 R) ........................... .............................................................................................................. Simulium iracouboense – Longitud ala 1,8-2,1 mm; vesícula sensorial del palpo maxilar más corta que la mitad de la longitud del palpómero .................................................... Simulium samboni 15(13) Sector basal de R con pelos; escudo amarillento con bandas mediana y sublaterales castaño amarillento claras ................................................. Simulium downsi – Sector basal de R sin pelos; escudo gris amarillento con bandas medianas y sublaterales castaño oscuro a negro ............................................................................. 16 16(15) Escudo generalmente amarillento con bandas medianas y sublaterales castaño claro a amarillo grisáceo o bien de castaño oscuro a negruzco sobre bandas mediana y sublaterales (Fig. 91 E) ....................................................... Simulium escomeli – Escudo amarillo grisáceo con bandas medianas y sublaterales castaño oscuro a negruzco (Fig. 91 F) ........................................................................................................ 17 17(16) Prolongación distal del paraprocto más corta que la mitad del ancho en la base (similar a Simulium dandretta) (Fig. 91 J) ........................................ Simulium zempoalense – Prolongación distal del paraprocto aproximadamente tan larga como el ancho en la base (Figs. 92 K-N) ..................................................................................................... 18 18(17) Banda mediana del escudo negra más ancha que las bandas plateadas submedianas, y bandas negras sublaterales no alcanzan el borde anterior (Fig. 91 F) .............................................................................................................. Simulium mangabeirai – Banda mediana negra del escudo aproximadamente del mismo ancho que las bandas plateadas submedianas, y las bandas negras sublaterales alcanzan o llegan muy cerca del borde anterior ....................................... Simulium longithallum, Simulium gonzalezherrejoni, Simulium trivittatum, Simulium bobpetersoni, Simulium veracruzanum Machos 1 –

Escudo uniformemente gris opaco sin cuñas plateadas .............................................. ................................................................................................ grupo BLANCASI sp.......2 Escudo con 1+1 cuñas plateadas (Figs. 93 A-G) ........ grupo ESCOMELI sp.......4

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Tergitos abdominales III-V negro aterciopelado, con borde anterior en el centro con una mancha clara (Fig. 98 P); setas de escudo esparcidas y no conspicuas con bajo aumento; genitalia con placa ventral contraida lateralmente cerca de la base (Fig. 98 W) ................................................................................................ Simulium blancasi – Tergitos abdominales III-V oscuros, a veces con áreas posteriores claras pero nunca sobre el borde anterior; setas del escudo muy abundantes, conspícuas con bajo aumento; genitalia con placa ventral no contraida lateralmente cerca de la base (Fig. 99 J) .................................................................................................................... 3 3(2) Basitarso posterior 4 veces más largo que ancho; calcipala muy reducida subtriangular (Fig. 100 A) .......................................................................... Simulium penai – Basitarso posterior menos de 4 veces largo que ancho; calcipala no tan reducida y redondeada apicalmente (como en la hembra, Fig. 99 D) ............. Simulium tenuipes 4(1) Escudo y abdomen amarillento pudiendo ser castaño grisáceo, pero por lo menos con alguna porción amarillo-anaranjado; escudo con cuñas plateadas solo en área anterior (Fig. 93 B) ............................................................................................................. 5 – Escudo y abdomen gris a negruzco; escudo con uñas plateadas de variable extensión (Figs. 93 A, C-G) ........................................................................................... 10 5(4) Cuerpo de la placa ventral subovoidal (Fig. 94 F) ...................................................... 6 – Cuerpo de la placa ventral de forma variable, no subovoidal .................................. 7 6(5) Sector basal de R con pelos ................................................................. Simulium callidum – Sector basal de R sin pelos ......................................................... Simulium pseudocallidum 7(5) Placa ventral con fuerte contracción lateral sobre la mitad del cuerpo, continuada por una porción angosta medio distal (Fig. 94 E) ....................... Simulium panamense – Placa ventral variable pero sin fuerte contracción lateral en el cuerpo (Figs. 94 C, H) ........................................................................................................................................... 8 8(7) Gonostilo subtrapezoidal con fuerte expansión sobre el lado externo subterminal (Fig. 93 N); placa ventral del cuerpo reducida,subtriangular (Fig. 94 H) y más angosta que la posición de las ramas .................................................... Simulium downsi – Gonostilo sin expansión lateral o subterminal (Figs. 93 K, L); cuerpo de la placa ventral más desarrollado, no subtrioangular y más ancho que la posición de las ramas (Fig. 94 C) ................................................................................................................ 9 9(8) Gonostilo subtriangular (Fig. 93 J); relación del largo/ancho del basitarso posterior = 7,8 ......................................................................................... Simulium samboni – Gonostilo subtrapezoidal (Fig. 93 K); relación del largo/ancho del basitarso posterior = 6,6 .................................................................................. Simulium iracouboense 10(4) Escudo con 1+1 banda corta plateada no llegando al área posterior grisácea (Fig. 93 A) ................................................................................................................................... 11 – Escudo con 1+1 banda larga plateada arribando al área posterior grisácea (Fig. 93 G) ................................................................................................................................... 19 11(10) Escudo con 1+1 áreas azul plateado triangulares u ovales, contiguas anteriormente con áreas laterales del mismo color (Fig. 93 A) ............................. 12 – Escudo con 1+1 áreas azul plateado no contiguas con áreas laterales del mismo color (Fig. 93 D) ............................................................................................................... 18

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12(11) Cuñas plateadas submedianas fuertemente afinadas posteriormente (Fig. 93 A), sobrepasando o no la mitad del escudo; gonostilo aproximadamente tan largo como ancho con el lado externo recto (Fig. 93 I); placa ventral subtriangular con ápice redondeado (Fig. 94 A) ........................................................................................ 13 – Cuñas plateadas submedianas generalmente sobrepasando la mitad del escudo; gonostilo expandido sobre el lado externo distal subterminal (Figs. 93 P, Q, S); placa ventral variable desde subtriangular, con o sin contracciones laterales, a subovoidal .......................................................................................................................... 15 13(12) Cuñas submedianas del escudo no llegando al área grisácea posterior (Fig. 93 A) .............................................................................................................................................. 14 – Cuñas submedianas del escudo bien afinadas posteriormente y llegando al área grisácea posterior .......................................................................... Simulium mediovittatum 14(13) Longitud del ala 1,9 mm ........................................................................... Simulium dugesi – Longitud del ala 1,6 mm ........................................................................... Simulium ochoai 15(12) Gonostilo menos de 2 veces más larga que ancho en la base, expandido distalmente, más ancho que en la base (Fig. 93 O); placa ventral sin contracciones laterales, con bordes redondeados (Fig. 94 I); relación largo/ancho en la base = 1,2 ..................... ................................................................................................................. Simulium dandrettai – Gonostilo generalmente más de 2 veces largo que ancho en la base, no expandido distalmente; placa ventral generalmente sin bordes redondeados; relación largo/ ancho en la base sobre 1,2 ............................................................................................. 16 16(15) Gonostilo subrectangular con los lados subparalelos (Fig. 93 I); placa ventral relativamente corta, sin contracciones laterales (Fig. 93 B) ....... Simulium bivittatum – Gonostilo subtrapezoidal, distalmente con fuerte curvatura y ápice suavemente aguzado (Fig. 93 N); placa ventral relativamente corta y ancha (Fig. 94 J) hasta más alargada con contracción lateral (Fig. 94 G) .............................................................. 17 17(16) Placa ventral relativamente corta con base prolongada basalmente (Fig. 94 J); relación largo/ancho = 2,3; longitud ala 1,9-2,1 mm ............... Simulium bobpetersoni – Placa ventral relativamente larga, con base no prolongada basalmente y lateralmente con suave contracción (Fig. 94 G); relación largo/ancho = 1,3-1,4; longitud ala 2,0-2,3 mm ......................................................................... Simulium escomeli 18(11) Placa ventral subtriangular sin contracción lateral (Fig. 94 D); gonostilo bien alargado distalmente con un largo y curvo espolón (Fig. 93 L); longitud del ala 1,82,1 mm ............................................................................................. Simulium haematopotum – Placa ventral con fuerte contracción lateral, con porciónn posterior a la contracción angosta, aproximadamente 1/3 del ancho del cuerpo (Fig. 94 K); gonostilo no muy alargado distalmente y espolón con moderado tamaño (Fig. 93 P); longitud ala 2,6-3,0 mm ........................................................... Simulium veracruzanun 19(10) Gonostilo aproximadamente 2/3 el ancho con respecto al largo (Fig. 93 R); placa ventral sin contracción lateral; borde distal del cuerpo subcircular .......................... ...............................................................................................................Simulium zempoalense – Gonostilo aproximadamente 1/2 el ancho con respecto al largo (Fig. 93 Q); placa ventral con o sin contracción y con diferente forma al dilema opuesto (Figs. 94 LN) ......................................................................................................................................... 20

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20(19) Relación largo/ancho del basitarso posterior = 3,8; longitud ala 2,5-2,8 mm; escudo con 1+1 cuñas claras no llegando al borde posterior (Fig. 93 F) ................ .............................................................................................................. Simulium mangabeirai – Relación largo/ancho del basitarso posterior >5,0; escudo con 1+1 cuña clara ancha llegando al borde posterior (Fig. 93 G) ........................................................... 21 21(20) Segmentos abdominales I-II amarillentos; longitud ala 2,5 mm ................................ ................................................................................................................. Simulium trivittatum – Segmentos abdominales I-II no amarillentos; longitud del ala 2,0-2,5 mm ........ 22 22(21) Escudo con 1+1 bandas laterales negruzcas no llegando a los lóbulos del postpronoto (Fig. 93 G); longitud ala 2,5 mm .................... Simulium gonzalezherrejoni – Escudo con 1+1 bandas laterales negruzcas llegando a los lóbulos del postpronoto; longitud alas 2,0 mm .............................................. Simulium longithallum Pupas 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(2) – 7(6) – 8(7)

Branquia con 20 ramas (Fig. 96 D) ................................................. Simulium panamense Branquia con 4-8 ramas .................................................................................................... 2 Branquia con 4 ramas (Fig. 96 M) .................................................................................. 3 Branquia con 6-8 ramas .................................................................................................... 6 Capullo bien desarrollado cubriendo la mayor parte del tórax y cabeza (Fig. 95 M); borde ventral anterior más adelantado que el dorsal ............... Simulium mangabeirai Capullo escasamente desarrollado no cubriendo la mayor parte del tórax y cabeza; borde dorsal anterior más adelantado que el ventral (Fig. 98 X) .............................. ................................................................................................ BLANCASI sp. grupo......4 Branquia con un corto tronco seguido por 2 ramas distinguibles con 2+2 filamentos (Fig. 98 Y) ............................................................................. Simulium blancasi Branquias con 4 filamentos aparentemente emergidos de un tallo muy corto común (Figs. 99 L, 100 C) ................................................................................................ 5 Frontoclipeo y porción expuesta del tórax con abundantes tubérculos; superficie del capullo rugosa y de color castaño cuando se lo examina en líquido ................. ..................................................................................................................... Simulium tenuipes Frontoclipeo y porción expuesta del tórax con muy esparcidos tubérculos (Figs. 100 D, E); capullo liso y de color castaño con brillo plomizo como un pergamino cuando se lo examina en líquido .............................................................. Simulium penai Branquias con 6 ramas ...................................................................................................... 7 Branquias con 8 ramas .................................................................................................... 10 Branquias con superficie rugosa; ápice romo con pigmentación castaño a negruzca (Figs. 96 I, J); branquias más cortas que el largo del capullo (Figs. 95 I, J) ................................................................................................................................................ 8 Branquias con superficie lisa; generalmente con ápice aguzado con pigmentación amarillenta; branquias tan largas o más largas que el largo del capullo (Fig. 95 N) ................................................................................................................................................ 9 Ramas secundarias externa e interna de la branquia más gruesas que otras ramas de la branquia (Fig. 96 I); tubérculos del frontoclípeo y tórax lisos; capullo sin elevaciones longitudinales (Fig. 95 I) .............................................. Simulium dandrettai

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– 9(7) – 10(6) – 11(10) – 12(10) – 13(12) – 14(12) – 15(14) – 16(15) – 17(16) –

Rama dorsal externa únicamente más gruesa que otras ramas de la branquia (Fig. 96 J); tubérculos del frontoclípeo y tórax rugosos; capullo con elevaciones longitudinales (Fig. 95 J) .......................................... Simulium sp. próximo a dandrettai Ramas primarias y secundarias de las branquias emergiendo muy próximas a la base (Fig. 96 O); tegumento dorsal del tórax con escasos tubérculos y elevaciones circulares chatas rugosas simulando tubérculos (Fig. 95 Q) ...... Simulium zempoalense Ramas primarias y secundarias de las branquias no emergiendo próximas a la base (Fig. 96 P); tegumento dorsal del tórax con tegumento liso y escasos tubérculos ................................................................................................................. Simulium trivittatum Branquias con tallo basal corto, con ramas primarias y secundarias emergiendo muy próximas a la base, con la rama primaria dorsal más gruesa que las otras (Figs. 96 E, F) .............................................................................................................................. 11 Branquias con ramas primarias y secundarias emergiendo no tan cerca de la base de un tallo basal más desarrollado y con las ramas primarias de aproximadamente del mismo diámetro ......................................................................................................... 12 Terciaria rama dorsal de la branquia notoriamente más gruesa y 1/3 más larga que las otras branquias (Fig. 96 F) ................................................... Simulium pseudocallidum Terciaria rama dorsal de la branquia solo un poco más gruesa y con similar longitud que las otras ramas (Fig. 96 E) ........................................... Simulium callidum Ramas dorsal secundaria y terciaria de la branquia más gruesas que las otras ramas (Figs. 96 N, O) .................................................................................................................. 13 Ramas dorsal secundaria y terciaria de la branquia del mismo diámetro que las otras ramas ......................................................................................................................... 14 Ramas dorsal secundaria y terciaria emergiendo lejos de la base (Fig. 96 N) ......... .......................................................................................................Simulium gonzalezherrejoni Ramas dorsal secundaria y terciaria emergiendo próximas a la base (Fig. 96 O) .............................................................................................................. Simulium longithallum Longitud de las branquias tan largas o más largas que el largo del capullo (Fig. 95 L) ............................................................................................................ Simulium veracruzanum Longitud de las branquias más cortas que el largo del capullo (Figs. 95 A-G) ......... 15 Tejido del capullo esparcido con trama muy abierta, sin borde anterior (Fig. 95 F) ................................................................................................................ Simulium iracoubense Tejido del capullo compacto con trama cerrada, con borde anterior (Figs. 95 A-E, G, H, K) ............................................................................................................................. 16 Tricomas cefálicos y torácicos más largos que la mitad del ancho basal del frontoclípeo (Fig. 95 R); borde anterior del capullo reforzado y con una prolongación dorsal anterior (Fig. 95 E) ............................................ Simulium samboni Tricomas cefálicos y torácicos no más largos que la mitad del ancho basal del frontoclípeo (Figs. 95 Q, T, U) borde anterior del capulloo no reforzado y sin una prolongación dorsal anterior .......................................................................................... 17 Capullo con pequeña prolongación dorsal y ventral (Fig. 95 G); frontoclípeo y tórax con escasos tubérculos (Fig. 95 T) .................................. Simulium haematopotum Capullo sin prolongación dorsal (Figs. 95 A, K); frontoclípeo y tórax desde abundantes a escasos tubérculos ................................................................................... 18

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18(17) Ramas branquiales relativamente apretadas basalmente, con disposición subparalela (Figs. 96 A, K) ............................................................................................. 19 – Ramas branquiales generalmente muy abiertas basalmente y con tendencia a convergir distalmente (Figs. 96 B, C, G, H) ................................................................ 20 19(18) Rama secundaria dorsal de la branquia más gruesa que la rama secundaria ventral; rama terciaria dorsal bifurcada a larga distancia de la base (Fig. 96 K); frontoclípeo y tórax con moderado número de tubérculos (Fig. 95 U) ....... Simulium bobpetersoni – Rama secundaria dorsal de la branquia con similar diámetro que la rama secundaria ventral; rama terciaria dorsal bifurcada a corta distancia de la base (Fig. 96 A); frontoclípeo y tórax con abundantes tubérculos (Fig. 95 P) ............... Simulium dugesi 20(18) Rama terciaria mediana de la branquia bifurcada desde corta distancia de la base (Fig. 96 B); frontoclípeo y tórax con escasos tubérculos y manchas cuticulares circulares rugosas no elevadas (Fig. 95 Q) ........................................... Simulium ochoai – Rama terciaria mediana de la branquia bifurcada a larga distancia de la base (Figs. 96 C, G, H); frontoclípeo y tórax con tubérculos desde abaundantes a escasos, en el último caso los tubérculos son elevados tan largos como anchos en la base ........ 21 21(20) Rama terciaria dorsal de la branquia bifurcada a la misma distancia o más lejos que la rama terciaria mediana (Fig. 96 H) ................................................... Simulium downsi – Rama terciaria dorsal de la branquia bifurcada por debajo de la rama terciaria mediana (Figs. 96 C, G) .................................................................................................. 22 22(21) Frontoclípeo y tórax con abundantes tubérculos (Fig. 56 S); longitud de la branquia 2,2-2,5 mm; longitud capullo 3,0-3,5 mm ........................................... Simulium escomeli – Frontoclípeo y tórax desde moderado número a escasos tubérculos; longitud de la branquia 1,5-2,1 mm; longitud capullo 2,4-2,7 mm ................................................. 23 23(22) Branquias proporcionalmente cortas 1,5-1,6 mm; relación largo branquia/capullo = 1,7 ................................................................................................. Simulium mediovittatum – Branquias proporcionalmente largas 2,0-2,1 mm; relación largo branquia/capullo = 1,2 ....................................................................................................... Simulium bivittatum Larvas 1 – 2(1) – 3(2) –

Hendidura postgenal no desarrollada (Figs. 99 M, 100 G) ......................................... .......................................................................................... Simulium tenuipes, Simulium penai Hendidura postgenal desarrollada con diferente profundidad ................................ 2 Hendidura postgenal escasamente desarrollada (Figs. 97 L, N); relación del largo puente hipostomial/hipostomio = 1,2-1,7 ................................................................... 3 Hendidura postgenal más desarrollada (Figs. 97 J, K, M); relación del largo puente hipostomial/ hipostomio = 0,5-1,1 ................................................................................ 6 Relación puente postgenal/hipostomio = 1,7; longitud del cuerpo de la larva madura 6,8 mm; apotoma cefálico con la mitad basal oscurecida, con una ventana mediana (Fig. 97 G) ......................................................................... Simulium mangabeirai Relación puente postgenal/hipostomio = 1,2-1,4; longitud del cuerpo de la larva madura 4,7-6,3 mm; apotoma cefálico escasamente oscurecido y sin una ventana mediana (Fig. 97 H) ........................................................................................................... 4

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Relación puente hipostomial/hipostomio = 1,4; anillo anal con alrededor de 78 hileras de ganchos .............................................................................Simulium zempoalense – Relación puente postgenal/hipostomio = 1,2; anillo anal con 66-70 hileras de ganchos ................................................................................................................................. 5 5(4) Relación de los artejos antenales proximal, mediano y distal = 1:1,3:1,5; anillo anal con alrededor de 66 hileras de ganchos ......................................... Simulium dandrettai – Relación de los artejos antenales proximal, mediano y distal = 1:1,5:2,0; anillo anal con alrededor de 70 hileras de ganchos ...................................... Simulium longithallum 6(2) Hipostomio más corto que el puente postgenal (Fig. 98 g); relación largo puente postgenal/hipostomio = 1,3 ................................................................. Simulium blancasi – Hipostomio tan largo o más largo que el puente postgenal; relación largo puente postgenal/hipostomio = 0,5-1,1 ..................................................................................... 7 7(6) Hipostomio más largo que el puente postgenal (Figs. 97 J, K); relación largo puente postgenal/hipostomio = 0,5-0,9 ...................................................................................... 8 – Hipostomio con similar largo o más corto que el puente postgenal (Fig. 97 M) ); relación largo puente postgenal/hipostomio = 1,0-1,1 ........................................... 14 8(7) Relación largo puente postgenal/hipostomio = 0,5-0,6; apotoma cefálico sin ornamentación ................................................................ Simulium dugesi, Simulium ochoai – Relación largo puente postgenal/hipostomio = 0,7-0,9; apotoma cefálico oscurecido en mitad basal, encerrando una ventana clara mediana (Figs. 97 B, C, E) ................... 9 9(8) Longitud del cuerpo 5,5-6,0 mm, antena con similar largo que el tallo del abanico cefálico; relación largo puente postgenal/hipostomio = 0,7 ......... Simulium escomeli – Longitud del cuerpo 4,3-5,4 mm; antena más larga que el tallo del abanico cefálico; relación largo puente postgenal/hipostomio = 0,8-0,9 ........................... 10 10(9) Anillo anal con 89 hileras de ganchos; artejo distal de la antena de similar longitud que el mediano (Fig. 97 O) ............................................................... Simulium panamense – Anillo anal con 63-77 hileras de ganchos; artejo distal de la antena más largo que el mediano (Fig. 97 P) ..................................................................................................... 11 11(10) Abanico cefálico con 52-56 rayos; longitud del cuerpo 5,0-5,5 mm ........................ ........................................................................................................... Simulium mediovattatum – Abanico cefálico con 31-48 rayos; longitud del curpo 4,1-5.0 mm ...................... 12 12(11) Abanico cefálico con 31-35 rayos; anillo anal con 74-91 rayo ................................... ........................................................................................................... Simulium haematopotum – Abanico cefálico con 38-48 rayos; anillo anal con 63-74 rayos ............................. 13 13(12) Esclerito anal con pelos y escamas; anillo anal con 65-74 hileras; papilas rectales con 1 divertículo en cada lóbulo ............................................ Simulium gonzalezherrejoni – Esclerito anal sin pelos ni escamas (Fig. 97 Q); anillo anal con 63 hileras de ganchos;papilas rectales con 2-3 divertículos en cada lóbulo (Fig. 97 S) ................. .............................................................................................................. Simulium iracouboense 14(7) Anillo anal con 100-115 hileras con 14-20 ganchos cada una; esclerito anal con abundantes pelos y escamas (Fig. 97 R); apotoma cefálico sin ornamentación (Fig. 97 D) ................................................................................................................................... 15 – Anillo anal con 60-90 hileras con 11-15 ganchos cada una; esclerito anal generalmente sin escamas; apotoma cefálico con o sin ornamentación .............. 16

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15(14) Apotoma cefálico con 30-32 rayos; esclerito lateral de la propata torácica con alrededor de 50 dientes; longitud máxima 5,5-6,0 mm .................. Simulium callidum – Apotoma cefálico con 40-42 rayos; esclerito lateral de propata torácica con alrededor de 37 dientes; longituud máxima 4,8-5,0 mm ..... Simulium pseudocallidum 16(14) Anillo anal con 60-62 hileras con 11 ganchos cada una ........... Simulium bobpetersoni – Anillo anal con 68-90 hileras con 13-15 ganchos cada una .................................... 17 17(16) Longitud del cuerpo 5,5-6,0 mm; esclerito anal con abundantes escamas .............. ............................................................................................................Simulium veracruzanum – Longitud del cuerpo 3,9-5,5 mm; esclerito anal sin escamas ................................. 18 18(17) Apotoma cefálico oscurecido subbasalmente con una ventana clara mediana longitudinal alargada y 1+1 transversa subbasal (Figs. 97 A, F); longitud del cuerpo 5,1-5,4 mm ................ Simulium downsi, Simulium samboni, Simulium trivittatum – Apotoma cefálico escasamente pigmentado, solamente un poco más oscurecido basalmente; longitud del cuerpo 3,7-4,5 mm ................................ Simulium bivittatum ESCOMELI species group Simulium (Psilopelmia) dugesi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 91 A, G, J; 92 A; 93 A, H; 94 A; 95 A, P; 96 A; 97 J.) Simulium (Lanea) dugesi VARGAS et al., 1946: 107; DALMAT, 1955: 59. Simulium (Psilopelmia) dugesi VARGAS & DÍAZ NÁJERA, 1957b: 182; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 48-50; CROSSKEY & HOWARD, 1997: 58; 2004: 53.

Female: Wing length 2.0-2.2 mm; scutum grayish with reddish brown median longitudinal stripe diffuse at sides (Fig. 91 A); with some illumination, 1+1 submedian anterior light subtriangular spots reaching anterior rim; scutum and metanotum dark gray; pleurae with pruinosity gray; legs yellowish brown with apex of femora and tarsomeres blackish; abdomen grayish with blackish subquadrate median spots on II-VI and 1+1 sublateral spots wider on III-VI tergites and sometimes on VII (Fig. 91 G); cibarium with several rows of minute spines except on median depressed area (Fig. 91 J); fronto-ocular triangle slightly longer than wide; basal sector of R glabrous; sensory vesicle of maxillary palp about half of palpomere length; claw without subbasal tooth; eighth sternite with about 20 setae on each side; paraproct short, truncate slightly acuminate dorsally (Fig. 92 A). Male: Wing length 1.9 mm; scutum black with 1+1 subtriangular silvery spots extending posteriorly as thin stripes surpassing half of scutum length (Fig. 93 A); abdomen blackish with 1+1 silvery spots on tergites II, VI, VII; gonostylus slightly acuminate on distal inner portion (Fig. 93 H); ventral plate short with gently curved border (Fig. 94 A). Pupa: Cocoon reinforced anterior border, with small anterior projection (Fig. 95 A), threads evident; length basally 2.9-3.4 mm, at dorsum 2.5-2.8 mm, gills 2.0-2.6 mm; frontoclypeus and thorax with numerous tubercles (Fig. 95 P); gill with 8 branches bifurcated not close to base (Fig. 96 A). Larva: Maximum length 4.6-5.2 mm; cephalic apotome with homogeneous coloration; ratio of proximal, medial and distal antennal segments = 1:1.3:1.7; cephalic

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A

C

B

G

F

H

J

D

E

I

L K N

M

O

Q

P

U

T

V

W

R

S

Figure 91. Female. A-F: Scutum; A: S. (Psilopelmia) dugesi; B: S. (P.) samboni; C: S. (P.) haematopotum; D: S. (P.) callidum; E: S. (P.) escomeli; F: S. (P.) mangabeirai. G-I: Abdomen: G: S. (P.) dugesi; H: S. (P.) iracouboense; I: S. (P.) escomeli. J-Q: Basal portion of cibarium; J: S. (P.) dugesi; K: S. (P.) bivittatum; L: S. (P.) samboni; M: S. (P.) ircouboense; N: S. (P.) panamense; O: S. (P.) pseudocallidum; P: S. (P.) trivittatum; Q: S. (P.) dandrettai. R-S: Sensory organ of palp; R: S. (P.) iracouboense; S: S. (P.) haematopotum. T-W: Claws; T: S. (P.) bivittatum; U: S. (P.) iracouboense; V: S. (P.) panamense; W: S. (P.) callidum.

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fan with about 40 rays; mandible with latero-mandibular process; postgenal cleft dome shaped; postgenal bridge about half of hypostomium length (Fig. 97 J); rectal papillae with only 3 single lobes; anal sclerite without hair or scales; anal ring with 70 rows of 15-16 hooks each. Distribution: Mexico: Nueva León, San Luis Potosí, Tamaulipas. Bionomics: Unknown. Discussion: Simulium dugesi is very close to S. mediovittatum and S. ochoai; differences are explained under those species. Simulium (Psilopelmia) ochoai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 95 B, Q; 96 B; 97 I.) Simulium (Lanea) ochoai VARGAS et al., 1946: 131-133, 169. Simulium (Psilopelmia) ochoai: VARGAS & DÍAZ NÁJERA, 1957b: 182; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 50-51; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

Female: Wing length 2.1 mm; scutum, pleurae, legs, and metanotum similar to those of S. dugesi; scutellum light yellowish brown; abdomen as in S. dugesi except segment II yellowish; cibarium, sensory vesicle of maxillary palp, basal sector of R and paraproct as in S. dugesi. Male: Wing length 1.6 mm; coloration of scutum and abdomen as in S. dugesi, but scutum submedian silvery stripes slightly shorter, not surpassing half of scutum length; gonostylus elongated on apical third, very similar to S. haematopotum. Pupa: Cocoon with anterior border reinforced, not projected anteriorly (Fig. 95 B), weave with evident threads; length along base 2.8-3.2 mm, along dorsum 2.7-2.9 mm, gills 1.5-2.0 mm; frontoclypeus and thorax with scarce tubercles (Fig. 95 Q); trichomes short, with 1-2 branches; gill with 8 branches arranged very open with primary branches very short (Fig. 96 B). Larva: Maximum length 4.5-4.8 mm; cephalic apotome light whitish yellow scarcely darkened basally with subtriangular or bell shape, but with light median window; antenna longer than cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.4:1.8; cephalic fan with about 42 rays; mandible with latero-mandibular process (Fig. 97 I); postgenal cleft as in S. dugesi; postgenal bridge longer than half of hypostomium length; lateral sclerite of proleg with 14-15 teeth arranged in 10-11 groups; anal sclerite with scarce hair and scales; anal ring with 64-70 rows of 14-15 hooks each; rectal papillae without diverticula. Distribution: Mexico: Veracruz. Bionomics: Unknown. Discussion: The adults are very similar to those of S. dugesi and S. mediovittatum, but the pupae are distinguishable. The first species has more numerous tubercles on the frontoclypeus and thorax, and both species have the pupal gill branching closer to the base, forming a bunch in S. dugesi.

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B

C

D

A

G E

J

F

M

H L N P

I

Q

K

O

Figure 92. Female. A-N: Cercus and paraproct; A: S. (Psilopelmia) dugesi; B: S. (P.) bivittatum; C: S. (P.) samboni; D: S. (P.) iracouboense; E: S. (P.) haematopotum; F: S. (P.) panamense; G: S. (P.) callidum; H: S. (P.) escomeli; I: S. (P.) downsi; J: S. (P.) dandrettai; K: S. (P.) bobpetersoni; L: S. (P.) veracruzanum; M: S. (P.) gonzalezherrejoni; N: S. (P.) trivittatum. O: Eighth sternite and gonapophyses of S. (P.) mediovittatum. PQ: Fronto-ocular triangle; P: S. (P.) iracouboense; Q: S. (P.) samboni.

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Simulium (Psilopelmia) mediovittatum KNAB (Figs. 92 O; 95 C; 97 K.) Simulium mediovittatum KNAB, 1915 a: 77; 1915b: 149; VARGAS & DÍAZ NÁJERA, 1954: 69. Simulium (Lanea) mediovittatum: VARGAS et al., 1946: 107. Simulium (Psilopelmia) mediovittatum: VARGAS & DÍAZ NÁJERA, 1957b: 154; PETERSON, 1993: 329; COSCARÓN, 1994: 31; COSCARÓN et al., 1996: 51-54; MOULTON, 1998: 63; CROSSKEY & HOWARD, 1997: 59; 2004: 53; ADLER et al., 2004: 352.

Female: Wing length 2.2 mm; frons and clypeus dark brown with grayish pruinosity; scutum light grayish brown with variable tones from orange to brown with median reddish-brown to black stripe, covered by decumbent yellowish pile; scutellum coloration as for scutum; metanotum brownish black; pleurae yellowish to brownish black; legs blackish with tibia apex and tarsomeres whitish; abdomen yellow-brown grayish to dark brown; tergites II-VI with median stripe of subquadrate brown matte spots and 1+1 lateral rectangular-shaped blackish spots with distal rim grayish; VIII-IX yellowish-gray brownish, pale and shiny; cibarium with 1+1 flat submedian lobes covered by acute teeth, as in S. dandrettai; sensory vesicle of maxillary palp about 1.5-1.8 of palpomere length; wing basal sector of R without hair; claw without subbasal tooth; paraproct truncate with small dorsal projection similar to that of S. dugesi; eighth sternite with 22-28 setae on each side; gonapophysis divergent with internal rim slightly sclerotized. (Fig. 92 O). Male: Wing length 1.8 mm; scutum black with 1+1 anterior subtriangular silvery spots continued posteriorly as thin stripes, surpassing half of scutum length, laterally grayish (Fig. 93 A); pilosity golden shiny; abdomen black with 1+1 silvery spots on tergites II, VI, VII; ratio of length/width of hind basitarsus = 6; gonostylus slightly acuminate on distal inner portion as in S. dugesi; ventral plate wider than long, slightly acuminate distally, very similar to S. haematopotum and S. trivittatum. Pupa: Cocoon slipper shaped, slightly elongated basally (Fig. 95 C), threads evident; length basally 2.5-2.7 mm, dorsally 2.2-2.5 mm, gills 1.5-1.6 mm; frontoclypeus and thorax with moderate to sparse tubercles, with rounded apical rim; frontal trichomes single and relatively short; gill with 8 branches, with primary ventral branch relatively elongated. Larva: Maximum length 5.0-5.5 mm; cephalic apotome as in S. escomeli, darkened median and basally with median and 1+1 lateral and subbasal light windows; mandible with 1 latero-mandibular process; antenna surpassing cephalic fan stem length; ratio of proximal, medial and distal segments = 1:1.3:1.5; cephalic fan with 52-56 rays; postgenal bridge slightly shorter than hypostomium length (Fig. 97 K); lateral sclerite of thoracic proleg with about 17 teeth; anal sclerite without hair or scales, similar to S. zempoalense; anal ring with 68-77 rows of 14-16 hooks each; rectal papillae with 3 single lobes; ninth segmente with 1 pair of ventral tubercles. Distribution: Mexico: Chihuahua, Coahuila, San Luis Potosi, Tamaulipas, Veracruz; USA: Texas. Bionomics: Simulium mediovittatum larvae live on aquatic vegetation of rivers and shallow streams. Females bite humans, livestock and wild mammals. Discussion: The closest species to S. mediovittatum are S. dugesi and S. ochoai. The first species can be differentiated by the greater number of frontoclypeal and thoracic tubercles. Simulium ochoai has gill branching nearer the base and more open. These small

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differences, plus the coloration and number of anal ring hook rows of S. mediovittatum, suggest that it would be useful to compare more material of those species and to also use additional nonmorphological studies. Simulium (Psilopelmia) bivittatum MALLOCH (Figs. 91 K, T; 92 B; 93 I; 94 B; 95 D; 96 C.) Simulium bivittatum MALLOCH, 1914: 31; VARGAS, 1945a: 120. Simulium (Psilopelmia) bivittatum: VARGAS & DÍAZ NÁJERA, 1958: 13; PETERSON, 1993: 308; COSCARÓN, 1987: 30; PETERSON & KONDRATIEFF, 1995: 35-36; COSCARÓN et al., 1996: 54, 55; 63; CROSSKEY & HOWARD, 1997: 58; 2004: 53; MOULTON, 1998: 50; ADLER et al., 2004: 349-350. Simulium idahoense TWINN, 1938: 50-51.

Female: Wing length 2.1-2-3 mm; frons and clypeus with silver pruinosity; scutum yellowish orange to reddish brown arranged in 1 median and 1+1 sublateral stripes separated by 1+1 submedian light gray whitish vittae, scutellum brown yellowish and metanotum black-brown grayish; abdomen grayish with dark brown subquadrate spots on median line and 1+1 rectangles laterally; legs yellow-grayish with apex of femora, tibiae and tarsomeres dark brown; cibarium with sclerotized rim with blunt teeth (Fig. 91 K); basal sector of R glabrous; claw without subbasal tooth (Fig. 91 T); eighth sternite with 20-23 setae; paraproct with short distal projection, with acute apex (Fig. 92 B). Male: Wing length 2.0-2.1 mm; scutum blackish to reddish brown covered with recumbent subbasal golden pile; with anterior light showing 1+1 anterior cuneiform silvery spots, not reaching half of scutum length; scutellum dark yellowish brown, pleurae dark grayish; metanotum brownish black, grayish pruinose; legs mostly dark brown; gonostylus subrectangular with small apical concavity (Fig. 93 I); ventral plate short without apical projection (Fig. 94 B). Pupa: Cocoon slightly projected anteriorly at base, anterior border reinforced, (Fig. 95 D), threads evident; length along base 2.4-2.7 mm, on dorsum 2.2-2.5 mm, gills 1.9-2.1 mm; frontoclypeus and thorax with tubercles relatively sparse elongated, as long as wide at base, with rounded apex and some of them acute; trichomes with 1 or 2 branches; gills with 8 branches, with primary and secondary branches short (Fig. 96 C). Larva: Maximum length 3.7-4.5 mm; cephalic apotome scarcely pigmented, only slightly darkened basally; antenna slightly longer than cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.4:1.6; cephalic fan with 30-38 rays; mandible with lateromandibular process; postgenal cleft dome shaped; postgenal bridge slightly shorter than hypostomium; anal sclerite without scales and very small hairs; anal ring with about 70 rows with 13-16 hooks each; rectal papillae without diverticula on lobes. Distribution: Canada: Alberta, Saskatchewan; Mexico: Chihuahua, Coahuila, San Luis Potosi, Tamaulipas, Veracruz, Zacatecas; U.S.A.: Arizona, Colorado, Idaho, Montana, Nebraska, New Mexico, Oregon, South Dakota, Texas, Utah, Washington, Wyomming. Bionomics: Simulium bivittatum breeds in rivers with bottoms of rocks and gravel in lowlands to low foothills. Females bite humans and livestock, and the attacks can make people stay indoors. Discussion: Based on genitalia and cibarium characters, S. bivittatum constitutes a group with S. mediovittatum, S. dugesi and S. ochoai, but the scutum coloration and

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Figure 93. Male. A-G: Scutum; A: S. (Psilopelmia) dugesi; B: S. (P.) iracouboense; C: S. (P.) escomeli; D: S. (P.) veracruzanum; E: S. (P.) zempoalense; F: S. (P.) mangabeirai; G: S. (P.) gonzalezherrejoni. H-S: Gonocoxite and gonostylus; H: S. (P.) dugesi; I: S. (P.) bivittatum; J: S. (P.) samboni; K: S. (P.) iracouboense; L: S. (P.) haematopotum; M: S. (P.) escomeli; N: S. (P.) downsi; O: S. (P.) dandrettai; P: S. (P.) veracruzanum; Q: S. (P.) mangabeirai; R: S. (P.) zempoalense; S: S. (P.) callidum.

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ornamentation of the adutls permit it to be differentiated. The presence of yellowish females and blackish males is similar to the condition for S. escomeli, but the other morphological characters allow identification. Simulium (Psilopelmia) samboni JENNINGS (Figs. 91 B, L; 92 C, Q; 93 J; 95 E, R; 97 A.) Simulium samboni JENNINGS, 1915: 67; FAIRCHILD, 1940: 704-705; VARGAS, 1942a: 240; RAMÍREZ PÉREZ, 1983: 67; SHELLEY et al., 1984: 155-156. Simulium (Psilopelmia) samboni: VARGAS & DÍAZ NÁJERA, 1957b, 356, 358:; COSCARÓN, 1987: 37; COSCARÓN et al., 1995: 292-294; COSCARÓN et al., 1996: 56-59; CROSSKEY & HOWARD, 1997: 59; 2004: 53; SHELLEY et al., 2002: 169. Simulium (Lanea) samboni VARGAS et al., 1946: 107; VARGAS & DÍAZ NÁJERA, 1948b: 343; DALMAT, 1955: 151-156. Simulium colvini DALMAT, 1952: 344-347; 131-135; VARGAS & DÍAZ NÁJERA, 1954: 62-63.

Female: Wing length 1.9-2-1 mm; scutum, scutellum and metanotum yellowish, the first with 1+1 silvery submedian stripes reaching posterior border (Fig. 91 B), and covered by sparse yellow pile; metanotum with anterolateral areas brownish with gray pruinosity; abdomen with segments I-II yellowish, III-VI with tergal plate and lateral areas dark brown but separated by narrow stripes determining 3 rows of subquadrate plates as in S. escomeli; pleural area light grayish; legs yellowish, darkened on tarsomeres and apex of hind tibia and femur; cibarium with 1+1 slight submedian elevations, covered with small teeth (Fig. 91 L); fronto-ocular triangle about as wide as high (Fig. 92 Q); basal sector of R with sparse hair arranged in 1 row; claw without subbasal tooth; eighth sternite with 14-16 setae; paraproct with short distal projection (Fig. 92 C). Male: Wing length 1.8-2.0 mm; coloration as in female, but with scutum dark and submedian silvery stripes not surpassing half of scutum length; abdomen with segments (III) IV-VI totally dark brown; gonostylus subrectangular, slightly longer than half of gonocoxite length (Fig. 93 J); ventral plate as in S. escomeli. Pupa: Cocoon with anterior border reinforced and small projection basal and dorsal (Fig. 95 E); weave compact with evident threads; length along base 2.2-2.5 mm, on dorsum 2.0-2.3 mm, gills 2.0-2.1 mm; frontoclypeus and thorax with moderate number of tubercles (Fig. 95 R), cephalic and thoracic trichomes bifid and elongated; gills with 8 branches, with short dorsal primary and secondary branches. Larva: Maximum length 5.0-5.2 mm; cephalic apotome with median and 1+1 subbasal windows (Fig. 97 A); antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9-1:1.3-1.5; cephalic fan with 34-36 rays; postgenal bridge about same length as hypostomium; anal sclerite with single hairs; anal ring with about 70 rows with 12-13 hooks each; ninth segment with pair of ventral tubercles; rectal papillae with 1-3 diverticula on each lobe. Distribution: Belice: Belice, Cayo, Stan Creek, Toledo; Guatemala: Altaverapaz, El Petén, Escuintla, San Marcos, Santa Rosa, Solalá, Totonicapán; Honduras: Cortés; Mexico: Chiapas, Tabasco, Oaxaca, Veracruz; Panama: Canal Zone, Chucumaque, Coclé; Trinidad; Venezuela: Anzoategui, Barinas, Carabobo, Federal District, Guarico, Miranda, Monagas; Portuguesa, Tachira, Yaracuy.

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Bionomics: Simulium samboni breeds in small to medium-sized creeks, with moderate to fast-flowing water, attached to aquatic vegetation. Females are zoophilic. Discussion: The gonostylus shape of S. samboni permits it to be separated from other S. (Psilopelmia) species. Simulium escomeli is similar but the elongated distal projection of the paraproct plus the subtrapezoidal gonostylus permit differentiation. Simulium (Psilopelmia) iracouboense FLOCH & ABONNENC (Figs. 91 H, M, R, U; 92 D, P; 93 B, K; 94 C; 95 F; 97 B, Q, S.) Simulium iracouboense FLOCH & ABONNENC, 1946b: 7-12; HAMADA & GRILLET, 2001: 29. Simulium (Psilopelmia) iracouboense: COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 59-60; SHELLEY et al., 1997: 30; CROSSKEY & HOWARD, 1997: 59; 2004: 53. Simulium sucamense NUNES DE MELLO, 1974: 41; PY-DANIEL, 1989a: 254. Simulium santaelenae RAMÍREZ PÉREZ & PETERSON, 1981: 161; RAMÍREZ PÉREZ, 1983: 20; PY-DANIEL, 1989a: 254.

Female: Wing length 1.5-1.6 mm; scutum, scutellum, metanotum and pleura brownyellowish, the first with 1+1 submedian and lateral whitish arrangement as in S. escomeli; abdomen brown-yellowish with 1 median and 1+1 sublateral blackish subquadrate spots (Fig. 91 H); legs brown-yellowish with apex of femora. tibiae and tarsomeres dark brown; fronto-ocular triangle wider than high (Fig. 92 P); sensory vesicle of maxillary palp about half as long as palpomere (Fig. 91 R); cibarium with 2 protuberances with acute teeth (Fig. 91 M); basal sector of R with hair; claw without subbasal tooth (Fig. 91 U); paraprocts subquadrate, rounded apically and well-developed basal lobe (Fig. 92 D). Male: Wing length 1.5 mm; scutum brown yellowish with yellow pile; with certain lighting, 1+1 submedian cuneiform spots on anterior third visible (Fig. 93 B); legs and abdomen as in female, but abdominal segments (III) VI-VII darker; gonostylus subtrapezoidal, and about 2/3 gonocoxite length (Fig. 93 K). Ventral plate wider than long, slightly acuminate apically (Fig. 94 C). Pupa: Cocoon with loose weave, with very obivous threads and dorsally slightly reinforced (Fig. 95 F); length basally 2.4-3.0 mm, dorsally 2.1-2.4 mm, gills 1.3-2.5 mm; frontoclypeus and thorax with smooth tubercles in moderate number; cephalic trichomes single, thoracic single or bifurcated; gills with 8 filiform branches, with primary and secondary branches arising close to base. Larva: Maximum length 4.1-4.5 mm; cephalic apotome with darkened median basal area circumscribing median subbasal windows (Fig. 97 B); antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.1-1.3:1.6-1.7; cephalic fan with 43-48 rays; postgenal bridge slightly shorter than hypostomium length; anal sclerite without scales and very sparse hair (Fig. 97 Q); anal ring with about 63 rows with 15-18 hooks each; rectal papillae with 2-3 diverticula on each lobe (Fig. 97 S). Distribution: Brazil: Amapa, Pará, Roraima; French Guiana; Surinam; Venezuela. Bionomics: Unknown. Discussion: The closest species to S. iracouboense is S. samboni, but it can be differentiated by the absence of hairs on the basal sector of R, presence of a frontoocular triangle, cephalic trichomes bifid, cocoon with compact weave and more

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Figure 94. Male. Ventral plate. A: S. (Psilopemia) dugesi. B: S. (P.) bivittatum. C: S. (P.) iracouboense. D: S. (P.) haematopotum. E: S. (P.) panamense. F: S. (P.) callidum. G: S. (P.) escomeli. H: S. (P.) downsi. I: S. (P.) dandrettai. J: S. (P.) bobpetersoni. K: S. (P.) veracruzanum. L: S. (P.) mangabeirai. M: S. (P.) gonzalezherrejoni. N: S. (P.) trivittatum.

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Figure 95. Pupa. A-N: General aspect; A: S. (Psilopelmia) dugesi; B: S. (P.) ochoai; C: S. (P.) mediovittatum; D: S. (P.) bivittatum; E: S. (P.) samboni; F: S. (P.) iracouboense; G: S. (P.) haematopotum; H: S. (P.) downsi; I: S. (P.) dandrettai; J: nr. S. (P.) dandrettai; K: S. (P.) bobpetersoni; L: S. (P.) veracruzanum; M: S. (P.) mangabeirai; N: S. (P.) zempoalense. O: Dorsal portion of thoracic integument of S. (P.) zempoalense. P-V: Frontoclypeus; P: S. (P.) dugesi; Q: S. (P.) ochoai; R: S. (P.) samboni; S: S. (P.) escomeli; T: S. (P.) haematopotum; U: S. (P.) bobpetersoni; V: S. (P.) panamense. W: Frontoclypeal tubercles, enlarged, of S. (P.) panamense.

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reinforced anterior border and larvae with a small number of primary rays and anal ring with more rows. Simulium (Psilopelmia) haematopotum MALLOCH (Figs. 91 C, S; 92 E; 93 L; 94 D; 95 G.) Simulium haematopotum MALLOCH, 1914: 61; DYAR & SHANNON, 1927: 26; FAIRCHILD, 1940: 704; IRIARTE, 1943: 191: LANE & VULCANO, 1943: 439: VARGAS, 1945a: 142. Simulium (Lanae) haematopotum: DALMAT, 1955: 144. Psilopelmia haematopotum: RUBTZOV & GARCÍA AVILA, 1972: 22. Simulium (Psilopelmia) haematopotum: COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 60; CROSSKEY & HOWARD, 1997: 59; 2004: 53. Simulium pseudohaematopotum HOFFMANN, 1930b: 293. Simulium boydi DE LEÓN, 1945: 75. Simulium marathrumi FAIRCHILD, 1940: 715.

Female: Wing length 2.0-2.2 mm; coloration generally brownish; scutum ornamented by median and 1+1 sublateral black stripes, separated by 1+1 thick submedian silvery pruinose vittae and bordered anteriorly, laterally and posteriorly by thin silvery pruinose area (Fig. 91 C), hair grayish to yellow in accordance with light position; scutellum and metanotum brownish; legs grayish, blackish on apex of femur, tibia and tarsomeres of fore and mid legs, and more expanded darker area on hind femur and tibia; abdomen grayish, with tergites I-II blackish with posterior border pale gray, III-VI with median row of subquadrate dark brown plates, and III-IV with 1+1 lateral row of subrectangular brown-grayish to blackish spots covering anterior 2/3 of tergites; segments V-VI laterally grayish pollinose; VIII-IX grayish brown shiny; cibarium with 1+1 short sublateral elevations, covered by teeth as is median depressed area, similar to S. bivittatum but with more acute teeth; sensory vesicle of maxillary palp shorter than half of palpomere (Fig. 91 S); basal sector of R glabrous; claw without subbasal tooth; eighth sternite with 28-30 setae; paraproct blunt apically and with large basal lobe (Fig. 92 E). Male: Wing length 1.8-2 mm; scutum black with 1+1 silvery pruinose cuneiform anterior stripes, extending about 1/4 of scutum length, but with changing light position, continued as thin stripes to posterior border; abdomen blackish with 1+1 lateral silvery spots on segments II, V-VII; gonostylus about as long as gonocoxite, becoming thin on distal 1/3 with enlarged apical spur (Fig. 93 L); ventral plate slightly acuminate distally (Fig. 94 D). Pupa: Cocoon compact with anterior border slightly reinforced and with small dorsal and ventral projections (Fig. 95 G); threads evident; length basally 2.8-3.0 mm, dorsally 2.4-2.5 mm; gills about 1.5 mm; frontoclypeus and thorax without or with very sparse tubercles; cephalic and thoracic trichomes single and bifid; gills with 8 branches, arranged as in S. escomeli. Larva: Maximum length 4.5-5.0 mm; cephalic apotome ornamentation very similar to that of S. escomeli; antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:0.9-1:1.3; postgenal cleft slightly short than hypostomium length; ninth segment with pair of ventral tubercles; anal sclerite without scales; anal ring with 74-80 rows, with 14-17 hooks each; rectal papillae with 1-3 diverticula on each lobe.

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Distribution: Colombia; Costa Rica: Valle Central; Cuba: Las Villas, Oriente, Pinar del Río; Guatemala: Altaverapaz, Chimaltenango, Chiquimula, El Petén, El Progreso, Escuintla, Guatemala, Huehuetenango, Izabal, Quezaltwenango, San Marcos, Santa Rosa; Guyana: Essequibo; Mexico: Chiapas, Colima, Guerrero, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Sinaloa and Veracruz; Panama: Canal Zone, Coclé, Colón, Panama; Puerto Rico: Río Piedras; Venezuela: Bolívar. Bionomics: Simulium haematopotum breeds in shallow water. Females are anthropophilic in Mexico and Guatemala. Discussion: Simulium haematopotum has female abdominal ornamentation and cibarium and paraproct shape similar to those of S. bivittatum, S. dugesi, S. ochoai, S. mediovittatum and S. samboni, but can be differentiated by the scutum and larval cephalic apotome. Simulium samboni shows similar scutum design but with yellowish coloration. Simulium marathrumi has similar ornamentation of the scutum and abdomen, and morphology of the wings and legs Based on the variability of these characters, S. marathrumi can be considered a synonym, but it needs to be confirmed with more material and the use of cytological or molecular studies. CROSSKEY & HOWARD (2004: 53) consider it a synonym of S. haematopotum. Simulium benjamini is very close to S. haematopotum, showing differences such as the elongated secondary dorsal and median pupal gill branches and a big subbasal tooth on the female claw, according to the drawings of DALMAT. These characters could represent variants, because in a pharate female paratype from the USNM, the toothed claw is not evident. We consider that S. benjamini could be a synonym of S. quadrivittattum, but it would be convenient to see more material in order to confirm if they are conspecific. Simulium (Psilopelmia) escomeli ROUBAUD (Figs. 91 E, I; 92 H; 93 C, M; 94 G; 95 S; 96 G; 97 E.) Simulium escomeli ROUBAUD, 1909: 428-429; SILVA FIGUEROA, 1917: 30; WYGODZINSKY, 1953b: 330-332. Psilopelmia rufidorsum ENDERLEIN, 1934b: 283. Simulium (Lanea) escomeli: VARGAS & DÍAZ NÁJERA, 1951: 152. Simulium (Psilopelmia) escomeli: VARGAS & DÍAZ NÁJERA, 1953a: 17-21; WYGODZINSKY, 1971: 2-12; COSCARÓN, 1987: 31; 1991: 266-267; SHELLEY et al., 1989: 97; COSCARÓN et al., 1996: 73-76; COSCARÓN et al., 1995: 289-295; COSCARÓN ARIAS, 2003: 198, 204, 209, 215; CROSSKEY & HOWARD, 1997: 58; 2004: 53; COSCARÓN ARIAS, 2003: 195.

Female: Wing length 2.3-2.8 mm; general coloration yellowish, some specimens brownish to blackish; frons and clypeus grayish pollinose; scutum yellow to blackish with 1+1 anterior scutal border and thick submedian silvery stripes reaching posterior area (Fig. 91 E); prescutellar area brownish; lateral and pleural areas brown with gray pruinosity; scutellum yellowish brown, metanotum dark brown with silvery pruinosity; legs yellowish, with apex of femora, tibiae and tarsomeres dark brown; abdomen light yellow, segment II with brown tergal anterior spot; III-VIII with 1 blackish tergal subquadrate median, 1+1 sublateral and 1+1 small lateral spots, except VII without median spot (Fig. 91 I); cibarium with 1+1 small lateral protuberances with thin teeth, and median portion depressed, smooth, similar to shape in S. trivittatum; sensory vesicle of maxillary palp about 1/3 of palpomere length; fronto-ocular triangle about as high as wide; basal sector of R glabrous; claw without subbasal tooth; paraproct strongly

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Figure 96. Pupa. Gills and thoracic trichomes. A: S. (Psilopelmia) dugesi. B: S. (P.) ochoai. C: S. (P.) bivittatum. D: S. (P.) panamense. E: S. (P.) callidum. F: S. (P.) pseudocallidum. G: S. (P.) escomeli. H: S. (P.) downsi. I: S. (P.) dandrettai. J: nr. S. (P.) dandrettai (also with filament very enlarged). K: S. (P.) bobpetersoni. L: S. (P.) veracruzanum. M: S. (P.) mangabeirai. N: S. (P.) gonzalezherrejoni. O: S. (P.) longithallum. P: S. (P.) trivittatum. Q: S. (P.) zempoalense.

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projected distally and with well-developed basal posterior protuberance (Fig. 92 H); eighth sternite with 20 setae on each side. Male: Wing length 2.2-2.6 mm; scutum velvety black with 1+1 silvery anterior submedian spots not surpassing anterior half of scutum length (Fig. 93 C) (some specimens with scutum orange brown to reddish); scutellum light brown and metanotum black with silvery pruinosity; abdomen black with segment I lighter, II, VI-VIII with 1+1 silvery lateral spots; gonostylus subtrapezoidal and slightly shorter than gonocoxite (Fig. 93 M); ventral plate slightly constricted laterally (Fig. 94 G). Pupa: Cocoon with reinforced anterior border straight to slightly projected dorsally; weave thin with evident threads; length basally 3.0-3.5 mm, dorsally 2.2-3.0 mm, gills 2.22.5 mm; frontoclypeus and thorax with numerous smooth tubercles (Fig. 95 S); gills with 8 filamentous branches, relatively open basally (Fig. 96 G). Larva: Maximum length 5.5-6.0 mm; cephalic apotome with centro-basal and 1+1 basal windows (Fig. 97 E); antenna as long as stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.3:1.1-1.6; cephalic fan with 33-38 rays; postgenal bridge 3/4 of hypostomium length; sclerite comb of thoracic proleg with about 30 teeth arranged in 11-12 groups; anal ring with 63-70 rows, with 10-13 hooks each; ninth segment without ventral tubercles; rectal papillae with 0-3 diverticula on each lobe. Distribution: Chile: Tarapacá; Colombia; Ecuador: Azuay, Cañar, Carchi, Chimborazo, Cotopaxi, Esmeraldas, Imbabura, La Unión, Los Ríos, Pichincha, Tungurahua; Peru: Ancash, Arequipa, Lima, Tacna. Bionomics: Simulium escomeli breeds in clear, fast flowing-streams of regular depth, attached to aquatic vegetation or stones. Females are anthropophilic and a dangerous pest in the valley of the Pacific desert. Discussion: Simulium escomeli is closest to S. downsi, having similar adult coloration and cibarial shape; some differences are in the paraproct, with a slight posterodorsal protuberance (Fig. 92 I), and the gonostylus with a more pronounced external median concavity (Fig. 93 N). Simulium (Psilopelmia) downsi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 92 I; 93 N; 94 H; 95 H; 96 H; 97 F, P.) Simulium (Lanea) downsi VARGAS et al., 1946: 126-129; DALMAT, 1955: 136-140. Simulium downsi: VULCANO, 1967: 10. Simulium (Psilopelmia) downsi: VARGAS & DÍAZ NÁJERA, 1957b: 154; COSCARÓN, 1995: 294-295, COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 76-78; CROSSKEY & HOWARD, 1997: 58; 2004: 53.

Female: Wing length 2.3 mm; general coloration yellow; scutum yellow with brownish pile, with 1+1 silvery submedian vittae reaching posterior border, and lateral border light gray, ornamented as in S. escomeli; scutellum light yellowish, metanotum brownish; pleurae light gray with brown episternum; legs yellowish with apex of femora, tibiae and tarsomeres blackish; abdomen with segments I-II yellowish, III-IX brownyellowish; tergites III-VIII with 5 longitudinal rows of blackish subquadrate spots as in S. escomeli: median row with wider spots, 1+1 sublateral and 1+1 lateral; the last with smaller spots; median spots absent on tergites VII-VIII; last segments shiny; cibarium with small

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A

B

E

F

J

M

C

G

K

D

O

H

P

I

N L

Q

R

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Figure 97. Larva. A-H: Cephalic apotome; A: S. (Psilopelmia) samboni; B: S. (P.) iracouboense; C: S. (P.) panamense; D: S. (P.) callidum; E: S. (P.) escomeli; F: S. (P.) downsi; G: S. (P.) mangabeirai; H: S. (P.) zempoalense. I: Mandible apex of S. (P.) ochoai. J-N: Hypostomium and postgenal cleft; J: S. (P.) dugesi; K: S. (P.) mediovittatum; L: S. (P.) mangabeirai; M: S. (P.) trivittatum; N: S. (P.) zempoalense. O-P: Antenna; O: S. (P.) panamense; P: S. (P.) downsi. Q-R: Anal sclerite; Q: S. (P.) iracouboense; R: S. (P.) pseudocallidum. S: Anal lobes and diverticula of rectal papillae of S. (P.) iracouboense.

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teeth on 1+1 submedian protuberances as in S. trivittatum; basal sector of R with 2-3 rows of hair; claw without subbasal tooth; paraprocts strongly acuminate mediodistally and with posterobasal protuberance (Fig. 92 I); eighth sternite with 20-28 setae on each side. Male: Wing length 2.1 mm; scutum coloration as in female, scutum with 1+1 submedian cuneiform stripes only reaching to half of scutum length; gonostylus with strong distal curvature (Fig. 93 N); ventral plate with body subtriangular (Fig. 94 H). Pupa: Cocoon with compact structure, slightly projected basally (Fig. 95 H); length basally 3.0-3.3 mm, dorsally 2.7-3.0 mm, gills 1.8-2.0 mm; frontoclypeus and thorax with moderate number of tubercles as in S. ochoai; trichomes single or bifid; gills with 8 branches, with branching not close to base (Fig. 96 H). Larva: Maximum length 5.3-5.5 mm; cephalic apotome darkened mediobasally with 1 light window medially and 1+1 subbasally (Fig. 97 F); antenna surpassing cephalic fan stem apex; ratio of proximal, medial and distal segments = 1:1-1.1:1.5 (Fig. 97 P); cephalic fan with 31-36 rays; postgenal bridge about same length as hypostomium; anal ring with about 70 rows, with 13-15 hooks each; rectal papillae with 4-6 diverticula on each lobe; anal sclerite without scales; comb of proleg lateral sclerite with 30-36 teeth. Distribution: Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Escuintla, Huehuetenango, Jalapa, Jutiapa, El Petén, El Progreso, El Quiché, Retalhuleo, San Marcos; Santa Rosa, Sololá, Suchitepequez; Mexico: Chiapas, Oaxaca; Venezuela: Miranda. Bionomics: Unknown. Discussion: Simulium escomeli is very close to S. downsi, but can be differentiated by the absence of hair on the basal sector of R, the darker female scutum, especially on the posterior area, the wider submedian silvery stripes, and the gonostylus without a distolateral outer expansion. The male of S. callidum is similar externally, but the ventral plate is subovoidal. Simulium (Psilopelmia) dandrettai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 91 Q; 92 J; 93 O; 94 I; 95 I, J; 96 I, J.) Simulium (Lanea) dandrettai VARGAS et al., 1946: 124-126; VARGAS & DÍAZ NÁJERA, 1948b: 323, 342-343. Simulium (Psilopelmia) dandrettai: VARGAS & DÍAZ NÁJERA, 1957b: 183; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 78-81; CROSSKEY & HOWARD, 1997: 58; 2004: 53.

Female: Wing length 2.4-3.0 mm; frons and clypeus with gray pruinosity; scutum brown with 1+1 submedian and 1+1 lateral whitish stripes bordering median and 1+1 sublateral brown to black vittae; whitish submedian vittae, in some illumination, shows dark transverse anterior spot, and with posterior light they join posteriorly, changing to gray-brown; scutum hair golden to silvery, according to light position; scutellum pale brown; metanotum dark brown grayish; legs brown yellowish with femorae, tibiae and tarsomeres dark brown to black; abdomen with tergites I-II brown yellowish with dark spot on median row and 1+1 slightly lighter laterally; VII-VIII shiny light brown grayish on median area and darker laterally; cibarium with median depression bordered by 1+1 submedian short projections apically covered by spines (Fig. 91 Q); fronto-ocular triangle higher than wide; basal sector of R without hair; claw without subbasal tooth; eighth sternite with 18-24 setae; paraproct short (Fig. 92 J).

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Male: Scutum black with 1+1 anterior submedian shiny silvery spots not surpassing 1/3 of scutum length, similar to S. veracruzanum in shape; in posterior illumination, with lighter subquadrate median posterior spot; abdomen black with 1+1 lateral silvery spots on tergites II-VI; ratio of length/width of hind basitarsus = 4; gonostylus very wide distally (Fig. 93 O); ventral plate rounded, without lateral constriction (Fig. 94 I). Pupa: Cocoon with weave consistent, reinforced on anterior margin (Fig. 95 I); length basally 2.7-3.0 mm, dorsally 3.0-3.1 mm, gills 3.0-3.2 mm; gills dark brown, with 6 branches, dorsal secondary and 2 dorsal tertiary branches longer and thicker than other branches (Fig. 96 I). Larva: Cephalic apotome dark on base, with light spots; ratio of proximal, medial and distal segments = 1:1.3:1.5-1.6; cephalic fan with 26-31 rays; postgenal cleft not deep, postgenal bridge longer than hypostomium, ratio of postgenal bridge/hypostomium = 1.2; anal sclerite with hair, but not scales; anal ring with 64-68 rows, with 13-14 hooks each. Distribution: Mexico: Distritop Federal, Mexico, Oaxaca, Veracruz. Bionomics: Simulium dandrettai breeds in small clear torrential creeks in mountain areas at 2000-3000 m, attached to aquatic vegetation and rocks. Female feedindg habits are unknown. Discussion: The pupal gill with 6 branches, thicker dorsal secondary and tertiary branches, apically enlarged gonostylus and rounded ventral plate permit S. dandrettai to be differentiated from other Neotropical S. (Psilopelmia) species. Material collected from the same place as S. dandrettai has cocoons with longitudinal low crest like folds (Fig. 95 J). The dorsal gill branches are not too much thicker than the other branches (Fig. 96 J) and the tubercles of the frontoclypeus and exposed portion of the thorax have a wrinkled apical border. COSCARÓN et al., 1996: 81 suggest that the material could be a different species, but it was not named as new until more material could be studied in order to confirm the differences; this point of view is maintained here. Simulium (Psilopelmia) zempoalense VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 93 R; 95 N, O; 96 Q; 97 H, N.) Simulium (Lanea) zempoaelense VARGAS et al., 1946: 107, 112,136, 138,168, 170-171. Simulium (Psilopelmia) zempoaelense: VARGAS & DÍAZ NÁJERA, 1957b: 183-184, 187-188; COSCARÓN, 1987: 32; COSCARÓN et al., 1996: 97-98; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

Female: Wing length 2.8-3.0 mm; thorax coloration similar to S. dandrettai; with lateral light sub-median vittae shown dark gray opaque; pleurae light gray; abdomen with dorsal spots ornamentation like S. escomeli; basal sector of R, claws and paraprocts as S. dandrettai but the last with basal lobe more prominent. Male: Wing length 2.5 mm; scutum black with 1+1 cuneiform vittae reaching thinly to posterior area; silvery pile; abdomen with 1+1 lateral silvery spots on tergites II, V-VI; hind basitarsus width and ventral plate shape as in S. dandrettai; gonostylus strongly acuminate apically (Fig. 93 R). Pupa: Cocoon with reinforced border (Fig. 95 N); length basally 3.6 mm, dorsally 3.2 mm, gills 2.5-3.6 mm; frontoclypeus and thorax with wrinkled platelets and sparse tubercles (Fig. 95 O); trichomes single to bifid; gills with 6 branches, with primary and secondary branches bifurcated near base (Fig. 96 Q) with branches about same width.

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Larva: Maximum length 6.0-6.5 mm; cephalic apotome light brown, without visible ornamentation (Fig. 97 H); ratio of proximal, medial and distal antennal segments = 1:1.3:1.8; cephalic fan with about 38-40 rays; postgenal cleft short, postgenal bridge longer than hypostomium length (Fig. 97 N); mandible with 2 mandibular processes; anal sclerite with numerous hairs, but not scales; anal ring with 76-80 rows, with 13-14 hooks each; rectal papillae with 9-12 diverticula on each lobe. Distribution: Mexico: Durango, Mexico, Morelos, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Simulium dandrettai is the closest species to S. zempoalense, with similar adults and pupae with wrinkled integument and 6 gill branches. There are small differences in the pupa such as the thicker dorsal external pro minence shown in S. dandrettai. It is necessary to carry out cytological and molecular studies of this species as well as other material close to S. dandrettai that might be considered possible new species (COSCARÓN et al., 1996). Simulium (Psilopelmia) trivittatum MALLOCH (Figs. 91 P; 92 N; 94 N; 96 P; 97 M.) Simulium trivittatum MALLOCH, 1914: 25, 30, 39, 68; DAMPF, 1927: 129; DYAR & SHANNON, 1927: 37; LANE & VULCANO, 1943: 439. Simulium (Psilopelmia) trivittatum: VARGAS & DÍAZ NÁJERA, 1957b: 154, 183, 185-186, 264, 360; PETERSON, 1993: 341; COSCARÓN et al., 1996: 94-97; CROSSKEY & HOWARD, 1997: 59; 2004: 53; MOULTON, 1998: 66; ADLER et al., 2004: 353. Simulium distinctum MALLOCH, 1913: 133-134 (preoc.). Simulium (Psilopelmia) mazzottii DÍAZ NÁJERA, 1979: 553-556.

Female: Wing length 2.0-2.5 mm; scutum ornamentation similar to that of S. bobpetersoni, with 3 brown-blackish longitudinal vittae delimited by 1+1 submedian and 1+1 lateral light yellowish stripes; on anterior border of sublateral vittae are 1+1 subtrianglular whitish shiny spots; scutellum and metanotum yellowish brown, the second changing to dark brownish black; legs yellowish, with apex of femora, tibiae and tarsomeres brown; abdomen yellowish with 5 rows of blackish subrectangular spots, 1 median and 2+2 at sides as is common in S. (Psilopelmia); median rows present from I to VI, lateral rows from III to VII; cibarium with median depression with sclerotized rim, flanked by 2 small protuberances with short spines (Fig. 91 P); basal sector of R glabrous; claw without subbasal tooth; eighth sternite with about 40 setae; paraproct with elongated distal projection (Fig. 92 N). Male: Wing length 1.8-2.3 mm; scutum dark brown to blackish with 1+1 yellowish submedian vittae reaching to posterior border; anterior border of submedian vittae with light grayish subcircular spot; abdomen with segments I-II yellow, the following blackish; sternites yellowish; gonostylus similar to that of S. escomeli but more elongated apically; ventral plate acuminate distally with rounded apex (Fig. 94 N). Pupa: Cocoon reinforced on anterior border, weave with threads evident; length basally about 3.0 mm, dorsally 2.7 mm, gills 2.7-3.0 mm; frontoclypeus and thorax with sparse tubercles; trichomes single to bifid; gills with 6 branches, with basal trunk and primary branches relatively short, median secondary branch elongated (Fig. 96 P). Larva: Maximum length 4.5 mm; cephalic apotome brownish basally, with light window medially. Antennae dark apically on medial and distal segments; ratio of proximal,

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medial and distal segments = 1:1.3:2; cephalic fan with 36 rays; postgenal bridge about same length as hypostomium (Fig. 97 M) Anal ring with about 68 rows, with 14 hooks each; rectal papillae with diverticulum on each lateral lobe. Distribution: Guatemala; Mexico: Coahuila, Tamaulipas; USA: Oklahoma, Texas. Bionomics: Simulium trivittatum breeds in shallow rivers and small ditches with trailing vegetation. Female feeding habits are not well known. There is only one citation of biting a human. Discussion: Simulium trivittatum is similar to S. gonzalezherrejoni, S. longithallum, S. mangabeirai, S. veracruzanum and S. bobpetersoni, but can be differentiated by the pupal gill of 8 branches, except S. mangabeirai, which has 4. As explained for S. bobpetersoni, some specimens of S. trivittatum can have a gill with 8 branches. Simulium (Psilopelmia) bobpetersoni COSCARÓN, IBÁÑEZ-BERNAL & COSCARÓN ARIAS (Figs. 92 K; 94 J; 95 K, U; 96 K.) Simulium (Psilopelmia) bobpetersoni COSCARÓN et al., 1996: 81-84; CROSSKEY & HOWARD, 1997: 58; 2004: 52; MOULTON, 1998: 66; ADLER et al., 2004: 353. Simulium (Lanea) trivittatum VARGAS, 1943: 363; VARGAS et al., 1946: 169. Simulium (Psilopelmia) trivittatum MALLOCH: DAMPF, 1927: 129; VARGAS & DÍAZ NÁJERA, 1957b: 183; DALMAT, 1955: 156-159.

Female: Wing length 2.2-2.9 mm; scutum with 3 blackish vittae as 1 median and 1+1 sublateral vittae flanked by 1+1 whitish submedian and 1+1 lateral stripes; scutellum and metanotum dark brown; abdomen with 1 median and 1+1 sublateral and lateral rows of dark brown subrectangular spots; cibarium with 1+1 small submedian protuberances with short spines similar to those of S. veracruzanum; basal sector of R glabrous; claw without subbasal tooth; eighth sternite with 28-30 setae; paraproct with elongated distal projection (Fig. 92 K). Male: Wing length 1.9-2.1 mm; scutum dark reddish brown, with 1+1 anterior submedian cuneiform white grayish stripes, generally reaching to half of scutum length but reaching to posterior border in some specimens; ratio of length/width of hind basitarsus = 7.5; gonostylus similar to that of S. escomeli but wider distally; ventral plate slightly projected distally (Fig. 94 J). Pupa: Cocoon with straight and reinforced anterior border (Fig. 95 K), length basally 2.7 mm, dorsally 2.8 mm, gills 2.2 mm; frontoclypeus and thorax with sparse tubercles (Fig. 95 U); trichomes short with 1-2 branches on frontoclypeus and 1-3 on thorax; gills with 8 branches, with primary branches relatively elongated (Fig. 96 K). Larva: Maximum length 5.5 mm; cephalic apotome darkened on median basal area, bell shaped, with 1 central and 1+1 basal light spots; antennae as long as or shorter than cephalic fan stem, ratio of proximal, medial and distal segments = 1:1.2:1.9; cephalic fan with about 28 rays; proleg lateral sclerite with about 22 teeth; anal sclerite without scales; anal ring with about 62 rows, with 11-12 hooks each; rectal papillae with 0-2 diverticula on each lobe. Distribution: Guatemala; Mexico: Aguascalientes, Chiapas, Chihuahua, Coahuila, Colima, Durango, Guanajuato, Guerrero, Jalisco, Michoacan, Morelos, Nueva León, Oaxaca, Puebla, San Luis Potosí, Tamaulipas, Veracruz. Bionomics: Unknown.

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Discussion: The closest species to S. bobpetersoni is S. veracruzanum, but this species has a shorter bifurcation of the primary and secondary branches and a ventral plate with the median distal portion more elongated. Males are similar to those of S. zempoalense but the pupal gill has 6 branches. Simulium trivittatum, of which Mexican material is considered to be S. bobpetersoni by COSCARÓN (1996), can be differentiated by the 8 pupal gill branches. ADLER et al., (2004) found S. trivittatum pupae with 8 branches; therefore, it would be useful to study more material and also to do cytological and molecular studies of this material. Simulium (Psilopelmia) veracruzanum VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 92 L; 93 D, P; 94 K; 95 L; 96 L.) Simulium (Lanea) veracruzanum VARGAS et al., 1946: 112, 133-136; DALMAT, 1951: 55-57; 1955: 33. Simulium (Psilopelmia) veracruzanum VARGAS & DÍAZ NÁJERA, 1957b: 184-187, 189; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 84-87; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

Female: Wing length 2.6-3.0 mm; scutum grayish with 1 median and 1+1 lateral dark brown vittae; with frontal light, appearing to join posteriorly and showing as silvery the 1+1 submedian light vittae; pile grayish yellow; scutellum light yellowish brown, metanotum dark grayish brown Pleura brownish yellow; legs yellowish gray with coxae, apex of hind femur, apex of tibiae and tarsomeres blackish; abdomen brown with tergites I-II yellowish, tergites II-VII grayish yellow with 5 rows of subquadrate dark brown grayish spots as 1 median, 1+1 sub-lateral and 1+1 lateral rows, as in S. escomeli. Basal sector of R, claw and cibarium as in S. trivittatum; paraproct very elongated distally (Fig. 92 L). Male: Wing length 2.7 mm; scutum with 1+1 silvery anterior cuneiform vittae reaching about half of scutum length (Fig. 93 D); scutellum and metanotum black; pleurae dark gray; legs brown-blackish with lighter areas on fore tibia, anterior half of middle tibia and tarsus, anterior third of hind tibia and anterior half of basitarsus; gonostylus with strong distal curvature (Fig. 93 P); ventral plate with deep lateral constrictions (Fig. 94 K). Pupa: Cocoon with reinforced anterior border (Fig. 95 L), length basally 3.3 mm, dorsally 2.8-3.0 mm, gills 3.0-3.4 mm; frontoclypeus and thorax with moderate number of smooth tubercles; cephalic trichomes with 1-2 branches, thorax with 2-5 branches; gills with 8 branches, with primary and secondary branches short (Fig. 96 L). Larva: Maximum length 5.5-6.0 mm; cephalic apotome mildly darkened basally, with light median and 1+1 subbasal areas; ratio of proximal, medial and distal antennal segments = 1:1.3-1.4:1.6-1.7; cephalic fan with 32-43 rays; postgenal bridge about same length as hypostomium; ninth segment with pair of ventral tubercles; anal sclerite with hair and abundant scales; anal ring with 72-76 rows, with 13-14 hooks each. Distribution: Belice; Guatemala: Acatenango, Chimaltenango; Mexico: Chiapas, Chihuahua, Coahuila; Durango, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Simulium veracruzanum females are similar to those of S. gonzalezherrejoni, S. longithallum, S. mangabeirai, S. bobpetersoni and S. trivittatum. The male ventral plate of S. veracruzanum with the strong lateral constrictions, allow it to be differentiated from those species. The primary and secondary pupal gill branches are very short and also help separate these species.

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Simulium (Psilopelmia) mangabeirai VARGAS (Figs. 91 F; 93 F, Q; 94 L; 95 M; 96 M; 97 G, L.) Simulium mangabeirai VARGAS, 1945b: 76. Simulium (Lanea) mangabeirai: VARGAS et al., 1946: 107; VARGAS, 1948: 309. Simulium (Psilopelmia) mangabeirai: VARGAS & DÍAZ NÁJERA, 1957b: 184; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 87-89; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

Female: Wing length 2.8-2.9 mm; scutum with 3 wide black vittae as 1 median and 2 sublateral vittae separated by 1+1 thinner light grayish submedian stripes; black sublateral vittae not reaching anterior border (Fig. 91 F); light submedian vittae, with lateral illumination, appears as shadow stripes, as in S. (Cerqueirellum) species; with posterior illumination, scutum has light grayish subquadrate area posteriorly; scutellum and metanotum brownish gray; pleura light brown-gray; legs brown-yellowish with blackish apex of femora, tibiae and tarsomeres; abdomen light grayish with 5 rows of blackish subquadrate spots on tergites from II-VII as in S. escomeli; cibarium, basal sector of R, claw and paraproct as in S. trivittatum, the latter with distal projection straight; eighth sternite with 24-28 setae. Male: Wing length 2.5-2.8 mm; scutum black with 1+1 anterior subtriangular spots, continued as thin stripes near posterior border (Fig. 93 F); hind basitarsus wide, ratio of length/width = 3.8; gonostylus without lateral projection and slight inner curvature (Fig. 93 Q); ventral plate with slight lateral constrictions (Fig. 94 L). Pupa: Cocoon slightly projected basally, with reinforced anterior border (Fig. 95 M), length basally 3.6-4.0 mm, dorsally 3.0-3.5 mm, gills 4.3-4.6 mm; frontoclypeus and thorax with moderate number of tubercles; trichomes with 1-2 branches; gills with 4 branches, longer than cocoon length, dorsal thicker than ventral branches (Fig. 96 M). Larva: Maximum length 6.5-7.0 mm; cephalic apotome slightly darkened on basal half, with median light window (Fig. 97 G); ratio of proximal, median and distal antennal segments = 1:1.4:1.5; cephalic fan with about 42 rays; postgenal cleft short, dome shaped (Fig. 97 L); postgenal bridge 1.7 longer than hypostomium length; lateral sclerite of thoracic proleg with 15-17 teeth arranged in 9-10 groups; anal sclerite with scales and hair; anal ring with about 89 rows, with 15-17 hooks each; rectal papillae with 8 diverticula on each lobe. Distribution: Mexico: Distritop Federal, Mexico, Michoacan, Morelos, Michoacán, Veracruz, Zempoala. Bionomics: Unknown. Discussion: Simulium mangabeirai is very similar, as adults, to S. trivittatum, S. veracruzanum, S. bobpetersoni and S. gonzalezherrejoni, but can be differentiated by its pupal gill of 4 branches. Also the ornamentation of the scutum in both sexes shows differences with the aforementioned species. Simulium (Psilopelmia) longithallum DÍAZ NÁJERA & VULCANO (Fig. 96 O.) Simulium (Psilopelmia) longithallum DÍAZ NÁJERA & VULCANO, 1961: 221; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 93-94; CROSSKEY & HOWARD, 1997: 59; 2004: 53; MOULTON, 1998: 61; ADLER et al., 2004: 351-352.

Female: Wing length 2.0-2.8 mm; scutum orange brown to brown-blackish with yellowish pile, 1+1 silvery-blue pruinose submedian vittae and lateral border; scutellum light

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brown-yellowish; pleural submedian vittae with yellowish brown pruinosity; legs yellowish with apex of femora, tibiae and tarsomeres dark brown; abdomen yellowish with dorsal dark spots arranged in median row on segments II-VII and 1+1 lateral rows on V-VIII; cibarium with median depression well sclerotized and laterally with rim covered by small teeth arranged as in S. trivittatum; basal sector of R without hair, and claws with subbasal tooth; eighth sternite with 25-28 setae; paraproct elongated and thin distally, similar to that of S. gonzalezherrejoni. Male: Wing length 2.0-2.6 mm; scutum velvety black with 1+1 silvery blue submedian vittae reaching posterior border, as in S. gonzalezherrejoni; scutellum and metanotum black; abdomen velvety black with golden pile; gonostylus half of gonocoxite length, with large posterolateral prominence, as in S. downsi; ventral plate wider than long, similar to that of S. downsi and S. gonzalezherrejoni. Pupa: Cocoon with small anterior dorsal and ventral projection, and reinforced anterior border; length basally 2.6-3.2 mm, dorsally 2.5 mm, gills 2.0-2.2 mm; frontoclypeus and thorax with tubercles of irregular size and in moderate number; trichomes with 2-3 branches; gill of 8 branches with primary, dorsal secondary and tertiary thicker than other branches (Fig. 96 O). Larva: Maximum length 5.0-5.8 mm; cephalic apotome darkened in median and basal areas, with light median longitudinal and 1+1 subbasal windows, very similar to that of S. escomeli; mandible with latero-mandibular process; antenna darker than stalk of cephalic fan; ratio of proximal, medial and distal segments = 1:1.2-1.3:1.8-2; cephalic fan with about 45 rays; postgenal cleft dome shaped; postgenal bridge slightly longer than hypostomium; lateral sclerite of thoracic proleg with about 22 teeth arranged in 5-7 groups; anal sclerite without scales; anal ring with about 68 rows, with 11-14 hooks each; rectal papillae with 1 diverticlum on each lobe. Distribution: Mexico: Jalisco, Morelos; USA: Arizona. Bionomics: Simulium longithallum breeds in small rivers with moderate to fast flow and sandy and rocky bottoms. Larvae are found on trailing vegetation, sticks and stones. Female feeding habits are unknown. Discussion: Simulium longithallum is closest to S. gonzalezherrejoni, as explained for that species. Simulium bobpetersoni is very similar in the thinner trunk and basal branches of the pupal gill. Simulium downsi is similar in the cibarium and genitalia of both sexes, but the adults are yellowish. Simulium (Psilopelmia) gonzalezherrejoni DÍAZ NÁJERA (Figs. 92 M; 93 G; 94 M; 96 N.) Simulium (Psilopelmia) gonzalezherrejoni DÍAZ NÁJERA, 1969: 22-28; COSCARÓN, 1987: 31; COSCARÓN et al., 1996: 89-93; CROSSKEY & HOWARD, 1997: 59; 2004: 53. Simulium (Psilopelmia) jerezense DÍAZ NÁJERA, 1969: 28-32. Simulium (Psilopelmia) netteli DÍAZ NÁJERA, 1969: 32-35. Simulium (Psilopelmia) letrasense DÍAZ NÁJERA, 1969: 35-36. Simulium (Psilopelmia) olympicus DÍAZ NÁJERA, 1969: 36-38. Simulium (Psilopelmia) vulcanoae DÍAZ NÁJERA, 1969: 38-40.

S. gonzalezherrejoni was redescribed by COSCARÓN et al., 1996 who established the synonyms. It shows very little difference from S. longithallum; the paraproct (Fig. 92 M),

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scutum ornamentation (Fig. 93 G) and ventral plate shape (Fig. 94 M) are similar. The biggest differences are in the more elongated primary gill branches (Fig. 96 N). It is necessary to obtain more material and, if possible, to use cytological and molecular methods to determine the status of these species. Distribution: Mexico: Aguascalientes, Zaacatecas. Bionomics: Simulium gonzalezherrejoni breeds in moderate- to fast-flowing creeks attached to trailing vegetation. Female feeding habits are undescribed. Simulium (Psilopelmia) panamense FAIRCHILD (Figs. 91 N, V; 92 F; 94 E; 95 V, W; 96 D; 97 C, O.) Simulium (Psilopelmia) panamense FAIRCHILD, 1940: 705. Simulium (Ectemnaspis) panamense: COSCARÓN, 1984: 314; 1987: 29. Simulium (Psilopelmia) panamense: COSCARÓN et al., 1995: 2297-298; COSCARÓN et al., 1996: 64-66; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

Female: Wing length 2.4 mm; scutum yellow-orange with 1+1 submedian silvery pruinose vittae, reaching scutellum base; lateral rim silvery; scutum hair brown-yellowish to blackish yellow, according to light position; scutellum yellow-lemon and metanotum yellow-brownish; legs yellowish, darkened on apex of femur, tibia and tarsomeres; abdomen without longitudinal rows of spots on tergites, as is typical in subgenus; tergites I-II brown to brown yellowish, the following dark brown, with light spot on tergite III; cibarium with median depression few evident and abundant teeth at sides until cornuae border (Fig. 91 N); basal sector of R with 1-2 rows of hair; claw with subbasal tooth (Fig. 91 V); eighth sternite with about 22 setae on each side; gonapophyses divergent, subovoidal with slightly sclerotized rim; paraprocts becoming slightly thin toward apex, and with well-developed basal process (Fig. 92 F). Male: Scutum yellow with 1+1 silvery anterior submedian cuneiform spots, scutellum, metanotum and abdominal tergites I-II yellowish, following abdominal tergites grayish brown; ratio of length/width of hind basitarsus = 4.4; gonostylus about as long as gonocoxite, with strong distal curvature, bigger than that of S. escomeli; ventral plate with large median process with parallel sides (Fig. 94 E). Pupa: Cocoon compact with reinforced anterior border, length basally 2.7-3.3 mm, dorsally 3.0-3.5 mm, gills 1.4-2.1 mm; frontoclypeus and thorax with wrinkled border (Fig. 95 W) numerous tubercles (Fig. 95 V); cephalic trichomes with 2-3 branches, most of them with curled apex, thoracic with 3-4 branches; gill with 20 branches, with branching close to base (Fig. 96 D). Larva: Maximum length 4.5-5.0 mm; cephalic apotome with median basal area brownish, with light median window (Fig. 97 C); antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1.4:1.4; shape as in figure 97 O; cephalic fan with 40-45 rays; mandible with 1 latero-mandibular process; postgenal cleft dome shaped; postgenal bridge about as long as hypostomium length; lateral sclerite of thoracic proleg with 26 teeth arranged in 9-10 groups; anal sclerite with scales and hair; anal ring with about 89 rows, with 14-16 hooks each; rectal papillae with 6 diverticula on each lobe, with 1 of these diverticula about 2 times thicker and slightly longer than the others. Distribution: Costa Rica; Panama: Coclé, Panama.

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Bionomics: Unknown. Discussion: Simulium panamense has similarites with other species with a banded yellow scutum, such as S. samboni and S. callidum, but the unique subtriangular and elongated paraproct, ventral plate with great median distal process, and pupal gill with 20 branches permit differentiation. Simulium panamense has characters very similar to those of S. (Ectemnaspis) species, such as in the cibarium, shape of the paraproct and ventral plate, presence of hair on the base of R and a subbasal tooth on the claw, creating some confusion at the supraspecific level. Simulium (Psilopelmia) callidum (DYAR & SHANNON) (Figs. 91 D, W; 92 G; 93 S; 94 F; 96 E; 97 D.) Eusimulium callidum DYAR & SHANNON, 1927: 16; LANEA & VULCANO, 1943: 437. Simulium (Lanea) callidum: DALMAT, 1955: 126-130. Simulium (Ectemnaspis) callidum: COSCARÓN, 1984: 315-316; COSCARÓN et al., 1995: 295-296. Simulium (Psilopelmia) callidum: VARGAS & DÍAZ NÁJERA, 1957b: 183; COSCARÓN, 1987: 30; COSCARÓN et al., 1996: 67-70; CROSSKEY & HOWARD, 1997: 58; 2004: 52. Eusimulium mooseri DAMPF, 1927: 127.

Female: Wing length 2.7-2.8 mm; scutum and scutellum yellowish, scutum with 1+1 silvery submedian thin vittae extended 2/3 to posterior margin (Fig. 91 D); metanotum brown-yellowish; pleurae yellow, darkened ventrally; legs yellowish; abdomen without rows of dark spots typical of subgenus, tergites of segments I-II and III yellowish, following segments dark brown, IV-V velvety, VI-IX shiny; cibarium with small median depression and lateral area slightly pronounced with small spicules, without teeth and spines, as in S. pseudocallidum; basal sector of R with 2 rows of hair; claw with subbasal tooth (Fig. 91 W); paraprocts subtriangular, becoming thin toward apex (Fig. 92 G). Male: Wing length 2.4-2.8; scutum yellow with 1+1 silvery anterior submedian cuneiform spots extending 1/3 of scutum length; metanotum and abdomen yellowish with abdominal segments III-IX grayish brown; ratio of length/width of hind basitarsus = 5.5; gonostylus relatively thick, about 3/4 of gonocoxite length (Fig. 93 S); ventral plate subovoidal, without lateral constrictions (Fig. 94 F). Pupa: Cocoon compact with threads evident, anterior border slightly reinforced with median basal projection; length basally 3.0-3.5 mm, dorsally 2.7-3.3 mm, gills 1.4-2.1 mm; frontoclypeus and thorax with abundant tubercles with smooth border; cephalic and thoracic trichomes with 2-3 long branches; gill with 8 branches, arising close to base and very open; first dorsal branch slightly thicker than others (Fig. 96 E). Larva: Maximum length 5.5-6.0 mm; cephalic apotome slightly darkened mediobasally (Fig. 97 D); antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal segments = 1:1:1.5-1.6; cephalic fan with 30-32 rays; postgenal bridge as long as hypostomium; lateral sclerite of thoracic proleg with about 50 teeth, arranged in 13-14 groups; anal sclerite with single hairs and abundant scales with numerous denticles similar to those of S. pseudocallidum; anal ring with about 105 rows, with 14-17 hooks each. Distribution: Belice; Colombia: Valle; Guatemala; Mexico: Cordoba, Morelos, Oaxaca, Veracruz.

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Bionomics: Simulium callidum breeds in slow-flowing streams. The larvae are found on fallen leaves and subaquatic vegetation. Females bite domestic mammals, birds and humans. Discussion: Simulium callidum has similarities with the yellowish species. Some characters distinguish it, such as the shorter and cuneiform submedian silvery stripes, cibarium with thick rim lacking teeth, paraproct subtriangular and ventral plate subovoidal. The presence of hair on the basal sector of R and a subbasal tooth on the claw, the presence of scales on the anal sclerite with several spiculae on the border, and the elevated number of hook rows on the anal ring also differentiate it. As for S. panamense, there are several characters shared with S. (Ectemnaspis) species, reinforcing the idea that there is a close relationship between S. (Ectemnaspis) and S. (Psilopelmia). Simulium (Psilopelmia) pseudocallidum DÍAZ NÁJERA (Figs. 91 O; 96 F; 97 R.) Simulium (Psilopelmia) pseudocallidum DÍAZ NÁJERA, 1965: 25-45; COSCARÓN, 1984: 316; 1987: 31; COSCARÓN et al., 1996: 70-73; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

This species is very close to S. callidum and here we mention only the differences. Female: Wing length 3.1-3.2 mm; metanotum black; abdomen with tergites I-V yellowish; cibarium generally without spicules on lateral prominences (Fig. 91 O); basal sector of R without hair. Male: Wing length 3.2 mm; ratio of length/width of hind basitarsus = 3.7. Pupa: Thoracic trichomes with 3-5 branches (Fig. 96 F). Larva: Cephalic fan with 40-42 rays; lateral sclerite of thoracic proleg with 35-38 teeth arranged in 12-14 groups; anal sclerite with scales and abundant hair (Fig. 97 R). Distribution: Mexico: Mexico, Oaxaca. Bionomics: Simulium pseudocallidum breeds in cold, fast-flowing creeks and is sympatric with S. veracruzanum and S. himmani. Female feeding habits are unknown. Discussion: The small differences could make S. pseudocallidum a variation of S. callidum. It is necessary to obtain more material in order to see how large the variation is among populations and also to do cytological and molecular studies to confirm its true identity. Simulium (Psilopelmia) travisi VARGAS, VARGAS L. & RAMÍREZ PÉREZ Simulium (Psilopelmia) travisi VARGAS et al., 1993: 175; CROSSKEY & HOWARD, 1997: 59; 2004: 53.

According to its description, S. travisi is very close to S. callidum and S. pseudocallidum; here we mention only the differences. Female: Cibarium without spicules on sublateral prominences. Male: Ventral plate with body laterally and basally slightly concave, contrary to convex shape in S. callidum and S. pseudocallidum. Pupa: Dorsal branch thicker and nearly 5 times longer than other filaments, which in the most similar species is only about 1.5 times longer. Larva: Ratio of proximal, medial and distal antennal segments= 1: 1.5: 2; cephalic fan with 40-43 rays. Distribution: Costa Rica: Alajuela, Heredia.

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Bionomics: Simulium travisi breeds in fast, turbulent streams at altitudes of 8001325 m, attaching to rocks and aquatic vegetation. No information is available on female feeding habits. Discussion: Simulium travisi is closest to S. callidum and S. pseudocallidum. Differences in the male ventral plate and length of the first dorsal pupal gill filament allow them to be separated. BLANCASI species group The S. (BLANCASI) species group is unique, characterized mainly by the uniformly gray leaden scutum in adults of both sexes, lack of denticles on the cibarium, simple claw of the female, elongated subcylindrical gonostylus and subtriangular and elongated ventral plate of the male, four long branches of the pupal gill and single rectal papillae of larvae. This combination of characters makes it very difficult to include the species in any known Simulium subgenus; the closest are S. (Ectemnaspis) and S. (Psilopelmia), two taxa with a tight relationship (COSCARÓN & MIRANDA ESQUIVELL, unpubl.). However, the similarities in the male and female grayish coloration and genitalia (gonostylus and ventral plate in male and paraproct in female) show a close relationship with S. (Psilopelmia), with great variation in coloration and morphology. Probably it is a desert lineage, with characters that distinguish it as a separate species group. Adult: Medium-sized (wing length, 2.2-3.2 mm); color of scutum in both sexes uniformly gray, except small area near anterior border; abdomen with conspicuous dark and light pattern; eighth and ninth tergites of female polished; female with frontoocular triangle well developed; cibarium glabrous between cornuae; mandibles and maxillae serrated on both edges; setae of scutum of uniform type, irregularly arranged, not forming lines or groups; pleural membrane and katepisternum bare; basal section of R bare in both sexes; R1 with setae and spines; fore tarsi narrow, approximately 8 times as long as wide; calcipala and pedisulcus present; hair on legs simple, not scale like; male genitalia with gonostylus about as long as gonocoxite, elongate, subconical distally, without projection at base, and with single apical spinule; ventral plate subtriangular, membranous, lacking median keel; median sclerite about as long as wide, with apical incision; endoparameres with several large hooks; gonapophysis small, glabrous; paraproct short, simple; spermatheca globular, not reticulate, its surface minutely punctate; internal spicules present. Pupa: Cocoon inverted slipper-shaped, lateral margins of anterior opening receding from dorsal to ventral surface; lateral apertures absent; gill with 4 long, slender filaments arising from 2 short bases or from common trunk; trichomes of head and thorax simple, very short; tergites II-V with 4+4 simple hooks, VII and VIII with spine combs; stemites V-VII each with 2+2 simple, bifid or trifid hooks; apex of abdomen lacking specialized setae, and with 1+1 apical tubercles. Larva: Abdomen gradually widening posteriorly, apex not truncate; central tubercles absent; head and cephalic fans normal; head pattern positive; antennae as long as stem of cephalic fan; medial antennal segment with 1 or 2 constrictions; apical tooth of mandible much larger than outer teeth; presence of 1 or 2 well-developed latero-mandibular

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processes; pre-apical teeth decreasing in size from first to third; 2 marginal teeth well developed; hypostomium with median and lateral teeth about equally prominent, intermediate teeth much smaller; hypostomial setae arranged in 1+1 single rows, these rows slightly diverging posteriorly from lateral margins of hypostomium; postgenal cleft either large or almost absent; cervical sclerite very small, isolated in neck membrane; lateral sclerite of proleg well developed, elongate subpentagonal; setae arranged in 1 row, clustered in groups in most cases formed of 3 or 4 long and short setae: overall size of setae increasing from upper to lower border of sclerite; cuticle of body bare; perianal setae present; rectal papillae composed of 3 simple lobes. Discussion: This group is formed of three closely knit species: S. blancasi WYGODZINSKY & COSCARÓN, S. tenuipes KNAB, and S. penai WYGODZINSKY & COSCARÓN. Simulium (Psilopelmia) blancasi WYGODZINSKY & COSCARÓN (Fig. 98.) Simulium blancasi WYGODZINSKY & COSCARÓN, 1970: 6-14; COSCARÓN, 1987: 55; 1991: 143.

Female: Wing length, 2.7-2.8 mm; frons and clypeus dull gray, with few scattered yellowish hairs; mouthparts piceous; maxillary palps grayish; antennae black, grayish pruinose; first, second, and base of third segment light orange; scutum dull gray; anterolateral angles and 1+1 anterior submedian spots silver-white; pleura and sterna with grayish pruinosity, faintly nacreous; pleural membrane dark brown; hair of scutum short, scattered, yellowish; scutellum yellowish brown, narrowly gray pruinose along margins; hair of scutellum from yellowish to black; metanotum glabrous, black, heavily gray pruinose; general color of legs yellowish white, dark pigment from piceous to black; pattern as in figures K-M; fore coxae light colored, mid and hind coxae dark; setae of legs colored with respect to background; anterior wing veins light grayish brown; setae at base of wing black; stem of halteres yellowish, knob silver-white; color pattern of abdomen as in figure A; first abdominal segment yellowish white, its fringe silver-white; second tergite blackish at center, sides with 1+1 very large silvery spots; third to fifth tergites velvety black, their center at anterior half with transverse dumbbell-shaped light-colored spot, pinkish or yellowish on third and fourth, silver-white on fifth segment that is larger than 2 before; these spots imperceptible in some specimens; sixth and seventh tergites silverwhite, dark only at extreme sides and in some cases also along posterior margin behind large silvery area, the latter occasionally indentate posteriorly at middle; eighth and ninth tergites piceous or black, not conspicuously lighter than anterior tergites; undersurface of abdomen grayish; fronto-ocular triangle (Fig. B) small, wider than deep; antennae 11 segmented, shape and proportions of segments as in figure C; frons as in figure D; cibarium as in figure E; maxillary palp as in figure F; apical segment as long as 2 preceding combined; third segment rather strongly widened; sensory vesicle (Fig. G) large, its diameter slightly larger than half width of third segment, its mouth wide; tubercles numerous; lacinia and mandibles serrate on both edges; lacinia (Fig. H) with 20-23 teeth, mandibles (Fig. I) with 33-35 teeth; scutum lacking scales, setae randomly distributed, not forming rows or groups; microtrichiae relatively long, of felt-like appearance. Sc and basal section of R glabrous. R1 with setae and spines arranged in 1 row. Rs with setae arranged in 1 row beginning slightly beyond base of vein (Fig. J); shape of legs and proportions of

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Figure 98. S. (Psilopelmia) blancasi. A-O: Female; A: abdomen, dorsal view; B: fronto-ocular triangle; C: antenna; D: frons; E: basal portion of cibarium; F: maxillary palp; G: sensory vesicle of maxillary palp; H: lacinia; I: mandible; J: wing anterior veins; K-M: fore, mid and hind leg, respectively; N: calcipala and tarsomere I; O: claw. P-W: Male; P: abdomen, lateral view; Q: sensory vesicle of maxillary palp; RT: fore, median and hind leg, respectively; U: calcipala and tarsomere I; V: gonocoxite and gonostylus; W: ventral plate. X-Z, a-c: Pupa; X: dorsal and lateral views; Y: gill; Z: frontoclypeus; a: cephalic trichomes enlarged; b: portion of thorax showing, at right, trichomes and tubercles with high magnification; c: abdominal chaetotaxy. d-g: Larva; d: cephalic apotome; e: antenna; f: mandibular teeth with 2 latero- mandibular processes; g: postgenal cleft and hypostomium.

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segments as in figures K-M; calcipala well developed, but not quite attaining level of pedisulcus (Fig. N); claws as in figure O; eighth stemite with about 25+25 setae; gonapophysis glabrous, inner border faintly sclerotized; setae and microtrichiae absent; cercus much wider than long, apex rounded; apex of paraproct subrectangular. Male: Wing length, 2.5-2.6 mm; frons and clypeus dull silver-gray; antennae black, faintly grayish pruinose; first, second, and basal third of third segments orange; eyes purple; scutum dull gray, faintly tinged with olivaceous; 1+1 submedian spots at anterior margin of mesonotum faintly silvery white; pleura and sterna gray; scutellum, metanotum and setae of thorax as in female; general color of legs yellowish white, dark pigment from piceous to blackish; pattern as in figures R-T; fore coxae light colored, mid and hind coxae dark; white reflections on leg segments absent; color of wings and halteres as in female; color of abdomen as in female, but all spots silver, and tergites VI and VII almost completely white (Fig. P); last segment of maxillary palp slightly longer than 2 preceding combined; sensory vesicle globular, its diameter equal to half width of third segment; structure of vesicle as in figure Q; chaetotaxy of thorax as in female; wing venation and chaetotaxy as in female; shape of legs and proportions of segments as in figures R-T, hind basitarsus 4.6 times as long as wide; calcipala as in figure U; setae of legs somewhat longer and more numerous than in female; gonocoxite wider than long; gonostylus approximately as long as gonocoxite, elongate conical, slightly over twice as long as wide, with 1 apical spine (Fig. V); ventral plate (Fig. W) subtriangular, membranous, only its short basal arms faintly sclerotized; keel absent; median sclerite broad, as long as wide, its apical incision occupying approximately 1/3 length of sclerite. Pupa: Cocoon (Fig. X) subconical, slipper-shaped, dorsal wall salient at middle, projecting somewhat beyond level of ventral border of anterior opening; ventral wall of cocoon delicate but complete; cocoon translucent, shining, threads perceptible but not coarse; color of cocoon light brown; length of cocoon along dorsal surface 3.0-3.5 mm, along ventral surface, 2.5 mm; length of pupa 2.8-3.0 mm, of gill up to 4.5 mm, viz., up to 1.5 times as long as body of pupa; gills composed of 4 forwardly directed slender filaments (Fig. Y); 2 primary branches arising from short common trunk; head and thorax of pupa with numerous small platelets, arranged in irregular groups as shown in figure Z; trichomes small, hair like, not branched (Fig. a); head with 2+2 frontal and 1+1 facial trichomes, occasionally only 1 frontal trichome on 1 side; dorsal surface of thorax on each side with 56 simple, very short, hair-like trichomes (Fig. b); abdomen as in Figure c; tergite I with few small setae; tergite II with 3+3 small, slender, simple spines accompanied laterally by characteristic group of 2 small spines and 1 long hair; tergites III and IV each with 4+4 simple hooks and few very short hairs; tergite V glabrous; anterior border of tergites VII and VIII with transverse row of 20-22 spiniform tubercles, these rows connecting laterally with areas of very small scalelike cuticular processes, the latter areas also present on tergites VI and IX; apex of abdomen with 1+1 distinct, slightly sclerotized tubercles; sternite IV with 1+1 groups composed of 2-4 simple short hairs; sternites V-VII with 2+2 hooks, closely approximated on V, more distant on VI and VII; most hooks bifid, occasionally trifid, outer ones frequently simple, especially on tergites VI and VII; sternites IV-VIII each with 1+1 groups of minute scalelike cuticular processes. Larva: Length of mature larva 6.0-6.5 mm; width of head capsule 0.7 mm; general body color whitish, greenish pigment dorsally and ventrally on anterior half of body; head

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yellowish white; pattern of cephalic apotome as in figure d; body integument glabrous, few short setae in caudal area; ventral tubercles absent; antennae as in figure e, faintly but distinctly pigmented throughout, medial segment lightest in color; medial segment with very deep constriction accompanied by unpigmented area; ratio of proximal, medial and distal segments = 1:1.1:1.25; cephalic fan with 40-45 rays; toothing of mandible as in figure f; hypostomium with anterior border heavily pigmented; median and lateral teeth prominent, intermediate teeth much smaller; sublateral teeth and lateral serrations very well developed; hypostomial setae arranged in single row, their number approximately 5 in each group; postgenal cleft deep, about as deep as postgenal bridge (Fig. g); anal sclerite as in S. tenuipes; anal ring with approximately 80 rows each composed of about 17 hooks; rectal papillae simple, composed of 3 lobes (Fig. h). Distribution: Chile: Tarapaca; Peru: Lima. Bionomics: Larval stages of S. blancasi live in small creeks about 1 m wide and 0.30 m deep, in crystalline waters, flowing over aquatic vegetation or stones, with a temperature of about 15 ºC. Discussion: The abdominal ornamentation of the female without transverse anterior light grayish spots on tergites III-V and the pupal gills without primary branches permit differentiation from S. tenuipes and S. penai, the closest species. Males of S. tenuipes also have a shorter ventral plate, without basal lateral constrictions. Simulium (Psilopelmia) tenuipes KNAB (Fig. 99.) Simulium tenuipes KNAB, 1914a: 21; SILVA FIGUEROA, 1917: 30; WYGODZINSKY & COSCARÓN, 1970: 1417; COSCARÓN, 1987: 19; 1991: 144-145.

Female: Wing length, 2.2-2.9 mm; color of head and thorax and their appendages as in S. blancasi. Abdomen (Fig. A) with tergites I and II as in S. blancasi, III-V dark velvety brown, VI and VII very light brown with 1+1 silvery white spots on VI, VIII and IX piceous Undersurface of abdomen grayish; fronto-ocular triangle, antennae, palpi, lacinia, sensory vesicle, mandible, and cibarium as in S. blancasi; basal portion of cibarium smooth (Fig. B); claws without subbasal tooth (Fig. C); wings and legs as in S. blancasi; calcipala about as long as wide (Fig. D); eighth sternite lightly pigmented, and with about 40 trichomes on each side (Fig. E); gonapophysis subtriangular and with abundant microtrichiae; genital fork with stem and lateral branches well sclerotized; spermatheca subovoidal (Fig. F); paraprocts subtriangular, slightly wider than long (Fig. G). Male: Wing length, 2.4-2.7 mm; color of head and thorax and their appendages very close to that of S. blancasi, but because of greater number of hairs on scutum, their golden color is more apparent; legs with tibiae not distinctly darkened at base; abdomen with color pattern very similar to that of female, but tergites VI and VII with 1+1 small sublateral silvery spots; shape of antennae, maxillary palp, and sensory vesicle as in S. blancasi; shape and proportions of segments of legs very similar to those of S. blancasi, but hind legs somewhat wider, hind basitarsus 3.7 times as long as wide; gonostylus elongated, subcylinderical, nearly as long as gonocoxite (Figs. H, I), ventral plate subtriangular (Fig. J); endoparameres with longer spines (Fig. I); median sclerite with median area translucent (Fig. I).

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Figure 99. S. (Psilopelmia) tenuipes. A-G: Female; A: abdomen, dorsal view; B: basal portion of cibarium; C: claw; D: calcipala and tarsomere I; E: eighth sternite, gonapophysis and genital fork; F: spermatheca; G: cercus and paraproct: right in posterior view, left mounted on slide. H-J: Male; H: gonocoxite and gonostylus, dorsal view; I: genitalia dorsal view showing gonocoxite and gonostylus, ventral plate, endoparameres and median sclerite; J: ventral plate. K-P: Pupa; K: lateral view; L: portion of thorax and gill; M-P: larva; M: postgenal cleft and hypostomium; N: lateral sclerite of thoracic proleg; O: anal sclerite; P: scales of anal sclerite area.

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Pupa: Color light brown, shape of cocoon as in figure K; length of cocoon along dorsal surface 2.3-2.9 mm, along ventral surface 1.8-2.0 mm; length of pupa proper 2.52.9 mm, of gills 2.1-2.5 mm, viz., not longer than body of pupa; gill composed of 4 forwardly and downwardly bent filaments (Figs. J, K) arising from short common trunk; trichomes and platelets of head as in S. blancasi; thorax with single trichomes (Fig. L); abdominal onchotaxy as in S. blancasi. Larva: Length of mature larva 6.0-6.8 mm; width of head capsule 0.65-0.70 mm; general body color whitish, with greenish pigment dorsally and on ventral surface of anterior half of body; antennae much as in S. blancasi; ratio of proximal, medial and distal segments = 1:1.1:1.2; cephalic fan with approximately 35 rays; primary teeth slender, spaced closely, secondary teeth more than half as long as primary ones; mandible much as in S. blancasi, with 2 or occasionally 3 serrations; hypostomium with 4 or 7 hypostomial setae in each row (Fig. M); postgenal cleft extremely reduced, almost imperceptible; lateral sclerites of thoraacic proleg as in figure N; anal sclerite with hair and small spicules (Figs. O, P); anal ring with about 80 rows each composed of 15-18 hooks. Distribution: Chile: Tarapacá; Peru: Tacna. Bionomics: Larvae and pupae were collected in ditches from leaves of Scirpus (Cyperaceae) and leaves of diverse aquatic plants, on branches of trees and on rocks. The water was slightly turbid, and the temperature was 16-18 °C. Simulium tenuipes attacks humans, mules, and horses. Discussion: This species can be most easily separated from S. blancasi in the larva, by its obsolete gular cleft, and in the pupa, by the secondary respiratory filaments that seemingly arise from a common trunk. The females and males of S. tenuipes have different abdominal ornamentation from S. blancasi and a distinct ventral plate shape, as indicated for that species. Simulium (Psilopelmia) penai WYGODZINSKY & COSCARÓN (Fig. 100.) Simulium penai WYGODZINSKY & COSCARÓN, 1970: 18; COSCARÓN, 1987: 19-20; 1991: 145.

Female: Wing length 2.8-3.2 mm; color of head and thorax and their appendages as in S. blancasi, except scutum and metanotum which are darker, contrasting more with pale yellow scutellum; pattern of abdomen similar to that of S. tenuipes but with posterior border of tergites III-V light colored, light-colored portion on V occupying entire posterior half; general structure of fronto-ocular triangle, antennae, maxillary palp with sensory vesicle, and cibarium as in S. blancasi; lacinia with approximately 20 teeth, mandible with 35 teeth; shape and pigmentation of segments of legs much as in S. blancasi, but calcipala much reduced, subtriangular; wings as in S. blancasi. Male: Wing length, 2.8 mm; color of head and thorax with their appendages as in S. blancasi, but scutum and metanotum darker; color pattern of abdomen unknown; structure of head and appendages and of wings as in S. blancasi; hind basitarsus 4.0-4.5 times as long as wide; calcipala (Fig. A) similar to that of female; genitalia similar to those of S. blancasi, but gonostylus (Fig. B) slightly wider. Pupa: Color and shape of cocoon much as in S. blancasi, but more closely woven and, when observed in liquid, somewhat parchment-like, viz., with lead like sheen; length of cocoon along dorsal surface 2.8-3.5 mm., along ventral surface 2.6-3.0 mm; length of

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C A

B

E D

G

F

Figure 100. S. (Psilopelmia) penai. A-B: Male; A: obsolescent calcipala and pedisulcus; B: gonostylus. C-E: Pupa; C: dorsal portion of thorax and gill; D: frontoclypeus; E: trichomes and tubercles of thorax with high magnification; G-F: larva; F: mandibular marginal teeth; G: cephalic capsule, ventral view.

body of pupa 3.4-3.7 mm., of gills 3.8-4.0 mm; gill either straight or downwardly curved (Fig. C); head (Fig. D) and thorax without or with only few scattered platelets (Fig. E); abdominal onchotaxy as in S. blancasi. Larva: Length of mature larva, 6.5-7.0 mm.; width of head capsule, 0.65-0.70 mm; overall color of body whitish; pattern of cephalic apotome as in S. blancasi; antennae similar to those of S. blancasi; ratios of proximal, medial and distal segments = 1:1.1:1.2; cephalic fan with 38-42 rays Primary teeth slender, not spaced closely, secondary teeth somewhat over half as long as primary ones; mandible and hypostomium much as in S. blancasi; marginal teeth as in figure F; postgenal cleft extremely reduced (Fig. G); anal sclerite as in S. tenuipes; anal ring with 76-80 rows, each composed of 13-17 hooks. Distribution: Chile: Antofagasta. Bionomics: The larvae and pupae were collected on aquatic plants in a small, slightly brackish stream. Discussion: Simulium penai seems to be closer to S. tenuipes than to S. blancasi. It agrees with the former in the structure of the gills and virtual disappearance of the postgenal cleft, a specialized character in this group. The pupae of S. tenuipes can be identified by the more abundant platelets and the adults by the bigger calcipala. Species inquirenda Simulium jacobsi DALMAT Simulium (Lanea) jacobsi DALMAT, 1953: 40; 1955: 149. Simulium (Psilopelmia) jacobsi: CROSSKEY & HOWARD, 1997: 59; 2004: 53.

This species was described based only on the larva that, according to the author, is similar to S. haematopotum and S. samboni. The characters described do not provide security

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for differentiating it from other species. It will be necessary to obtain more material of other stages to determine its identity; until then, we maintain S. jacobsi as species inquirenda. Simulium (Aspathia) ENDERLEIN (Figs. 101-112; 150.) Aspathia ENDERLEIN, 1935: 359 (as genus). Type-species: Simulium hunteri MALLOCH, 1914: 59. Jalacingomyia PY-DANIEL, in PY-DANIEL & MOREIRA SAMPAIO, 1994: 127, 129-137. Type species: Simulium anduzei VARGAS & DÍAZ NÁJERA, 1948b: 328.

Adults: Female and male blackish. Frontal pruinosity dull blackish to shining black; scutum blackish with color pattern variable, but generally with 2+2 shining silvery nacreous longitudinal stripes to 1+1 subquadrate diffuse silvery pruinose anterior spots in females and usually 1+1 silvery diagonal anterior stripes in males, with some variation, sometimes with greenish blue iridescence or silvery pruinose areas; female pleural membrane and katepistemum bare, sometimes shining; abdomen blackish without special ornamentation, tergites V-IX shining; male with typical 1+1 silvery spots on both sides. Wing with basal section of R bare. Base of cibarium smooth but sometimes with small nodular granulations or with median projection that can be covered with small denticles at apex. Frons slightly convergent below. Fronto-ocular triangle deep. Arms of furcastemum with conspicuous projections. Sensory vesicle of palpus well developed, about 1/3 to 1/2 length of palpomere. Legs generally with petaloid hair; tarsal claws with strong basal tooth. Female gonapophysis distally truncate to bluntly rounded; paraprocts about as long as wide at base, slightly acuminate ventrally or rounded distally as are cerci. Genital fork with lateral branches relatively short but apodemes elongate and acute distally, spermatheca well developed, with inner spiculae. Male gonocoxite subquadrate; gonostylus about 1.5 times as long as gonocoxite, subparallel-sided, apex rounded, with subterminal spine, and internal basal process frequently strong and as long as transverse diameter of gonostylus basally; ventral plate generally with narrow curved body that is interspecifically variably shaped, sometimes hairy, and narrower than basal arms; parameres large, with numerous robust hooks. Pupa: Cocoon generally slipper shaped, with distinct threads, sometimes with reinforced edge and projecting dorsally, covering only gill bases. Gill with 5-130 filamentous branches to membranous with 3-24 terminal branches. Frontoclypeus with 2+2 frontal and 1+1 facial trichomes, absent on cephalic, epicranial, genal and ocular areas. Thorax with 5 multibranched to 25-140 simple or bifurcated trichomes on disc. Tubercles of frontoclypeus and thorax variable, smooth to rugose, abundant to sparse or absent. Abdominal onchotaxy with following cuticular processes dorsally on each side of tergites: I with 1 short median and 1 long sublateral seta; II with 3 median, 3 submedian (that are more robust, like small hooks in some species), and 3 lateral setae; III and IV with 4 large hooks near posterior margin and 1 median, 1 submedian and 2 or 3 lateral setae on each tergite; V with 3 short submedian setae and small scales on anterior edge present also on following posterior tergites; VI with 2 submedian and 2 lateral setae and 24 small spines on anterior edge as on tergites VII-VIII that have 8-10 and 14-16 spines, respectively; IX with short and robust distal hook. Ventrally, sternites I-II without setae; III with 4 submedian and 1 lateral seta; IV with submedian simple or bifurcated hook plus

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2 submedian and 3 lateral setae; V with 2 submedian trifurcated hooks and 2 setae plus 2 lateral setae; VI-VII with submedian bifurcated or trifurcated and 1 sublateral simple to bi- or trifurcate hooks plus 1 submedian and 1 or 2 sublateral setae. III-VII with small rugosities like scales on submedian area; VIII with 1 or 2 sublateral setae. Larva: Cephalic apotome variable, without unusual ornamentation, frequently with positive spots on median longitudinal area and 1+1 anterior groups with subbasal position, or with negative spots and gradually darkened basally. Postgenal cleft shallow, internal edge rounded or subacute, postgenal bridge generally longer than height of hypostomium. Hypostomial teeth well differentiated, median tooth at same level as lateral teeth, 3-9 lateral serrations on each side with 4-9 lateral setae. Mandible normal; inner teeth 8-12, generally in 2 rows; 2 marginal teeth present, the posterior one very small; lateral mandibular process generally present. Antenna generally longer than stem of cephalic fan. Antennal segments medial and distal with transverse striations evenly sclerotized, without light areas and medial segment generally longer than proximal and distal. Cephalic fan with 29-45 primary rays. Cervical sclerites small and free. Body shape normal. Thoracic cuticle bare. Sclerite of proleg with 30-36 teeth, generally arranged in groups. Anal sclerite X-shaped, without struts, generally with simple and sparse hair, rarely with scales. Anal ring generally with 66-90 rows but may be up to 109, with 12-15 hooks on each. Rectal papillae with 3 lobes and 0 to 13 branches on each lobule. Body without hair or scales. Body generally with ventral tubercle on ninth segment. This subgenus in the Neotropical realm is composed of 25 species, which occur at elevations of 500-3700 m. Several species are anthropophilic and vectors of the agent of human onchocerciasis. The Neotropical species are in the METALLICUM and HUNTERI species groups (CROSSKEY & HOWARD, 1997, 2004). KEYS

TO

NEOTROPICAL SIMULIUM (ASPATHIA)

SPECIES

Females Simulium covagarciai, S. kompi, S. ruizi and S. sandyi are excluded due to incomplete information. 1 – 2(1)

– 3(2)

Scutum with 1+1 anterior grayish spots not continued as light grayish vittae ..... 2 Scutum with 3 light nacreous or silvery stripes (Figs. 101 A, F) between 5 blackish vittae (Figs. 101 B, G) ....................................................................................................... 5 Scutum with 1+1 whitish subtrapezoidal anterior spots (Fig. 101 H); cibarium with median basal area smooth, without median process or reinforcement on edge (Fig. 102 C); paraprocts scarcely produced ventrally, with rounded edge (Fig. 103 D) .....................................................................................................................Simulium putre Scutum with 1+1 whitish anterior spots not subtrapezoidal; cibarium with median process or reinforced on edge; paraprocts produced ventrally and gradually acuminate at apex (Figs. 103 J, K) .................................................................................. 3 Scutum with 1+1 anterior subtriangular silvery spots; cibarium without median process or only slightly elevated medially (Fig. 102 G) ............................................... ............................................................................................. Simulium hechti, Simulium wygoi

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– 6(5) – 7(6) – 8(7) – 9(6) – 10(9) – 11(10) – 12(10) – 13(12) – 14(12) –

473

Scutum with 1+1 anterior subcircular silvery spots or arranged as continuous anterior band; cibarium with median process (Fig. 102 O) ...................................... 4 Small species, wing length about 3.0 mm; cibarium with sparse denticles (Fig. 102 O) .................................................................................................................. Simulium bustosi Larger species, wing length about 3.8 mm; cibarium without denticles .................. .................................................................................................................. Simulium costalimai Scutum, when viewed with lateral illumination, with pollinose iridescent area; paraprocts produced ventrally and acuminate at apex (Fig. 103 B) .......................... .................................................. Simulium metallicum, Simulium horacioi, Simulium jobbinsi, Simulium morae, Simulium puigi, Simulium racenisi Scutum, when viewed with lateral illumination, not iridescent; paraprocts produced or not produced ventrally .............................................................................. 6 Relatively large species; wing length 3.3-4.0 mm; cibarium smooth, without denticles, papillae or median process; sensory vesicle of palp less than 1/2 length of palpomere ...................................................................................................................... 7 Medium to small species; wing length less than 3.2 mm; cibarium smooth or not; sensory vesicle of palp generally 1/2 or more length of palpomere ..................... 9 Wings with hair on basal section of R ............................................... Simulium anduzei Wings without hair on basal section of R .................................................................... 8 Wing length 4.0 mm; sensory vesicle of palp slightly less than 1/2 length of palpomere ............................................................................................... Simulium marquezi Wing length 3.4 mm; sensory vesicle of palp 1/2 length of palpomere ................. ...................................................................................................................... Simulium tricorne Cibarium smooth; wing length 2.0-2.5 mm; subbasal tooth of tarsal claw absent or very small ................................................................................................. Simulium spilmani Cibarium with denticles or spherules; wing length 2.7-3.0 mm; subbasal tooth of tarsal claw well developed (Fig. 102 M) ...................................................................... 10 Cibarium without median process, with nearly straight edge; paraproct shorter than wide at base (Fig. 103 G) ...................................................................................... 11 Cibarium with median process (Figs. 102 J-L); paraprocts about as long as wide .............................................................................................................................................. 12 Cibarium with papillae (Fig. 102 N); paraproct not acuminate distally; sensory vesicle of palp about 1/2 length of palpomere .................................. Simulium parrai Cibarium with small acuminate denticles (Fig. 102 F); paraproct gradually acuminate distally (Fig. 103 G); sensory vesicle of palp less than 1/2 length of palpomere (Fig. 101 L) ............................................................ Simulium matteabranchium Paraproct rounded distally; sternite VIII with 29-35 setae ..................................... 13 Paraproct acuminate distally; sternite VIII with 20-25 setae .................................. 14 Inner edges of gonapophyses divergent below .................................... Simulium piperi Inner edges of gonapophyses parallel .............................................. Simulium jacumbae Inner edges of gonapophyses divergent below .................................. Simulium iriartei Inner edges of gonapophyses parallel to gradually divergent below ........................ .......................................................................................................................Simulium hunteri

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Males The following species are not considered due to lack of information: Simulium covagarciai, S. sandyi, S. racenisi, S. kompi. 1 – 2(1) – 3(2) – 4(3) – 5(3) – 6(5) – 7(5) –

8(1) – 9(8) –

Ventral plate subquadrate, not laterally flattened (Figs. 106 A-L) ........................... 2 Ventral plate subconical, subcircular to subcylindrical, gradually flattened from base (Figs. 106 M-Q) ......................................................................................................... 8 Gonostylus with inner basal process long and slender; scutum with 1+1 anterior short subtriangular silvery spots (Fig. 104 I) .......... Simulium jobbinsi, Simulium piperi Gonostylus with inner basal process relatively short and stout, frequently wrinkled and dentate apically; scutum variable; 1+1 short anterior to long convergent stripes diagonally directed to midline (Figs. 104 A, B) .............................................. 3 Scutum with 1+1 submedian and 1+1 sublateral silvery spots or bands (Figs. 104 L-N) ...................................................................................................................................... 4 Scutum with 1+1 silvery spots of variable shape, subtriangular to subquadrate or subrhomboidal, or stripe-like ........................................................................................... 5 Median process of ventral plate wide basally (Fig. 106 G) ................. Simulium puigi Median process of ventral plate narrow basally (Fig. 106 L) ............ Simulium hechti Scutum with 1+1 silvery diagonal spots projecting posteriorly (Figs. 104 A, B, L) ................................................................................................................................................ 6 Scutum with 1+1 silvery spots not diagonal but nearly transverse (Fig. 104 D) ................................................................................................................................................ 7 Ventral plate with narrow, short median process (Fig. 106 I); scutum with 1+1 short and basally stout silver spots (Fig. 104 J) ................................... Simulium parrai Ventral plate with medium to robust process (Figs. 106 A, B); scutum with 1+1 elongate and distally slender silver spots (Figs. 104 A, B) .......................................... ................................................. Simulium metallicum, Simulium horacioi, Simulium spilmani Ventral plate with robust median process edged by cleft, and lateral margins curved (Figs. 106 D, E); gonostylus inner basal process short and smooth (Fig. 105 F) ........................................................................ Simulium marquezi, Simulium tricorne Ventral plate with medium-sized median process not edged by cleft, and lateral margins straight (Figs. 106 C, H, K); gonostylus inner basal process medially long and dentate distally (Figs. 105 G, K) ................................................................................ ........ Simulium bustosi, Simulium iriartei, Simulium jacumbae, Simulium morae, Simulium wygoi Scutum without isolated silver spots or stripes, entire disc silvery margined (Fig. 104 K) ...................................................................... Simulium costalimai, Simulium anduzei Scutum with silver spots or stripes ................................................................................ 9 Scutum with 1+1 submedian anterior subquadrate to subrhomboidal silvery spots joining silver lateral area; body of ventral plate as wide basally as median transverse diameter (Fig. 106 M) ...........................................................Simulium hunteri Scutum with 1+1 submedian anterior silvery subtriangular to subtrapezoidal spots or diagonally curved bands (Figs. 104 E, G), not joining silver lateral area; body of ventral plate narrower basally than median transverse diameter (Figs. 106 O-Q) ................. 10

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10(9)

Gonostylus with distinct curvatures in lateral view (Fig. 105 E); body of ventral plate strongly flattened laterally from base, subcylindrical (Fig. 106 Q) .................. ..........................................................................................................................Simulium putre – Gonostylus with subparallel straight sides in lateral view; body of ventral plate not strongly flattened laterally, subconical (Figs. 106 J, P) ............................................. 11 11(10) Gonostylus inner basal process shorter than basal transverse diameter .................. .......................................................................................................................... Simulium ruizi – Gonostylus inner basal process longer than basal transverse diameter (Fig. 105 H) ...................................................................................................... Simulium matteabranchium Pupae 1 – 2(1) – 3(2) – 4(3) – 5(3)

– 6(2) – 7(6) – 8(6) – 9(1)

Gill stout and membranous (Figs. 107 H, J, 109 A-C, G, I) ..................................... 2 Gill filamentous (Figs. 107 A, C, F) ............................................................................... 9 Gill without filaments; frontoclypeus and thorax with tubercles ............................ 3 Gill with flexible filaments (Figs. 109 A, E, I); frontoclypeus and thorax without tubercles ............................................................................................................................... 6 Gill branches digitiform, without small tubuliform secondary branches; cocoon without anterior projection (Fig. 107 H) ....................................................................... 4 Gill branches tubuliform with small tubuliform secondary branches (Figs. 109 B, C); cocoon with (Fig. 107 I) or without anterior projection ..................................... 5 Gill with 3 branches directed anteriorly, with annulations; cocoon without dorsal carina ............................................................................................................. Simulium kompi Gill with 2 branches directed anteriorly and 1 directed posteriorly, without annulations (Fig. 107 H); cocoon with dorsal keel ........................... Simulium tricorne Gill with 2 globose, basally subcylindrical branches that are progressively narrower distally and main branch subdivided distally into 7 small and narrower branches (Fig. 109 B); abdominal tergites I and II with tubercles ............................ ................................................................................................................... Simulium marquezi Gill arborescent, with 25 relatively narrow terminal branches emerging from 3 short, stout primary branches (Fig. 109 C); abdominal tergites I-II without tubercles ....................................................................................................... Simulium sandyi Gill with main branch subdivided distally (Fig. 107 J) ............................................... 7 Gill subdivided basally into 4 large branches ............................................................... 8 Gill distally with main branch and basally with only 1 short branch (Fig. 109 E) .......................................................................................................................... Simulium ruizi Gill with robust stalk expanded distally, with several small and short branches and basally with 3 short branches (Fig. 109 A) ......................... Simulium matteabranchium Gill branches with pseudosegmentation, 2 branches directed anteriorly and 2 ventrally (Fig. 109 I) .................................................................................. Simulium parrai Gill branches without pseudosegmentation, the largest directed anteriorly, with small subtriangular protuberances and 2 ventral subglobose protuberances .......... ................................................................................................................. Simulium covagarciai Gill terminal branches greater than 8.5 mm; frontoclypeus and thorax with simple trichomes (Figs. 108 G, H) ............................................................................................. 10

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Gill terminal branches less than 3.5 mm; frontoclypeus and thorax with simple or multibranched trichomes ................................................................................................ 12 Gill with 3 stout primary branches, subdivided into 7-10 main branches and 85100 small terminal branches (Fig. 108 H); thorax with about 50 trichomes on each side ............................................................................................................... Simulium iriartei Gill with about 110-135 branches (Fig. 108 G); thorax with 70-100 trichomes on each side ............................................................................................................................. 11 Frontoclypeus and thorax without tubercles; cocoon with dorsal exterior projection ............................................................................................... Simulium costalimai Frontoclypeus and thorax with moderate number of tubercles; cocoon with straight anterior edge ................................................................................Simulium hunteri Gill with 5 branches (Fig. 108 E) ....................................................... Simulium spilmani Gill with 6-30 branches ................................................................................................... 13 Gill with 6 branches about same length (Fig. 108 A); cocoon with straight edge (Figs. 107 A, C) ................................................................................................................. 14 Gill with 10-30 branches; cocoon generally with anterodorsal projection or aperture circle-shaped anteriorly................................................................................... 16 Frontoclypeal and thoracic trichomes with 2-6 branches. (Figs. 107 B, 108 A), thorax with 5 trichomes on each side .......................................................................... 15 Frontoclypeus and thorax with simple trichomes, those of thorax number 60-85 on each side (Figs. 107 C-E) ......................................................................Simulium putre Cocoon flattened, subcircular in dorsal view .................................... Simulium horacioi Cocoon normal, not flattened (Fig. 107 A) .................................................................... .... Simulium metallicum, Simulium jobbinsi, Simulium morae, Simulium puigi, Simulium racenisi Cocoon flattened, subcircular, with anterior edge fused ventrally (Fig. 107 F); gill with 10-16 terminal branches ........................................................................................ 17 Cocoon with normal shape, not flattened, with or without anterodorsal projection; gill with 10-30 branches ............................................................................. 19 Gill with 10 terminal branches, with short stem at base and 4 primary branches; thorax with 25 trichomes on each side, their diameter diminishing gradually toward apex ................................................................................................ Simulium bustosi Gill with 2 or 3 main branches, with 9-11 narrower flexible branches; thorax with 5-120 trichomes that are very stout in their basal 1/3 (Fig. 107 K) ..................... 18 Gill with 3 main branches; thorax with 100-200 trichomes; thorax smooth (Fig. 108 J) .............................................................................................................. Simulium hechti Gill with 2 main branches; thorax with 5 trichomes on each side; thorax wrinkled (Fig. 108 I) .................................................................................................... Simulium wygoi Cocoon with anterodorsal projection; thorax with 25-65 trichomes on each side .............................................................................................................................................. 20 Cocoon without anterodorsal projection; thorax with 5+5 trichomes on each side ..................................................................................................................... Simulium anduzei Gill with 10 terminal branches; thorax with 50-65 trichomes .......... Simulium piperi Gill with 25 terminal branches (Fig. 108 F); thorax with 25-35 trichomes ............. ................................................................................................................... Simulium jacumbae

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Larvae Not included: Simulium costalimai, S. covagarciai, S. kompi, S. matteabranchium, S. morae, S. racenisi and S. sandyi. 1

Postgenal cleft only indented or scarcely developed; ratio of hypostomium length/postgenal bridge = 0.4-0.7 (Figs. 112 A-D) .................................................... 2 – Postgenal cleft scarcely to well developed; ratio of hypostomium/postgenal bridge = 0.8-1.4 .................................................................................................................. 5 2(1) Medial antennal segment shorter than distal, and proximal segment generally longer or equal to medial (Fig. 111 B) ........................................................................... 3 – Medial antennal segment longer than distal, and proximal segment shorter than medial (Fig. 111 L) ............................................................................................................. 4 3(2) Rectal papillae lobes without diverticula (Fig. 111 G) .......................Simulium putre – Rectal papillae lobes with several diverticula ............................................................. .................................................................................. Simulium anduzei, Simulium jacumbae 4(2) Anal disc with about 106 rows; inner teeth of mandible 13-15 ........................... ................................................................................................................... Simulium marquezi – Anal disc with 80 or less rows; inner teeth of mandible 10-12 ................................. .......................................................................................... Simulium piperi, Simulium tricorne 5(1) Hypostomium larger than postgenal bridge ................................................................. 6 – Hypostomium as short as or shorter than postgenal bridge (Figs. 112 B, C, G) ................................................................................................................................................ 7 6(5) Ratio of hypostomium/postgenal bridge = 1; anal ring with about 90 rows of hooks ............................................................................................................... Simulium ruizi – Ratio of hypostomium/postgenal bridge = 1.2-1.4; anal ring with about 78-80 rows of hooks ............................................................ Simulium parrai, Simulium spilmani 7(5) Scales on anal sclerite present ........................................................................................... ........................ Simulium metallicum, Simulium jobbinsi, Simulium horacioi, Simulium puigi – Scales on anal sclerite absent ........................................................................................... 8 8(7) Medial antennal segment shorter than proximal and distal segments ...................... ....................................................................................................................... Simulium bustosi – Medial antennal segment longer than proximal and sometimes distal segment as well (Figs. 111 H, N) ......................................................................................................... 9 9(8) Cephalic apotome without spots ............................................................. Simulium hechti – Cephalic apotome with median and submedian positive or negative spots ........ 10 10(9) Rectal papillae with 13-14 diverticula on each lobe ............................ Simulium wygoi – Rectal papillae with 5-8 diverticula on each lobe ...................................................... 11 11(10) Ratio of proximal, medial and distal antennal segments = 1:1.1:1.2 ........................ ....................................................................................................................... Simulium iriartei – Ratio of proximal, medial and distal antennal segments = 1:2.1-2.2:1.3-1.5 .......... .......................................................................................................................Simulium hunteri

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CLAVES

DE ESPECIES DE

SIMULIUM (ASPATHIA)

Hembras Simulium covagarciai, S. kompi, S. ruizi y S. sandyi excluidos por escasa información. 1 – 2(1)

– 3(2) – 4(3) – 5(1) – 6(5) – 7(6) – 8(7) – 9(6)

Escudo con 1+1 manchas grisáceas no continuadas como bandas claras ............ 2 Escudo con 3 bandas nacaradas claras o plateadas (Figs. 101 A, F) entre 5 bandas negras (Figs. 101 B, G) ...................................................................................................... 5 Escudo con 1+1 manchas blanquecinas subtrapezoidales anteriores (Fig. 101 H); cibario con área media basal lisa, sin prolongación mediana o reforzada el borde (Fig. 102 C); paraproctos escasamente desarrollados ventralmente, con borde redondeado (Fig. 103 D) ............................................................................Simulium putre Escudo con 1+1 manchas blanquecinas no subtrapezoidales; cibario con un proceso mediano o reforzado sobre el borde; paraproctos con desarrollo ventral y aguzado en el ápice (Figs. 103 J, K) ............................................................................... 3 Escudo con 1+1 mancha plateada subtriangular; cibario sin una prolongación mediana o solo ligeramente elevado medianamente (Fig. 102 G) ............................. ............................................................................................. Simulium hetchi, Simulium wygoi Escudo con 1+1 manchas plateadas subcirculares o dispuestas como una banda continua anterior; cibario con una pequeña prolongación (Fig. 102 O) ................ 4 Especies pequeñas, largo ala 3,0 mm; cibario con pequeños dientes esparcidos (Fig. 102 O) ................................................................................................ Simulium bustosi Especies grandes, largo ala aproximadamente 3,8 mm; cibario sin dientes ............ .................................................................................................................. Simulium costalimai Escudo visto con iluminación lateral con área polinosa iridiscente; paraproctos desarrollados ventralmente y aguzados en el ápice (Fig. 103 B) ................................ .... Simulium metallicum, Simulium horacioi, Simulium morae, Simulium puigi, Simulium racenisi Escudo visto con iluminación lateral no iridiscente; paraproctos desarrollados o no ventralmente .................................................................................................................. 6 Especies relativamente grandes; largo ala 3,3-4,0 mm; cibario liso, sin dentículos, papilas o prolongación mediana; vesícula sensorial del palpo menos de 1/2 del largo del palpómero ........................................................................................................... 7 Especies medianas a pequeñas; largo ala menos de 3,2 mm; cibario liso o no; vesícula sensorial del palpo generalmente la mitad o más del largo del palpómero ................................................................................................................................................ 9 Alas con pelos en sector basal de R .................................................... Simulium anduzei Alas sin pelos en sector basal de R ................................................................................ 8 Largo ala 4,0 mm; vesícula sensorial del palpo ligeramente menor de 1/2 del largo del palpómero ........................................................................................ Simulium marquezi Largo ala 3,4 mm; vesícula sensorial del palpo 1/ 2 del largo del palpómero ....... ...................................................................................................................... Simulium tricorne Cibario liso; largo del ala 2,0-2,5 mm; diente subbasal de la uña ausente o muy pequeño .................................................................................................... Simulium spilmani

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Cibario con dentículos o esférulas; largo ala 2,7-3,0 mm; diente subbasal de la uña bien desarrollado (Fig. 102 M) ...................................................................................... 10 Cibario sin prolongación mediana con bordes casi rectos; paraprocto más corto que ancho en la base (Fig. 103 G) ................................................................................ 11 Cibario con una prolongación mediana (Figs. 102 J-L); paraprocto aproximadamente tan largo como ancho .................................................................... 12 Cibario con papilas (Fig. 102 N); paraprocto no aguzado distalmente; vesícula sensorial del palpo aproximadamente la mitad del largo del palpómero ................. ........................................................................................................................ Simulium parrai Cibario con pequeños dientes aguzados (Fig. 102 F); paraprocto afinándose escasamente distalmente (Fig. 103 G); vesícula sensorial del palpo menos de la mitad del largo del palpómero (Fig. 101 L) .......................... Simulium mattebranchium Paraprocto redondeado distalmente; VIII esternito con 29-35 cerdas ................ 13 Paraprocto aguzado distalmente; VIII esternito con 20-25 cerdas ....................... 14 Borde interno de las gonapófisis divergente hacia abajo ................... Simulium piperi Borde interno de las gonapófisis paralelo ....................................... Simulium jacumbae Borde interno de las gonapófisis divergente hacia abajo ................. Simulium iriartei Borde interno de las gonapófisis paralelo a gradualmente divergente hacia abajo .......................................................................................................................Simulium hunteri

Machos Simulium covagarciai, S. sandyi, S. racenisi y S. kompi no se incluyen por falta de información. 1 – 2(1) –

3(2) – 4(3) – 5(3) –

Placa ventral subcuadrada, no achatada lateralmente (Figs. 106 A-L) ................... 2 Placa basal subcónica, subcircular a subcilíndrica, gradualmente achatada desde la base (Figs. 106 M-Q) ......................................................................................................... 8 Gonostilo con una prolongación basal larga y delgada; escudo con 1+1 mancha anterior plateada subtriangular (Fig. 104 I) ............ Simulium jobbinsi, Simulium piperi Gonostilo con una prolongación basal relativamente corta y robusta, frecuentemente rugosa y dentada apicalmente; escudo variable; generalmente con 1+1 banda plateada anterior, convergente, diagonalmente dirigidas hacia la línea media (Figs. 104 A, B) ....................................................................................................... 3 Escudo con 1+1 submediana y 1+1 sublateral mancha o banda plateada (Figs. 104 L-N) ...................................................................................................................................... 4 Escudo con 1+1 mancha plateada de forma variable, subtriangular a subcuadrada o subromboidal, o como una banda .............................................................................. 5 Prolongación mediana de la placa ventral ancha basalmente (Fig. 106 G) ............. .......................................................................................................................... Simulium puigi Prolongación mediana de la placa ventral angosta basalmente (Fig. 106 L) ........... .........................................................................................................................Simulium hechti Escudo con 1+1 manchas plateadas diagonales prolongándose posteriormente (Figs. 104 A, B, L) .............................................................................................................. 6 Escudo con 1+1 mancha plateada no diagonal prolongándose posteriormente (Fig. 104 D) ......................................................................................................................... 7

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Placa ventral con una prolongación, mediana, corta y angosta (Fig. 106 I); escudo con 1+1 manchas plateadas cortas y robustas (Fig. 104 J) ............... Simulium parrai – Placa ventral con una prolongación mediana a robusta (Figs. 106 A, B)); escudo con 1+1 manchas plateadas, alargadas y afinadas distalmente (Figs. 104 A, B) .... ................................................. Simulium metallicum, Simulium horacioi, Simulium spilmani 7(5) Placa ventral con una prolongación mediana robusta bordeada por una hendidura, y los lados curvos (Figs. 106 D-E); prolongación interna basal del gonostilo corta y lisa (Fig. 105 F) .................................................... Simulium marquezi, Simulium tricorne – Placa ventral con una prolongación mediana no bordeada por una hendidura y lados rectos (Figs. 106 C, H, K); prolongación interna basal del gonostilo larga y dentada distalmente (Figs. 105 G, K) .............................................................................. ........ Simulium bustosi, Simulium iriartei, Simulium jacumbae, Simulium morae, Simulium wygoi 8(1) Escudo sin manchas o bandas aisladas, todo el disco marginalmente plateado (Fig. 104 K) ...................................................................... Simulium costalimai, Simulium anduzei – Escudo con manchas o bandas plateadas ..................................................................... 9 9(8) Escudo con 1+1 mancha plateada anterior subcuadrada o subromboidal uniéndose al área lateral plateada; cuerpo de la placa ventral tan ancho como el diámetro mediano transversal (Fig. 106 M) .........................................Simulium hunteri – Escudo con 1+1 mancha plateada anterior subtriangular o subtrapezoidal o bandas diagonales curvas (Figs. 104 E, G) no unidas al área lateral plateada; cuerpo de la placa ventral más angosto basalmente que el diámetro mediano transversal (Figs. 106 O-Q) ............................................................................................ 10 10(9) Gonostilo con distintas curvaturas en vista lateral (Fig. 105 E); cuerpo de la placa ventral fuertemente achatado lateralmente desde la base, subcilíndrico (Fig. 106 Q) .....................................................................................................................Simulium putre – Gonostilo con lados rectos subparalelos en vista lateral; cuerpo de la placa ventral no fuertemente achatado lateralmente, subcónico (Figs. 106 J, P) ........................ 11 11(10) Gonostilo con prolongación interna basal más corta que el diámetro basal transvesal ........................................................................................................ Simulium ruizi – Gonostilo con prolongación interna basal más larga que el diámetro basal transversal (Fig. 105 H) ........................................................... Simulium matteabranchium Pupas 1 – 2(1) – 3(2) – 4(3)

Branquias robustas y membranosas (Figs. 107 H, J, 109 A-C. G, I) ....................... 2 Branquias filamentosas (Figs. 107 A, C, F) ................................................................... 9 Branquias sin filamentos; frontoclípeo y tórax con tubérculos ............................... 3 Branquias con filamentos flexibles (Figs. 109 A, E, I); frontoclípeo y tórax sin tubérculos ............................................................................................................................ 6 Branquias con ramas digitiformes sin pequeñas ramas secundarias tubuliformes; capullo sin prolongación anterior (Fig. 107 H) ........................................................... 4 Branquias con ramas digitiformes con pequeñas ramas secundarias tubuliformes (Figs. 109 B, C); capullo con (Fig. 107 I) o sin anterior prolongación ................... 5 Branquia con 3 ramas dirigidas anteriormente, con anillaciones; capullo sin carena dorsal ............................................................................................................ Simulium kompi

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Branquia con 2 ramas dirigidas posteriormente y 1 anteriormente, sin anillaciones (Fig. 107 H); capullo con carena dorsal .............................................. Simulium tricorne 5(3) Branquia con ramas basalmente globosas, subcilíndricas que se angostan progresivamente distalmente, y la rama principal subdividida distalmente en 7 ramas pequeñas y angostas (Fig. 109 B); tergitos abdominales I y II con tubérculos ................................................................................................................... Simulium marquezi – Branquia arborescente con 25 ramas terminales relativamente angostas emergiendo de 3 ramas primarias cortas y robustas (Fig. 109 C); tergitos abdominales I-II sin tubérculos .............................................................. Simulium sandyi 6(2) Branquia con rama principal subdividida distalmente (Fig. 107 J) .......................... 7 – Branquia subdividida basalmente en 4 grandes ramas ............................................... 8 7(6) Branquia con la rama principal distalmente y basalmente con 1 corta rama únicamente (Fig. 109 E) .............................................................................. Simulium ruizi – Branquia con un robusto tallo expandido distalmente, con varias ramas cortas y pequeñas y basalmente con 3 cortas ramas (Fig. 109 A) .................................... ...................................................................................................... Simulium matteabranchium 8(6) Ramas de las branquias con pseudosegmentaciones, 2 dirigidas anteriormente y 2 ventralmente (Fig. 109 I) .......................................................................... Simulium parrai – Ramas de las branquias sin pseudosegmentaciones, la más larga dirigida anteriormente con pequeñas protuberancias subtriasngulares y 2 protuberancias ventrales subglobosas ......................................................................... Simulium covagarciai 9(1) Ramas terminales de la branquia superando los 8,5 mm; frontoclípeo y tórax con tricomas simples (Figs. 108 G, H) ................................................................................ 10 – Ramas terminales de la branquia menores de 3,5 mm; frontoclípeo y tórax con tricomas simples o multirramosas ................................................................................ 12 10(9) Branquia con 3 ramas primarias robustas, subdividida en 7-10 ramas principales con 85-100 ramas terminales pequeñas (Fig. 108 H); tórax con alrededor de 50 tricomas de cada lado ............................................................................... Simulium iriartei – Branquias con alrededor de 110-135 ramas (Fig. 108 G); tórax con 70-100 tricomas de cada lado ...................................................................................................... 11 11(10) Frontoclípeo y tórax sin tubérculos, capullo con prolongación dorsal anterior ..... .................................................................................................................. Simulium costalimai – Frontoclípeo y tórax con moderado número de tubérculos; capullo con borde anterior recto ..............................................................................................Simulium hunteri 12(9) Branquias con 5 ramas (Fig. 108 E) ................................................... Simulium spilmani – Branquias con 6-30 ramas .............................................................................................. 13 13(12) Branquias con 6 ramas de aproximadamente el mismo largo (Fig. 108 A); capullo con borde anterior recto (Figs. 107 A, C) ................................................................... 14 – Branquias con 10-30 ramas; capullo generalmente con prolongación dorsal anterior o abertura en forma de círculo ...................................................................... 16 14(13) Tricomas del frontoclípeo y tórax con 2-6 ramas (Figs. 107 B, 108 A), tórax con 5 tricomas de cada lado ...................................................................................................... 15 – Tricomas de frontoclípeo y tórax simples; aquellos del tórax en número de 60-85 de cada lado (Figs. 107 C-E) .....................................................................Simulium putre

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15(14) Capullo achatado, subcircular en vista dorsal ................................... Simulium horacioi – Capullo normal no achatado (Fig. 107 A ) ..................................................................... ........ Simulium metallicum, Simulium jabbinsi, Simulium morae, Simulium puigi, Simulium racenisi 16(13) Capullo achatado, subcircular con borde anterior unido ventralmente (Fig. 107 F); branquia con 10-16 ramas terminales .......................................................................... 17 – Capullo con forma normal, achatado con o sin prolongación dorsal anterior; branquia con 10-30 ramas .............................................................................................. 19 17(16) Branquia con 10 ramas terminales, con un corto tallo en la base y 4 ramas primarias; tórax con 25 tricomas de cada lado, con el diámetro disminuyendo gradualmente hacia el ápice .................................................................... Simulium bustosi – Branquia con 2-3 ramas principales con 9-11 ramas angostas y flexibles, con 5-120 tricomas que son muy robustos en su 1/3 basal (Fig. 107 K) ............................... 18 18(17) Branquias con 3 ramas principales; tórax con 100-200 tricomas; tórax liso (Fig. 108 J) .............................................................................................................. Simulium hechti – Branquias con 2 ramas principales; tórax con 5 tricomas de cada lado; tórax rugoso (Fig. 108 I) ...................................................................................... Simulium wygoi 19(16) Capullo con prolongación dorsal anterior; tórax con 25-65 tricomas de cada lado .............................................................................................................................................. 20 – Capullos sin proyección anterior dorsal; tórax con 5+ 5 tricomas de cada lado ..................................................................................................................... Simulium anduzei 20(19) Branquia con 10 ramas terminales; tórax con 50-65 tricomas .......... Simulium piperi – Branquia con 25 (Fig. 108 F) ramas terminales; tórax con 25-35 tricomas ............ ................................................................................................................... Simulium jacumbae Larvas No incluidas Simulium costalimai, S. covagarciai, S. kompi, S. matteabranchium, S. morae, S. racenisi y S. sandyi. 1 – 2(1) – 3(2) – 4(2) – 5(1)

Hendidura postgenal no evidente o escasamente desarrollada; relación largo hipostomio/puente postgenal = 0,4-0,7 (Figs. 112 A-D) .......................................... 2 Hendidura postgenal escasa a bien desarrollada; relación largo hipostomio/ puente postgenal = 0,8-1,4 .............................................................................................. 5 Artejo antenal mediano más corto que el distal, y el artejo proximal generalmente más largo o igual al mediano (Fig. 111 B) .................................................................... 3 Artejo antenal mediano más largo que el distal, y el artejo proximal generalmente más corto que el mediano (Fig. 111 L) ......................................................................... 4 Lóbulos de las papilas rectales sin divertículos (Fig. 111 G) ..............Simulium putre Lóbulos de las papilas rectales con varios divertículos ................................................ .................................................................................... Simulium anduzei, Simulium jacumbae Disco anal con alrededor de 106 hileras de ganchos; mandíbula con 13-15 dientes internos .................................................................................................... Simulium marquezi Disco anal con 80 o menos hileras de ganchos; mandíbula con 10-12 dientes internos ........................................................................... Simulium piperi, Simulium tricorne Hipostomio más largo que el puente postgenal .......................................................... 6

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Hipostomio tan corto o más corto que el puente postgenal (Figs. 112 B, C, G) ....... 7 Relación largo hipostomio/puente postgenal = 1; anillo anal con alrededor de 90 hileras de ganchos ........................................................................................ Simulium ruizi – Relación largo hipostomio/puente postgenal = 1.2-1.4; anillo anal con alrededor de 78-80 hileras de ganchos .................................... Simulium parrai, Simulium spilmani 7(5) Escamas en el esclerito anal presente .............................................................................. ........................ Simulium metallicum, Simulium jobbinsi, Simulium horacioi, Simulium puigi – Escamas en el esclerito anal ausente .............................................................................. 8 8(7) Artejo antenal mediano más corto que el proximal y distal ............ Simulium bustosi – Artejo antenal mediano más largo que el proximal y a veces también del distal (Figs. 111 H, N) .................................................................................................................. 9 9(8) Apotoma cefálico sin manchas ................................................................. Simulium hechti – Apotoma cefálico con manchas medianas y submedianas positivas o negativas .............................................................................................................................................. 10 10(9) Papilas rectales con 13-14 divertículos sobre cada lóbulo ................. Simulium wygoi – Papilas rectales con 5-8 divertículos sobre cada lóbulo ........................................... 11 11(10) Relación de los artejos antenales proximal, mediano y distal = 1:1.1:1.2 ................ ....................................................................................................................... Simulium iriartei – Relación de los artejos antenales proximal, mediano y distal = 1:2.1-2.2:1.3-1.5 .......................................................................................................................Simulium hunteri Simulium (Aspathia) anduzei VARGAS & DÍAZ NÁJERA Simulium (Simulium) anduzei VARGAS & DÍAZ NÁJERA, 1948b: 328-330; 1949: 296-298; 1957b: 264; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 59; COSCARÓN et al., 1999: 554. Simulium (Aspathia) anduzei: ADLER et al., 2004: 365-366. Jalacingomyia anduzei: PY-DANIEL & MOREIRA SAMPAIO, 1994: 101. Simulium (Simulium) patziciaense TAKAOKA & TAKAHASI, 1982: 63.

Female: Wing length 3.3 mm. Head blackish, frons and clypeus black; clypeus pollinosity shining blue. Thorax black with median and 1+1 whitish submedian vittae, subtriangular anteriorly, and 1+1 lateral silvery stripes. Abdominal tergites black with pale median spots on T II-V. Cibarium smooth. Basal section of R haired. Tarsal claw with subbasal tooth. Gonapophysis distally truncated and with parallel internal sides, paraproct short, gradually acuminate ventrally. Male: Wing length 3.3 mm. Head blackish, clypeus gray pollinose. Thorax blackish, only ornamented dorsally by lateral silver stripe. Abdominal tergites black with long black hair; T II-V with 1+1 ovoid sublateral shining gray spots. Gonostylus with prominent internal basal process smooth distally; ventral plate subconical, becoming gradually narrower toward apex, similar to S. costalimai and S. hunteri. Pupa: Cocoon slipper-shaped, without anterior projection but with anterior collar. Gill length 4.5-4.7 mm, with 17-19 branches originating from long stem. Frontoclypeus and thorax with spiculae on surface. Frontoclypeus with 1+1 frontal and 1+1 facial simple trichomes. Thorax with 5+5 simple trichomes. Abdominal sternites VI-VII with only 1 simple or bifurcated posterior hook on each side (from PY-DANIEL, 1994).

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Larva: Length of body 7.8-8.0 mm. Body with 1+1 ventral tubercles posteriorly. Cephalic apotome with positive spots. Cephalic fan with 27-31 primary rays. Antennae longer than stem of cephalic fan. Ratio of segments proximal, medial and distal = 1:0.9-1.0:1.0-1.2. Mandible with 6-10 inner teeth and 3 lateral mandibular processes. Hypostomium with 13-15 teeth and 6-7 lateral serrations, lateral setae 5-8; postgenal cleft rounded, shallow. Ratio of hypostomium/postgenal bridge = 0.5. Anal sclerite with hair, no scales. Anal ring with 65-74 rows of 11-12 hooks each. Rectal papillae with 9-13 diverticula on each lobe. Distribution: Mexico: Distrito Federal, Oaxaca, Veracruz; USA: Arizona, New Mexico. Bionomics: This species breeds in spring-fed streams at an altitude of about 2000 m, attached to fallen leaves and stones. Female feeding habits are unknown. Discussion: The female of S. anduzei is similar to that of S. spilmani and S. tescorum, but those species have a bare basal section of R. The male is similar to S. costalimai, but this species has about 102 gill branches. We accept the synonymy of S. patziciaense proposed by ADLER et al. (2004: 365). Simulium (Aspathia) bustosi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Fig. 102 O.) Simulium (Simulium) bustosi VARGAS et al., 1946: 138-140; VARGAS & DÍAZ NÁJERA, 1957b: 264; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 59; COSCARÓN et al., 1999: 555.

Female: Wing length 2.8-3.0 mm. General coloration blackish, clypeus and pleurae gray pollinose. Scutum black with golden hair and 1+1 small anterior silvery subcircular spots. Abdomen black. Frons parallel sided; fronto-ocular triangle about as long as wide. Palpus sensory vesicle a little shorter than 1/2 length of palpomere. Basal portion of cibarium with small median process and sparse denticles (Fig. 102 O). Ratio of length/ width of posterior basitarsus = 7.3. Paraproct shorter than wide at base, distal edge gradually acuminate, directed ventrally; sternite VIII with about 18-20 hairs on each side; gonapophysis weakly produced and curved distally; spermatheca well developed. Male: Wing length 3.1 mm. Scutum velvety black with 1+1 short anterior silver spots, connected with silver lateral band that extends to posterior margin. Gonostylus narrow, subcylindrical, with internal basal process robust and dentate on apex. Ventral plate wider than high, with short median process, shape similar to that of S. metallicum. Pupa: Cocoon slipper-shaped, with collar present; length basally 3.6-4.0 mm, dorsally 3.0 mm. Frontoclypeus and exposed portion of thorax with small and sparse tubercles; thorax with 25-30 simple trichomes. Gill filaments 3.7 mm long, with 10 branches, short stem at base, and 4 primary branches whose first and second dorsal branches are narrower than others. Larva: Length 5.7-6.3 mm. Cephalic apotome with 1+1 median longitudinal and 2+2 submedian positive spots. Ratio of hypostomium/postgenal bridge = 0.8-0.9; hypostomium with 5+5 lateral setae and 3-4 lateral serrations. Cephalic fans with 37-43 primary rays. Ratio of segments proximal, medial and distal = 1:0.9:0.9-1.0, palpomere robust. Mandible with 8 inner teeth, second marginal tooth well separated from first; 2 anterolateral setae present. Comb of proleg with about 36 teeth. Anal sclerite with hair, no scales. Anal ring with 66-72 rows of 12-13 hooks each. Rectal papillae lobes without diverticula. Distribution: Mexico: Oaxaca, Veracruz.

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Bionomics: Unknown. Discussion: The number of gill branches (10) is similar to that for S. piperi, but the dorsal projection of the cocoon, 50-54 thoracic trichomes, vittae on the female scutum, and smooth apex of the inner basal process of the male gonostylus allow the separation of these 2 species. Simulium (Aspathia) costalimai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 103 K; 104 K; 106 N.) Simulium (Simulium) costalimai VARGAS et al., 1946: 108, 140-142; VARGAS & DÍAZ NÁJERA, 1957b: 264, 372373; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 556. Simulium (Simulium) jacumbae: VARGAS et al., 1943: 289; (not DYAR & SHANNON).

Female: Wing length 3.8 mm. Blackish, frons bright pollinose. Scutum black with golden pilosity and 1+1 anterior whitish spots connected laterally with silvery marginal area. Tarsal claw with subbasal tooth well developed. Cibarium with median process not covered by denticles. Paraproct shorter than high, produced ventrally and gradually acuminate distally (Fig. 103 K); gonapophysis longer than wide at base, truncated apically; sternite VIII with 28-30 hairs on each side. Male: Wing length 3.0 mm. Scutum velvety black without spots or vittae, with golden pilosity and edged by grayish iridescent pollinosity on lateral, posterior and anterior margins (Fig. 104 K). Ratio of length/width of hind basitarsus = 3.4. Abdominal tergites II, IV-VIII with 1+1 silver pollinose spots. Gonostylus subcylindrical, with internal-basal process well developed, not denticulate apically. Ventral plate subconical, 1.5 times longer than wide, gradually projected distally (Fig. 106 N). Pupa: Cocoon with thick silk reinforced on edge, with very long dorsal projection curved ventrally; basal length 4.5-5.0 mm, dorsal length 5.5-6.0 mm. Gill 1.2-1.3 mm long. Frontoclypeus and thorax without tubercles. Thoracic trichomes simple, elongate, about 80-100 on each side. Gill filaments with about 110-135 branches, with irregular branching. Abdominal sternite IV with bifurcated hooks. Larva: Unknown. Distribution: Mexico: Morelos, Oaxaca. Bionomics: Unknown. Discussion: The high number of gill terminal branches (over 100) is similar to that of S. hunteri, but the cocoon’s straight anterior edge and only about 70 thoracic trichomes in S. hunteri differentiates these 2 species. Simulium (Aspathia) covagarciai RAMÍREZ PÉREZ, YARZABAL, TAKAOKA, TADA & RAMÍREZ Simulium covagarciai RAMÍREZ PÉREZ et al., 1982: 41-44. S. (Simulium) covagarciai COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 556.

(Based on original description). Female (pharate): Frons and vertex blackish with nacreous pruinosity and silver hair. Thorax black with golden hair, color pattern unknown. Abdomen blackish, tergites VI-

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VIII shining. Fronto-ocular triangle as long as wide. Sensory vesicle of palp 1/3 length of palpomere. Cibarium base smooth but with small, sclerotized, rounded papillae. Paraproct 1/2 as long as wide at base. Male (pharate): Thorax and abdomen velvety black, thorax with golden hair; abdominal tergites II, VI-VIII with 1+1 silver lateral spots. Gonostylus with median constriction, with internal basal process. Ventral plate about as long as wide, with short median process. Pupa: Cocoon threads thick and conspicuous. Basal length 4.75 mm, dorsal length 4.50 mm. Frontoclypeus and thorax without tubercles, frontoclypeus with 1+1 facial and 2+2 frontal, simple, filiform trichomes. Thorax with about 40 simple, filiform trichomes. Gill membranous, with 4 subcylindrical, smooth dorsal lobes, 2 ventral lobes subglobose, with some hair, the frontal largest, with small subtriangular protuberances and some long hair basally. Larva: Unknown. Distribution: Venezuela: Amazonas, Sierra de Parima (950 m). Bionomics: Unknown. Discussion: The closest species is S. matteabranchium, but S. covagarciai can be differentiated by the gills of the pupa. Simulium (Aspathia) hechti VARGAS, MARTINEZ-PALACIOS & DÍAZ NÁJERA (Figs. 104 M, N; 105 J; 106 L, 107 K; 108 J.) Simulium (Simulium) hechti VARGAS et al., 1946: 142; VARGAS & DÍAZ NÁJERA, 1957b: 264, 372-373; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 557. Simulium (Aspathia) hechti: ADLER et al., 2004: 366. Simulium (Simulium) sayi: VARGAS et al., 1943: 288; (not DYAR & SHANNON).

Female: Wing length 3.3 mm. Blackish, clypeus silvery pollinose, scutum black with 1+1 anterior whitish triangle-shaped spots and silvery edging posteriolaterally. Abdomen dark brownish with 1+1 spot on tergite II. Basal portion of cibarium smooth, weakly reinforced on edge. Frons slowly convergent, fronto-ocular triangle about as long as wide at base. Sensory vesicle of palp near half of palpomere length. Ratio of length/width of hind basitarsus = 6.0. Paraprocts about as long as wide at base, slightly acuminate distally. VIII stemite with 25-26 setae; gonapophysis with convex margin, inner margins parallel and reinforced. Male: Wing length 3.1 mm. Scutum reddish brown with 1+1 median ovoid, and 1+1 diagonal cuneiform spots (Figs. 104 M, N), anterior stripes silvery pollinose. Hind basitarsus wide, ratio of length/width = 3.0. Gonostylus subcylindrical with strongly wrinkled internal basal process denticulate apically (Fig. 105 J). Ventral plate wider than high, slightly flattened, with short and wide subconical median process (Fig. 106 L). Pupa: Cocoon with marginal collar and projected antero ventrally, threads well defined. Basal length 3.4-4.2 mm, dorsal length 2.0-2.5 mm. Gill length 25-30 mm. Frontoclypeus and exposed portion of thorax with abundant tubercles. Trichomes of head and thorax simple, with peculiar thickening on basal 1/3 and filiform on distal 3/4 (Fig. 107 K), 100-120 on each side of thorax. Gill with 3 main filiform branches bearing 911 narrower flexible branches (Fig. 108 J).

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Larva: Cephalic apotome without ornamentation or spots. Postgenal cleft short, ratio of hypostomium/postgenal bridge = 0.8; hypostomium with 7-8 lateral setae and 57 lateral serrations. Ratio of segments proximal, medial and distal = 1:1.5:1.3. Cephalic fan with 32-34 primary rays. Mandible with 8 inner teeth and 1 lateral mandibular process. Comb of proleg with about 28-32 teeth. Anal sclerite with sparse hair, no scales. Anal ring with about 75-80 rows of 11 -13 hooks each. Rectal papillae with 6 diverticula on each lobe, with distal lobule larger than others. Distribution: Mexico: Durango, Distrito Federal, Morelos; USA: Arizona, New Mexico. Bionomics: Simulium hechti breeds in mountainous areas. Female feeding habits are unknown. Discussion: The most similar species is S. wygoi, especially in the female, but the pupal gill with two main branches and wrinkled thorax with five trichomes on each side in S. wygoi will differentiate them. Simulium (Aspathia) horacioi OKAZAWA & ONISHI Simulium (Simulium) horacioi OKAZAWA & ONISHI, 1980: 167-173; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 69; 2004: 63; GRILLET & BARRERA, 1997: 197, 200, 206; COSCARÓN et al., 1999: 558. Simulium horacioi: TAKAOKA, 1982: 49. ?Simulium metallicum (cytoform “H”) CONN et al., 1989: 1217.

This species is very close to S. metallicum. In S. horacioi, small differences occur: the paraprocts are shortest distally, the median process of the ventral plate is more projected ventrally, the pupal cocoon is flattened and rounded, and the cephalic apotome of the larva has positive spots and segments VII-IX are blackish brown dorsally. Simulium horacioi may correspond to any cytotype of the S. metallicum species complex. It would be useful to do complementary studies. Distribution: Guatemala; Mexico: Chiapas, Oaxaca; Venezuela: Carabobo. Bionomics: The species breeds in small rivers, tributaries and channels in mountainous areas at about 1300 m. It is an anthropophilic and zoophilic species. Simulium (Aspathia) hunteri MALLOCH (Figs. 102 K; 106 M; 108 G.) Simulium (Simulium) hunteri MALLOCH, 1914: 59; PETERSON & KONDRATIEFF, 1995: 39; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 559-560. Simulium (Aspathia) hunteri: ADLER et al., 2004: 366. Simulium (Simulium) lassmanni VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA, 1946: 149, 174; VARGAS & DÍAZ NÁJERA, 1957b: 264; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65.

Female: Wing length 3.1 mm. General coloration blackish. Scutum ornamented with silvery lyre-shaped stripes; 1+1 anterior, subtriangular silvery spots when viewed with other illumination angles. Abdomen blackish, last segment shining. Frons convergent below. Sensory vesicle of palp less than 1/2 length of palpomere. Fronto-ocular triangle about as long as wide. Basal portion of cibarium with denticulate median process (Fig. 102 K). Ratio of length/width of hind basitarsus = 6.0. Paraprocts about as long as wide with

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curved posterior edge. Stemite VIII with 20-25 hairs on each side; gonapophysis truncated, with parallel inner margins. Male: Wing length 3.0 mm. Scutum blackish, with yellow pilosity and 1+1 silvery anterior subrhomboidal spots meeting laterally, lateral and posterior margins silvery. Legs blackish with fore coxa and hind basitarsus whitish on basal half. Ratio of length/width of hind basitarsus = 4.7. Abdomen black, with 1+1 silvery spots on tergites II and VII. Gonostylus curved, subcylindrical, with internal-basal process as long as transverse diameter of gonostylus at base and not dentate apically. Ventral plate subconical-elongate, slightly longer than wide at base (Fig. 106 M). Pupa: Cocoon with straight, reinforced anterior edge, loosely woven with well– defined threads. Basal length 4.3 mm, dorsal length 3.6 mm. Frontoclypeus and thorax with moderate number of tubercles arranged in groups; trichomes simple and elongate, those of thorax about 70 on each side. Gill arborescent, with about 130 branches, branching from 7 primary branches (Fig. 108 G), length about 1.2-1.5 mm. Larva: Length 5.8 mm. Cephalic apotome with median longitudinal and 1+1 discontinuous sublateral positive spots. Ratio of hypostomium/postgenal bridge = 0.8. Postgenal bridge 1.2 times height of hypostomium; hypostomium with 7-9 lateral setae and 3-4 lateral serrations. Ratio of segments proximal, medial and distal = 1:2.1-2.2:1.31.5. Cephalic fan with about 35-38 primary rays, combs of rays with 7 intermediate minor teeth alternating with 1 bigger tooth. Inner mandibular teeth 10-11. Proleg comb with 28 teeth. Anal sclerite with hair, crochet circlet with 72-76 rows of 13-14 hooks. Rectal papillae with 5 diverticula on each lobe. Distribution: Canada: Alberta, British Columbia, Saskatchewan, Yukon; Mexico: Distrito Federal, Michoacan; USA: Alaska, Arizona, California, Colorado, Idaho, New Mexico, Nevada, Oregon, South Dakota, Texas, Utah, Washington, Wyoming. Bionomics: The species breeds in small creeks, frequently with fast current. Females bite livestock and humans. Discussion: The closest species is S. costalimai; differences are discussed under that species. Simulium (Aspathia) iriartei VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 105 G; 106 H; 108 H; 111 N.) Simulium iriartei: VARGAS et al., 1946: 144-146, 173-174; VARGAS & DÍAZ NÁJERA, 1957b: 264; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 560. Simulium (Aspathia) iriartei: ADLER et al., 2004: 366-367. Simulium hirtipes: VARGAS, 1943a: 143; (not FRIES).

Female: Wing length 3.0 mm. Body blackish. Scutum with 1+1 anterior light grayish, subtriangular spots continued as longitudinal, very narrow median stripes. Frons convergent below, fronto-ocular triangle about as long as wide. Sensory vesicle of palp more than 1/2 length of palpomere. Tarsal claw with subbasal tooth well developed. Ratio of length/width of hind basitarsus = 6.5. Basal portion of cibarium with median process covered by denticles. Paraprocts longer than wide at base; gonapophyses reinforced on inner margins and gradually divergent below; stemite VIII with 21-25 hair.

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Male: Wing length 2.8 mm. Head blackish, clypeus shining whitish. Scutum black with 1+1 subrhomboidal anterior silvery spots with green hair, edged laterally by green metallic pollinose stripe. Gonostylus with internal-basal process well developed, wrinkled and dentate distally (Fig. 105 G). Ventral plate subquadrate basally, horseshoe-like, with small median process, subacute distally (Fig. 106 H). Pupa: Cocoon with reinforced anterior margin and strongly projected dorsally, with conspicuous threads. Basal length 4.4-5.0 mm, dorsal length 4.2-5.0 mm. Gill 2.0 mm long, arborescent, with about 85-100 terminal filaments branching basally from 3 stout primary branches, subdivided 3-6 times to end (Fig. 108 H). Frontoclypeus and thorax with sparse tubercles, trichomes simple and elongate; thorax with about 50 trichomes on each side. Scales on abdominal stemites III-VIII conspicuous as small spines. Larva: Length 6.3 mm. Cephalic apotome gradually darkened basally, with 1 median anterior, 1 subbasal and 1+1 submedian positive spots. Postgenal bridge 1.2 times longer than height of hypostomium, ratio of hypostomium/postgenal bridge = 0.8, hypostomium with 7 lateral setae and 4-6 lateral serrations. Antennae longer than stem of cephalic fan. Ratio of segments proximal, medial and distal =1:1.1: 1.1-1.2 (Fig. 111 N). Cephalic fan with 32-34 primary rays. Mandible with 10-11 internal teeth. Thoracic proleg comb with 22-24 teeth arranged in 6-8 groups. Anal sclerite with hair, no scales; anal ring with 72-76 rows of 13-14 hooks each; rectal papillae with 5-8 diverticula on each lobe. Distribution: Mexico: Distrito Federal, Durango, Guerrero, Oaxaca, Veracruz; USA: Arizona, New Mexico. Bionomics: Simulium iriartei breeds in small creeks in mountainous areas. Female feeding habits are unknown. Discussion: Simulium hunteri and S. costalimai are the closest species. The first can be separated from S. iriartei by the elongate and apically smooth inner basal process of the gonostylus; the second species can be separated by the female scutum with 1+1 anterior grayish spots. Both of these similar species can be distinguished from S. iriartei in the pupal stage by having a gill with more than 100 terminal branches. Simulium (Aspathia) jacumbae DYAR & SHANNON (Figs. 102 J; 106 K; 108 F.) Simulium jacumbae DYAR & SHANNON, 1927: 44-45, VARGAS & DÍAZ NÁJERA, 1948b: 333-334. Simulium (Aspathia) jacumbae: ADLER et al., 2004: 367. Simulium (Simulium) jacumbae: VARGAS et al., 1946:146; DALMAT, 1951: 54; 1955: 208-213; VARGAS & DÍAZ NÁJERA, 1957b: 264, 376-377; STONE & BOREHAM, 1965: 170; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 561-562. Simulium guatemalense DE LEÓN, 1945: 75.

Female: Wing length 2.5-2.8 mm. General coloration dark grayish brown. Scutum ornamentation with lyre-shaped pattern as in S. metallicum, with gray pollinosity and golden to silver hair. Legs blackish, with basal half of median and hind tibiae whitish. Abdomen black, tergites V-VIII shining. Frons convergent below, fronto-ocular triangle as long as wide. Basal portion of cibarium with median process, wrinkled on anterior edge (Fig. 102 J). Palpal sensory vesicle 1/2 length of palpomere. Sc and basal section of R not haired. Tarsal claw with subbasal tooth well developed. Paraprocts as long as wide,

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rounded distally. Stemite VIII with 29-35 setae. Gonapophyses with straight distal edge, internal margins sclerotized and parallel-sided. Male: Wing length 2.6 mm. General coloration as in female. Scutum similar to that of S. metallicum but 1+1 anterior triangles smaller, hair golden to tin color. Hair on legs and first abdominal segment yellowish brown. Ratio of length/width of hind basitarsus = 4.3. Abdomen black with 1+1 silvery spots on tergites II, V-VII. Gonostylus about twice length of gonocoxite, subcylindrical, acuminate distally, with internal basal process narrow, almost as long as transverse diameter of gonostylus at base and dentate on edge; ventral plate subquadrate basally, gradually elongate distally and about as long as wide, with sparse hair (Fig. 106 K). Pupa: Cocoon with anterodorsal process, loosely woven and with threads visible. Basal length 3.0-3.3 mm, dorsal length 3.6-4.2 mm. Gill length 2.0 mm, with 24-25 filaments, stem short, with 3 primary branches: ventral primary branch short and stouter, with 2 large secondary branches as stout as other primary branches (Fig. 108 F). Frontoclypeus and thorax with sparse tubercles, cephalic and thoracic trichomes simple, those of thorax about 23-35 on each side. Hooks of stemites VI-VIII bi- or trifurcated. Larva: Length 6.7 mm. Cephalic apotome with median and submedian positive spots. Postgenal cleft shallow. Ratio of hypostomium/postgenal bridge = 0.4-0.6. Hypostomium with 4 lateral setae, and 5-7 lateral serrations. Antennae longer than base of cephalic fans. Ratio of segments proximal, medial and distal = 1:0.9:1.2, without evident transverse striations. Cephalic fan with 55-60 primary rays. Inner mandibular teeth in number 8-10 in 2 series, with 2 lateral mandibular processes. Anal sclerite without hair or scales; anal ring with about 70 rows of 12 hooks each; rectal papillae with 1 -3 diverticula on each lobe. Distribution: Guatemala; Mexico: Chiapas, Hidalgo, Jalisco, Michoacan, Oaxaca, Veracruz; USA: Arizona, California, Colorado, Kansas, Nebraska, Nevada, Texas, Utah, Wyoming. Bionomics: The species breeds in lowlands, in small streams. Females bite livestock and humans. Discussion: Simulium piperi is closest to S. jacumbae but the latter can be differentiated by the female inner margins of the gonapophyses being parallel and the pupal gill having 25 terminal branches. Simulium (Aspathia) jobbinsi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA Simulium (Simulium) jobbinsi VARGAS et al., 1946: 147-149; VARGAS & DÍAZ NÁJERA, 1957b: 264, 378379; DALMAT, 1951: 54; 1955: 213; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 562-563.

Female: Wing length 2.6 mm. General color blackish, with ornamentation on thorax and abdomen similar to that of S. metallicum (with iridescent pruinosity on thorax and 1+1 silver spots on tergite II). Frons with subparallel sides; fronto-ocular triangle as long as wide at base. Cibarium smooth. Sensory vesicle of palp about 1/2 length of segment. Tarsal claw with subbasal tooth well developed; ratio of length/width of hind basitarsus = 5.8. Basal portion of R without hair. Paraproct longer than wide, gradually acuminate. Stemite VIII with 37-40 hairs on each side. Gonapophyses truncate, inner margins reinforced and divergent posteriorly.

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Male: Wing length 2.4 mm. Color black, very similar to S. metallicum but 1+1 laterofrontal cuneiform silvery spots shorter (not prolonged posteriorly). Legs slightly darker than in S. metallicum. Gonostylus subcylindrical, curved, with well-developed internal basal process almost as long as transverse diameter of gonostylus at base; ventral plate wide, with short median process very similar to that of S. metallicum. Pupa: Cocoon with straight edge, loosely woven. Basal length 3.3-3.5 mm, dorsal length 3.0-3.2 mm; gills 4.0-4.3 mm. Frontoclypeus and exposed portion of thorax with abundant rounded tubercles. Gill with 6 filaments, stem short, and with well-defined primary branches. Cephalic trichomes with 3-4 branches. Larva: Length 6.2-6.4 mm. Cephalic apotome with small gradually darkened median longitudinal spot. Ratio of hypostomium/postgenal bridge = 0.9; hypostomium with 4 to 5 lateral setae on each side and 6-8 lateral serrations. Antenna typical of the subgenus; ratio of segments proximal, medial and distal = 1:1.1:1. Cephalic fan with 40-45 primary rays. Mandible with 9-10 inner teeth. Anal sclerite with hair and scales basally; anal ring with 76-80 rows of 13-14 hooks each. Ninth sternite with 2 tubercles. Rectal papillae with 3 lobes, each with 6 diverticula. Distribution: Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Guatemala, Huehuetenango, Quezaltenango, San Marcos, Sacatepequez, Solola, Suchitepequez, Totomicapan; Mexico: Chiapas, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Simulium jobbinsi is very close to S. metallicum, separated only by small differences in the male: the ventral plate and silvery diagonal stripes on the thorax are shorter. Females cannot be separated from those of S. metallicum and S. horacioi (TAKAOKA, 1982). Molecular and cytological studies would be useful to assure the validity of this species. Simulium (Aspathia) kompi DALMAT Simulium (Simulium) kompi DALMAT, 1951: 38; 1955: 218; COSCARÓN, 1987: 32; CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 563.

(Diagnosis based on DALMAT, 1955) (female and male pharate). Female: Paraproct higher than long, gradually acuminate distally; cercus with convex edge; gonapophysis with internal edge parallel, distal edge straight. Male: Gonostylus parallel-sided, with small bifid terminal spur; subbasal process small; ventral plate longer than wide, gradually acuminate and haired distally, without strong lateral constriction. Pupa: Cocoon slipper-shaped, without anterior collar, aperture reinforced. Gill thick and membranous, composed of 3 inflated, tube-like branches, with transverse striations, the dorsal one measuring 2.1 mm, the middle one 1.8 mm and the ventral one 1.3 mm. Larva: Length 6.2 mm. Cephalic apotome darkened, with 1 basal median and 1+1 sublateral elongate spots. Cephalic fan with 28 primary rays. Antennae as long as stalk of cephalic fan, with transverse striations on proximal and medial segments. Medial segment longer than palpomere, and medial and distal subequal. Hypostomium with 6+6 lateral serrations. Postgenal cleft shallow, not acuminate apically. Thoracic proleg comb sclerite with 38-40 teeth. Anal sclerite with hair, no scales; anal ring with 78-82 rows of hooks.

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Distribution: Guatemala: Chimaltenango; Mexico: Chiapas. Bionomics: Unknown. Discussion: The three-lobed pupal gill branches are similar to those of S. kompi and S. tricorne, but the latter have no annulations and the branches have a different orientation. There are slight differences in the female, such as the distally truncate cerci and gonapophyses divergent below. Simulium (Aspathia) marquezi VARGAS & DÍAZ NÁJERA (Figs. 104 D; 105 F; 106 D; 107 I; 109 B; 111 L; 112 A.) Simulium (Simulium) marquezi VARGAS & DÍAZ NÁJERA, 1957a: 37; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 564-565. Simulium vandalicum: VARGAS et al., 1946: 109 (larva); (not DYAR & SHANNON).

Female: Wing length 4.0 mm. General coloration blackish. Scutum ornamented with lyre-shaped markings and silvery posterolateral margin, pleurae brownish, with silver pollinosity. Legs blackish, with basal 1/3 of middle basitarsus and basal 2/3 of hind basitarsus whitish. Abdomen blackish, with 1+1 silvery spots on tergites II-IV, tergites VVIII shining. Frons relatively wide; fronto-ocular triangle as long as wide at base. Sensory vesicle of palp a little shorter than 1/2 length of palpomere. Basal portion of cibarium smooth. Sc with 1-3 hairs, basal section of R bare. Ratio of length/width of hind basitarsus = 8; tarsal claw with subbasal tooth well developed. Paraprocts rounded distally, shorter than wide at base, cerci truncate. Stemite VIII with 15-20 setae on each side; gonapophysis truncate distally, with internal edge divergent and more sclerotized. Male: Wing length 3.8 mm. Scutum dark brown, with pilosity and silver pruinosity on lateral and posterior margins, prolonged into subapical diagonal bands separated from each other by black band at midline (Fig. 104 D). Pleura as in female. Abdomen dark brown or black, concolorous with thorax, with median silvery pollinose spots on tergites II-VII when viewed from behind. Gonostylus subcylindrical, with strong internal basal process, without denticles on edge (Fig. 105 F), ventral plate wider than long, with anterolateral angular projections and small rounded median projections (Fig. 106 D). Pupa: Cocoon with dorsal projection keel-shaped, loosely woven, anterior edge reinforced (Fig. 107 I). Basal length 5.0-5.2 mm, dorsal length 5.5-6.3 mm. Frontoclypeus and exposed portion of thorax with abundant tubercles irregularly arranged and with small spiculae on surface, frontoclypeal trichomes simple and small, difficult to see. Thorax with about 30-33 short, curved trichomes. Gill membranous, with 2 globose subcylindrical basal branches bifurcated at median distal portion and becoming narrower at 1/4-1/3 of length distally, main dorsal branch subdivided distally into 7 small, narrower branches (Fig. 109 B). Abdominal onchotaxy evident, tergites I and II with tubercles. Larva: Length 6.8-7.3 mm. Cephalic apotome with 1+1 submedian spots not well defined, diffuse brownish area present mediobasally. Hypostomium with 9+9 lateral setae and 6-7 lateral serrations; Postgenal cleft reduced (Fig. 112 A); ratio of hypostomium/ postgenal bridge = 0.6. Antennae with second segment wrinkled (Fig. 111 L); ratio of segments proximal, medial and distal = 1:1.3:1. Cephalic fan with 40-42 primary rays, several small teeth, with 1 longest, alternating on combs. Inner mandibular teeth 13-15,

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with secondary tooth row, marginal tooth with 1 large cusp and another smaller one at base. Thoracic proleg comb with about 22 teeth, not arranged in groups. Anal sclerite with abundant setae, no scales; crochet circlet with 106 rows of 15 hooks each. Distribution: Mexico: Chihuahua, Distrito Federal, Morelos. Bionomics: Unknown. Discussion: Simulium tricorne is very similar morphologically, but the small body size, proportionally shorter sensory vesicle of the palp in the female, absence in the pupa of small tubuliform gill branches and cocoon with a straight anterior edge permit separation. Simulium (Aspathia) matteabranchium ANDUZE (Figs. 101 L; 102 F; 103 F-H; 105 H, I; 106 P; 107 J; 109 A.) Simulium matteabranchia ANDUZE, 1947: 374; VARGAS & DÍAZ NÁJERA, 1948b: 333-334; PERRUOLO LANETI, 1971: 397; RAMÍREZ PÉREZ, 1971: 290; 1983: 53. Simulium (Simulium) matteabranchia: COSCARÓN, 1987: 33. Simulium (Simulium) matteabranchium: CROSSKEY, 1987: 471; CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 565-566.

Female: Wing length 2.7-2.8 mm. Body blackish. Frontoclypeus silvery pollinose. Thorax with lyre-shaped pattern; scutum with decumbent hair, some darker and stronger than others. Abdomen black with 1+1 silvery spots on tergite II. Legs dark brownish, hind tibia whitish on basal 3/4; ratio of length/width of hind basitarsus = 7.5; tarsal claw with subbasal tooth well developed. Frons parallel-sided, moderately wide; fronto-ocular triangle about as long as wide. Cibarium with anterior edge not elevated and with abundant small sclerotized papillae at base (Fig. 102 F). Palpal sensory vesicle slightly shorter than 1/2 length of palpomere (Fig. 101 L). Sc and base of R bare. Stemite VIII with 17-21 setae; gonapophysis curved distally, with internal edge sclerotized and 5+5 setae (Fig. 103 F); paraprocts shorter than wide at base, gradually acuminate distally (Fig. 103 G); genital fork with long, acuminate apodemes (Fig. 103 H). Male: Wing length 2.3 mm. Body blackish; scutum black with 1+1 anterior subtriangular silvery spots. Legs brownish black, lighter only on anterior half of second and third basitarsi. Sensory vesicle of palp more than 1/2 length of palpomere. Ratio of length/width of hind basitarsus = 3.6-3.7. Gonostylus relatively narrow with large internal basal process slightly longer than height of gonocoxite (Fig. 105 H). Endoparameres with strong hooks and poorly defined median sclerite (Fig. 105 I). Ventral plate subconical, laterally flattened, acuminate distally (Fig. 106 P). Pupa: Cocoon with reinforced edge, tightly woven, without visible threads (Fig. 107 J). Gill membranous, with 3 branches over thorax and head produced anteriorly as thick stalk distally expanded into several lobules and with thin, filiform branches (Fig. 109 A). Frons with 2+2 frontal and 1+1 simple facial trichomes; frontoclypeus and exposed portion of thorax without tubercles. Thorax with about 52-104 trichomes on each side, very narrow beyond middle, curved and simple, and relatively wide. Abdominal tergite I with sparse acuminate tubercles medially. Larva: Ratio of hypostomium/postgenal bridge = 1.1. Antenna as long as stalk of cephalic fans. Mandible with 11 inner teeth.

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H A

B

J

G

F

C

E K

D

I

L Figure 101. Female. A-E: S. (Aspathia) metallicum; A-B: scutum with different illumination; C-E: anterior, middle and posterior legs, respectively. F-G: S. (A.) piperi, scutum with different illumination. H-J: S. (A.) putre; H: scutum; I: maxillary palp; J: sensory vesicle of palp. K: S. (A.) wygoi, sensory vesicle of palp. L: S. (A.) matteabranchium, sensory vesicle of maxillary palp.

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Distribution: Colombia: Cundinamarca; Mexico(?); Venezuela: Aragua, Carabobo, Distrito Federal, Merida, Monagas, Tachira, Trujillo. Bionomics: Simulium matteabranchium is an anthropophilic species and occurs between 700 and 1800 m, according to RAMÍREZ PÉREZ (1971). Discussion: Simulium parrai is the most closely related species and is differentiated by the female scutum not ornamented with vittae, cibarium base with papillae, paraproct rounded distally, and pupa with the gill having 4 large lobes: 2 basal and 2 anterior. Simulium (Aspathia) metallicum BELLARDI (Figs. 101 A-E; 102 S; 103 A, B; 104 A-C; 105 A-D; 106 A, B; 107 A, B; 108 A; 110; 111 M; 112 C.) Simulium metallicum BELLARDI, 1859: 13; DYAR & SHANNON, 1927: 41; FAIRCHILD, 1940: 712; VARGAS, 1942a: 234-236; VARGAS et al., 1946: 175; DALMAT, 1955: 220-223; VARGAS & DÍAZ NÁJERA, 1957b: 264; LEWIS & LEE POTTER, 1964: 99; WYGODZINSKY, 1971: 12-14; RAMÍREZ PÉREZ, 1971: 275; 1983: 54; 1984: 79; SHELLEY & ARZUBE, 1985: 328. Simulium (Simulium) metallicum OKAZAWA & ONISHI, 1980: 167-179; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 566-568; SHELLEY et al., 2002: 171-176; COSCARÓN ARIAS, 2003: 196, 204, 208, 215. Simulium (Aspathia) metallicum: ADLER et al., 2004: 365 (as species group). Simulium riveti ROUBAUD, 1906b: 108; WYGODZINSKY, 1953b: 332; SHELLEY & ARZUBE, 1985: 328. Simulium nitidum MALLOCH, 1912: 652. Simulium versicolor LUTZ & NUÑEZ TOVAR, 1928: 47. Simulium avidum HOFFMANN, 1930a: 51; 1930b: 293. Simulium violacescens ENDERLEIN, 1934b: 285. “A”-”K” CONN et al., 1989 (cytoform). “L” ARTEAGA & MUÑOZ DE HOYOS, 1999 (cytoform). “X” MILLEST, 1989 (thesis) (cytoform).

Female: Wing length 1.9-2.3 mm. Head blackish, frons and clypeus grayish pollinose; frons parallel-sided. Scutum black, with 3 longitudinal black stripes separated by 2 narrower light ones plus 1+1 submedial grayish pollinose stripes in frontal view, that change to black when viewed from behind as do lateral margins (Figs. 101 A, B), in lateral view scutum has shining metallic greenish iridescence, hair of scutum black and recumbent; scutellum blackish brown; pleurae blackish gray. Legs brownish black, ornamentation as in figures 101 C-E. Abdomen blackish brown, tergites II, IV-VIII with 1+1 silvery pollinose spots that become iridescent when viewed with lateral light, tergites VI-VIII shining. Sensory vesicle of palp about 1/3 length of palpomere. Cibarium smooth. Basal section of R bare. Calcipala and pedisulcus as in figure 102 S; tarsal claw with subbasal tooth well developed. Ratio of length/width of hind basitarsus about 6. Stemite VIII with 27-30 hairs on each side; gonapophyses with subparallel sides and posterior edge truncate; spermatheca well developed (Fig. 103 A); paraproct subtriangular, gradually acuminate from base to apex, longer than wide at base; cercus with rounded edge (Fig. 103 B). Male: Wing length 2.0-2.3 mm. General color blackish. Scutum velvety black with silvery anterior diagonal bands from anterolateral band to midline, but not quite reaching

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Figure 102. Female. A-E: S. (Aspathia.) putre. A: front; B: fronto-ocular triangle; C: basal portion of cibarium; D: furcasternum; E: subbasal claw tooth. F: S. (A.) matteabranchium, cibarium. G-I: S. (A.) wygoi; G: cibarium; H: frons; I: subbasal claw tooth. J: S. (A.) jacumbae, cibarium. K: S. (A.) hunteri, cibarium. L-M: S. (A.) piperi; L: cibarium, M: claw. N: S. (A..) parrai, cibarium. O: S. (A.) bustosi, cibarium. P-R: S. (A.) sandyi; P: frons; Q-R: subbasal claw tooth in different orientations. S: S. (A.) metallicum, calcipala and pedisulcus.

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Figure 103. Female. A-B: S. (Aspathia) metallicum. A: eighth sternite and gonapophysis, spermatheca, genital fork; B: paraproct and cercus in different position. C-E: S. (A.) putre; C: eighth sternite and gonapophysis; D: paraproct and cercus; E: genital fork. F-H: S. (A.) matteabranchium; F: eighth sternite and gonapophysis; G: paraproct and cercus; H: genital fork. I-J: S. (A.) wygoi; I: eighth sternite and gonapophysis; J: paraproct and cercus. K: S. (A.) costalimai; paraproct and cercus.

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midline and not joined (Figs. 104 A, B) in lateral light, silver areas become iridescent. Legs similar to those of female; ratio of length/width of hind basitarsus about 4.8-5.4. Abdomen black, with 1+1 silvery spots on tergites II, IV-IX (Fig. 104 C). Gonocoxite and gonostylus as in figures 105 A-C; gonostylus with well-developed internal basal process and rounded edge; ventral plate subquadrate, with short median process (Figs. 106 A, B); endoparameral organ with robust spines (Fig. 105 D). Pupa: Cocoon basal length 3.6-4.0 mm, dorsal length 2.4-3.2 mm; body length 3.2 mm; gills 3.6-4.0 mm long; cocoon slipper-shaped, loosely woven, slightly reinforced anteriorly (Fig. 107 A). Frontoclypeus and thorax with abundant tubercles. Frontoclypeus with 1+1 facial and 2+2 bifid trichomes (Fig. 107 B); thorax with 5+5 bifid or trifid trichomes on disc (Fig. 108 A). Gill with 6 elongate filaments, with 3 short primary branches emerging from short stem. Abdominal chaetotaxy as in figure 110. Larva: Length 4.8-5.6 mm. Head light yellowish brown; cephalic apotome slightly brownish pigmented midbasally, with negative spots. Antenna longer than stem of cephalic fan; ratio of proximal, medial and distal segments = 1:1.5-1.6:0.9-1.2 (Fig. 111 M). Cephalic fan with 35-40 primary rays. Mandible with 9-10 inner teeth and 2 lateral mandibular processes. Postgenal cleft shallow (Fig. 112 C); ratio of hypostomium/ postgenal bridge = 0.9:1.0-1.1; hypostomium with 4+4 lateral setae and 4-5 lateral serrations. Sclerite of thoracic proleg with 18-22 teeth arranged in 5-6 groups. Anal sclerite with sparse hair and abundant spiculae; anal ring with 80 rows of 12-14 hooks each; rectal papillae with 3-4 diverticula on each lobe. Distribution: Colombia: Cundinamarca, Magdalena, Valle; Costa Rica; Belize; Cayo, Toledo; Ecuador: Azuay, Carchi, Chimborazo, Esmeraldas, Imbibura, Napo, Pichincha, Tungurahua, Zamora; Guatemala: Altaverapaz, Bajaverapaz, Chimaltenango, Chiquimula, El Peten, El Progreso, El Quiche, Escuintla, Guatemala, Huehuetenango, Jutiapa, Quetzaltenango, Retalhuleu, Sacatepequez, San Marcos, Santa Rosa, Solola, Suchitepequez; Jamaica; Mexico: Chiapas, Chihuahua, Colima, Durango, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, San Luis Potosi, Sinaloa, Sonora, Tabasco, Veracruz; Panama: Cocle, Colon; Peru: Huanuco; Trinidad; Venezuela: Anzoategui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcon, Guarico, Lara, Merida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Tachira, Trujillo, Yaracuy, Zulia. Bionomics: Simulium metallicum breeds in small creeks frequently in shaded areas and also in rivers, attached to submerged vegetation and fallen leaves, in slow to fast currents. Females attack humans and livestock and are considered vectors of Onchocerca volvulus. Discussion: Simulium metallicum is a widely distributed species complex and shows some morphological variation. Cytological studies indicate that it is a species complex of at least 7 cytospecies. CONN (1988) established cytotypes “A” and “B” from Guatemala and Mexico, “C” from Colombia, “D” and “E” from Venezuela, “F”, “J” and “K” from Panama, “G” from Costa Rica, “H” from Guatemala, Mexico, Panama, and “I” from Guatemala and Mexico. ARTEAGA & MUNOZ DE HOYOS (1999) found 2 cytotypes in Colombia, and MILLEST (1990) established differences between cytotypes and larval morphology. The closest species are S. horacioi, S. jobbinsi and S. racenisi; differences from each species are explained under those species. A detailed explanation of similarities with those species was given by SHELLEY et al. (2002: 174-175).

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Figure 105. Male. Gonocoxite and gonostylus. A-D: S. (Aspathia) metallicum; A: with ventral plate, endoparameral organ and median sclerite. D: endoparameres, median sclerite and aedeagal membrane in different position. E: S. (A.) putre. F: S. (A.) marquezi. G: S. (A.) iriartei. H-I: S. (A.) matteabranchium; H: gonocoxite, gonostylus; I: endoparameral organ and median sclerite. J: S. (A.) hechti. K: S. (A.) wygoi. L: S. (A.) spilmani.

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Simulium (Aspathia) morae RAMÍREZ PÉREZ, RASSI & RAMÍREZ Simulium (Simulium) morae RAMÍREZ PÉREZ et al., 1977: 167; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 569. Simulium morae: RAMÍREZ PÉREZ et al., 1982: 48-50; 1983: 59; RAMÍREZ PÉREZ, 1983: 98.

(Based on original description). Female: Vertex and frontoclypeus blackish, bright nacreous. Scutum dark, with golden hair, ornamentation similar to that of S. metallicum but without shining iridescence. Pleurae blackish with nacreous pollinosity. Abdomen with tergite II nacreous, the following tergites shining black with golden pilosity. Fronto-ocular triangle narrower than high at base. Sensory vesicle of palp about 1/2 length of segment. Basal portion of cibarium elevated medially, with abundant teeth. Tarsal claw with subbasal tooth well developed. Paraproct subtriangular, shorter than wide at base. Male: General coloration blackish as in female. Thorax black, with golden pilosity and with 1+1 anterior submedian dark triangles edged by nacreous areas. Gonostylus subcylindrical, acuminate distally, with robust internal basal process; ventral plate shorter than wide, subquadrate, with median haired process. Pupa: Cocoon length 4.0 mm, with loose, irregular weave. Frontoclypeus and thorax with tubercles abundant medially. Cephalic trichomes with 2-3 branches, thoracic trichomes with 2-5 branches. Gill composed of 6 branches with short stem and primary branches. Larva: Unknown. Distribution: Venezuela: Amazonas, Dpto. of Atabapo, Sierra Parima. Bionomics: Unknown. Discussion: This species is close to S. metallicum and S. racenisi but is differentiated in the female by a medially elevated cibarium with small denticles, which is unique among Neotropical Simulium (Aspathia) species, and in the male by a narrower gonostylus. Simulium (Aspathia) parrai VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 102 N; 104 J; 106 I; 109 H, I.) Simulium (Simulium) parrai VARGAS et al., 1946: 151-175; VARGAS & DÍAZ NÁJERA, 1948b: 336; 1957b: 264; DALMAT, 1951: 55; 1955: 230; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 65; 2004: 60; COSCARÓN et al., 1999: 569-570.

Female: Wing length 3.0 mm. Scutum blackish not ornamented, covered with dense silvery pilosity. Sensory vesicle of palp about 1/2 length of palpomere. Frons gradually convergent below, fronto-ocular triangle about as long as wide. Basal portion of cibarium with abundant papillae in middle (Fig. 102 N). Scutum without special ornamentation, with dark, strong hair intermixed with paler, weaker hair. Sc and basal section of R without hair. Ratio of length/width of hind basitarsus = 6.0. Tarsal claw with subbasal tooth well developed. Paraproct slightly shorter than wide, not acuminate distally. Stemite VIII with 25-29 hairs on each side; gonapophysis truncate, with slightly diverging, reinforced inner edge; spermatheca oval-shaped; genital fork with small submedian process on lateral branches.

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Figure 106. Male. Ventral plate. A-B: S. (Aspathia) metallicum, in different position. C: S. (A.) wygoi. D: S. (A.) marquezi. E: S. (A.) tricorne. F: S. (A.) piperi. G: S. (A.) puigi. H: S. (A.) iriartei. I: S. (A.) parrai. J: S. (A.) spilmani. K: S. (A.) jacumbae. L: S. (A.) hechti. M: S. (A.) hunteri. N: S. (A.) costalimai. O: S. (A.) ruizi. P: S. (A.) matteabranchium. Q: S. (A.) putre.

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Male: Wing length 2.7 mm. Scutum reddish brown, with pale golden iridescent pilosity and 1+1 anterior silvery diagonal cuneiform spots; lateral and posterior margins with silver pollinosity (Fig. 104 J). Ratio of length/width of hind basitarsus = 4.1. Gonostylus subcylindrical, with strong internal basal process; ventral plate body subquadrate, with short median process (Fig. 106 I). Pupa: Basal length 4.1 mm, dorsal length 3.0 mm; gill about 1.5 mm long. Cocoon flattened, anterior rim reinforced with conspicuous threads. Frontoclypeus and thorax without tubercles; thorax with 7-12 simple trichomes (Fig. 109 H). Gill membranous and pseudoannulated, with 1 basal and 4 large distal lobes with abundant and long filaments on median basal area (Fig. 109 I). Abdominal tergite II with 3 submedian setae developed as small hooks. Larva: Length 5.2-5.6 mm. Cephalic apotome with diffuse ornamentation on basal half, 1+1 spots and median basal darkened area present. Postgenal cleft miter-shaped. Postgenal bridge slightly shorter than height of hypostomium; ratio of hypostomium/postgenal bridge = 1.1; hypostomium with 6-7 lateral setae and 3-4 lateral serrations. Cephalic fan with 40-44 primary rays. Ratio of proximal, medial and distal segments = 1:1.1:0.8. Mandible with 9-11 inner teeth arranged in 2 rows, with 1 lateral mandibular process. Anal sclerite with hair, no scales; anal ring with 78 rows of 14-15 hooks each; rectal papillae not seen. Distribution: Guatemala: Chimaltenango, Solola; Mexico: Oaxaca, Veracruz. Bionomics: Unknown. Discussion: This species is similar to S. matteabranchium. Simulium covagarciai is similar in the globose membranous respiratory organ, but the unique morphology and disposition of the lobules in S. covagarciai allows differentiation. Simulium (Aspathia) piperi DYAR & SHANNON (Figs. 101 F, G; 102 L, M; 104 I; 106 F.) Simulium piperi DYAR & SHANNON, 1927: 38. Simulium (Aspathia) piperi: ADLER et al., 2004: 367. Simulium (Simulium) piperi PETERSON & KONDRATIEFF, 1995: 39; CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 572-573. Simulium knowltoni TWINN, 1938: 53. Simulium stonei STAINS & KNOWLTON, 1943: 277. Simulium sayi DYAR & SHANNON, 1927: 40.

Female: Wing length 3.0 mm. General coloration grayish brown. Head brownish, frons and clypeus grayish pollinose; frons gradually converging, fronto-ocular triangle about as high as wide. Sensory vesicle of palp about 1/2 length of palpomere. Basal portion of cibarium smooth, with small median prominence (Fig. 102 L) Thorax blackish brown with light gray pilosity; scutum when viewed from behind with silvery lyre-shaped pattern delimited by median and 1+1 sublateral black stripes (Fig. 101 F), and with anterior illumination, 1+1 subtriangular light grayish spots, followed by 1+1 black vittae that reach posterior edge (Fig. 101 G); metanotum velvety grayish brown. Abdomen brown with 1+1 silvery spots on tergite II, tergites VI-VII shining. Wing veins yellowish brown, with brown hair, Sc with 3-4 hairs. Legs brown, darker on apices of tibiae and tarsi, ratio of length/width of hind basitarsus = 6.1, tarsal claw with subbasal tooth well developed (Fig. 102 M). Stemite VIII strongly sclerotized, about 27-29 setae on each side;

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gonapophyses with distal edge truncate, inner edges more sclerotized, divergent below; paraprocts rounded distally, shorter than wide at base. Male: Wing length 2.8 mm. General coloration brownish, darker than female. Scutum blackish brown with grayish yellow decumbent hair, when viewed with lateral light from front, with 1+1 silvery triangular spots about 1/6 length of scutum (Fig. 104 I); in other lights, scutum silvery laterally and posteriorly. Ratio of length/width of hind basitarsus = 3.5. Gonostylus subcylindrical, twice as long as gonocoxite; internal-basal process long and smooth; ventral plate wider than high, with broad median process gradually acuminate distally (Fig. 106 F). Pupa: Cocoon projected dorsally; basal length 4.5-4.7 mm, dorsal length 4.0 mm, threads conspicuous. Gill with 10 very open branches and with short basal stem, 4 primary branches of about same length present. Thorax with 50-65 simple hairs. Frontoclypeus and dorsal portion of thorax with moderate number of small and smooth tubercles. Frontoclypeus with 1+1 facial and 2+2 frontal simple trichomes; thorax with 50-54 simple trichomes. Larva: Cephalic apotome with positive spots edged by diffuse pigmentation; cephalic fan with 35-38 primary rays; ratio of proximal, medial and distal segments = 1:1.3-1.5:1.0-1.1. Ratio of hypostomium/postgenal bridge = 0.5; hypostomium with 4-5 lateral setae and 8-9 lateral serrations. Inner mandibular teeth 10-12. Anal sclerite with hair, no scales. Comb of thoracic proleg with 12-15 teeth, not grouped. Anal ring with 7276 rows of 14-15 hooks each; rectal papillae lobes with 5-7 diverticula each. Distribution: Canada: Alberta, British Columbia, Saskatchewan; Mexico: Distrito Federal; USA: Arizona, California, Colorado, Idaho, Montana, Nebraska, New Mexico, Oregon, South Dakota, Texas, Utah, Washington, Wyoming. Bionomics: Simulium piperi breeds in small turbulent streams attached to aquatic vegetation. Females attack livestock and also bite humans. Discussion: Simulium jacumbae is the most closely related species; differences between them were discussed under S. jacumbae. Variation in morphology (e.g., gill branches numbering 9-13) suggests that S. piperi is a species complex (ADLER et al., 2004). Simulium (Aspathia) puigi VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 104 L; 106 G; 108 B, C.) Simulium (Simulium) puigi VARGAS et al., 1946: 153-155, 176; VARGAS & DÍAZ NÁJERA, 1957b: 191; TAKAOKA & TAKAHASI, 1982: 63; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004: 63; MOULTON, 1996: 741; COSCARÓN et al., 1999: 573. Simulium (Aspathia) puigi: ADLER et al., 2004: 368.

Female: Wing length 3.1 mm. Body color blackish. Scutal ornamentation lyreshaped, very similar to S. metallicum, appearing metallic green iridescent when viewed with lateral illumination. Legs predominantly yellowish. Abdomen dark dorsally, tergite II with 1+1 silvery pollinose spots, tergites VII-IX shining cinereous. Basal portion of cibarium smooth. Paraprocts 1/2 as long as wide, rounded distally; gonapophysis subcircular, with sclerotized inner margin. Stemite VIII with 26-30 hairs on each side. Male: Wing length 3.0 mm. Scutum velvety blackish with 1+1 suboval median and 1+1 diagonal anterior silver spots, joining basally with lateral and posterior marginal silver

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band (Fig. 104 L). Legs blackish, whitish basally on fore tibia, on basal half of middle tarsus, and hind basitarsus; ratio of length/width of hind basitarsus = 3.2. Abdomen blackish with 1+1 spots on tergite II, tergites VI-VII with 1+1 silvery greenish pollinose spots. Gonostylus subcylindrical, with internal basal process slightly developed; ventral plate subquadrate, as long as wide at base with wide median process basally (Fig. 106 G). Pupa: Cocoon slipper-shaped, heavy, with texture well defined and strongly reinforced on anterior edge, slightly projected ventrally; basal length 4.2 mm, dorsal length 3.4 mm; gill length 4.0-5.5 mm. Frontoclypeus and exposed portion of thorax with abundant large, elevated tubercles, surface wrinkled, and bearing spinules. Facial trichomes with 3 branches. Thoracic trichomes with 3-5 long branches on disc and 2-4 simple ones on lateral edge. Gill with 6 branches, with short stem and relatively short primary branches (Figs. 108 B, C). Larva: Length 6.0-6.5 mm. Cephalic apotome without spots or special ornamentation. Ratio of hypostomium/postgenal bridge = 0.8; hypostomium with 4-5 lateral setae and 2-4 lateral serrations on each side. Cephalic fan with 28-36 primary rays. Postgenal cleft medially developed, subtriangular; ratio of hypostomium/postgenal bridge = 0.8-1.0. Ratio of proximal, medial and distal segments = 1:1.2-1.3:1.1. Mandible with 8-10 inner teeth. Anal sclerite with scales and hair; anal ring with 64 rows of 11-12 hooks each; rectal papillae lobes with 5-8 diverticula each. Distribution: Costa Rica; Guatemala; Mexico: Aguascalientes, Chiapas, Oaxaca, Veracruz; USA: New Mexico. Bionomics: This species breeds in creeks with moderate flow and aquatic vegetation, from lowlands to 2700 m of altitude. Female feeding habits are unknown. Discussion: Simulium puigi is very difficult to separate from other species of Simulium (Aspathia). The wider spindle-shaped posterior basitarsus, the silver thoracic spots meeting with the lateral and posterior silvery area, the shorter basal process on the gonostylus in the male and absence of ornamentation on the cephalic apotome of the larva help identify S. puigi. Simulium (Aspathia) putre COSCARÓN & MATTA (Figs. 101 H, J; 102 A-E; 103 C-E; 104 E-H; 105 E; 106 Q; 107 C-E; 108 D; 111 A-G; 112 D-F.) Simulium putre COSCARÓN & MATTA, 1982: 52-54; 1987; CROSSKEY & HOWARD, 1997: 81; 2004: 60. Simulium (Simulium) putre COSCARÓN, 1987: 33; 1991: 268-269; COSCARÓN et al., 1999: 574-575; CROSSKEY & HOWARD, 2004: 60.

Female: General coloration blackish. Frons and clypeus blackish, frons shining nacreous, clypeus with silver pollinosity. Scutum blackish, when viewed with lateral illumination with 1+1 anterior subtrapezoidal silvery spots and marginal silvery pollinosity laterally and posteriorly (Fig. 101 H), with shifting light, becoming black; hair of scutum tin colored. Frons convergent below (Fig. 102 A); fronto-ocular triangle about as long as tall (Fig. 102 B). Maxillary palp as in figure 101 I. Cibarium smooth basally (Fig. 102 C). Sensory vesicle of palp about 1/2 length of palpomere (Fig. 101 J). Leg hair filiform; tarsal claw with well-developed subbasal tooth (Fig. 102 E). Gonapophyses parallel-sided along inner margins and with sclerotized margin truncate distally (Fig. 103 C); paraproct gradually produced ventrally, cerci with rounded edge (Fig. 103 D); genital fork as in figure 103 E.

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Figure 107. Pupa. A-B: S. (Aspathia) metallicum; A: lateral view; B: frontoclypeus. C-E: S. (A.) putre; C: lateral view; D: frontoclypeus; E: thoracic trichomes. F-G: S. (A.) wygoi; F: dorsal view; G: frontoclypeus. H: S. (A.) tricorne. I: S. (A.) marquezi. J: S. (A.) matteabranchium. K: S. (A.) hechti, thoracic trichomes.

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Figure 108. Pupa. Gill and thoracic trichomes. A: S. (Aspathia) metallicum. B-C: S. (A.) puigi. D: S. (A.) putre. E: S. (A.) spilmani. F: S. (A.) jacumbae. G: S. (A.) hunteri. H: S. (A.) iriartei. I: S. (A.) wygoi. J: S. (A.) hechti.

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Male: Wing length 2.8 mm. Scutum black, when viewed with lateral illumination with 1+1 silvery anterior spots and marginal silver pollinosity laterally and posteriorly (Figs. 104 E, F), with the same but more inclined illumination, color changes to black, with light from front, sublateral elongate silvery spots appear (Fig. 104 G). Abdomen black with 1+1 silvery spots on tergites II, V-VII. Sc and basal section of R bare. Pigmentation of hind leg as in figure 104 H; ratio of length/width of posterior basitarsus = 3.5. Gonostylus without internal-basal process but with high lateral keel from base to middle length and lateral submedian sinuosity, apically flattened with elongate depression on inner side where small subapical spur is inserted (Fig. 105 E); body of ventral plate flattened and curved, with ventral setae (Fig. 106 Q); parameres with well-developed teeth, median sclerite subrhomboidal. Pupa: Cocoon basal length 3.5-3.7 mm, dorsal length 3.0 mm, loosely woven; like chenille (Fig. 107 C). Frontoclypeus and thorax with moderate to abundant tubercles (Fig. 107 D), and simple slender hair (Fig. 107 E); frontoclypeus with 2+2 frontal and 1+1 facial trichomes; thorax with about 60-85 trichomes on each side. Gill grayish, with 6 filaments, stem and primary branches short (Fig. 108 D). Abdomen with terminal hooks relatively little developed. Larva: Maximum length 6.5 mm. General aspect as in figure 112 F. Head yellowish brown, body greenish gray. Cephalic apotome with positive spots (Fig. 111 A). Antenna longer than stem of cephalic fans; ratio of proximal, medial and distal segments = 1:0.8:1; proximal and medial segments with conspicuous transverse striations (Figs. 111 B-C). Mandible with 10 inner and 12 acuminate marginal teeth (Fig. 111 D). Cephalic fan with 3034 primary rays. Postgenal cleft very short, subtriangular (Fig. 112 D); ratio of hypostomium/postgenal bridge = 0.5; hypostomium teeth as in figure 112 E, lateral setae 3-4 and 5-7 lateral serrations on each side. Proleg sclerite with about 20 comb teeth arranged in groups and in 2 rows (Fig. 111 E). Anal sclerite with hair, no scales (Fig. 111 F); anal ring with 72 rows of 12-14 hooks each; rectal papillae with 3 simple lobes (Fig. 111 G). Distribution: Chile: Arica, Parinocota, Tarapaca. Bionomics: Simulium putre inhabits creeks at 3500 m elevation, where larvae and pupae are attached to aquatic vegetation. Discussion: Simulium putre is an isolated species and has the southernmost distribution of this subgenus. It has a unique female and male scutal ornamentation and male gonostylus morphology. The pupa, with 6 filamentous gill branches, is similar to that of S. metallicum, but the presence of trichomes with 2-6 branches and the 5+5 thoracic trichomes permit separation. Simulium (Aspathia) racenisi RAMÍREZ PÉREZ Simulium racenisi RAMÍREZ PÉREZ, 1971: 332; 1983: 66. Simulium (Simulium) racenisi COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 69; 2004: 63; COSCARÓN et al., 1999: 575. Simulium sp. G. 1962 LEWIS & IBAÑEZ ALDECOA, 1962: 449; LEWIS, 1963: 62.

(Based primarily on original description). Female: Body color blackish. Frontoclypeus nacreous pollinose; fronto-ocular triangle slightly wider than high. Sensory vesicle of palp about 1/3 length of segment. Basal portion of cibarium smooth. Scutum ornamentation very similar to that of S. metallicum. Scutellum dark

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brownish. Pleurae blackish, with silver pollinosity. Legs blackish, with femur and fore tibia, base of hind femur and most of middle and hind basitarsi whitish. Abdomen blackish, tergites VVIII shining; tergite II with 1+1 silvery spots. Tarsal claw with subbasal tooth well developed. Paraproct about as long as high, with truncate edge; gonapophysis wider than tall. Male: Gonostylus subcylindrical, with rounded apex and short intemal-basal process; ventral plate wider than tall. Pupa: Gill with 6 branches. Frontoclypeus with 3+3 bifid trichomes. Thorax with 56 bifid or trifid trichomes on each side. Larva: Cephalic apotome with small 1+1 median longitudinal and 1+1 sublateral spots; postgenal bridge as long as hypostomium. Distribution: Venezuela: Miranda, Monagas. Bionomics: Unknown. Discussion: Simulium racenisi is very close to S. metallicum, but according to RAMÍREZPEREZ, S. racenisi has longer primary and secondary gill branches, a shorter median process on the ventral plate, a stouter gonostylus distally in the male, and shorter paraprocts in the female. Because we were unable to examine specimens, we are uncertain about the validity of this species, particularly in view of the differences in genitalic characters, which depends on how they are viewed. Simulium (Aspathia) ruizi VARGAS & DÍAZ NÁJERA (Figs. 106 O; 109 E-G.) Simulium (Simulium) ruizi VARGAS & DÍAZ NÁJERA, 1948b: 330; 1957b: 265, 392; COSCARÓN, 1987: 33; CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 576-577.

Female: Wing length 2.6 mm. Body color blackish. Frons and clypeus blackish, clypeus with dirty white pruinosity. Cibarium with small rounded denticles. Sensory vesicle of palp nearly 1/2 as long as segment. Scutum black, when seen from behind with frontal light, there is 1 slender median blackish stripe and 1+1 wider submedian silvery pollinose vittae anteriorly, lateral and posterior margins silvery pollinose, when seen in frontal view, the color is inverted, with 1+1 submedian vittae separated by 0.25 of width of one, and 1+1 sublateral vittae nearly twice as wide as submedian bands of silver pollinosity. Scutellum yellowish brown. Abdomen concolorous with thorax, tergite II silvery pollinose. Legs predominantly yellowish brown. Paraprocts half-moon-shaped, nearly as long as basal width; stemite VIII with 16-18 hairs on each side. Male: Wing length 3.0 mm. Body blackish. Scutum black with 1+1 diagonal curved bands not joining silvery marginal band. Legs blackish, with basal 1/3 of hind basitarsus yellowish; posterior basitarsus wide, ratio of length/width = 2.7-3.2. Abdomen black, with 1+1 silvery spots on tergites I-VII. Gonostylus subcylindrical, with internal basal process smooth and shorter than basal transverse diameter; ventral plate subconical, 1.3 times longer than wide, with strong basal branches (Fig. 106 O). Pupa: Cocoon projected dorsally with anterior edge reinforced, tissue compact with conspicuous threads. Frontoclypeus and thorax without tubercles. Trichomes simple, curved, about 45-50 on each side. Gill membranous, haired and subcylindrical, with short basal and 3 apical branches (Fig. 109 E), and filaments simple (Fig. 109 F) or with several branches (Fig. 109 G).

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Figure 109. Pupa. Gill and thoracic trichomes. A: S. (Aspathia) matteabranchium. B: S. (A.) marquezi. C-D: S. (A.) sandyi; C: gill; D: thoracic trichome and tubercles. E-G: S. (A.) ruizi; E: gill; F-G: appendages of gill. H-I: S. (A.) parrai; H: thoracic sclerite with gill insertion and trichomes; I: frontal view.

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Figure 110. Pupa. Abdominal chaetotaxy of S. (Aspathia) metallicum.

Larva: Length 6.5 mm. Cephalic apotome scarcely darkened subbasally with 1 median and 1+1 submedian elongated spots. Postgenal cleft subtriangular, acuminate cephalad. Ratio of hypostomium/postgenal bridge = 1.1; hypostomium with subdental platform, 8-9 lateral setae and 2-3 lateral serrations on each side. Ratio of antennal segments proximal, medial and distal = 1:1.3:0.8. Cephalic fan with about 35-40 primary rays. Mandible with 911 inner teeth arranged in 2 rows, second marginal tooth very close to first. Thoracic proleg comb with 30 teeth arranged in about 7 groups with 1+1 well defined apodemes. Anal sclerite with simple hair, no scales; anal ring with 90 rows of 15 hooks each. Distribution: Mexico: Oaxaca, Veracruz. Bionomics: Unknown. Discussion: This species is very close to S. matteabranchium. The peculiar membranous and flexible filaments of the gill are similar to those of S. matteabranchium, S. parrai and S. covagarciai, but the subdivision of branches allows differentiation from other Neotropical species of the subgenus Aspathia.

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Simulium (Aspathia) sandyi COSCARÓN, IBÁÑEZ BERNAL & COSCARÓN ARIAS (Figs. 102 P-R; 109 C, D.) Simulium (Simulium) sandyi COSCARÓN et al., 1999: 577-578; CROSSKEY & HOWARD, 2004: 60.

Female (Description based on pharate female): Frons wide (Fig. 102 P), frontoocular triangle about as long as high. Basal portion of cibarium smooth, without denticles or spherules. Sensory vesicle of palp 1/2 length of palpomere. Basal section of R bare. Tarsal claw with subbasal tooth well developed (Figs. 102 Q, R). Male: Unknown. Pupa: Cocoon loosely woven and covered by detritus. Frontoclypeus and thorax with abundant tubercles, with spiculate surface (Fig. 109 D). Thorax with 16-19 simple, robust trichomes on each side. Gill membranous, without hair, with 25 terminal branches emerging from 3 short, stout primary branches, secondary branches digitiform, and terminal portion subdivided (Fig. 109 C). Larva: Unknown. Distribution: Mexico: Distrito Federal, Lagunas de Zempoala. Bionomics: Unknown. Discussion: The pupa of S. marquezi is closest to S. sandyi, but the gill has 2 robust basal branches and only 7 terminal branches, and thus is easily separated. Simulium (Aspathia) spilmani STONE (Figs. 105 L, 106 J; 108 E .) Simulium (Simulium) spilmani STONE, 1969: 313; CROSSKEY, 1987: 474; CROSSKEY & HOWARD, 1997: 69; 2004: 60; COSCARÓN et al., 1999: 578.

(Diagnosis based on original description). Female: Wing length 2.0 mm. Frons shining black, convergent below. Sensory vesicle of palp about 0.4 times length of palpomere. Basal portion of cibarium flat, unarmed. Scutum black with 1+1 submedian and 1+1 sublateral silvery pollinose stripes, anterior portion of sublateral stripes iridescent. Abdominal tergite II with 1+1 iridescent spots, tergites II-V velvety black, tergites VI-VIII shining black. Wing with Sc and basal section of R bare. Stemite VIII, gonapophysis, paraprocts and cerci unknown. Male: Scutum black with 1+1 oblique silvery stripes on frontal declivity, broad blue to gold velvety lateral stripe, iridescent blue on anterior edge. Leg hair filiform and petaloid; ratio of length/width of posterior basitarsus = 6. Gonostylus spatuliform, with thin subapical spur; subbasal process short, with abundant denticles (Fig. 105 L); ventral plate wide basally, with narrow median process edged by deep emargination (Fig. 106 J). Pupa: Body length 2.2 mm. Head and thorax with small tubercles, sparser on thorax; head with 2+2 bifid trichomes, thorax with 5-6 simple to bifid and 1+1 simple trichomes on each side. Gill 1.5-1.8 mm long, with 5 terminal branches, stalk short, with primary dorsal and ventral branches short and bifurcated, lateral branch simple, length similar to secondary branches (Fig. 108 E). Larva: Length 5.0-5.5 mm. Cephalic apotome darkened mediobasally, subtriangular, without positive head spots. Postgenal cleft deep, with anterior edge subacute; ratio of hypostomium/postgenal bridge = 1.1; hypostomium with 4+4 lateral serrations and 5

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setae on each side. Antennae transversally striated on proximal and medial segments; ratio of antennal segments proximal, medial and distal = 1:1.2-1.3:0.8-0.9. Mandible with 10-11 inner teeth and 1 lateral mandibular process, secondary marginal teeth very small. Cephalic fan with 38-40 primary rays. Thoracic proleg comb with 24-26 teeth arranged in 8-9 main groups. Anal sclerite with scarce hair, no scales; anal ring with 75-80 rows with 14 hooks each; rectal papillae with 3 simple lobes. Distribution: Dominica. Bionomics: Unknown. Discussion: The unique 5 branches of the gill of the pupa is a useful character to differentiate S. spilmani from other species of the subgenus Aspathia. The scutal ornamentation of the female and shape of the male genitalia place S. spilmani close to S. metallicum. Simulium (Aspathia) tricorne DE LEÓN (Figs. 106 E; 107 H.) Simulium tricornis DE LEÓN, 1945: 68. Simulium (Simulium) tricornis: VARGAS et al., 1946: 155-176; VARGAS & DÍAZ NÁJERA, 1957b: 265, 398; DALMAT, 1955: 234; COSCARÓN, 1987: 33. Simulium (Simulium) tricorne CROSSKEY & HOWARD, 1997: 66; 2004: 60; COSCARÓN et al., 1999: 580.

Female: Wing length 3.4 mm. Body blackish. Scutum black with dark golden and blackish pilosity lyre-shaped, scutum hair of different sizes. Abdomen blackish. Legs black, with coxa, anterior side of fore tibia, basal 1/2 of tibia and middle basitarsus, basal 1/2 of tibia, hind basitarsus and second tarsomere yellowish; posterior basitarsus wide, ratio of length/width = 6.5. Sensory vesicle of palp 1/4 length of palpomere. Frons convergent. Basal portion of cibarium smooth. Cerci wide, with truncate distal edge; paraprocts 1/2 length of basal width; sternite VIII with about 17-22 setae; gonapophyses divergent below, with internal margin more sclerotized. Male: Wing length 3.1 mm. Body blackish. Scutum with 1+1 cuneiform spots indented toward middle, lateral margin edged by silvery band; abdomen black, with 1+1 silver spots on tergite II, IV-VI. Ratio of length/width of posterior basitarsus = 3.7. Gonostylus subcylindrical, with short internal-basal process apically smooth and shorter than transverse diameter of gonostylus; ventral plate about as long as wide with short median process edged by submedian depressions; basal branches very robust (Fig. 106 E). Pupa: Cocoon basal length 3.0-3.5 mm, dorsal length 2.8-3.0 mm, with anterior edge reinforced, projected anteroventrally, tissue dense and threads conspicuous, general aspect as in figure 107 H. Frontoclypeus and exposed portion of thorax with sparse, small, irregularly arranged tubercles, superficially with microscopic spiculae as seen in S. sandyi. Thorax with about 25-40 simple trichomes, 2-3 of median anterior area about twice as large as others. Gills membranous, with 3 strong subcylindrical branches. Abdominal tergites I and II with flat, multicuspidate tubercles. Larva: Length 6.0-6.2 mm. Cephalic apotome with positive spots arranged in 2 groups: basal-median longitudinal spot and 1+1 submedian spots. Postgenal cleft subquadrate to rounded. Ratio of hypostomium/postgenal bridge = 0.7; hypostomium with 7-8 lateral setae and 5-6 lateral serrations on each side. Cephalic fan with 26-32 primary rays. Ratio of

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antennal segments proximal, medial and distal = 1:1.3:1.2. Mandible with 10-12 inner teeth in 2 series, small platform present below median tooth. Thoracic proleg sclerite with 27 teeth. Anal sclerite without hair or scales; anal ring with 76-78 rows of 12-14 hooks each; rectal papillae with 9-12 lobules each (DALMAT, 1955: 239). Distribution: Guatemala: Chimaltenango, Guatemala, Huehuetenango, San Marcos, Solola, Totonicapán; Mexico: Chiapas. Bionomics: Unknown. Discussion: The pupal gill with three branches is similar to that of S. kompi, but the branches are all orientated in same direction in S. kompi, permitting differentiation. Simulium (Aspathia) wygoi COSCARÓN, IBAÑEZ BERNAL & COSCARÓN ARIAS (Figs. 101 K; 102 G-I; 103 I, J; 105 K; 106 C; 107 F, G; 108 I; 111 H-K; 112 G, H.) Simulium (Simulium) wygoi COSCARÓN et al., 1999: 581; 582; CROSSKEY & HOWARD, 2004: 60.

Female (pharate): Scutum dark brown, with 1+1, anterior light grayish spots. Frons convergent below (Fig. 102 H); fronto-ocular triangle deep. Sensory vesicle of palp large, about 1/2 length of palpomere (Fig. 101 K). Cibarium scarcely elevated medially, bearing spherules (Fig. 102 G). Basal section of R bare. Tarsal claw with small subbasal tooth (Fig. 102 I); ratio of length/width of hind basitarsus = 6.5; hair of legs filiform. Thorax with decumbent hair. Stemite VIII with about 25 hairs on each side; gonapophyses with parallel sides on inner margin and rounded distally (Fig. 103 I); paraproct about as long as wide, not acuminate, but with small process on apex (Fig. 103 J); spermatheca with rows of spiculae. Male (pharate): Sensory vesicle about 1/3 length of palpomere. Hind basitarsus wide, ratio of length/width = 3.6. Gonocoxite as in figure 105 K; gonostylus subcylindrical, with internal-basal process relatively short, covered with acuminate denticles; ventral plate wide, subquadrate, with small, curved median process (Fig. 106 C); paramere with robust hooks. Pupa: Cocoon basal length 5.0 mm, dorsal length 3.3 mm, body length 4.4 mm, gill length 3.0 mm. Cocoon flattened, translucent, subcircular in dorsal view, thorax free dorsally, fused anteriorly into ring, anterior edge reinforced along thoracic edge and enlarged anteriorly at gill insertion (Fig. 107 F). Frontoclypeus and thorax with abundant tubercles and conspicuously rugose (Fig. 107 G), similar to S. parnassum MALLOCH. Frontal (2+2) and facial (1+1) trichomes simple; thorax with 5+5 simple trichomes. Gill with short stem and 2 main branches with 9-10 smaller, and flexible branches: 3-4 branches on stem, and 3 on each large branch, some can be subdivided into 2 or 3 branches, gill base with sclerotized apodeme (Fig. 108 I). Abdominal tergite II with stout setae resembling small hooks. Tergite V with 4 setae, the largest on margin, without anterior scales; hooks on sternites V-VII large, bifid. Larva: Cephalic apotome with negative spots and slightly darkened mediobasally (Fig. 112 H). Postgenal cleft scarcely developed (Fig. 112 G). Ratio of hypostomium/ postgenal bridge = 0.7-0.9; hypostomium with 5-8 lateral setae and 4+4 lateral serrations on each side. Antenna as in figure 111 H; ratio of proximal, medial and distal segments = 1:1.3-1.4:1.1-1.2. Mandible with 8-10 inner teeth and 2 lateral mandibular processes (Fig. 111 I), second marginal tooth small and variably shaped (Fig. 111 J). Cephalic fan with 38-

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Figure 111. Larva. A-G: S. (Aspathia) putre; A: dorsal view of head; B: antenna; C: Portion of antenna under high magnification; D: dental portion of mandible; E: comb of thoracic proleg sclerite; F: anal sclerite; G: rectal papillae. H-K: S. (A.) wygoi; H: antenna; I: dental portion of mandible; J: mandibular marginal teeth; K: rectal papillae. L: S. (A.) marquezi, antenna. M: S. (A.) metallicum, antenna. N: S. (A.) iriartei, antenna.

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H Figure 112. Larva. A-D: postgenal cleft and hypostomium; A: S. (Aspathia) marquezi. B: S. (A.) iriartei. C: S. (A.) metallicum. D: S. A(.) putre. E-F: S. (A.) putre; E: hypostomium anterior edge; F: general structure of larva. G-H: S. (A.) wygoi; G: postgenal cleft and hypostomium; H: frontoclypeus.

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42 primary rays. Thoracic proleg sclerite with 27-28 teeth. Anal sclerite with sparse hair and few scales; anal ring with 80 rows of 13-14 hooks each; rectal papillae with 13-15 diverticula each, one larger than others on each lobe (Fig. 111 K). Distribution: Mexico: Distrito Federal. Bionomics: Unknown. Discussion: This species is similar to S. parnassum MALLOCH, based on the clypeal and thoracic rugosity of the pupa. The gill morphology is similar to that of S. hechti but S. hechti has 3 main branches. Simulium (Hemicnetha) ENDERLEIN (Figs. 113-117; 151.) Hemicnetha ENDERLEIN, 1934a: 190 (as genus). Type-species: Simulium paynei (VARGAS) 1942a: 246, new name for Hemicnetha mexicana ENDERLEIN, 1934a: 190. Dyarella VARGAS et al., 1946: 105, as subgenus of Simulium. Type-species: Simulium mexicanum BELLARDI, 1862: 6.

Female: Species medium-sized to large, wing length 2.6-4.5 mm. Frons relatively wide, convergent, with gray pollinosity. Fronto-ocular triangle deep. Scutum brown to blackish, pollinose, frequently with 1+1 submedian longitudinal grayish stripes appearing as light lyre shape; tonality of vittae changing with different illumination; pilosity homogeneous to grouped and simulating scales. Abdomen brownish. Cibarium basally smooth, medially with well sclerotized cornuae, with or without small spiculae, as also central area which sometimes appear over pharyngeal membrane. Sensory vesicle of maxillary palp big, frequently half of palpomere length. Sc with hair, sometimes reduced only to basal area. Basal sector of R generally without hair. Legs without petaloid hair. Calcipala well developed; claws with big subbasal tooth. Eighth sternite with median depression. Gonapophysis well developed; frequently 2 times longer than wide, subtrianglular to subtrapezoidal with abundant microtrichiae and internal edge well sclerotized; big paraprocts subquadrangular, longer than wide, with distal edge straight to slightly curved or rounded; cerci mostly with distal edge flattened. Genital fork robust with anterior conical acuminate apodemes on lateral branches. Spermatheca subtrapezoidal with light neck connecting with spermathecal duct. Male: Coloration as in female but darker. Gonocoxite wider than high; gonostylus generally 2 times longer than gonocoxite; gonostylus flattened with median lateral protuberances, generally 2 times longer than wide, rounded apically and with small subapical spur. Ventral plate wider than high with keel projected apically as median process to about as long as wide or flattened laterally. Endoparameres with big bases and strong teeth. Pupa: Cocoon with compact weave, generally with anterior edge elevated, shoe shaped, sometimes with festoons protecting gills. Gills relatively thick, branching close to base with short basal trunk, filaments in number 6-90, externally frequently with striated annulations and spiculae, with rounded to acute apices. Frontoclypeus very wide in males with basal reinforcement over facial trichomes. Frontoclypeus and exposed portion of thorax generally without tubercles, but present and very small on facial area and posterior edge of thorax. Cephalic and thoracic trichomes small and generally single in number,

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with 2+2 frontal, 1+1 facial and 5+5 or 6+6 thoracic. Distal abdominal tergites without spine comb and no terminal spur. Larva: Body wider distally and flattened posterodorsally. Cephalic apotome with punctiform spots median and sublateral generally not evident. Hypostomium generally with frontal edge curved, with median tooth longer than lateral teeth. Mandible with anterior internal teeth big and showing several rows of teeth; marginal teeth in number of 1-2 with second tooth very small. Antenna with medial article bigger than proximal and distal. Anal ring with 200-500 rows with 22-73 hooks each. Rectal papillae with 5-41 diverticulae on each lobe. Simulium paynei was created by VARGAS for Hemicnetha mexicana ENDERLEIN because the latter corresponded to the genus Simulium and is a homonym of Simulium mexicanum BELLARDI, which also is a Simulium (Hemicnetha) species. Simulium (Hemicnetha) consists of 23 species in 4 species groups (COSCARÓN, 1987; CROSSKEY & HOWARD, 1997, 2004) plus another one not included. The subgenus is widely distributed in Central America, reaching the Nearctic Region with 4 species and South America with 7 species. The subgenus Simulium (Hemicnetha) is morphologically similar to some other Simulium taxa, showing relationships not only with the Neotropical fauna, including Hearlea, Trichodagmia and Thyrsopelma, but also the Afrotropical fauna such as Freemaniellum, Anasolen and Xenosimulium (MIRANDA ESQUIVEL & COSCARÓN, 2001, 2004), suggesting Gondwanian ancestors. There are two species from northern Venezuela, Simulium oviedoi and Simulium rivasi that were included in Hemicnetha (RAMÍREZ PÉREZ, 1971). These species have common characters with Hemicnetha, such as the smooth cibarium, deep fronto-ocular triangle, absence of hair on the basal sector of R, shape of the pupal cocoon and gill and reinforcement on the frontoclypeal base, characters that also are similar in Hearlea. Simulium oviedoi and Simulium rivasi have characters that differ from the known HEMICNETHA species groups, such as the gonapophysis acute apically and about as long as wide at the base (Fig. 118 C), subtriangular paraprocts (Fig. 118 C), gonostylus shorter than the gonocoxite, with slight distal curvature (Figs. 118 F, H), ventral plate subtriangular, without a median carina (Figs. 118 E, G) (closer to Ectemnaspis species) and larval body with dorsal curvature subterminally (Fig. 119 H ), which differs from the characteristic flat shape typical of Hemicnetha. These differences from Hemicnetha are considerable, but this subgenus offers the closest fit. In order to facilitate the arrangement of the Simulium Neotropical species, the OVIEDOI species group is included in Hemicnetha. The Simulium (HEMICNETHA) species groups are characterized as follows: PAYNEI species group: gonapophysis 2 times longer than wide, blunt apex with abundant microtrichiae and not sclerotized internally (Fig. 113 C); paraproct subrectangular, with anterior edge not concave. Gonostylus 2.1-2.2 times longer than wide at base, lateral sides with slight protuberances (Fig. 114 A); ventral plate with moderate body length and thick median process with abundant hair (Fig. 114 B). Cocoon well elevated anteriorly with collar of festoons (Figs. 115 A-D); frontoclypeus and exposed portion of thorax without tubercles, only diminutive tubercles on facial area; thorax dorsally generally with gelatinous cover of variable disposition according to species (Figs. 115 L-N). Postgenal bridge about as long as hypostomium. Gills with 8 branches. Hypostomium

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frontal edge curved (Fig. 117 N). Anal ring with about 220-360 rows of 40 hook each: Simulium paynei, Simulium rubrithorax, Simulium biuxinisa, Simulium virgatum, Simulium bricenoi. BRACHYCLADUM: Gonapophysis subtriangular, acute apex, nearly 2 times longer than width with sparse microtrichiae, moderate number of hairs and well sclerotized along inner margin (Fig. 113 Q); paraprocts membranous without microtrichiae and low number of hairs, apex rounded with small concavity posteriorly (Fig. 113 M). Cibarium with median area bearing small teeth (Fig. 113 J). Gonostylus relatively short with lateral protuberances, ratio of length/width about 1.5. Cocoon with aperture elevated, without festoons; frontoclypeus and thorax without tubercles, except on facial area; gill with 6-15 branches. Hypostomium anterior edge straight with median teeth generally of same length or longer than lateral teeth (Fig. 117 Q). Postgenal bridge longer than hypostomium. Anal ring with 134-190 rows with 25-28 hooks each. Simulium brachycladum, Simulium pulverulentum, Simulium cristalinum, Simulium guerrerense, Simulium lobatoi, Simulium solarii. MEXICANUM: Large variation in characters. Gonapophysis about as long as wide at base, up to 2 times longer than wide, distally acute to blunt (Figs. 113 K, V). Paraproct subrectangular, with abundant microtrichia and no anterior concavity (Fig. 113 R). Gonostylus about 3 times longer than width with slight lateral protuberances (Fig. 114 I); ventral plate about as long as wide, without median process but flattened laterally and with high keel (Fig. 114 J). Cocoon with aperture moderately to well elevated, no festoons; frontoclypeus and thorax with or without tubercles. Gill with 8-90 branches. Postgenal bridge frequently shorter than hypostomium; hypostomium front slightly curved with median teeth higher than lateral teeth. Postgenal cleft apex sharp. Anal ring with 200-550 rows with 27-55 hooks. Simulium mexicanum, Simulium seriatum, Simulium smarti, Simulium freemani, Simulium earlei, Simulium keenani, Simulium yepocapense, Simulium hinmani, Simulium hieroglyphicum. OVIEDOI: Eighth sternite well sclerotized on the center; gonapophysis subtriangular, relatively short with acute apex (Fig. 118 C) ratio of length/width at base = 0.7-0.9. Paraprocts subtriangular, slightly shorter than wide at base (Figs. 118 A, B). Genital fork with anterolateral process strong and well sclerotized (Fig. 118 D). Male ratio of length/ width of hind basitarsus = 4. Gonostylus shorter than gonocoxite subcylindrical with slight distal curvature and apical spur; ventral plate about as long as wide at base without any keel or lateral entrances (Figs. 118 E, G); endoparameres with strong teeth. Cocoon shoe shaped, relatively low anteriorly with reinforced anterior edge (Fig. 119 A). Frontoclypeus reinforced basally and very wide in males, nearly same shape as female, frontoclypeus and thorax with abundant granulose tubercles (Figs. 119 B, E), frequently very acute (Fig. 119 D) and multibranched trichomes (Fig. 119 G). Gill with 6 thick relatively short branches emerging form short and thick trunk (Figs. 119 C, F). Abdomen with apical spur small. Larval body with slight dorsal subterminal curvature and without ventral tubercles (Fig. 119 H). Cephalic apotome with positive spots. Antennae short and robust, only reaching half length of cephalic fan stem; medial article longer than others. Cephalic fan with 20-25 rays with thick teeth all of similar length (Fig. 119 J). Mandible with 2 marginal teeth of similar size (Fig. 119 K). Postgenal bridge as long as hypostomium length. Lateral sclerite of thoracic proleg with 32-45 teeth (Fig. 119 N). Anal sclerite with abundant hair, but no scales. Anal ring with 100-120 rows with 17-18 hooks each. Rectal papillae with 20-26 diverticula on each lobe.

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KEYS

TO

SIMULIUM (HEMICNETHA)

SPECIES

Females 1

Gonapophysis shorter than width at base (Fig. 118 C); paraprocts subtriangular and apically blunt (Figs. 118 A, B) .............................Simulium oviedoi, Simulium rivasi – Gonapophysis longer than width at base (Figs. 113 C, K, Q, T, V); paraprocts subrectangular and apically rounded or curved (Figs. 113 C, F, L, M, R, U) ....... 2 2(1) Scutum without evident longitudinal stripes; gonapophysis subtrapezoidal, with blunt or rounded apex, relatively short (Fig. 113 Q) ratio of length/width at base = 1.0-1.1 ............................................................................................................................... 3 – Scutum with 1+1 longitudinal light gray submedian stripes, generally joined posteriorly, forming lyre shape (Figs. 113 D, G), with different illumination, color changes from silvery to blackish; gonapophysis generally subtriangular and acuminate apically (Figs. 113 C, K.); ratio of length/width at base = 1.3-3.0 ................................................................................................................................................ 4 3(2) Wing length 4.0-4.4 mm; basal portion of cibarium with spiculae on median portion ..........................................................................................................Simulium smarti – Wing length 3.1-3.4 mm; basal portion of cibarium without spiculae on median portion (Fig. 113 B) .......................................... Simulium mexicanum, Simulium seriatum 4(2) Gonapophysis subtrapezoidal, with blunt apex (as Fig. 113 Q) .............................. 5 – Gonapophysis subtrapezoidal, with acute (Fig. 113 K) to blunt apex (Fig. 113 C) ................................................................................................................................................ 7 5(4) Gonapophysis relatively short, ratio of length/width at base = 1.0; paraproct with apical edge mostly straight ................................................................... Simulium freemani – Gonapophysis relatively long, ratio of length/width at base = 1.8-2.7; paraproct with apical edge curved .................................................................................................... 6 6(5) Scutum hair arranged homogeneously; ratio of length of palpomere/sensory vesicle = 2.3 ..............................................................................................Simulium keenani – Scutum hair arranged in groups; ratio of length of palpomere/sensory vesicle = 1.9 ..................................................................................................... Simulium hieroglyphicum 7(4) Gonapophysis with apex acute (Figs. 113 K, V) ......................................................... 8 – Gonapophysis with apex blunt (Fig. 113 C) ............................................................... 15 8(7) Paraproct membranous on posterior 3/4, without microtrichia and with sparse setae only on edge .............................................................................. Simulium cristalinum – Paraproct with variable sclerotization on posterior 3/4, but covered by microtrichia and relative abundance of setae (Figs. 113 L, M) ................................ 9 9(8) Small species, wing length generally less than 2.8 mm; ratio of palpomere length/ sensory vesicle = 1.6-1.8 ................................................................................................ 10 – Medium to big species, wing length 2.8-4.3; ratio of palpomere length/sensory vesicle = 2.2-2.5 ................................................................................................................ 13 10(9) Sc with hair at least on basal 3/4 of vein ................................................................... 11 – Sc with hair only on basal 1/4 of vein ........................................................................ 12 11(10) Scutum with submedian vittae joining posteriorly, forming lyre shape (Fig. 113 G); wing length 2.8-3.3 mm ................................................................ Simulium brachycladum

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Scutum with submedian vittae extending only to 2/3 of scutum, not joining posteriorly; wing length 2.4-2.9 mm ..................................................... Simulium lobatoi 12(10) Scutum dark brownish, without ornamentation changing in different illumination ....................................................................... Simulium pulverulentum, Simulium guerrerense – Scutum brown orange, with submedian vittae forming lyre shape in some illumination .................................................................................................. Simulium solarii 13(9) Big species, wing length 4.0-4.3 mm ...................................................Simulium hinmani – Medium-sized species, wing length 2.4-2.9 mm ........................................................ 14 14(13) Sc with hair all along vein ......................................................................... Simulium earlei – Sc with hair only on basal half of vein ......................................... Simulium yepocapense 15(7) Without hair on Sc; ratio of gonapophysis length/width = 2.2 .......................... .................................................................................................................... Simulium bricenoi – With hair on Sc; ratio of gonapophysis length/width = 1.4-1.6 ........................... 16 16(15) Gonapophysis with inner edge bearing microtrichia ..................... Simulium biuxinisa – Gonapophysis with inner edge not bearing microtrichia ............................................ ................................................. Simulium paynei, Simulium rubrithorax, Simulium virgatum Males Not included: Simulium biuxinisa (unknown) and S. keenani (lack of characters). 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(3) – 7(6) – 8(7)

Gonostylus shorter than gonocoxite (Figs. 118 F, H); ventral plate subtriangular and without carina (Figs. 118 E, G) ...........................Simulium oviedoi, Simulium rivasi Gonostylus longer than gonocoxite (Figs. 114 A, E, I, M, N); ventral plate not subtriangular and with carina (Figs. 114 B, C, F, H, J, L, O) .................................... 2 Ventral plate with strong median distal projection (Figs. 114 B, C, F, L) .............. 3 Ventral plate without strong median distal projection (Figs. 114 J, O) ................ 10 Ratio of length of gonostylus/gonocoxite = 1.4-1.8 ................................................ 4 Ratio of length of gonostylus/gonocoxite = 1.9-2.2 ................................................ 6 Ratio of gonostylus length/width = 1.3-2.0 ................................................................ 5 Ratio of gonostylus length/width = 2.4-2.9 ............................... Simulium rubrithorax Ventral plate body without 1+1 concavities on posterior edge (Fig. 114 F), ratio of gonostylus length/width = 1.7 .................................................... Simulium brachycladum Ventral plate body with 1+1 deep concavities on posterior edge, ratio of gonostylus length/width = 1.8-2.1 ........................... Simulium lobatoi, Simulium solarii Ratio of hind basitarsus length/width = 3.8-5.6 ........................................................ 7 Ratio of hind basitarsus length/width = 5.7-7.2 ........................................................ 9 Gonostylus with slight outer lateral protuberances; ratio of length of gonostylus/ gonocoxite = 1.6 ........................................................................................ Simulium paynei Gonostylus with well-developed outer lateral protuberances; ratio of length of gonostylus/gonocoxite = 1.8-1.9 ..................................................................................... Ventral plate body with 1+1 submedian concavities posteriorly (Fig. 114 C); scutum with dark pollinose stripes not reaching anterior edge; ratio of length/ width of hind basitarsus = 5.6 ............................................................ Simulium virgatum

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Ventral plate body straight posteriorly without submedian concavities; scutum with dark pollinose stripes reaching anterior edge; ratio of length/width of hind basitarsus = 3.8-4.0 .................................................................................. Simulium briceroi Ratio of hind basitarsus length/width = 5.7 ........................... Simulium pulverulentum Ratio of hind basitarsus length/width = 7.2 ............................... Simulium cristalinum Ventral plate body narrow, flattened laterally, appearing as axe in lateral view ......... 11 Ventral plate body not narrow or flattened laterally ................................................ 13 Ratio of length of gonostylus/gonocoxite = 2.0; ratio of gonostylus length/ width = 2.3 ..................................................................................... Simulium hieroglyphicum Ratio of length of gonostylus/gonocoxite = 2.2-2.3; ratio of gonostylus length/ width = 2.4-2.9 ................................................................................................................. 12 Ratio of hind basitarsus length/width = 3.3 .......................................Simulium smarti Ratio of hind basitarsus length/width = 3.8 ................................... Simulium freemani Ratio of hind basitarsus length/width = 7.2 ................................Simulium guerrerense Ratio of hind basitarsus length/width = 3.1-5.0 ...................................................... 14 Wing length 4.2-4.4 mm; ratio of hind basitarsus length/width = 5.0 .................... .....................................................................................................................Simulium hinmani Wing length 2.7-3.4 mm; ratio of hind basitarsus length/width = 3.1-4.8 ......... 15 Ratio of gonostylus length/width 2.3-2.5 .................................................................. 16 Ratio of gonostylus length/width 1.9/2.1 ................................................................. 17 Wing length 2.7-2.9 mm ....................................................................... Simulium seriatum Wing length 3.0-3.3 mm ................................................................... Simulium mexicanum Wing length 3.1 mm; ratio of gonostylus length/width = 2.1 ......... Simulium earlei Wing length 3.4 mm; ratio of gonostylus length/width = 1.9 Simulium yepocapense

Pupae 1 – 2(2) – 3(2) – 4(3) – 5(4) – 6(1)

Cocoon with festoons edging entire aperture (Figs. 115 A-D) ................................ 2 Cocoon without festoons edging entire aperture or festoons very reduced (Figs. 115 E-J) ................................................................................................................................ 6 Frontoclypeus and thorax without tubercles; thorax wrinkled (Fig. 116 E) ........... ...................................................................................................................... Simulium bricenoi Frontoclypeus and thorax with tubercles (Fig. 115 Q); thorax smooth or wrinkled ................................................................................................................................................ 3 Festoons of anterior edge of aperture joined with thin weave; gills not surpassing height of festoons (Fig. 115 B) .......................................................... Simulium biuxinisa Festoons of anterior edge of aperture free, without intermixed weave; gills surpassing or not height of festoons ............................................................................. 4 Without gelatinous material on thoracic dorsum ............................ Simulium virgatum With thick gelatinous material on thoracic dorsum (Figs. 115 L-N) ...................... 5 Thoracic thick gelatinous material with lateral entrances appearing as 4 branches (Fig. 115 M); frontoclypeus uniformly light brown ............................ Simulium paynei Thoracic thick gelatinous material with entrance only on median longitudinal line (Fig. 115 L); frontoclypeus darker on anterior area (Fig. 115 O) .......... Simulium rubrithorax Gill with 6 branches .......................................................................................................... 7

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Gill with 8 or more branches ........................................................................................ 10 Cocoon very elevated anteriorly (Fig. 115 E); frontoclypeus and exposed portion of thorax without tubercles or with only sparse tubercles ... Simulium brachycladum Cocoon medially elevated anteriorly (Figs. 115 F; 119 A); frontoclypeus and exposed portion of thorax with abundant tubercles (Figs. 119 B, E) or sparse and very dark with much sclerotization (Fig. 115 S) .......................................................... 8 Gills relatively short (Fig. 116 G), length 0.7-0.8 mm; frontoclypeus with tubercles only on base (Fig. 115 S) .................................................................. Simulium cristalinum Gills relatively long (Figs. 119 C, F), length 1.4-3.0 mm; frontoclypeus with abundant tubercles (Figs. 119 B, E) ............................................................................... 9 Frontoclypeal and thoracic tubercles acuminate, like spines (Fig. 119 D); gill length 1.4-1.7 mm ..................................................................................... Simulium oviedoi Frontoclypeal and thoracic tubercles not acuminate, granulose (Fig. 119 G); gill length 2.8-3.0 mm ....................................................................................... Simulium rivasi Gill with 8 branches ........................................................................................................ 11 Gill with more than 10 branches .................................................................................. 12 Frontoclypeus with moderate abundance of tubercles on frontal area, gills free; gill length 2.0-2.1 mm ........................................................................... Simulium freemani Frontoclypeus without tubercles on frontal area, gills protected by cocoon; gill length 0.9-1.8 mm ..................................................................................... Simulium lobatoi Gill with 10 branches ...................................................................................................... 13 Gill with more than 10 branches .................................................................................. 15 Cocoon not covering gills ............................................................ Simulium pulverulentum Cocoon covering gills ...................................................................................................... 14 Cocoon aperture cylindrical with same height anteriorly and posteriorly; cocoon length basally 2.3-2.7 mm, gills 0.8-1.0 mm; thoracic trichomes single ................... ................................................................................................................ Simulium guerrerense Cocoon aperture more elevated anteriorly than posteriorly; cocoon length basally 3.5-4.0 mm, gills 1.4-1.5 mm; thoracic trichomes with 2-5 branches ....................... ................................................................................................................ Simulium yepocapense Gill with 12 branches ...................................................................................................... 16 Gill with more than 12 branches .................................................................................. 17 Exposed portion of thorax with tubercles arranged homogeneously (Fig. 116 H); frontoclypeus with abundant tubercles ......................................... Simulium mexicanum Exposed portion of thorax arranged in 2+2 longitudinal stripes joined anteriorly (Figs. 116 I-J); frontoclypeus without abundant tubercles or with very sparse tubercles (Fig. 115 T) ............................................................................ Simulium seriatum Gill with 15 branches ..................................................... Simulium solarii, Simulium earlei Gill with 16 or more branches ...................................................................................... 18 Gill with 16 branches .................................................. Simulium earlei, Simulium keenani Gill with 18 or more branches ...................................................................................... 19 Gill with 18 branches; thorax not dorsally wrinkled .......................... Simulium smarti Gill with 60-90 branches (Fig. 116 L); thorax dorsally wrinkled ............................... ........................................................................................................... Simulium hieroglyphicum

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Larvae Not included: Simulium freemani, S. keenani and S. lobatoi. 1 – 2(1)

Postgenal bridge longer than hypostomium length (Fig. 117 P) .............................. 2 Postgenal bridge shorter or same as hypostomium length ....................................... 7 Hypostomium frontal edge almost straight, with median tooth at same level as lateral teeth; postgenal cleft arc shaped apically (Fig. 117 P) ................................... 3 – Hypostomium frontal edge convex, with median tooth extended beyond level of lateral teeth (Fig. 117 N); postgenal cleft with arc to acute shape apically ........... 6 3(2) Anal ring with about 134 rows of hooks; larva maximum length 4.5-4.8 mm ...... ............................................................................................................ Simulium pulverulentum – Anal ring with about 160-220 rows of hooks; larva maximum length 6.0-9.0 mm ................................................................................................................................................ 4 4(3) Cephalic apotome with negative spots (Fig. 117 F); rectal papillae with 5 diverticula on each lobe ................................................................ Simulium brachycladum – Cephalic apotome with positive spots; rectal papillae with 6-9 diverticula on each lobe ........................................................................................................................................ 5 5(4) Anal ring with 190-220 rows, and 28-30 hooks each; maximum larva length 9.0 mm ................................................................................................................. Simulium earlei – Anal ring with 160 rows, and 22 hooks each; maximum larva length 5.8 mm ....... .................................................................................................................Simulium guerrerense 6(2) Postgenal cleft subtriangular with acute apex; anal ring with 500-550 rows of hooks; maximum larva length about 13.2 mm ....................... Simulium hieroglyphicum – Postgenal cleft arc shaped with blunt apex; anal ring with 225 rows of hooks; maximum larva length about 8.0 mm .................................................Simulium hinmani 7(1) Maximum larva length 10.5-12.6 mm; anal ring with 280-455 rows of hooks ........ 8 – Maximum larva length 6.0-9.5 mm; anal ring with 100-320 rows of hooks ....... 10 8(7) Postgenal bridge as long as hypostomium length (Fig. 117 O) ........ Simulium biuxinisa – Postgenal bridge shorter than hypostomium length (Figs. 117 N, R) .................... 9 9(8) Anal ring with 450-460 rows of hooks; rectal papillae with 17-20 diverticula on each lobe ......................................................................................................Simulium smarti – Anal ring with 280-346 rows of hooks; rectal papillae with 9-15 diverticula on each lobe ................................................................ Simulium paynei, Simulium rubrithorax 10(7) Anal ring with 100-170 rows of hooks ....................................................................... 11 – Anal ring with 210-320 rows of hooks ....................................................................... 13 11(10) Antennae with thick articles (Fig. 119 I); second marginal tooth as long as first (Fig. 119 K) ...................................................................... Simulium oviedo, Simulium rivasi – Antennae with articles relatively thin; second marginal tooth less than half as long as first marginal tooth (Fig. 117 L) ............................................................................... 12 12(11) Maximum larva length 5.8-6.3 mm; cephalic apotome with negative spots (Fig. 117 G); postgenal bridge as long as hypostomium length (Fig. 117 Q); ratio of proximal, medial and distal antennal articles = 1:1.6-1.7:1.0 ................................ ................................................................................................................ Simulium cristalinum

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Maximum larva length 8.2-8.9 mm; cephalic apotome with positive spots; postgenal bridge shorter than hypostomium length; ratio of proximal, medial and distal antennal articles = 1: 1.2:0.8 ......................................................... Simulium solarii Postgenal bridge as long as hypostomium length (Fig. 117 N) .............................. 14 Postgenal bridge shorter than hypostomium length (Fig. 117 R) .......................... 15 Cephalic apotome with positive spots; rectal papillae with 10-17 diverticula on each lobe .............................................................................................. Simulium mexicanum Cephalic apotome with negative spots; rectal papillae with 7-8 diverticula on each lobe ............................................................................................................ Simulium virgatum Mandible with only 1 big marginal tooth (Fig. 117 M); maximum larva length 6.57.0 mm ...................................................................................................... Simulium seriatum Mandible with 1 big marginal tooth and another small tooth; maximum larva length 7.1-10.7 mm .......................................................................................................... 16 Maximum larva length 9.1-10.7 mm; ratio of proximal, medial and distal antennal articles = 1: 1.7:1.2 ................................................................................... Simulium bricenoi Maximum larva length 7.0 mm; ratio of proximal, medial and distal antennal articles = 1:1.9-2:1.4-1.5 ................................................................... Simulium yepocapense DE ESPECIES DE

S. (HEMICNETHA)

Hembras 1 – 2(1) –

3(2) – 4(2) – 5(4)

Gonapófisis más cortas que anchas en la base (Fig. 118 C); paraproctos subtriangulares y apicalmente romos (Figs. 118 A, B) ................................................. ............................................................................................Simulium oviedoi, Simulium rivasi Gonapófisis más largas que anchas en la base (Figs. 113 C, K, Q, T, V); paraproctos subrectangulares y apicalmente redondeados o curvados (Figs. 113 C, F, L, M, R, U) ...................................................................................................................... 2 Escudo sin bandas longitudinales bien evidentes; gonapófisis subtrapezoidales con ápice romo o redondeado y relativamente cortas (Fig. 113 Q); relación largo/ ancho en la base = 1,0-1,1 ............................................................................................... 3 Escudo con 1+1 bandas submedianas gris claro, generalmente unidas posteriormente en forma de lira (Figs. 113 D, G), con diferente iluminación cambia el plateado por coloración negruzca; gonapófisis generalmente con forma subtriangular y aguzadas apicalmente (Figs. 113 C, K); relación del largo/ancho en la base = 1,3-3,0 ................................................................................................................. 4 Largo ala 4,4 mm; porción basal del cibario con espículas sobre la porción mediana ........................................................................................................Simulium smarti Largo ala 3,1-3,4 mm; porción basal del cibario sin espículas sobre la porción mediana (Fig. 113 B) ........................................ Simulium mexicanum, Simulium seriatum Gonapófisis subtrapezoidal con ápice romo ( como en Fig. 113 Q) ...................... 5 Gonapófisis subtrapezoidal con ápice agudo (Fig. 113 K) hasta con ápice romo (Fig. 113 C) .......................................................................................................................... 7 Gonapófisis relativamente cortas; relación del largo/ancho en la base = 1,0; borde apical del paraprocto mayormente recto ........................................... Simulium freemani

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Gonapófisis relativamente largas; relación del largo/ancho en la base = 1,8-2,7; borde apical del paraprocto curvo ................................................................................. 6 Pelos del escudo dispuestos homogéneamente; relación largo palpómero/vesícula sensorial = 2,3 ...........................................................................................Simulium keenani Pelos del escudo dispuestos en grupos; relación largo palpómero/vesícula sensorial = 1,9 ................................................................................ Simulium hieroglyphicum Gonapófisis con ápice aguzado (Figs. 113 K, V) ........................................................ 8 Gonapófisis con ápice romo (Fig. 113 C) ................................................................... 15 Paraprocto membranoso sobre la 3/4 porción posterior, sin microtriquias y escasas cerdas solo sobre el borde ................................................. Simulium cristalinum Paraprocto con variable grado de esclerotización sobre la 3/4 porción posterior, pero cubierta por microtriquias y abundantes cerdas (Figs. 113 L, M) .................. 9 Especies pequeñas; largo ala generalmente por debajo de 2,8 mm; relación del largo del palpómero/vesícula sensorial = 1,6-1,8 ..................................................... 10 Especies medianas a grandes; largo ala 2,8-4,3; relación del largo del palpómero/ vesícula sensorial 2,2-2,5 ................................................................................................. 13 Sc con pelos por lo menos en el 3/4 basal de la vena ............................................. 11 Sc con pelos solo en el 1/ 4 basal de la vena ............................................................ 12 Escudo con 1+1 banda submediana uniéndose posteriormente dando aspecto de lira (Fig. 113 G); largo ala 2,8-3,3 mm ...................................... Simulium brachycladum Escudo con 1+1 banda submediana alcanzando solamente los 2/3 del escudo; no uniéndose posteriormente; largo ala 2,4-2,9 mm ............................... Simulium lobatoi Escudo castaño oscuro, sin cambio de ornamentación con diferente iluminación ........................................................................ Simulium pulverulentum, Simulium guerrerense Escudo castaño naranja, mostrando con alguna iluminación las bandas submedianas en forma de lira ................................................................. Simulium solarii Especies grandes, largo ala 4,0-4,3 mm .............................................Simulium hinmani Especies medianas, largo ala 2,4-2,9 mm .................................................................... 14 Sc con pelos todo a lo largo de la vena .................................................. Simulium earlei Sc con pelos solo en la mitad basal ................................................ Simulium yepocapense Sin pelos sobre la Sc; relación largo/ancho en la base de gonapófisis = 2,2 ......... ...................................................................................................................... Simulium bricenoi Con pelos sobre la Sc; relación largo/ancho en la base de gonapófisis = 1,4-1,6 .............................................................................................................................................. 16 Borde interno de gonapófisis con microtriquias ............................ Simulium biuxinisa Borde interno de gonapófisis sin microtriquias ............................................................ ................................................. Simulium paynei, Simulium rubrithorax, Simulium virgatum

Machos Simulium biuxinisa y S. keenani no incluidas por desconocimiento de caracteres. 1

Gonostilo más corto que el gonocoxito (Figs. 118 F, H); placa ventral subtriangular y sin carena (Fig. 118 G) .....................Simulium oviedoi, Simulium rivasi

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– 2(1) – 3(2) – 4(3) – 5(4) – 6(3) – 7(6) – 8(7) – 9(6) – 10(2) – 11(10) – 12(11) – 13(10) – 14(13) – 15(14) – 16(15) –

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Gonostilo más largo que el gonocoxito (Figs. 114 A, E, I, M, N); placa ventral no subtriangular y con carena (Figs. 114 B, C, F, H, J, L, O) ......................................... 2 Placa ventral con fuerte prolongación medio distal (Figs. 114 B, C, F, L) ............ 3 Placa ventral sin fuerte prolongación medio distal (Figs. 114 J, O) ...................... 10 Relación largo gonostilo/gonocoxito = 1,4-1,8 .......................................................... 4 Relación largo gonostilo/gonocoxito = 1,9-2,2 .......................................................... 6 Relación largo/ancho del gonostilo = 1,3-2,0 ............................................................. 5 Relación largo/ancho del gonostilo = 2,4-2,9 ............................ Simulium rubrithorax Cuerpo de placa ventral sin 1+1 concavidad sobre el borde posterior (Fig. 114 F): relación largo/ancho del gonostilo = 1,7 ................................. Simulium brachycladum Cuerpo de la placa ventral con 1+1 concavidad profunda sobre el borde posterior; relación largo/ancho del gonostilo = 1,8-2,1 ........ Simulium lobatoi, Simulium solarii Relación largo/ancho de basitarso posterior 3,8-5,6 .................................................. 7 Relación largo/ancho de basitarso posterior 5,7-7,2 .................................................. 9 Gonostilo con escasas protuberancias látero-externas; relación largo del gonostilo/gonocoxito = 1,6 ..................................................................... Simulium paynei Gonostilo con protuberancias latero-externas bien desarrolladas; relación largo del gonostilo/gonocoxito = 1,8-1,9 ............................................................................... 8 Cuerpo de la placa ventral con 1+1 concavidad posterior submediana (Fig. 114 C); bandas polinosas oscuras del escudo no llegan al borde anterior; relación largo/ ancho del basitarso posterior = 5,6 .................................................... Simulium virgatum Cuerpo de la placa ventral recto posteriormente, sin concavidad submediana; bandas polinosas oscuras del escudo llegan al borde anterior; relación largo/ ancho del basitarso posterior = 3,8-4,0 ............................................... Simulium bricenoi Relación largo/ancho del basitarso posterior = 5,7 ............... Simulium pulverulentum Relación largo/ancho del basitarso posterior = 7,2 ................... Simulium cristalinum Cuerpo de placa ventral angosto aplastado lateralmente semejando un hacha antigua ................................................................................................................................ 11 Cuerpo de placa ventral no angosto, ni aplastado lateralmente ............................. 13 Relación del largo gonostilo/gonocoxito = 2,0; relación largo/ancho de gonostilo = 2,3 ................................................................................................. Simulium hieroglyphicum Relación del largo gonostilo/gonocoxito = 2,2-2,3; relación largo/ancho del gonostilo = 2,4-2,9 ........................................................................................................... 12 Relación largo/ancho del basitarso posterior = 3,3 ...........................Simulium smarti Relación largo/ancho del basitarso posterior = 3,8 ....................... Simulium freemani Relación largo/ancho del basitarso posterior = 7,2 ....................Simulium guerrerense Relación largo/ancho del basitarso posterior = 3,1-5,0 .......................................... 14 Largo ala 4,2-4,4 mm; relación largo/ancho del basitarso posterior = 5,0 ............. .....................................................................................................................Simulium hinmani Largo ala 2,7-3,4 mm; relación largo/ancho del basitarso posterior = 3,1-4,8 ......... 15 Relación largo/ancho del gonostilo 2,3-2,5 ............................................................... 16 Relación largo/ancho del gonostilo 1,9-2,1 ............................................................... 17 Largo ala 2,7-2,9 mm ............................................................................ Simulium seriatum Largo ala 3,0-3,3 mm ........................................................................ Simulium mexicanum

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17(15) Largo ala 3,1 mm ; relación gonostilo largo/ancho = 2,1 ................. Simulium earlei – Largo ala 3,4 mm; relación gonostilo largo/ancho = 1,9 ......... Simulium yepocapense Pupas 1 –

Capullo con festones bordeando totalmente la abertura (Figs. 115 A-D) ............. 2 Capullo sin festones bordeando totalmente la abertura o muy reducidos (Figs. 115 E-J) ........................................................................................................................................ 6 2(1) Frontoclípeo y tórax sin tubérculos, tórax rugoso (Fig. 116 E) ........ Simulium bricenoi – Frontoclípeo y tórax con tubérculos (Fig. 115 Q); tórax liso o rugoso ................. 3 3(2) Festones del borde anterior de la apertura unidos por un tejido fino; branquias no sobrepasando la altura de los festones (Fig. 115 B) ...................... Simulium biuxinisa – Festones del borde anterior de la apertura libres, sin tejido entremezclado; branquias sobrepasando o no la altura de los festones .............................................. 4 4(3) Sin membrana gelatinosa sobre el dorso del tórax ......................... Simulium virgatum – Con gruesa membrana gelatinosa sobre el dorso del tórax (Figs. 115 L-N) ........ 5 5(4) Tórax con gruesa membrana gelatinosa con entradas laterales presentando como 4 ramas (Fig. 115 M); frontoclípeo castaño claro homogéneo ......... Simulium paynei – Tórax con gruesa membrana gelatinosa con una sola entrada sobre la línea media longitudinal (Fig. 115 L); frontoclípeo más oscuro sobre el área anterior (Fig. 115 O) .......................................................................................................... Simulium rubrithorax 6(1) Branquias con 6 ramas ...................................................................................................... 7 – Branquias con 8 ó más ramas ........................................................................................ 10 7(6) Capullo muy elevado anteriormente (Fig. 115 E); frontoclípeo y porción expuesta del tórax sin o con escasos tubérculos ...................................... Simulium brachycladum – Capullo medianamente elevado anteriormente (Figs. 115 F; 119 A); frontoclípeo y porción expuesta del tórax con abundantes tubérculos (Figs. 119 B, E) o muy escasos oscuros, con gran esclerotización (Fig. 115 S) .............................................. 8 8(7) Branquias relativamente cortas (Fig. 116 G); largo 0,7-0,8 mm; frontoclípeo con tubérculos solo en la base (Fig. 115 S) .......................................... Simulium cristalinum – Branquias relativamente largas (Figs. 119 C, F); largo 1,4-3,0 mm; frontoclípeo con abundantes tubérculos (Figs. 119 B, E) ................................................................. 9 9(8) Frontoclípeo y tórax con tubérculos aguzados como espinas (Fig. 119 D); largo branquia 1,4-1,7 mm ................................................................................ Simulium oviedoi – Frontoclípeo y tórax con tubérculos no aguzados y granulosos (Fig. 119 G); largo branquia 2,8-3,0 mm .................................................................................. Simulium rivasi 10(6) Branquia con 8 ramas ...................................................................................................... 11 – Branquia con 10 o más ramas ....................................................................................... 12 11(10) Frontoclípeo con tubérculos de área frontal en moderada abundancia; branquias libres; largo branquia 2,0-2,1 mm ....................................................... Simulium freemani – Frontoclípeo sin tubérculos en área frontal; branquias protegidas por capullo; largo branquias 0,9-1,8 mm .................................................................... Simulium lobatoi 12(10) Branquias con 10 ramas .................................................................................................. 13 – Branquias con más de 10 ramas ................................................................................... 15 13(12) Capullo no cubre las branquias ................................................... Simulium pulverulentum

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– Capullo cubriendo las branquias ................................................................................... 14 14(13) Abertura del capullo en forma cilíndrica con la misma altitud anterior y posterior; largo capullo en la base 2,3-2,7 mm; branquias 0,8-1,0 mm; tricomas torácicos simples ...................................................................................................Simulium guerrerense – Abertura del capullo más elevada anterior que posteriormente; largo capullo en la base 3,5-4,0 mm; branquias 1,4-1,5 mm; tricomas torácicos con 2-5 ramas .......... ................................................................................................................ Simulium yepocapense 15(12) Branquia con 12 ramas ................................................................................................... 16 – Branquia con más de 12 ramas ..................................................................................... 17 16(15) Porción expuesta del tórax con tubérculos distribuidos homogéneamente (Fig. 116 H); frontoclípeo con abundantes tubérculos ........................ Simulium mexicanum – Porción expuesta del tórax con tubérculos dispuestos en 2+2 bandas longitudinales unidas anteriormente (Figs. 116 I, J); frontoclípeo sin abundantes tubérculos o muy escasos (Fig. 115 T) .............................................. Simulium seriatum 17(15) Branquia con 15 ramas .................................................. Simulium solarii, Simulium earlei – Branquia con 16 o más ramas ....................................................................................... 18 18(17) Branquia con 16 ramas ............................................... Simulium earlei, Simulium keenani – Branquia con 18 o más ramas ....................................................................................... 19 19(18) Branquia con 18 ramas .............................................................................Simulium smarti – Branquia con 60-90 ramas (Fig. 116 L); tórax dorsalmente rugoso .......................... ........................................................................................................... Simulium hieroglyphicum Larvas No incluidas: Simulium freemani, S. keenani y S. lobatoi. 1 – 2(1) – 3(2)

4(3) – 5(4) –

Puente postgenal más largo que el hipostomio (Fig. 117 P) .................................... 2 Puente postgenal más corto que el hipostomio .......................................................... 7 Borde frontal del hipostomio casi recto, con diente mediano al mismo nivel que el diente lateral; ápice de hendidura postgenal con forma de arco (Fig. 117 P) ....... 3 Borde frontal del hipostomio convexo, con diente mediano sobre el nivel del diente lateral (Fig. 117 N); ápice de abertura postgenal con forma de arco o aguzado ................................................................................................................................. 6 Anillo anal con alrededor de 134 hileras de ganchos; máximo largo de larva 4,54,8 mm .............................................................................................. Simulium pulverulentum Anillo anal con 160-220 hileras de ganchos; máximo largo de larva 6,0-9,0 mm ................................................................................................................................................ 4 Apotoma cefálico con manchas negativas (Fig. 117 F); papilas rectales con 5 divertículos en cada lóbulo .......................................................... Simulium brachycladum Apotoma cefálico con manchas positivas; papilas rectales con 6-9 divertículos en cada lóbulo .......................................................................................................................... 5 Anillo anal con 190-220 hileras y con 28-30 ganchos en cada una; máximo largo de larva 9,0 mm ........................................................................................... Simulium earlei Anillo anal con 160 hileras y con 22 ganchos en cada una; máximo largo de larva 5,8 mm ...................................................................................................Simulium guerrerense

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6(2) – 7(1) – 8(7) – 9(8) – 10(7) – 11(10) – 12(11)

– 13(10) – 14(13) – 15(13) – 16(15) –

Hendidura postgenal subtriangular, con ápice aguzado; anillo anal con 500-550 hileras de ganchos; máximo largo de larva alrededor de 13,2 mm ........................... ........................................................................................................... Simulium hieroglyphicum Hendidura postgenal en forma de arco con ápice romo; anillo anal con 225 hileras de ganchos; máximo largo de larva alrededor de 8,0 mm ............. Simulium hinmani Máximo largo de larva 10,5-12,6 mm; anillo anal con 280-455 hileras de ganchos ................................................................................................................................................ 8 Máximo largo de larva 6,0-9,5 mm; anillo anal con 100-320 hileras de ganchos .............................................................................................................................................. 10 Puente postgenal tan largo como el largo del hipostomio (Fig. 117 O) .................. ................................................................................................................... Simulium biuxinisa Puente postgenal más corto que el largo del hipostomio (Figs. 117 N, R) ........... 9 Anillo anal con 450-460 hileras de ganchos; papilas rectales con 17-20 divertículos sobre cada lóbulo .......................................................................................Simulium smarti Anillo anal con 280-346 hileras de ganchos; papilas rectales con 9-15 divertículos sobre cada lóbulo ................................................. Simulium paynei, Simulium rubrithorax Anillo anal con 100-170 hileras de ganchos ............................................................... 11 Anillo anal con 210-320 hileras de ganchos ............................................................... 13 Antena con artejos gruesos (Fig. 119 I); segundo diente marginal tan largo como el primero (Fig. 119 K) .................................................Simulium oviedoi, Simulium rivasi Antena con artejos relativamente finos; segundo diente marginal menos de la mitad del primer diente marginal (Fig. 117 L) ........................................................... 12 Máximo largo de larva 5,8-6,3 mm; apotoma cefálico con manchas negativas (Fig. 117 G); puente postgenal tan largo como el hipostomio (Fig. 117 Q); relación de los artejos antenales proximal, mediano y distal = 1:1,6-1,7:1,0 ....... ................................................................................................................ Simulium cristalinum Máximo largo de larva 8,2-8,9 mm; apotoma cefálico con manchas positivas; puente postgenal más corto que el hipostomio; relación de los artejos antenales proximal, mediano y distal = 1:1,2:0,8 .................................................. Simulium solarii Puente postgenal tan largo como el hipostomio (Fig. 117 N) ............................... 14 Puente postgenal más corto que el hipostomio (Fig. 117 R) ................................. 15 Apotoma cefálico con manchas positivas; papilas rectales con 10-17 divertículos sobre cada lóbulo ............................................................................... Simulium mexicanum Apotoma cefálico con manchas negativas; papilas rectales con 7-8 divertículos en cada lóbulo .............................................................................................. Simulium virgatum Mandíbula con solo un diente marginal grande (Fig. 117 M); máximo largo de larva 6,5-7,0 mm ..................................................................................... Simulium seriatum Mandíbula con un gran diente marginal y otro pequeño; máximo largo de larva 7,1-10,7 mm ...................................................................................................................... 16 Máximo largo de larva 9,1-10,7 mm; relación de artejos antenales proximal, mediano y distal = 1: 1,7:1,2 .................................................................. Simulium bricenoi Máximo largo de larva 7,0 mm; relación de artejos antenales proximal, mediano y distal = 1:1,9-2,1:1,4-1,5 ................................................................... Simulium yepocapense

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PAYNEI species group Simulium (Hemicnetha) paynei VARGAS (Figs. 113 D; 115 M, P.) Simulium (Eusimulium) paynei VARGAS, 1942a: 246 (n. nom. for Simulium mexicanum ENDERLEIN, 1934a: 190-191, homonym of Simulium mexicanum BELLARDI, 1862: 6). Simulium (Hemicnetha) paynei VARGAS & DÍAZ NÁJERA, 1957b: 175, 176, 178, 179, 232, 240, 241, 322; WYGODZINSKY, 1958: 134-140; COSCARÓN, 1987: 36; IBÁÑEZ-BERNAL, 1992: 212-228; COSCARÓN ARIAS, 2003: 200, 205-208, 213; ADLER et al., 2004: 372; CROSSKEY & HOWARD, 1997: 38; 2004: 30. Hemicnetha paynei PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Simuloium (Dyarella) virgatum: VARGAS et al., 1946: 111; DALMAT, 1955: 169-170. Simulium mathesoni VARGAS, 1943b: 360-363. Simulium bilimekae SMART, 1944: 132. Simulium (Dyarella) acatenangoensis DALMAT, 1951: 31-38. Simulium (Hemicnetha) conviti RAMÍREZ PÉREZ & VULCANO, 1973: 376-379. ?Simulium mutucuna NUNES DE MELLO & VIEIRA DA SILVA, 1974: 69-71.

Female: Wing length 3.6-3.9mm. Frons and clypeus brownish with grayish pruinosity. Scutum light brown with 1+1 submedian brown-reddish lyre-shaped stripes (Fig. 113 D) that with different light position become thicker and darker; humeral area lighter; pilosity decumbent, pale gray with hair arranged in groups not well defined. Scutellum and metanotum brown, the latter with velvety aspect. Pleurae brown, grayish pollinose. Legs light grayish brown with darker areas brown-blackish. Abdomen dark brown. Cibarium with small serrations on lateral cornuae, wing chaetotaxy and genitalia similar to those of S. rubrithorax. Ratio of palpomere/sensorial vesicle 1.5-1.7. Male: 3.1-3.6 mm. Scutum dark brown to reddish brown with 1+1 silvery submedian stripes joining posterior gray pollinose area, showing variation in accordance with light position. Legs as in female but more darkened; ratio of length/width of hind basitarsus = 4.3-4.4. Abdomen dark brown. Genitalia as in S. rubrithorax. Pupa: Cocoon and gill shape as in S. rubrithorax. Length basally about 3.7 mm, gills 2.5 mm. Frontoclypeus with abundant platelets on facial area (Fig. 115 P). Thorax dark brown dorsally, smooth on exposed portion and with small acuminate tubercles, relatively abundant on posterior portion. Thorax dorsally covered by gelatinous substance, with lateral entrances appearing as 4 branches (Fig. 115 M). Larva: Maximum length 10.4 mm. Cephalic apotome slightly darkened on basal 2/3, with positive spots on median longitudinal area and 1+1 sublateral spots. Ratio of proximal, medial and distal antennal articles = 1:1-1.2:0.9-1.1. Cephalic fan with 55-65 rays. Hypostomium with 7-8 lateral setae on each side. Postgenal bridge slightly shorter than hypostomium length. Lateral sclerite of thoracic proleg with 54-63 teeth. Anal ring with 326344 rows with 47-57 hooks each. Rectal papillae with 9-15 diverticula on each lobe. Distribution: Colombia: Cundinamarca; Ecuador: Pichincha; Guatemala: Chimaltenango, Huehuetenango, Sacatepiquez, Santa Rosa, Totonicapan; Mexico: Chiapas, Colima, Durango, Guerrero, Jalisco, Mexico, Michoacan, Morelos, Nuevo León, Oaxaca, Potosí, Puebla, San Luis, Sinaloa, Veracruz; Peru: Lima, Junin; Venezuela: Aragua,

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I

A

D B

G

H

O

J V P C T N M

L

U

Q

K F S R

E

Figure 113. Female. A-C: S. (Hemicnetha) rubrithorax; A: scutum; B: cibarium, C: genitalia ventral view, showing eighth sternite, gonapophysis, cercus and paraproct (at left cercus and paraproct mounted on slide), genital fork and spermatheca. D: S. (H.) paynei, scutum. E-F: S. (H.) biuxinisa; E: palpomere and sensory vesicle; F: cercus and paraproct. G-L: S. (H.) brachycladum; G: scutum with frontal illumination, H: scutum with posterior illumination; I: fronto-ocular triangle; J: cibarium; K: eighth sternite and gonapophysis; L: cercus and paraproct. M: S. (H.) pulverulentum, cercus and paraproct. N: S. (H.) cristalinum, scutum hairs. O-S: S. (H.) seriatum; O: cibarium; P: claw; Q: eighth sternite and gonapophysis; R: cercus and paraproct; S: genital fork. T-U: S. (H.) smarti; T: eighth sternite and gonapophysis; U: cercus and paraproct. V: S. (H.) yepocapense, eighth sternite and gonapophysis.

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Carabobo, Federal District, Guarico, Mérida, Táchira, Nueva Esparta; USA: Arizona, Oklahoma, Texas. Bionomics: Simulium paynei breeds in small and torrential creeks, primarily attached to rocks in water of 10-25 °C. Females bite cows and horses (DALMAT, 1955). Discussion: The closest species are S. rubrithorax and S. biuxinisa; the differences are explained under each of those species. Simulium bricenoi shows great similarity, but the wrinkled thorax of the pupa allows separation. Simulium (Hemicnetha) rubrithorax Figs. 113 A-C; 114 A, B; 115 A, L, O; 116 A; 117 A, D, H, K, N. Simulium rubrithorax LUTZ, 1909: 132; 1910: 221-224; MAIA-HERZOG et al., 1984: 342-354. Simulium (Hemicnetha) rubrithorax: COSCARÓN, 1987: 36; PY-DANIEL 1989a: 255; CROSSKEY & HOWARD, 1997: 38; 2004: 31. Simulium magnum LANE & PORTO, 1940: 192 (major, incorrect spelling). Eusimulium rubrithorax LANE & VULCANO, 1943: 436. Hemicnetha rubrithorax: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118; STRIEDER, 2004: 130-131.

Female: Wing length 3.0-4.0 mm. Scutum brown-orange to reddish brown with posterior 1/3 dark grayish, anteriorly with 1+1 submedian silver stripes light grayish, wider anteriorly, and lateral rim grayish silvery (Fig. 113 A); with changing light position, submedian light vittae extend posteriorly in lyre shape as in S. paynei but slightly darkened. Scutum pilosity silvery to tin colored, not homogeneously distributed. Scutellum and metanotum brown-grayish, the latter velvety. Pleurae dark brown with grayish pruinosity. Legs yellowish with apical portion of segments darker. Abdomen yellowish brown with dark spots on tergal plates I-V; tergites VI-VIII shinny. Cibarium with central area smooth without spiculae, cornuae well sclerotized (Fig. 113 B). Ratio of palpomere length/palp sensory vesicle = 1.9. Basal portion of R without hair. Eighth sternite with about 10 setae on each side; gonapophysis about 2 times longer than wide at base, blunt apically; paraproct wide, subquadrate, with abundant microtrichia (Fig. 113 C); genital fork with 1+1 robust subcylindrical posterior sublateral process. Male: Wing length 3.0-3.5 mm. Scutum yellowish brown to blackish, with anterior 1/3 silvery gray, with gray pruinosity edge laterally and posteriorly; with different illumination, submedian light vittae show lyre shape. Metanotum as in female. Abdomen with tergal plates velvety brown-blackish; silvery pruinose on anterior edge and 1+1 big lateral spots on tergite II and 1+1 lateral spots on tergites VI-VII. Ratio of length/width of hind basitarsus = 4.1. Gonostylus subcylindrical, slightly acuminate distally and with weak apical spur (Fig. 114 A); ratio of length of gonostylus/gonocoxite = 1.6; ratio of length/width of gonostylus = 2.4-2.9. Ventral plate with robust median process with parallel sides (Fig. 114 B); median process subrectangular; endoparameres with abundant and strong hooks; aedeagal membrane with abundant spiculae. Pupa: Cocoon shoe shaped with collar of festoons joined apically, protecting gills and sometimes larval exuviae (Fig. 115 A), length basally 3.5-4.0 mm, dorsally about 5.0 mm; gills 2.5-3.1 mm. Frontoclypeus and thorax with small tubercles, only present on facial area of frontoclypeus (Fig. 115 O) and on posterior edge of thorax; trichomes small

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D

K

G E I

M B F N

C

L

H

J

O

Figure 114. Male. A-B: S. (Hemicnetha) rubrithorax; A: genitalia ventral view showing gonocoxite, gonostylus, ventral plate and endoparameres; B: ventral plate. C: S. (H.) virgatum, ventral plate. D-F: S. (H.) brachycladum; D: scutum; E: gonocoxite and gonostylus ventral view; F: ventral plate. G-H: S. (H.) cristalinum; G: gonostylus dorsal view; H: ventral plate. I-K: S. (H.) seriatum; I: gonocoxite and gonostylus dorsal view; J: ventral plate; K: endoparameres and median sclerite. L: S. (H.) pulverulentum, ventral plate. M: S. (H.) smarti, gonocoxite and gonostylus dorsal view. N-O: S. (H.) earlei; N: gonocoxite and gonostylus ventral view; O: ventral plate.

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and single. Dorsum of thorax covered by thick gelatinous material only separated on median line (Fig. 115 L) (material is destroyed by potash). Gill with 8 filamentous branches, smooth with blunt apex and bifurcated close to base (Fig. 116 A). Larva: Maximum length 11.0-12.5 mm. Aspect as in figure 117 A; color dark greenish gray. Head dark brown, except anterolateral portion of frontoclypeus; cephalic apotome with small dark spots on midline and mediolaterally (Fig. 117 D). Antennae about as long as cephalic fan stem; shape as in figure 117 H; ratio of proximal, medial and distal articles = 1:1.2-1.7:1-1.1. Palps well sclerotized. Mandibular teeth as in figure 117 K, with big marginal tooth generally accompanied by another small tooth. Hypostomium with curved edge and median tooth higher than lateral teeth (Fig. 117 N); lateral setae 812; postgenal cleft deep subpentagonal, medially acuminate; postgenal bridge slightly shorter than hypostomium length. Cephalic fan with about 70 rays. Lateral sclerite of thoracic proleg with about 110 teeth arranged in groups in 1 or 2 rows. Anal sclerite well sclerotized with hair, but not scales. Anal ring with about 320 rows of 40-60 hooks each. Rectal papillae with about 10 diverticula on each lobe. Distribution: Argentina: Misiones; Bolivia: Cochabamba; Brazil: Bahia, Goiás, Minas Gerais, Rio Grande do Sul, Rio de Janeiro, Roraima, São Paulo; Guyana; Venezuela: Aragua, Merida, Miranda Yaracuy. Bionomics: This species breeds in very fast streams that are clear to slightly muddy. The larvae and pupae are attached to rocks and stones in large groups covering a great portion of the surface. Females bite horses ( LANE & PORTO, 1940; as magnum). Discussion: Simulium paynei is the closest species, but S. paynei has a darker male thorax, a little narrower gonostylus, and the gelatinous material of the pupal thorax has lateral entrances appearing as 4 branches, which is very easy to see in material maintained in alcohol. Simulium biuxinisa is differentiated by the presence in the pupa of abundant tubercles on the exposed portion of the thorax, the gill totally protected by the cocoon, and the gelatinous material on the thorax leaving free the median area; in the female by the larger sensory vesicle of the palp; and in the larva by the postgenal bridge as long as the hypostomium height. Simulium bricenoi is close but can be separated by the wrinkled surface of the thoracic pupal dorsum. Simulium virgatum can be differentiated by the male gonostylus with a narrower median process. The scarce differences with S. paynei, S. biuxinisa and S. virgatum suggest the need to do cytological and molecular studies in order to resolve species identities. Simulium (Hemicnetha) biuxinisa COSCARÓN & IBÁÑEZ-BERNAL (Figs. 113 E, F; 115 B, N, Q; 116 C, D; 117 O.) Simulium (Hemicnetha) biuxinisa COSCARÓN & IBÁÑEZ-BERNAL, 1993: 64; CROSSKEY & HOWARD, 1997: 38; 2004: 31.

Female (pharate): General coloration brown grayish similar to Simulium paynei. Scutum with wide median and 1+1 sublateral reddish brown vittae separated by 1+1 narrow submedian light grayish stripes in lyre shape. Scutellum and metanotum brown grayish, metanotum velvety. Pleurae dark brown. Abdomen brown-yellow grayish. Legs yellowish with apex of femora, tibiae and tarsomere blackish. Fronto-ocular triangle deep; frons with abundant long hair. Sensory vesicle of maxillary palp surpassing half length of

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Figure 115. A-K: Pupa in lateral view; A: S. (Hemicnetha) rubrithorax; B: S. (H.) biuxinisa; C: S. (H.) virgatum; D: S. (H.) bricenoi; E: S. (H.) brachycladum; F: S. (H.) cristalinum; G: S. (H.) pulverulentum; H: S. (H.) mexicanum; I: S. (H.) seriatum; J: S. (H.) smarti; K: S. (H.) earlei. L-N: Pupa without cocoon showing gelatinous mass over scutum; L: S. (H.) rubrithorax; M: S. (H.) paynei; N: S. (H.) biuxinisa. OU: Frontoclypeus; O: S. (H.) rubrithorax; P: S. (H.) paynei; Q: S. (H.) biuxinisa; R: S. (H.) brachycladum; S: S. (H.) cristalinum; T: S. (H.) seriatum; U: S. (H.) smarti.

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palpomere (Fig. 113 E). Sc with 7-9 hairs. Cibarium and gonapophysis similar to those of S. rubrithorax. Paraproct slightly wider than length, wider than in S. rubrithorax (Fig. 113 F). Male: Unknown. Pupa: Cocoon with collar of 10-12 festoons not joined apically and covering gills (Fig. 115 B). Length basally 4.5-4.8 mm dorsally 6.2-6.3 mm, gills 2.2-2.5mm. Frontoclypeus with abundant rounded platelets (Fig. 115 Q). Exposed portion of thorax with abundant tubercles, wrinkled on dorsum, acuminate in those laterally and posteriorly positioned (Fig. 116 D); gelatinous material with 4 branches (Fig. 115 N). Gill with 8 branches (Fig. 116 C). Larva: Maximum length 11.4-12.6 mm. Cephalic apotome dark brown, mostly uniform in coloration, with small negative spots midlongitudinally and 1+1 median sublaterally and subbasally. Postgenal bridge about same length as hypostomium (Fig.117 O). Antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal articles 1:1.5-1.6:1-1.1. Mandible with 1 well-developed marginal tooth sometimes with additional very small teeth. Cephalic fan with 56-59 rays. Lateral sclerite of thoracic proleg with 52-80 teeth arranged in 22-26 groups. Anal ring with 350-362 rows, with 39-42 hooks each. Anal sclerite with hair, but not scales. Rectal papillae with 15-17 diverticula on each lobe. No hair on body integument. Distribution: Mexico: Oaxaca. Bionomics: This species breeds in torrential creeks with fast, crystalline and relatively cold water, attached to rocks, in mountainous areas with mesophilic vegetation. Female feeding habits are unknown. Discussion: The species is very close to S. paynei and S. rubrithorax. Differences were treated under those species. Simulium bricenoi is similar, but can be separated by the wrinkled dorsal portion of the thorax. Simulium (Hemicnetha) virgatum COQUILLETT (Figs. 114 C; 115 C; 116 B.) Simulium virgatum COQUILLETT, 1902: 97. Simulium (Dyarella) virgatum: VARGAS et al., 1946: 111; DALMAT, 1955: 194-199. Simulium (Hemicnetha) virgatum: VARGAS & DÍAZ NÁJERA, 1957b: 154, COSCARÓN, 1987: 36; PETERSON et al., 1988: 84; PETERSON & KONDRATIEFF, 1995: 34-35; CROSSKEY & HOWARD, 1997: 38; 2004: 31; IBÁÑEZ-BERNAL, 1992: 228-244; ADLER et al., 2004: 374; SHELLEY et al., 2002: 148-153. Simulium cinereum BELLARDI, 1859: 13. Simulium tephrodes SPEISER, 1904: 148. Simulium hippovorum MALLOCH, 1914: 28-29. Simulium rubicundulum KNAB, 1915b: 178-179. Simulium virgatum chiapanense HOFFMAN, 1930b: 293.

Female: Wing length 3-3.7. General coloration brownish. Scutum brown-grayish with recumbent golden to blackish pilosity. Ornamentation similar to that of S. rubrithorax. Changing light direction shows 1+1 submedian vittae wider anteriorly with lyre shape, dark brown to silvery, edged by median and lateral areas gray pruinose to brownish. Scutellum and metanotum dark brown to black with grayish pruinosity. Pleura gray to blackish with grey pruinosity. Legs light yellowish grey with dark brown areas on apex of tibiae, femora and tarsomeres. Abdomen brownish with tergites I-V velvety black; VI-IX shiny black.

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Figure 116. A-M: Pupal gills and portion of thorax dorsally; A: S. (Hemicnetha) rubrithorax; B: S. (H.) virgatum; C-D: S. (H.) biuxinisa; E: S. (H.) bricenoi; F: S. (H.) brachycladum; G: S. (H.) cristalinum; H: S. (H.) mexicanum; I-J: S. (H.) seriatum, K: S. (H. ) smarti; L: S. (H.) hieroglyphicum (from PETERSON et al., 1988); M: S. (H.) earlei.

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Genitalia as in S. rubrithorax but gonapophysis more rounded apically. Cibarium smooth with small spiculae centrally. Basal sector of R bare; Sc with 10-16 hairs. Sensory vesicle of maxillary palp surpassing half of palpomere length; ratio of palpomere/vesicle length = 1.8. Male: Wing length 2.5-3.5mm. General coloration brownish black. Scutum yellowish brown with ornamentation as in Simulium rubrithorax, changing coloration in accordance with light position. Legs as in female. Abdominal segments I-IX velvety brown with 1+1 silvery pruinose spots on tergites II-VI. Genitalia as in S. rubrithorax but ventral plate with median process thinner (Fig. 114 C). Pupa: Cocoon with collar, festoons joined on superior edge and covering most of gills (Fig. 115 C). Basal length 4.5-6.0 mm, dorsally 4.0-5.0 mm, height considering festoons 2.4 mm, gills 1.9-2.3 mm. Frontoclypeus with abundant rounded to pointed tubercles. Cephalic and thoracic trichomes single and weakly developed. Gill with 8 branches, showing well-defined basal trunk and very short primary branches (Fig. 116 B); branches with minute striations on surface and rounded apex. Larva (based on IBÁÑEZ-BERNAL, 1992): Maximum length about 9.0 mm. Cephalic apotome ornamentation similar to that of rubrithorax but with median longitudinal and sublateral median and basal spots negatives. Ratio of proximal, medial and distal antennal articles = 1:1.2:0.9; cephalic fan with about 45 rays. Hypostomium and postgenal cleft as in S. rubrithorax. Anal ring with 220-255 rows of hooks. Rectal papillae with 7-8 diverticula on each lobe. Distribution: Mexico: Colima, Chiapas, Chihuahua, Durango, Michoacán, Morelos, Oaxaca, Tabasco, Veracruz; USA: Arizona, Colorado, New Mexico, South Dakota, Texas, Utah, Wyoming. Bionomics: This species breeds in streams at 900-2500 m, on rocks and aquatic vegetation. Females bite livestock and domestic animals. Discussion: Simulium virgatum is very similar to other species of the Simulium PAYNEI species group. Differences are few and often can be considered population variations. SHELLEY et al. (2002) gave a detailed explanation of the synonymical history and the possibilities of four different cytotypes that MUHAMMAD (1988) described, but according to PETERSON & KONDRATIEFF (1995), might correspond to different species because of incorrect determination of the material. The differences are not consistent enough to justify that the Neotropical material of S. virgatum is conspecific with S. paynei and S. rubrithorax (SHELLEY et al., 2002). ADLER et al. (2004) recognized two species in North American material formerly considered S. virgatum: S. hippovorum and S. virgatum sensu stricto. A cytological and molecular study based on morphologically well-identified material of the species in the S. PAYNEI species group is needed. Simulium (Hemicnetha) bricenoi VARGAS , M ARTÍNEZ P ALACIOS & D ÍAZ N ÁJERA (Figs. 115 D; 116 E.) Simulium (Dyarella) bricenoi VARGAS et al., 1946: 115-118, 177. Simulium (Hemicnetha) bricenoi: VARGAS & DÍAZ NÁJERA, 1957b: 174, 175, 177-179, 310; COSCARÓN, 1987: 36; IBÁÑEZ-BERNAL, 1992: 65-81; ADLER et al., 2004: 371; CROSSKEY & HOWARD, 1997: 38; 2004: 31.

Female: Wing length 4.0 mm. Frons clypeus dark brown with grayish pollinosity. Scutum brown-grayish with hair light golden; in frontal illumination, with 1 median and 1+1 sublateral

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grayish vittae, margined by dark brown 1+1 submedian lyre-shaped and thin lateral stripes; with changing light position, grayish vittae become blackish, and thin marginal stripes become light gray and wider anteriorly. Scutellum light brown. Legs yellowish with dark brown on apex of tibiae, femora and tarsomeres. Abdomen brownish with tergites III-V lighter, VI-VIII shiny. Cibarium, claws and genitalia as in S. rubrithorax, with gonapophysis slightly narrower, and ratio of length/width 2.2. Sensory vesicle of maxillary palp nearly half of palpomere length, ratio of length of palpomere/sensory vesicle = 1.8. Sc without hair. Male: Wing length about 3.7 mm. Scutum blackish with ornamentation as in Simulium paynei. Abdomen dark brown, II-VIII with 1+1 lateral pruinose silvery spots. Ratio of length/width of hind basitarsus = 3.8-4.0. Genitalia as in S. rubrithorax but gonostylus more pronounced with outer subapical curvature and narrower median process of basal plate. Pupa: Cocoon with collar of festoons joined distally (Fig. 115 D); length basally 4.5 mm, height 3.0 mm, gills 1.8 mm. Frontoclypeus as in S. rubrithorax, with platelets only on basal area. Exposed portion of thorax smooth, but with rugosity as shown in figure 116 E, and small tubercles laterally and posteriorly. Cephalic and thoracic trichomes single and diminutive. Gill with 8 branches arising from short basal trunk as in S. rubrithorax. Larva: Maximum length 9.1-10.7 mm. Cephalic apotome with positive spots arranged discontinuously on midlongitudinal line, 1+1 sublateral and 1+1 latero-basal spots. Ratio of proximal, medial and distal antennal articles = 1:1.8:1-1.3. Cephalic fan with 45-58 rays. Hypostomium and postgenal cleft as in S. rubrithorax. Anal ring with 260-270 rows with 5560 hooks on each row. Rectal papillae with 10-18 diverticula on each lobe. Distribution: Mexico: Distrito Federal, Durango, Mexico, Morelos, Veracruz; USA: Arizona, New Mexico. Bionomics: The species breeds in clear and cold water of mountainous creeks 1-3m wide, at altitudes over 1700 m, with sandy and rocky bottoms. Female feeding habits are unknown. Discussion: Simulium bricenoi is very close to S. rubrithorax, S. paynei and S. biuxinisa in the adult ornamentation and general aspects of the pupa, but S. bricenoi has unique rugosity on the dorsal portion of the thorax. Simulium hieroglyphicum has similar rugosity on the thorax, but has 60-90 gill branches. BRACHYCLADUM species group Simulium brachycladum LUTZ & PINTO (Figs. 113 G-L; 114 D-F; 115 E, R; 116 F; 117 F, J, P.) Simulium brachycladum LUTZ & PINTO, in PINTO, 1931: 690; MAIA-HERZOG et al., 1984: 341-350, 353-354. Simulium (Hemicnetha) brachycladum COSCARÓN, 1987: 36; 1991: 274-275; CROSSKEY & HOWARD, 1997: 37; 2004: 30. Simulium brevibranchium LUTZ & MACHADO, 1915: 46 (nomen nudum).

Female: Wing length 2.8-3.3 mm. General coloration brownish gray. Scutum brown with gray pollinosity and abundant pile light gray intermixed with blackish hair; in frontal illumination, with 1+1 submedian silver, narrow lyre-shaped vittae (Fig. 113 G) that with posterior light become darker especially anteriorly (Fig. 113 H). Scutellum and metanotum brownish gray. Pleurae dark brown, grayish pollinose. Legs brownish yellow with apical

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portion of segments dark brown. Abdomen brownish yellow with tergal plate of segments III-IX dark brown. Fronto-ocular triangle deep (Fig. 113 I). Sensory vesicle of maxillary palp big, ratio of palpomere/sensory vesicle length = 1.6. Cibarium with abundant small spiculae on median area (Fig. 113 J). Sc with hair on basal 3/4 or more. Eighth sternite well sclerotized with about 25 setae; gonapophysis subtriangular, acute distally, with internal edge well sclerotized, with abundant hair and sparse microtrichia (Fig. 113 K); ratio of length/ width at base = 1.8. Paraprocts membranous and rounded apically (Fig. 113 L). Genital fork well sclerotized and similar to that of S. rubrithorax in shape. Male: Wing length 2.6-3.1. Scutum darker brown-grayish with pile tin colored. Ornamentation shape as in figure 114 D. Abdomen dark brown with 1+1 big silvery spots on tergite II, only on anterior margin in tergites III-V and most developed laterally on VI-VII. Ratio of length/ width of hind basitarsus = 6. Gonostylus relatively wide with strong protuberances laterally on median portion (Fig. 114 E); ratio of length/wide = 1.5; ratio of gonostylus/gonocoxite length = 1.7. Ventral plate with median process slightly wider distally (Fig. 114 F). Pupa: Cocoon shoe shaped, leaving free gills or protecting only gill base (Fig. 115 E). Length basally 3.0-4.0 mm, dorsally 2.7-3.8 mm; maximum length 3.8-4.5 mm; gills 1.5 mm. Frontoclypeus and exposed portion of thorax smooth with only some on facial area and sparse acute tubercles on posterior area of thorax (Fig. 115 R). Cephalic and thoracic trichomes short, single or bifid. Gill with 6 branches short and acute distally, arising from short basal trunk (Fig. 116 F). Larva: Maximum length 7.5-8.5 mm. Cephalic apotome darkened on basal 2/3 and with negative spots (Fig. 117 F). Hypostomium with straight edge with lateral teeth at same level as median teeth; postgenal cleft medially developed with postgenal bridge longer than hypostomium length, postgenal cleft subpentagonal (Fig. 117 P); ratio of postgenal bridge/hypostomium length = 1.2. Cephalic fan with about 45 rays. Antenna with relatively thick articles (Fig. 117 J); ratio of proximal, media; and distal articles = 1:1.5-1.6:0.9-1. Mandible with 1 thin marginal tooth. Anal ring with 190 rows with 31-35 hooks each. Rectal papillae with 5 diverticula on each lobe. Distribution: Brazil: Bahia, Minas Gerais, Pernambuco, São Paulo. Bionomics: This species breeds in torrential creeks with a range of low to high discharge, rocky bottoms and clear water, attached to stones and consolidated sediment. Female feeding habits are unknown. Discussion: A similar species is S. cristalinum whose differences are treated under that species. Simulium pulverulentum is also very close but can be differentiated by the pupal gill with 10 branches. Simulium (Hemicnetha) cristalinum COSCARÓN & PY-DANIEL (Figs. 113 N; 114 G, H; 115 F, S; 116 G; 117 B, G, L, Q, T, V.) Simulium (Hemicnetha) cristalinum COSCARÓN & PY-DANIEL, 1989: 313-318; CROSSKEY & HOWARD, 1997: 37; 2004: 30. Hemicnetha cristalina PY-DANIEL & SAMPAIO, 1995: 118.

Female (pharate): General coloration brownish. Scutum dark brown with gray pollinosity and thin and thicker hair intermixed (Fig. 113 N) and arranged in groups.

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Figure 117. A-V: Larva. A-C: general aspect in lateral view; A: S. (Hemicnetha) rubrithorax; B: S. (H.) cristalinum; C: S. (H.) seriatum. D: S. (H.) rubrithorax, head dorsal view. E-G: cephalic apotome; E: S. (H.) seriatum; F: S. (H.) brachycladum; G: S. (H.) cristalinum. H-J: antenna; H: S. (H.) rubrithorax; I: S. (H.) seriatum; J: S. (H.) brachycladum. K-M: mandibular teeth; K: S. (H.) rubrithorax; L: S. (H.) cristalinum; M: S. (H.) seriatum. N-R: postgenal cleft, hypostomium and frontal portion of hypostomium; N: S. (H.) rubrithorax; O: S. (H.) biuxinisa; P: S. (H.) brachycladum; Q: S. (H.) cristalinum; R: S. (H.) seriatum; S: S. (H.) seriatum, lateral sclerite of thoracic proleg; T: S. (H.) cristalinum, anal sclerite. U-V: rectal papillae; U: S. (H.) seriatum; V: S. (H.) cristalinum.

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Scutum in anterior illumination with 1+1 anterior submedian subtriangular silvery pollinose spots, continued posteriorly as vittae with lyre shape; with changing light position, vittae get thinner and darker, and lateral edges become silvery. Scutellum light brown, metanotum brownish gray. Abdomen dark brown. Cibarium with small teeth on central area. Sensory vesicle of maxillary palp shorter than half of palpomere length; ratio of palpomere/sensory vesicle length = 1.8. Eighth sternite well sclerotized medially with about 24-28 setae on each side. Gonapophysis subtriangular, acute distally, shorter than in S. brachycladum. Paraproct membranous, rounded apically, with sparse setae, no microtrichia. Male (pharate): Coloration as in female. Ratio of length/width of posterior basitarsus about 7.2. Gonostylus relatively short with moderate mediolateral inner and outer protuberances (Fig. 114 G), ratio of length/width = 1.8. Ratio of length gonostylus/gonocoxite = 2.1. Ventral plate with short and relatively thick median process (Fig. 114 H). Pupa: Cocoon shoe shaped, aperture lower anteriorly and covering most of gill (Fig. 115 F). Length basally 3.5-3.8 mm, dorsally 2.0-2.3 mm; gills 0.7-0.8 mm. Frontoclypeus and exposed portion of thorax smooth, with small tubercles on facial and posterior area of thorax; frontoclypeus darkened by strong sclerotization (Fig. 115 S); cephalic and thoracic trichomes simple and small. Gill with 6 branches very short, with transverse striations and slightly thin but not acute apices (Fig. 116 G). Larva: Maximum length 5.8-6.3 mm. Body shape and coloration as in figure 117 B. Cephalic apotome with posterior 2/3 darkened with median longitudinal, sublateral median and sublateral posterior negative spots (Fig. 117 G). Antenna surpassing length of cephalic fan stem, with ratio of proximal, medial and distal articles = 1:1.61.7:1. Cephalic fan with 36-40 rays. Mandibular teeth as in figure 117 L. Hypostomium frontal edge straight with median tooth longer than lateral teeth. Postgenal cleft subtriangular, with postgenal bridge as long as hypostomium length (Fig. 117 Q). Lateral plate of thoracic proleg sclerite with about 72 teeth. Anal sclerite with hair and scales (Fig. 117 T). Anal ring with 156-170 rows with 25-29 hooks each. Rectal papillae with 5-6 diverticula on each lobe (Fig. 117 V). Distribution: Brazil: Roraima. Bionomics: This species was collected in a medium-size, torrential creek, with rocky bottom and clear, temperate water. Discussion: The closest species is S. brachycladum. It can be differentiated by several small characters: in the female by the smaller teeth on the median area of the cibarium and sensory vesicle of the maxillary palp longer than half of the palpomere, in the male by the shorter gonostylus with more developed lateral protuberances and the median process of the ventral plate being thinner and more elongated, in the pupa by the larger size and higher cocoon aperture and gill branches about two times longer, and in the larva by the greater number of hook rows and the hypostomium lateral teeth longer than the median tooth. It would be useful to have more material, and to do karyological and molecular studies to evaluate the status of these species.

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Simulium (Hemicnetha) pulverulentum KNAB (Figs. 113 M; 114 L; 115 G.) Simulium pulverulentum KNAB, 1914b: 177-178; FAIRCHILD, 1940: 716-717; VARGAS, 1943b: 363; DALMAT, 1955: 190-194. Simulium (Hemicnetha) pulverulentum VARGAS & DÍAZ NÁJERA, 1957b: 154, 174, 176-178, 180, 263, 324; RAMÍREZ PÉREZ, 1983: 64-65; COSCARÓN, 1987: 37; IBÁÑEZ-BERNAL, 1992: 178-195; CROSSKEY & HOWARD, 1997: 37; 2004: 30; SHELLEY et al., 2002: 146-148.

Female: Wing length 2.3-2.7. Scutum dark brown-grayish to blackish with gray pollinosity and abundant gray groups of recumbent hair. Ornamentation similar to that of S. brachycladum and changeable with light position. Scutellum dark brown and metanotum blackish. Pleurae blackish with gray pollinosity. Legs brownish yellow with darkened areas of segments dark brown. Abdomen dark brown with tergites III-V velvety and VI-IX shiny; tergite II with 1+1 grayish spots. Cibarium with small teeth on median area. Ratio of palpomere/sensory vesicle length = 2.1. Sc with sparse hair on basal half. Gonapophysis subtriangular, acute apically, as in S. brachycladum but basally wider. Paraprocts subquadrangular, membranous, with sparse hair and microtrichia (Fig. 113 M). Male: Wing length 2.0-2.5 mm. Scutum dark brown to blackish with light gray pollinosity and golden hair, showing changeable ornamentation with light position, but very similar to S. brachycladum. Scutellum, metanotum and pleurae as in female. Abdomen dark brown to blackish with silver pruinosity on 1+1 large areas of tergites II and VI, and small areas on VII-VIII. Ratio of length/width of posterior basitarsus = 5.7. Gonostylus with well-developed lateral inner and outer protuberances, similar to that of S. brachycladum. Ratio of length/width of gonostylus = 1.8; ratio of length of gonostylus/ gonocoxite = 2. Ventral plate with elongate median process (Fig. 114 L). Pupa: Cocoon shoe shaped, relatively low anteriorly leaving gills free (Fig. 115 G). Length basally 2.5-3.5 mm, maximum length 4.1 mm; on dorsum 2-3mm. Frontoclypeus and exposed portion of thorax smooth, except facial area of frontoclypeus and posterior area of thorax with small tubercles. Cephalic and thoracic trichomes short and single. Gill with 10 branches relatively short; filaments with superficial annulations, with spiculae and pointed apices. Larva: Maximum length 4.9-5.5 mm. Cephalic apotome with negative spots. Antenna surpassing apex of cephalic fan; ratio of proximal, medial and distal antennal articles = 1:1.1:0.6. Cephalic fan with about 44 rays. Hypostomium with straight edge and median tooth longer than lateral teeth, postgenal cleft arc shaped. Postgenal bridge about same length as hypostomium, anal ring with about 140 rows of 25-28 hooks each. Rectal papillae with 6-10 diverticula on each lobe. Distribution: Belize; Costa Rica; El Salvador; Guatemala; Honduras; Mexico: Chiapas, Guerrero, Hidalgo, Michoacan, Morelos, Oaxaca, San Luis Potosi, Veracruz; Panama; Venezuela. Bionomics: This species breeds in creeks and rivers with fast currents, rocky bottoms, and clear or moderately muddy, warm water (16-21 °C). Larvae and pupae are found on aquatic vegetation. Females bite cows, mules and horses. Discussion: The most similar species are S. brachycladum, S. cristalinum and S. guerrerense. The first two species can be separated by their pupal gill with 6 branches.

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Simulium guerrerense has 10 branches and the apex of the postgenal cleft is arc shaped, but it has a pupal cocoon with an elevated aperture covering the branches, a male ventral plate with a shorter median process and a larva with positive spots on the cephalic apotome and lateral teeth of the hypostomium longer than the median tooth. These small differences can be intraspecific variations (SHELLEY et al., 2002). Simulium (Hemicnetha) lobatoi LUNA DIAS, HERNÁNDEZ, MAIA-HERZOG & SHELLEY Simulium lobatoi LUNA DIAS et al., 2004: 37-42.

(Based on original description). Female: Wing length 2.4-2.9 mm. Scutum dark brown with recumbent whitish hair interspersed with brown hair; in anterior illumination, with 1+1 submedian whitish vittae as in S. rubrithorax but only on 2/3 of scutum length; humeri brown, silvery pruinose; in posterior light, with median silver pruinose vittae and median dark brown line and 1+1 rounded anterior pruinose spots, as commonly seen in S. rubrithorax and S. brachycladum. Scutellum and metanotum dark brown, metanotum silvery pruinose. Pleurae brown, silvery pruinose. Basal section of R and Sc haired. Legs brown with apex of femur, tibiae and tarsomeres dark brown. Abdomen dark brown with 1+1 lateral silver pruinose spots. Cibarium smooth. Claws with subbasal tooth. Gonapophysis subtriangular, similar to that of S. smarti but slightly narrower apically; paraproct subquadrangular, weakly sclerotized on margin, membranous apically and with prominent hair. Male: Wing length 2.0-2.7 mm. Scutum orange on anterior 1/3 and darker posteriorly, with yellowish pile and semi-erect brown hair. With anterior illumination, vittae very similar to those of S. brachycladum; with posterior light, vittae dark brown. Sc without hair. Abdomen dark brown to black, with pruinosity on anterolateral margins of tergite II and 1+1 laterally on tergites III-VIII. Gonostylus similar to that of S. cristalinum. Ratio of gonostylus/ gonocoxite = 1.7; ratio of gonostylus length/width = 1.8. Ventral plate with median process moderately developed, similar to that of S. cristalinum, but more acuminate apically. Pupa: Cocoon with aperture anteriorly elevated at same height as posteriorly, very similar to that of S. seriatum, protecting branches. Length basally 3.9-6.0 mm, dorsally 2.53.6 mm; gills 0.9-1.8 mm. Frontoclypeus with tubercles on facial region. Gill with 8 branches arising from short trunk, branches stout, pointed apically as in S. brachycladum, edges weakly crenate and without spicules. Larva: Unknown. Distribution: Brazil: Goiás, Mato Grosso. Bionomics: The species breeds in fast currents of small to medium (0.5-10 m) clearwater streams, attached to rocks and dead leaves. Females bite humans. Discussion: Females of S. rubrithorax and S. brachycladum have similar scutal coloration, but in these species the 1+1 submedian vittae join posteriorly, whereas in S. lobatoi, they diverge posteriorly. In those species, the gonapophyses and paraprocts are longer, and the gonapophyses are more acute apically in S. brachycladum. The median process of the male ventral plate is also more elongated; a similar species in this character is S. cristalinum. The pupa gill of S. brachycladum has 6 branches. Simulium rubrithorax has a similar number of branches that are longer and not pointed apically, and the cocoon has conspicuous festoons.

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Simulium (Hemicnetha) guerrerense VARGAS & DÍAZ NÁJERA Simulium (Hemicnetha) guerrerense VARGAS & DÍAZ NÁJERA, 1956: 51-56; 1957b: 153, 174, 176, 177, 179, 180, 262, 316; COSCARÓN, 1987: 35-36; IBÁÑEZ-BERNAL, 1992: 111-127; CROSSKEY & HOWARD, 1997: 38; 2004: 30.

Female: Wing length 2.5-2.7mm. Scutum brownish, with golden pile; with frontal illumination, 1+1 submedian light gray pollinose stripes wider frontally and curved posteriorly in shape of lyre, anterior portion connected laterally with thin marginal concolorous stripes; thin blackish stripe along midline; with posterior illumination, 1 median and 1+1 sublateral wide grayish vittae emarginated by 1+1 submedian and 1+1 lateral thin dark grayish pollinose stripes. Scutum yellowish, metanotum brown. Legs yellowish with most segments apically dark brown. Abdomen brown with tergite I light brown, tergite II and 1+1 on V with whitish pruinosity, shiny. Sc with hair on basal 1/3. Cibarium with small spiculae on median area; sensory vesicle of maxillary palp surpassing half of palpomere length. Paraproct membranous, and with scarce microtrichia as in S. pulverulentum. Male: Wing length 2.1-2.4 mm. Scutum dark brown to black with vittae as in female. Scutellum, metanotum, legs and abdomen as in female. Ratio of length/width of hind basitarsus = 4.2. Ratio of gonostylus/gonocoxite = 1.4; ratio of gonostylus length/width = 1.5. Gonostylus shape as in S. cristalinum; ventral plate similar to that of S. cristalinum but wider distally. Pupa: Cocoon shoe shaped, very high anteriorly covering gill. Length basally: 2.3-2.7 mm; anterior height 1.5-1.9 mm, gill length 0.8-1.0. Frontoclypeus and exposed portion of thorax smooth, with small tubercles on facial area and posterior portion of thorax; trichomes single and small. Gill with 10 branches emerging from short trunk. Larva: Maximum length 5.5-6.0 mm. Cephalic apotome with positive spots. Antenna as long as cephalic stem; ratio of proximal, medial and distal segments = 1:1.3:0.8. Postgenal cleft rounded apically; postgenal bridge slightly longer than hypostomium length. Cephalic fan with about 47 rays. Anal ring with about 160 rows with 22 hooks each. Rectal papillae with 6 diverticula on each lobe. Distribution: Mexico: Guerrero. Bionomics: Unknown. Discussion: Simulium pulverulentum has similar adult coloration and pupal gill, but it can be separated by the elongate median process of the ventral plate and short anterior portion of the cocoon that leaves free the gill branches. Studies with more material and using cytological and molecular techniques would be useful in order to assure the identity of S. guerrerense. Simulium (Hemicnetha) solarii STONE Simulium (Dyarella) solarii STONE, 1948: 402-404. Simulium (Hemicnetha) solarii: VARGAS & DÍAZ NÁJERA, 1957b: 175, 176, 178, 180, 263, 328; COSCARÓN, 1987: 37; IBÁÑEZ-BERNAL, 1992: 212-228; ADLER et al., 2004: 373-374; CROSSKEY & HOWARD; 1997: 37; 2004: 30.

(Based on IBÁÑEZ-BERNAL, 1992). Female: Wing length 2.5-2.8 mm. Scutum brown-orange with grayish yellow pollinosity, 1+1 submedian and sublateral longitudinal areas delimited by 1 dark and thin

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stripe, 1+1 submedian wider anteriorly, and 1+1 laterally; with anterior illumination, submedian darker vittae conspicuous as lyre shape. Scutellum and metanotum with coloration similar to scutum. Legs yellowish with apex of femora, tibiae and tarsomeres dark brown. Abdomen brown with gray pollinosity on posterior margin of segments IIIVI and velvety brown on tergal plates; VII-IX shiny. Cibarium with lateral margins of cornuae and median area of pharyngeal membrane with spiculae. Sc with hair on basal half. Gonapophysis acute distally; relatively short and wide medially. Male: Wing length 2.3-2.5 mm. Scutum dark brown with hair light golden and grayish pollinosity, 1+1 submedian and 1+1 sublateral areas delimited by 1+1 submedian and 1+1 lateral thin and dark stripes. Scutellum and metanotum similar to scutum; scutellum shiny. Legs as in female. Abdomen dark brown with silvery pruinose spots on tergites, medially on tergite I, and 1+1 laterally on tergites II, V-VII. Ratio of length/ width of hind basitarsus = 4.8. Gonostylus very similar to that of S. brachycladum and S. cristalinum. Ratio of length/width = 1.9-2.1; ratio of length gonostylus/gonocoxite = 2.02.2. Ventral plate short with median elongated process as in S. brachycladum. Pupa: Cocoon shoe shaped with anterior edge moderately lower leaving pupal gills free; weave relatively thin, translucent. Length basally 3.0-3.5 mm, height of anterior edge 1.0-1.7 mm. Frontoclypeus and exposed portion of thorax smooth, with tubercles on facial area of frontoclypeus, and posterior area of thorax; cephalic and thoracic trichomes single. Gill with 15 filamentous branches bearing transverse striations with spiculae, and acute apex. Larva: Maximum length 8.2-8.9 mm. Cephalic apotome with positive and negative spots. Ratio of antennal articles proximal, medial and distal = 1:1.2:0.8. Ratio of hypostomium/postgenal bridge = 1.4. Cephalic fan with about 62 rays. Anal ring with about 164 rows of hooks. Distribution: Mexico: Nueva Leon, San Luis Potosi, Tamaulipas; USA: Texas, New Mexico. Bionomics: Simulium solarii lives in torrential streams and rivers. Female feeding habits are poorly known; they have been collected from horses. Discussion: The female is very close to those of S. guerrerense and S. pulverulentum in genitalia morphology, but in these species the scutum is darker. The male is externally similar to that of S. virgatum but differs in the ventral plate, lacking laterodistal protuberances and having a gonostylus with lower lateral curvature. The greater number of gill branches differs from the condition in the aforementioned species. Some pupae of S. earlei with 15 gill branches can produce confusion, but differences in genitalia, as mentioned for that species, allow separation. MEXICANUM species group Simulium (Hemicnetha) mexicanum BELLARDI (Figs. 115 H; 116 H.) Simulium mexicanum BELLARDI, 1862: 6; VARGAS, 1941a: 119-121; 1942a: 236-237. Simulium (Hemicnetha) mexicanum: VARGAS & DÍAZ NÁJERA, 1957b: 174-177, 179, 236-237; RAMÍREZ PÉREZ, 1983: 57-59, COSCARÓN, 1987: 36; COSCARÓN & COSCARÓN ARIAS, 2000a: 187; SHELLEY

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et al., 2002: 144-145; IBÁÑEZ-BERNAL, 1992: 144-161; CROSSKEYS & HOWARD, 1997: 38; 2004: 30; HERNÁNDEZ & SHELLEY, 2005: 6. Simulium (Dyarella) mexicanum: VARGAS et al., 1946: 164-179; DALMAT, 1955: 185-190. Simulium aureopunctatum MALLOCH, 1914: 27. Simulium placidum KNAB, 1915c: 281. Simulium lugubre LUTZ & NUÑEZ TOVAR, 1928: 46. Eusimulium turgidum HOFFMANN, 1930b: 293, 298-300, 303.

Female: Wing length 3.0-3.4mm. General coloration dark brownish. Head dark brown, frons and clypeus brownish with grayish pollinosity; antennae and legs dark brown; scape and pedicel yellowish brown. Scutum reddish brown to blackish pollinose without change of ornamentation in different light positions, with grouped gray to golden iridescent scales, depending on illumination. Scutellum and metanotum dark brown to black; pleura dark grayish brown, pollinose. Legs light brownish, black on tarsomeres and brown on apices of femora and tibiae. Abdomen blackish; tergite II with 1+1 submedian silvery spots and posterior edge dark brown; III-V velvety, VI-IX shiny and waxy. Frons convergent below, front-ocular triangle deep. Smooth cibarium base and cornuae branches, without spiculae. Sensory vesicle of maxillary palp about 1/3 of palpomere length. Basal sector of R without hair; Sc with hair all along vein. Ratio of length/width of posterior basitarsus = 5.6; claw with strong subbasal tooth. Genitalia: paraproct robust, subrectangular with abundant microtrichia, cercus rounded on outer edge; eighth sternite with median transverse sclerotized spot, gonapophysis about as long as wide, subovoidal with rounded apical edge. Genital fork with robust branches and distal apodemes. Male: Wing length 3.0-3.3 mm. Coloration similar to that of female but darker; scutum blackish brown, pollinose, without special ornamentation and golden pilosity. Abdomen brown with 1+1 silvery, pollinose, sublateral spots on tergite II, IV-VII. Basal sector of R without hair. Ratio of length/width of hind basitarsus = 3.1-3.8. Gonocoxite subtrapezoidal, gonostylus about 2 times gonocoxite length, with slightly rounded lateral projection and thin apical spur; ratio of length/width of gonostylus = 2.3-2.5; endoparameres with strong teeth; basal plate wider than high with median carina and abundant setae. Pupa: Cocoon shoe shaped, projecting anteriorly with anterior edge reinforced (Fig. 115 H); thick silk without visible threads. Length basally 3.5-4.0 mm, dorsally 2.6-3.4 mm, gill branches length 2.0-2.3 mm. Frontoclypeus with few platelets mostly on basal edge and laterally, reinforced basally with some wrinkles; trichomes single with 1+1 facial and 2+2 frontal. Thorax with exposed area homogeneously covered by tubercles (Fig. 116 H). Gill with 12 filamentous branches, emerging from 3 short primary branches that originate from short basal trunk. Filaments closely arranged in bunch (Fig. 116 H). Larva (based on IBÁÑEZ-BERNAL, 1992): Maximum length 6.5-8.6 mm. Coloration light brown grayish, darkened on dorsum and lighter ventrally. Cephalic apotome light brown, slightly darkened at base, and with median longitudinal, 1+1 submedian and subbasal positive spots. Antennae longer than base of cephalic fans; ratio of proximal, medial and distal articles 1:1.1-1.3:0.5-.0.6. Postgenal cleft deep; ratio of hypostomium/ hypostomial bridge = 1:1.3-1.5. Cephalic fan with about 57-63 rays. Anal ring with about 320-340 rows, with 44-47 hooks each. Rectal papillae with 10-14 diverticula on each lobe.

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Distribution: Belize; Bolivia; Colombia; Costa Rica; Ecuador; El Salvador; Guatemala; Honduras; Mexico: Chiapas, Oaxaca, Veracruz; Panama; Trinidad, Tobago; Venezuela. Bionomics: Simulium mexicanum breeds in small streams with warm water (16-21 °C), attached to rocks and aquatic vegetation. It is a zoophilic species (DALMAT, 1955), biting livestock and domestic birds, but not humans (SHELLEY et al., 2002). Discussion: Simulium seriatum is the closest species. Differences are explained under that species. The female of S. mexicanum is similar to that of S. smarti and the male to that of S. smarti, S. freemani and S. bricenoi. Differences in the pupa allow identification. Simulium smarti has 18 gill branches, S. bricenoi has rugosity on the exposed area of the thorax and S. freemani has 8 gill branches. HERNÁNDEZ & SHELLEY (2005) synonymized S. mexicanum and S. seriatum with S. tarsatum MACQUART, 1845, based on the female holotype (not male as published) from Colombia, without locality information. As in Simulium (Hemicnetha), there are several species with females that cannot be differentiated by morphological characters. We prefer to maintain the validity of S. mexicanum and S. seriatum, pending future studies of the other life stages from the same area. Simulium (Hemicnetha) seriatum KNAB (Figs. 113 O-S; 114 I-K; 115 I, T; 116 I, J; 117 C, E, I, M, R, S, U.) Simulium seriatum KNAB, 1914b: 81. Simulium (Hemicnetha) seriatum: COSCARÓN, 1987: 36; COSCARÓN & COSCARÓN ARIAS, 2000a: 182-187; COSCARÓN ARIAS, 2003: 200, 203, 208, 213; CROSSKEY & HOWARD, 1997: 38; 2004: 30. HERNÁNDEZ & SHELLEY, 2005: 6. Trichodagmia laticalx ENDERLEIN, 1934b: 291. Trichodagmia latidigitus ENDERLEIN, 1936: 129.

Because S. seriatum is very close to S. mexicanum, we will include only the differences. The figures presented are in accordance with the description of S. mexicanum. Female and Male: Similar to S. mexicanum; coloration slightly darker, sensory vesicle of maxillary palp longer, with ratio of vesicle/palpomere length = 1.7-3.0, and cibarium with small spicules at base of cornuae (Fig. 113 O). Claw as in figure 113 P; female genitalia as in figures 113 Q-S and male genitalia as in 114 I-K. Pupa: Frontoclypeus without or with very few tubercles (Fig. 115 T) and exposed portion of thorax with tubercles arranged in 2+2 longitudinal stripes, leaving other areas smooth (Figs. 116 I, J); cocoon thinner, allowing pupa to be seen, not reinforced on aperture border, and with small dorsal projection (Fig. 115 I). Gill with filaments relatively open and with length of 1.6-1.8 mm. (Fig. 116 I). Larva: General aspect as in figure 117 C; maximum length surpassing S. mexicanum length (8.7-9.6 mm); cephalic apotome as in figure 117 E; postgenal cleft deep with ratio of length hypostomium/postgenal bridge = 1.3-1.5 (Fig. 117 R); cephalic fan with 55-60 rays; antennae with proximal article relatively short (Fig. 117 I), with ratio of articles 1:1.41.7:0.7-1. Mandibular teeth as in figure 117 M. Lateral sclerite of thoracic proleg as in figure 117 S. Anal ring with 200-320 rows of hooks and fewer diverticula; rectal papillae (9-14 on each lobe) (Fig. 117 U).

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Distribution: Argentina: Jujuy; Bolivia: Beni; Colombia: Valle; Ecuador: Canar, Chimborazo, (Esmeraldas), Pichincha; Peru: Cuzco, Lima; Trinidad; Venezuela: Bolivar, Distrito Federal, Táchira. Bionomics: Immature stages breed in fast, clear streams, attached to stones. Female feeding behavior is unknown. Discussion: As previously explained, S. seriatum and S. mexicanum are very similar and the adults are practically inseparable. The pupae show several differences. The pupa of S. mexicanum has a thicker cocoon, with a reinforced border and lower aperture without an anterior projection, the gills are a little longer and arranged in a tighter bunch, and the frontoclypeus and exposed portion of the thorax have thick tubercles homogeneously disposed without bare areas. The larvae show small differences, such as more hooks in the anal ring. Material studied from Argentina, Ecuador (states of Pichincha, Chimborazo and Canar),Venezuela (Bolivar and Tachira) and Trinidad show the characteristic pupa of S. seriatum and those of Guatemala and Mexico are as typically described for S. mexicanum. We consider these differences justification for maintaining S. seriatum as a valid species. Simulium (Hemicnetha) smarti VARGAS (Figs. 113 T, U; 114 M; 115 J, U; 116 K.) Simulium (Eusimulium) smarti VARGAS, 1946: 332-335; DALMAT, 1955: 199-204. Simulium (Hemicnetha) smarti: VARGAS & DÍAZ NÁJERA, 1957b: 174, 175, 177, 178, 180, 263, 326; COSCARÓN, 1987: 36; IBÁÑEZ-BERNAL, 1992: 195-212; CROSSKEY & HOWARD, 1997: 38; 2004: 30.

Female: Wing length 4.0-4.4 mm. Scutum dark brown with grayish pollinosity, without special ornamentation; hair dark brown. Scutellum brownish, slightly lighter than scutum. Metanotum as for scutum, but with posterior edge light brownish gray. Legs yellowish with apex of femora, tibiae and tarsomeres brownish. Abdomen brownish gray. tergites II-V velvety with tergal plates darkened, VI-VIII shiny. Cibarium basally with spiculae on cornuae edge and on median area. Sensory vesicle of maxillary palp half of palpomere length. Sc with hair along entire length. Gonapophysis relatively short with rounded apex and nearly as wide at base as length (Fig. 113 T), half of distal surface with microtrichia; paraproct relatively short, nearly as wide at base as length, with rounded apex and entire surface with microtrichia (Fig. 113 U). Male: Wing length 3.9-4.2 mm. Scutum dark brown grayish with 1+1 thin submedian and lateral gray pollinose stripes, covered by golden groups of recumbent hair. Scutellum, metanotum and legs as in female. Abdomen dark brownish, tergites VVII with 1+1 lateral silvery pruinosity. Ratio of length/width of hind basitarsus = 3.3. Gonostylus elongated, about 2.5-2.9 longer than wide, with slight outer lateral protuberance (Fig. 114 M). Ratio of length of gonostylus/gonocoxite = 2.3. Ventral plate short, narrow with elevated keel. Pupa: Cocoon shoe shaped with basal projection (Fig. 115 J); length at base 3.8-4.9 mm, height of anterior portion 2.7-3.2 mm. Gills 2.6 mm, mostly covered by cocoon. Frontoclypeus and exposed portion of thorax with moderate number of tubercles, small and rounded (Fig. 115 U); trichomes short and single. Gills with 18 branches, branching close to base, with primary and secondary branches very short (Fig. 116 K).

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Larva: Maximum length 1.9-2.5 mm. Cephalic apotome with positive spots arranged along midline, and with 1+1 submedian and basal spots. Ratio of proximal, medial and distal antennal articles = 1:1.5:0.6. Cephalic fan with 45-55 rays. Postgenal cleft acuminate apically, postgenal bridge slightly shorter than hypostomium; ratio of hypostomium/ postgenal bridge = 1.2. Lateral sclerite of thoracic proleg with about 74 teeth. Anal ring with about 450-460 rows of hooks. Rectal papillae with 17-20 diverticula on each lobe. Distribution: Guatemala: Chimaltenango, El Quiché, Huehuetenango, Sacatepequez, Solalá; Mexico: Chiapas, Oaxaca. Bionomics: Simulium smarti breeds in small waterfalls and torrential streams of clear water (0-23 °C), attached to rocks and aquatic vegetation. Females bite livestock (DALMAT, 1955). Discussion: The adults of S. mexicanum and S. smarti are very similar, not showing a pronounced keel on the basal plate of the males and with no median group of spiculae on the female cibarium. The major differences are in the pupa, with that of S. mexicanum having more tubercles on the frontoclypeus and thorax, and the gill having 12 branches. Simulium (Hemicnetha) hieroglyphicum PETERSON, VARGAS & RAMÍREZ PÉREZ (Fig. 116 L.) Simulium (Hemicnetha) hieroglyphicum PETERSON et al., 1988: 76; CROSSKEY & HOWARD, 1997: 38; 2004: 30.

(Based on original description). Female: Wing length 4.0-4.5 mm. Scutum brownish with silvery pollinose margins; with posterior illumination, 3 slender dark stripes extending from anterior to posterior edge; scutum covered by golden groups of recumbent hair. Scutellum yellowish brown with golden yellow and black setae. Metanotum with silver pollinosity. Pleurae and abdomen brownish black. Length of palpomere/sensory vesicle = 1.9. Cibarium with spiculae on central area; Sc with hair covering most of vein. Paraproct and gonapophysis as in S. mexicanum. Male: Wing length 3.9-4.2 mm. Scutum, scutellum and metanotum as in female. Abdomen brownish black dorsally, yellowish laterally; tergites II-VII with 1+1 silvery pollinose lateral spots. Gonostylus similar to that of S. seriatum, but with rather thin median lateral protuberances; ratio of length/width = 2.3; ratio of length of gonostylus/ gonocoxite = 2. Ventral plate with elevated median keel as in S. smarti. Pupa: Cocoon shoe shaped but lower anteriorly than in other Simulium (Hemicnetha) species, projected anteroventrally and with 2-4 festoons. Length basally 6.3-7.0 mm, gills 3.0 mm. Frontoclypeus and exposed portion of thorax without tubercles but with strong rugosity as in S. bricenoi; trichomes single. Gill very unique with short trunk and 1 thick basal branch projecting anteriorly and posteriorly over head and thorax, respectively, with 60-90 small thin branches emerging from basal branch (Fig. 116 L). Larva: Maximum length 13.0-13.5 mm. Cephalic apotome with positive spots. Antenna surpassing length of cephalic fan stem; ratio of proximal, medial and distal articles = 1:2:0.6. Cephalic fan with 52-59 rays. Postgenal bridge slightly longer than hypostomium length. Anal ring with 500-550 rows of 50-55 hooks each. Rectal papillae with 20-41 diverticula on each lobe. Distribution: Costa Rica: Alajuela.

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Bionomics: The species breeds in shaded torrential streams about 3 m wide with a stone bottom. Larvae are found on rocks and aquatic vegetation. Females have not been found biting humans. Discussion: The adult is similar in coloration and genitalia to those of S. mexicanum, S. seriatum and S. smarti but the pupa of S. hieroglyphicum has rugosity on the frontoclypeus and thorax and the gill has 60-90 branches. The pupa of S. bricenoi has similar rugosity but the gill has 8 branches, and the cocoon is anteriorly more elevated. Simulium (Hemicnetha) freemani VARGAS & DÍAZ NÁJERA Simulium (Dyarella) freemani VARGAS & DÍAZ NÁJERA, 1949: 289-292. Simulium (Hemicnetha) freemani VARGAS & DÍAZ NÁJERA, 1957b: 153, 175, 177-179, 263, 314-315; COSCARÓN, 1987: 35; IBÁÑEZ-BERNAL, 1992: 98-111; CROSSKEY & HOWARD, 1997: 38; 2004: 30; ADLER et al., 2004: 370.

Female: Wing length 4.0 mm. Scutum dark brown with yellowish pile, with 1 median wide brown vitta separated medially by thin dark brown stripe, and 1+1 light brown sublateral vittae edged by dark gray pollinosity that also covers lateral margins and prescutellum area. Scutellum light yellowish. Legs yellowish with apex of median and hind femora and tarsomeres dark brown. Abdomen blackish with tergal plates of segments III-V brownish. Sensory vesicle of maxillary palp half of palpomere length. Cibarium with small spiculae on central area. Gonapophysis subtrapezoidal and rounded apically, with inner edge lacking microtrichia, as in S. seriatum; paraproct relatively short, about as long as wide. Male: Wing length 4.0-4.2 mm. Scutum dark brown, edged all around except anteromedially with gray pollinosity; covered by yellow hair with green shiny metallic color. Scutellum light yellowish. Ratio of length/width of hind basitarsus = 3.8. Gonostylus elongated with median internal prominence well developed; ratio of length/width = 2.6-2.8. Ventral plate with prominent median carina, appearing in outline like primitive ax. Pupa: Cocoon shoe shaped, without expansion on anterior edge, gill free. Length basally 2.7-3.1, height in front 2.0-2.2 mm, gills 2.0-2.1mm. Frontoclypeus and exposed portion of thorax with rounded tubercles moderately abundant on frontal area, and more numerous on thorax; facial area with smaller tubercles. Cephalic area and thorax with trichomes single. Gill with 8 branches arising from short basal trunk. Larva: Undescribed from Neotropical area. ADLER et al. (2004) show dorsally positive spots on cephalic apotome, and deep and acute postgenal cleft, with postgenal bridge shorter than hypostomium length. Distribution: Mexico: Guanajuato, Guerrero, Michoacan, Oaxaca; USA: Arizona, New Mexico, Utah. Bionomics: The immatures were collected in rocky streams at altitudes of 16002000 m. The female feeding habits are unknown. Discussion: The closest species is S. smarti, having males with a similar laterally compressed ventral plate, but that species has a wider hind basitarsus and a pupal gill with 18 branches.

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Simulium (Hemicnetha) earlei VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Fig. 115 K.) Simulium (Dyarella) earlei VARGAS et al., 1946: 118-120; DALMAT, 1955: 176-181. Simulium (Hemicnetha) earlei VARGAS & DÍAZ NÁJERA, 1957b: 175-177, 180, 256, 312; COSCARÓN, 1987: 35; IBÁÑEZ-BERNAL, 1992: 81-98; SHELLEY et al., 2002: 141-144; CROSSKEY & HOWARD, 1997: 37; 2004: 30. ?Simulium (Hemicnetha) dehnei FIELD, 1969a: 162.

Female: Wing length 2.6-3.8 mm. Scutum yellowish brown to dark brown with gray pollinosity, with frontal light 1+1 submedian and lateral light grayish vittae running 2/3 of scutum length; with posterior light scutum darker and submedian vittae get thinner and blackish; scutum covered by grouped whitish recumbent hair. Scutellum yellowish to dark brown and metanotum dark brown. Pleura brown with grayish pollinosity. Legs yellowish with middle and hind femur apices, tibiae and tarsomeres dark brown. Abdomen blackish, tergites I-II with grayish pruinosity; tergal plates of segments III-V light brown, VI-IX shiny. Cibarium with spiculae on central area and over pharyngeal membrane. Ratio of length of sensory vesicle of maxillary palp/palpomere = 2.5. Paraproct typical for subgenus; gonapophysis subtriangular, elongated with acute apex, similar to that of S. brachycladum. Male: Wing length 2.8-3.7 mm. Scutum dark brown with golden recumbent hair; with posterior light, 1+1 elongated submedian light grayish pollinose stripes separated on median longitudinal line by thin blackish stripe running to anterior 2/3 of scutum length; scutellum, metanotum pleura and legs as in female. Abdomen blackish with 1+1 silver pruinose spots on tergites II-IV, VI-VII. Ratio of length/width of hind basitarsus = 4.8. Gonostylus with conspicuous prominences on lateral edges of median area, as in S. seriatum, ratio of length/width = 2.1, ratio of length/width of gonostylus/gonocoxite = 3.2. Ventral plate wide, membranous with abundant hair, as in S. seriatum. Pupa: Cocoon shoe shaped with strong anteromedian projection leaving most of gill free (Fig. 115 K). Length basally 3.5-4.5 mm; dorsally 3.0-3.6 mm. Gills: 1.3-1.6 mm. Frontoclypeus and exposed portion of thorax mostly smooth, with tubercles on facial area of frontoclypeus, fewer on anterior and posterior lateral areas of thorax. Cephalic trichomes single, thoracic with 2-5 branches. Gill with 16 (15-17) branches emerging from short trunk with very short primary branches; branches filamentous, acute apically, striated and covered by spiculae. Larva: Maximum length 8.7-9.6 mm. Cephalic apotome with positive spots. Ratio of proximal, medial and distal antennal articles = 1:1.6-1.8:0.8-0.9. Postgenal cleft subcircular; postgenal bridge as long as or longer than hypostomium, ratio of hypostomium/ postgenal bridge length = 0.8-1.0. Cephalic fan with 40-51 rays. Anal ring with 200-230 rows with 27-30 hooks each. Rectal papillae with 7-10 diverticula on each lobe. Distribution: Belize; Costa Rica; Guatemala; Mexico: Colima, Distrito Federal, Guerrero, Mexico, Michoacan, Morelos, Nueva Leon, Oaxaca, Veracruz. Bionomics: Simulium earlei was collected in localities between 200 and 1900 m of altitude in fast-flowing streams with water temperatures of 13-25 °C and pH of about 7. The larval stages were found attached to rocks and aquatic vegetation. There are no records of females biting humans; in Belize they are zoophilic (SHELLEY et al., 2002). Discussion: The adult scutum ornamentation and genitalia have similarities with S. brachycladum, S. guerrerense, S. keenani, S. solarii and S. yepocapense. Differences from S. keenani

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are explained under that species. The other species are easy to separate by their lower number of gill branches, but there are specimens of S. earlei with 15 branches that can be confused with S. solarii; the latter species has a cocoon that is lower anteriorly, gonapophyses that are wider medially, and a ventral plate that is shorter and with a long median process. Simulium dehnei is included here as a synonym with doubts, as presented by CROSSKEY & HOWARD (2004). This species is known only from a female from Panama and the type material is lost. From the description, S. dehnei is similar to S. earlei, S. pulverulentum and S. yepocapense, but as is common in this subgenus, females closely resemble one another. Without other stages, it is difficult to establish the species identity, and S. dehnei could be positioned as species inquirendae. Simulium (Hemicnetha) keenani FIELD Simulium (Hemicnetha) keenani FIELD, 1969a: 157-162; COSCARÓN, 1987: 30; CROSSKEY & HOWARD, 1997: 38; 2004: 30.

(Diagnosis based on original description). Female: Scutum brownish with silvery submedian vittae lyre shaped, and with recumbent silver hair intermixed with some erect black hair; scutellum light brown, metanotum brown blackish. Sensory vesicle of maxillary palp slightly shorter than palpomere; ratio of sensory vesicle/palpomere length = 2.3. Cibarium without denticles. Sc with hair surpassing basal 3/4 of vein. Genitalia as in S. rubrithorax with paraproct slightly shorter. Male: Scutum brownish, with some illumination nacreous; scutellum brown, metanotum brownish black, as in female Genitalia as in S. earlei, with ventral plate subtrapezoidal and gonostylus with median lateral prominence well developed. Pupa: Gill with 16 filamentous branches emerging very close to short basal trunk. Larva: Unknown. Distribution: Panama: Canal Zone. Bionomics: Unknown. Discussion: Simulium earlei is very similar, especially in the number of pupal gill branches and the male genitalia. The principal difference is in the gonapophysis, which is more elongated in S. earlei. The absence of type material causes more doubts about the species validity; the name might be a synonym of S. earlei. More topotypical material and cytological and molecular studies are needed for better identification. Simulium (Hemicnetha) hinmani VARGAS, MARTINEZ-PALACIOS & DÍAZ-NÁJERA Simulium (Dyarella) hinmani VARGAS et al., 1946: 120-122, 178. Simulium (Hemicnetha) hinmani: VARGAS & DÍAZ NÁJERA, 1957b: 153, 155-177, 179, 180, 263; COSCARÓN, 1987: 36; CROSSKEY & HOWARD, 1997: 38 ; 2004: 30; IBÁÑEZ-BERNAL, 1992: 127-144.

Female: Wing length 4.0-4.3 mm. Scutum brown-orange with golden hair; in frontal illumination, with median and 1+1 sublateral lighter vittae, separated by 1+1 submedian darker pollinose stripes, as in S. brachycladum; with posterior illumination, median and submedian vittae become darker, scutellum yellowish; metanotum dark brown. Legs as in

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S. guerrerense. Abdomen brownish gray with silver pollinose spot on tergite II, and 1+1 smaller spots on III-V. Cibarium with small spiculae on median area, as in S. brachycladum. Sensory vesicle of maxillary palp slightly shorter than half of palpomere length. Sc with hair surpassing level of branching of R2+3 and R4+5. Gonapophysis elongated with acute apex, as in S. brachycladum. Paraprocts as in S. brachycladum, but with abundant microtichia on half distal. Male: Wing length 4.2-4.4 mm. Scutum coloration as in female, changing intensity of coloration on submedian and sublateral stripes with change of light position. Legs as in female. Abdomen dark brown grayish with 1+1 grayish pollinose spots on tergites II, IVVII. Gonostylus long, 1.5-2.2 times gonocoxite length, and with slight lateral protuberances, as in S. bricenoi; ventral plate wide, similar to that of S. seriatum. Pupa: Cocoon shoe shaped, with strong anterior elevation, covering about half of basal gill. Length basally 3.6-3.7 mm high on anterior portion 1.6-1.9 mm. Frontoclypeus smooth, exposed portion of thorax with numerous acute, short spine-like tubercles on median dorsal area. Cephalic trichomes single and short, thoracic trichomes single but relatively long. Gill with 10 branches, emerging from short basal trunk; filaments striated and acute apically. Larva: Maximum length about 8.0 mm. Cephalic apotome with positive spots. Cephalic fan with about 45 rays. Ratio of proximal, medial and distal antennal articles = 1:1.4:0.8. Postgenal cleft relatively short with arc shape apex; ratio of postgenal bridge/ hypostomium = 1.5. Anal ring with about 225 rows of hooks. Rectal papillae with about 15 diverticula on each lobe. Distribution: Mexico: Chiapas, Distrito Federal, Mexico, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Simulium yepocapense is the closest species, with similar coloration and female and male genitalia, anteriorly elevated cocoon, and number of gill branches. However, in the pupa it lacks acute tubercles on the thoracic dorsum and multibranched thoracic trichomes, and in the larva, it has a deeper acute postgenal cleft. Cytological and molecular studies would help resolve specific differences. Simulium guerrerense and S. pulverulentum also have 10 gill branches. The first species also has the cocoon elevated anteriorly and no tubercles on the thorax, the male ventral plate narrower distally, and the female gonapophysis shorter and wider medially. Simulium pulverulentum has a lower anterior portion on the cocoon an absence of tubercles on the pupal thorax, a ventral plate with a median process and a shorter gonapophysis. Simulium (Hemicnetha) yepocapense DALMAT (Fig. 113 V.) Simulium (Dyarella) yepocapense DALMAT, 1949: 548-553; 1955: 204-208; VARGAS & DÍAZ NÁJERA, 1954: 60. Simulium (Hemicnetha) yepocapense: VARGAS & DÍAZ NÁJERA, 1957b: 154, 175-177, 179, 180, 332, 333; COSCARÓN, 1987: 36; CROSSKEY, 1987: 453; PETERSON et al., 1988: 85; IBÁÑEZ-BERNAL, 1992: 244-260; CROSSKEY & HOWARD, 1997: 38; 2004: 31. Simulium (Dyarella) ardeni DALMAT, 1953: 35-40; 1955: 171-176.

Female: Wing length 3.4 mm. Scutum brownish gray with hair light gray-golden and blackish intermixed; coloration changing with illumination: with frontal light, 1+1 submedi-

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an and lateral dark grayish pollinose stripes, as in S. brachycladum; with posterior light, submedian vittae thinner and anterior area darker. Scutellum and metanotum brownish gray. Legs yellowish brown with most of apices of segments brown-grayish. Abdomen brownish, pale on segments I-II, VI-IX reddish brown. Cibarium as in S. brachycladum; sensory vesicle of maxillary palp shorter than half of palpomere length; Sc with hair on basal half. Gonapophysis elongate and apically acute (Fig. 113 V), paraprocts rounded. Male: Wing length 3.4 mm. Scutum dark brownish gray with golden and blackish pile, with 1+1 submedian gray pollinose bands separated medially by thin longitudinal stripe, and lateral margin of scutum whitish pruinose. Abdomen brown with 1+1 gray pruinose spots on tergites II, IV-VI. Ratio of length/width of hind basitarsus = 3.8-4.0. Gonostylus elongate with inner and outer curvature, as in S. seriatum; ratio of length/width = 1.9; ratio of length of gonostylus/gonocoxite = 2.1-2.2. Ventral plate wide, as in S. earlei. Pupa: Cocoon shoe shaped, very elevated anteriorly, covering gills, length basally 3.54.0 mm, maximum height 1.8 mm, gills 1.3-1.5 mm. Frontoclypeus and exposed portion of thorax without tubercles, except small platelets on facial area of frontoclypeus and posterolateral area of thorax. Head trichomes single, thoracic trichomes with 2-5 branches. Gill with 10 branches emerging from short basal trunk; filaments striated and acute apically. Larva: Maximum length 7.0-8.0 mm. Cephalic apotome with positive spots; antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal antennal articles = 1:1.9-2:1.4-1.5. Cephalic fan with about 45 rays. Postgenal cleft acute apically, postgenal bridge slightly shorter than hypostomium length. Anal ring with about 220 rows of hooks. Rectal papillae with 11 diverticula each. Distribution: Guatemala: Chimaltenango, El Quiche, Huehuetenango, Suchitepequez; Mexico: Chiapas, Durango. Bionomics: According to DALMAT (1955), this species breeds in small torrential creeks attached to rocks and aquatic vegetation from sea level to 2400 m. Females have zoophilic habits. Discussion: Simulium yepocapense is a peculiar species with characters that permit identification. The female is similar in coloration and genitalia to S. brachycladum and S. cristalinum. The male is close to S. earlei in the shape of the ventral plate and to S. bricenoi in the shape of the gonostylus, but these species have 16 and 8 pupal gill branches, respectively. The pupal gill with 10 branches is similar to that of S. pulverulentum and S. guerrerense but in the former species, the cocoon does not cover the gills, and in the latter species, the aperture is cylindrical with the same height anteriorly and posteriorly. OVIEDOI species group Simulium oviedoi RAMÍREZ PÉREZ (Figs. 118 A-F; 119 A-D, H-O.) Simulium oviedoi RAMÍREZ PÉREZ, 1971: 363; 1983: 66; COSCARÓN, 1987: 27; CROSSKEY & HOWARD, 1997: 82; 2004: 77.

Female: Wing length 3.0 mm. Scutum blackish with 1 median and 1+1 sublateral longitudinal darker areas. Scutellum brownish; metanotum dark brown, pollinose. Legs brownish gray with apical portions of femur, tibia and tarsomeres dark brown. Abdomen

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blackish, with 1+1 silvery spots on tergite II. Eighth sternite and gonapophysis as in S. rivasi; paraproct about as long as wide at base (Fig. 118 A ); genital fork as figure 118 D. Male: Wing length 2.6 mm. Scutum coloration as in female. Abdominal tergites II, VVII with 1+1 silvery spots. Ventral plate with very short concavity basally (Fig. 118 E). Gonostylus slightly shorter than gonocoxite (Fig. 118 F); ratio of length/width at base = 1.8. Pupa: Cocoon aperture relatively lower basally (Fig. 119 A), length basally 3.2 mm, dorsally 3.0-3.4 mm, maximum length 4.0-4.4 mm; gills 1.4-1.7 mm. Frontoclypeus and exposed portion of thorax with tubercles acuminate, spine-like (Figs. 119 B, D); cephalic trichomes with 1-3 branches, thoracic trichomes with 2-4 branches. Gills as in figure 119 C. Larva: Maximum length 7.0-8.0 mm. General aspect as in figure 119 H. Antenna with very thick articles (Fig. 119 I); ratio of proximal, medial and distal articles = 1:1.82.4:2-2.4. Comb of cephalic fan rays with similar teeth length (Fig. 119 J). Mandible as in figure 119 K. Postgenal cleft subtriangular (Fig. 119 L); hypostomium median tooth slightly longer than lateral teeth (Fig. 119 M). Lateral sclerite of thoracic proleg as in figure 119 N, Anal sclerite with abundant hair (Fig. 119 O), anal ring with 100-160 rows. Distribution: Venezuela: Merida. Bionomics: Unknown. Discussion: This species is very close to S. rivasi, which has rounded and granulose pupal tubercles. Simulium hinmani also has acuminate tubercles. Simulium rivasi RAMÍREZ PÉREZ (Figs. 118 B-D, G, H; 119 E-G.) Simulium rivasi RAMÍREZ PÉREZ, 1971: 359-360; COSCARÓN, 1987: 27; CROSSKEY, 1987: 485; CROSSKEY & HOWARD, 1997: 82; 2004: 77. Simulium rivai and Simulium ribasi: incorrect spellings.

Female: Size and coloration as in S. oviedoi but with more evident light, thin median and 1+1 submedian vittae. Paraproct slightly curved apically (Fig. 118 B). Eighth sternite with about 30 setae on each side, gonapophysis slightly sclerotized on internal margin (Fig. 118 C); ratio of length/width at base = 1; genital fork as in figure 118 D. Male (pharate): Genitalia as in S. oviedoi but gonostylus proportionately longer (Fig. 118 H) and ventral plate with deeper concavity at base (Fig. 118 G). Pupa: Cocoon similar to that of S. oviedoi. Length basally 5.6-6.2 mm, gills 2.8-3.0 mm. Frontoclypeus and exposed portion of thorax with abundant granulose tubercles (Figs. 119 E, G); trichomes with 2-5 branches. Gills as in figure 119 F. Larva: Maximum length 9.0-9.5 mm. Mandible and cephalic fan as S. oviedoi. Ratio of proximal, medial and distal antennal articles = 1:2.7:2.8. Anal ring with about 120 rows with 20-23 hooks each. Distribution: Venezuela: Merida. Bionomics: Unknown. Discussion: Simulium rivasi is very similar to S. oviedoi, as described for that species. The small differences between these species, based on specimens from the same place, suggest that it would be useful to obtain more material from different localities and to do cytological and molecular studies to assure the identity of the species and their supraspecific taxonomic position.

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C A

B D

H

G

F E

Figure 118. A-D: Female; A: S. (Hemicnetha) oviedoi, cercus and paraproct. B-D: S. (H. ) rivasi; B: cercus and paraproct; C: eighth sternite and gonapophysis; D: genital fork. E-H: Male. E-F: S. (H.) oviedoi; E: ventral plate; F: gonocoxite and gonostylus. G-H: S. (H.) rivasi; G: ventral plate; H: gonocoxite and gonostylus.

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A B

J

C

I

D F M

G

E L N

O

K H Figure 119. A-G: Pupa; A-D: S. (Hemicnetha) oviedoi; A: lateral view; B: frontoclypeus; C: anterodorsal portion of thorax and gill; D: thoracic tubercles and trichomes enlarged. E-G: S. (H.) rivasi; E: frontoclypeus; F: anterodorsal portion of thorax and gill; G: thoracic tubercles and trichomes enlarged. H-O: Larva; S. (H.) oviedoi; H: lateral view; I: antenna; J: comb of cephalic fan ray; K: mandibular teeth; L: postgenal cleft and hypostomium; M: hypostomium teeth of anterior margin; N: lateral sclerite of thoracic proleg; O: anal sclerite.

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Species inquirendae Simulium tarsale WILLISTON Simulium tarsale WILLISTON, 1893: 268-269; STONE, 1969: 316-317; HERNÁNDEZ & SHELLEY, 2005: 5-6. Simulium clavipes MALLOCH, 1914: 40-41.

STONE’s (1969) redescriptions of females and males of this species from Dominica, which are in accordance with the type material, and the later study of types by HERNÁNDEZ & SHELLLEY (2005), indicate similarities with other Simulium (Hemicnetha) species. The absence of knowledge of the larval stage, and in particular the pupa, makes it impossible to decide if S. tarsale is conspecific with some other well-known species, such as S. earlei, S. bricenoi, S. mexicanum or S. virgatum, with which it is similar. An important difference is the absence in S. tarsale of subbasal tooth on the female claw and the blunt anterolateral apodemes on the lateral branches of the genital fork. We prefer to maintain Simulium tarsale as species inquirenda, pending more information. Distribution: Dominica; Guadeloupe. Simulium falculatum ENDERLEIN Simulium falculatum ENDERLEIN, 1929: 327 (= Simulium coffeae VARGAS, 1945a: 4). Simulium (Hemicnetha) falculatum: HERNÁNDEZ & SHELLEY, 2005: 3-5.

This species is externally similar to S. paynei and without knowledge of the genitalia, it was synonymized with doubt (COSCARÓN, 1987). Later, ADLER et al. (2004: 373) indicated that S. falculatum was distinct from S. paynei and removed it from synonymy. HERNÁNDEZ & SHELLEY (2005), through genitalia studies, found differences and agreed with the revalidation of S. falculatum. Their digital illustrations (Figs. 50, 69, 78) show a short gonapophysis and paraproct and, in particular, an elongated stem of the genital fork, resembling the characters of S. pictipes HAGEN and some Simulium (Hearlea) species. The absence of knowledge of the male and larva does not permit placement of S. falculatum in a Simulium subgenus. Without more information we consider it as species inquirenda. Simulium tarsatum MACQUART Simulium tarsatum MACQUART, 1845 (1844): 20. Simulium (Hemicnetha) tarsatum: HERNÁNDEZ & SHELLEY, 2005: 6.

According to HERNÁNDEZ & SHELLEY (2005), this species, known only from the poorly preserved female holotype (not male, as originally published) from Colombia, without a specific locality, is similar to S. mexicanum with which it was synonymized. As mentioned under S. mexicanum, we consider it better to maintain the well-known species name, S. mexicanum, until other studies with more material, including all life stages of S. tarsatum, preferably from the type area, can be examined.

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Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Figs. 120-127; 152.) Simulium (Hearlea) VARGAS et al., 1946: 104. Type-species: Simulium canadense HEARLE, 1932: 14-15.

Female: Wing length 2.5-5.0 mm. General coloration blackish brown; scutum blackish with 1+1 silvery vittae sometimes joining 1+1 anterior submedian subtriangular or subquadrate spots, and bordered laterally and posteriorly by silver pruinose areas. Legs yellowish to grayish brown, with apical portions dark brown to black. Frons convergent below, fronto-ocular triangle deep, about as long as wide; maxillary palpus with sensory vesicle elongate, nearly 1/2 length of basal palpomere. Cibarium broadly U-shaped, with basal portion often thickened, medially cleft, and without strong teeth, although often with wrinkles or small serrations on pair of sublateral subovoidal prominences. Basal sector of R bare. Sc with 6-18 setae. Hind basitarsus with length/width ratio 5.7-6.0; claw with small, subbasal tooth. Gonapophysis surpassing edge of sternite VIII, subtriangular, with blunt apex and gradually concave internal margins; cerci generally subquadrate, Paraprocts low, sparsely haired, truncate distally, with abundant short hair; genital fork with stout rods, apically expanded arms bearing strong anteriorly directed apodemes; spermatheca ovoid, with internal spicules. Male: Wing length 2.6-5.0 mm. General coloration similar to that of female, differing as follows: scutum generally velvety black with anterior 1/3 with whitish pruinosity. Sc frequently bare; hind basitarsus with length/width ratio of 2.7-3.6. Gonocoxite subquadrate, with small protuberance on external margin; gonostylus thin, elongate, 1/3 longer than gonocoxite, with small haired subbasal carina posteriorly, and 1 subapical spine; ventral plate generally subquadrate, with narrow median carina; endoparameres with wide base and numerous strong hooks; median sclerite subrectangular. Pupa: Cocoon slipper or shoe shaped, typically coarsely woven, no threads evident, with or without reinforced anterior edge; length basally 3.5-4.5 mm (range = 2.4-7.0 mm). Gill inflated, gradually flattened, pseudosegmented, and lacking secondary branches to cylindrical, smooth, with secondary branches. Frontoclypeus gradually swollen basally. Frontoclypeus and anterodorsum of thorax smooth or covered with abundant impressed platelets or rounded or pointed elevated granules. Frontoclypeus with 2+2 frontal and 1+1 facial setae; thorax with 3(4)+3(4) dorsocentral and 2+2 dorsolateral stout trichomes. Abdominal chaetotaxy as in figure 125. Setae of abdominal tergite I relatively short, hairlike, and with small platelets along anterior edge; setae of abdominal tergites II-VI short and slender. Tergites II-IV with 4+4 retrorse hooks, those of tergite II smaller; tergites VVIII with minute spine-combs. Tergite X with or without terminal hooks. Sternites IV-VII with small, wrinkled areas. Larva: Body length 6.5-12.0 mm. Distal portion of body typically flattened dorsally and gradually swollen ventrally (Fig. 127 G); body without evident setae. Cervical sclerites free or adjoining posterior edge of postgena. Antenna longer than stalk of cephalic fan, with proximal article elongate and deeply incised, creating segmented appearance; medial article longer than proximal and distal. Cephalic apotome faintly darkened basally, typically lacking discernible head spots. Cephalic fan with 39-54 primary rays. Mandible with strong lateral process, robust intermediate and internal teeth, and 2 thin marginal

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teeth. Postgenal cleft bluntly to acutely tapered apically; hypostomial bridge generally shorter than hypostomium height, ratio of hypostomium/hypostomial bridge = 0.71.5:1.0; anterior margin of hypostomium curved or straight, with median tooth longer than corner teeth. Anal sclerite sometimes encircling posterior circlet, and distal portion of abdomen with accessory plates. Anal papillae of 3 lobes, with 15-78 secondary lobules. Posterior circlet with 102-400 rows of 16-27 hooks. Bionomics: The immature stages are found typically on stones or vegetation in torrenticolous streams with temperatures of 10-21 °C and at altitudes of 1000-3000 m. Feeding habits of females are unknown. Phylogenetic analyses show two well-differentiated clades within Simulium (Hearlea), named JUAREZI species group and CAROLINAE species group. Hearlea closely resembles other Neotropical subgenera, particularly Hemicnetha ENDERLEIN, Trichodagmia ENDERLEIN and Thyrsopelma ENDERLEIN (COSCARÓN, 1987). These subgenera have recently been included in the same subgeneric group with Freemaniellum CROSSKEY, Anasolen ENDERLEIN, Xenosimulium CROSSKEY, and Metomphalus ENDERLEIN (MIRANDAESQUIVEL, 1999; MIRANDA-ESQUIVEL & COSCARÓN, 2001). These morphological similarities incline us to believe that Hearlea might belong to a Gondwanian lineage that likely passed into Central America or Mexico (from South America?) during the late Cretaceous or early Paleocene, presumably during a brief period of contact. KEYS

TO

SIMULIUM (HEARLEA)

SPECIES

Females Not included are Simulium nigrocornis and S. temascalense, which are inseparable from other species, and S. chiriquiense and S. paracarolinae which are unknown. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(4)

Scutum with silvery submedian vittae joining 1+1 anterior silvery spot (Figs. 120 A, B) ...................................................................................................................................... 2 Scutum with silvery submedian vittae not joining 1+1 anterior silvery spots (Figs. 120 K, L) .............................................................................................................................. 7 Wing length 2.5-2.7 mm ......................................................................... Simulium estevezi Wing length > 3.0 mm ...................................................................................................... 3 Cibarium without 1+1 spindle-shaped phalanges in space between arms .............. ...................................................................................................................... Simulium ayrozai Cibarium with 1+1 spindle-shaped phalanges in space between arms (Fig. 120 M) ................................................................................................................................................ 4 Scutum with anterior submedian 1+1 silvery spots subquadrate (Figs. 120 A, B) ................................................................................................................................................ 5 Scutum with anterior submedian 1+1 silvery spots subtriangular (Figs. 120 K, L) ................................................................................................................................................ 6 Posterior edge of tergites II-V grayish ............................................ Simulium canadense Posterior edge of tergites II-V not grayish ......................................... Simulium juarezi Cibarium with anterior rim thin (Fig. 120 C); wing length 3.1 mm .......................... ................................................................................................................... Simulium delatorrei

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Cibarium with anterior rim thick (Fig. 120 M); wing length 3.4-3.6 mm ................ ................................................ Simulium dalmati, Simulium ethelae, Simulium larvispinosum 7(1) Cibarium with acute indentation in space between arms (Fig. 120 G) .................. 8 – Cibarium with shallow indentation in space between arms (Fig. 120 M) ............ 11 8(7) Scutum with anterior 1+1 submedian silver spots subtrapezoidal to subquadrate ................................................................................................................................................ 9 – Scutum with anterior 1+1 submedian silver spots subtriangular .............................. ........................................................................................................ Simulium microbranchium 9(8) Wing length 2.7-3.1 mm ................................................................................................. 10 – Wing length 3.4 mm ............................................................................ Simulium menchacai 10(9) Wing length 2.7-2.8 mm; scutum with anterior submedian 1+1 silver spots subtrapezoidal ....................................................................................... Simulium capricorne – Wing length 2.9-3.1 mm; scutum with anterior submedian 1+1 silver spots subquadrate ................................................................................................. Simulium burchi 11(7) Wing length about 5.0 mm ............................................................... Simulium contrerense – Wing length less than 4.0 mm ....................................................................................... 12 12(11) Wing length 3.7-3.9 mm; scutum with anterior submedian silvery spots subtrapezoidal ......................................................................................... Simulium carolinae – Wing length < 3.2 mm; scutum with anterior submedian silvery spots subtriangular .......................................................... Simulium gorirossiae, Simulium johnsoni Males No male key is presented due to lack of characters. Pupae Unknown: Simulium paracarolinae. 1 – 2(1) – 3(2) – 4(3) – 5(3)

Gill branches cylindrical, smooth, and with 4 or more small secondary branches (Fig. 124). Cocoon generally slipper-shaped (Figs. 124 A, J) .................................... 2 Gill branches flattened or subcylindrical, with noticeable wrinkling or annulations, and lacking small secondary branches (Figs. 122, 123). Cocoon more typically shoe-shaped (Figs. 122 A, J, L; 123 A, G) .................................................................... 7 Secondary gill branches smaller than primary anterodorsal branch. Ventral branch with 2 secondary acuminate processes; anterior edge of cocoon complete, lacking noticeable threads .............................................................................................................. 3 Secondary gill branches as large as primary anterodorsal branch (Figs. 124 J, K). Ventral branch with 6-7 secondary acuminate processes; anterior edge of cocoon ragged, with noticeable threads ......................................................... Simulium menchacai Gill with primary anterodorsal branch acuminate (Figs. 124 F, G) ........................ 4 Gill with primary anterodorsal branch blunt (Figs. 124 A-E, H, I) ........................ 5 Frontoclypeus and thorax with sparse granules ............................Simulium gorirossiae Frontoclypeus and thorax without granules ............................... Simulium temascalense Frontoclypeus and thorax with abundant granules. Cocoon with reinforced anterior edge ........................................................................................... Simulium carolinae

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– 6(5) – 7(1) – 8(7) – 9(8) – 10(8) – 11(8) – 12(7) – 13(12) – 14(12) – 15(14) – 16(15) – 17(14) –

Frontoclypeus and thorax without elevated granules. Cocoon without, or with only slightly, reinforced anterior edge ........................................................................... 6 Secondary gill branches strongly spined apically (Figs. 124 H, I). Frontoclypeus without platelets ...................................................................................... Simulium johnsoni Secondary gill branches blunt or weakly spined apically (Figs. 124 A-D). Frontoclypeus and thorax with impression of minute non-elevated granules especially on edge of disc ............................................................ Simulium larvispinosum Gill with 2 branches .......................................................................................................... 8 Gill with 3 branches ........................................................................................................ 12 Cocoon shoe-shaped (Figs. 122 A, J, L) ........................................................................ 9 Cocoon mostly slipper-shaped (Figs. 122 D, F) ........................................................ 11 Dorsal gill branch 1.8 times longer than ventral branch and strongly curved (Figs. 122 J, K) ....................................................................................... Simulium microbranchium Dorsal gill branch 0.6-1.0 times as long as ventral branch and only mildly curved (Figs. 122 B, L-N) ............................................................................................................. 10 Gill protruding well beyond anterior margin of cocoon (Figs. 122 L-N); apex of gill branches blunt .................................................................................Simulium nigricorne Gill largely concealed by cocoon (Figs. 122 A, B); apex of gill branches acuminate ...................................................................................................................... Simulium ayrozai Dorsal gill branch acuminate, subequal in thickness to ventral branch, and only slightly curved apically (Figs. 122 F-I) ............. Simulium ethelae, Simulium chiriquiense Dorsal gill branch blunt, twice as thick as ventral branch, and strongly curved apically (Figs. 122 D, E) ...................................................................... Simulium capricorne Frontal gill branch subglobose, inflated, tuberculated, not annulated (Figs. 123 N, O) ......................................................................................................................................... 13 Frontal gill branch subcylindrical, scarcely or not tuberculated, annulated ........ 14 Two frontal gill branches inflated, ventral branch with ventral area membranous; another ventrolateral branch small and subannulated ................. Simulium contrerense Only 1 frontal gill branch inflated, plus 1 ventrolateral and another dorsolateral branch thinner and subannulated (Figs. 123 N, O) ........................... Simulium juarezi Dorsal gill branch attenuated distally (Figs. 122 C; 123 C-I) .................................. 15 Dorsal gill branch thickened and capitate distally (Figs. 123 A, B, J-M) .............. 17 Dorsal gill branch with small ventral protuberance at about midlength (Figs. 123 C, G) ................................................................................................................................... 16 Dorsal branch lacking ventral protuberance (Fig. 122 C) ........... Simulium canadense Dorsal gill branch with blunt apex (Figs. 123 C-F). Frontoclypeus with platelets laterally and medially with only punctuation (Fig. 121 H) ........... Simulium delatorrei Dorsal gill branch with acuminate apex (Figs. 123 G-I). Frontoclypeus with platelets on entire surface (Fig. 121 I) ................................................. Simulium estevezi Dorsal gill branch globose apically, with well-developed tubercles. Cocoon shoeshaped (Figs. 123 A) .................................................................................. Simulium burchi Dorsal gill branch with 3 apical prominences, lacking tubercles (Figs. 123 J-M). Cocoon slipper-shaped (Figs. 123 J) ....................................................Simulium dalmati

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Larvae Simulium nigricorne and S. chiriquiense are unknown. 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(4) – 7(6) – 8(1) – 9(8) –

Hypostomium with anterior edge convex and with intermediate teeth above level of corner teeth (Fig. 126 J). Distal portion of abdomen with accessory plates (Figs. 127 B-E) .................................................................................................................... 2 Hypostomium with anterior edge straight and with intermediate teeth below level of corner teeth (Fig. 126 B). Distal portion of abdomen without accessory plates ................................................................................................................................................ 8 Distal portion of abdomen with only 1+1 accessory dorsolateral plates; ventral arm short, not encircling base of posterior circlet (Figs. 127 B, C); anal ring with about 400 rows of hooks ..............................................................Simulium paracarolinae Distal portion of abdomen with 2+2 accessory dorsolateral plates (Figs. 127 D, E); anal sclerite ventral arm encircling base of posterior circlet (Figs. 127 D-I); anal ring with < 300 rows of hooks .............................................................................. 3 Distal portion of abdomen without ventrolateral accessory plates .......................... .................................................................................................................... Simulium carolinae Distal portion of abdomen with stout ventrolateral accessory plates (Figs. 127 D, H, J) ....................................................................................................................................... 4 Lobes of accessory ventrolateral plate shallowly incised (Figs. 127 H, I [v1]). Segment VIII without ventrolateral tubercles ............................................................. 5 Lobes of accessory ventrolateral plate deeply incised (Fig. 127 D [v1]). Segment VIII with well-sclerotized ventrolateral tubercles (Figs. 127 D-F [vp]) ................. 6 Cephalic fan with 43-48 primary rays; area before to anal sclerite with 1+1 dorsal subrectangular flattened accessory plates (Figs. 127 H, I [df]) ................... ............................................................................................................ Simulium larvispinosum Cephalic fan with 56 primary rays; area before to anal sclerite without dorsal accessory plates ....................................................................................... Simulium johnsoni Accessory ventrolateral plates of distal portion of abdomen with anteriorly directed acuminate processes; anal ring with 310 rows of hooks ............... Simulium menchacai Accessory ventrolateral plates of distal portion of abdomen with anterolaterally directed acuminate processes; anal ring with 242-282 rows of hooks ................... 7 Accessory ventrolateral plates of distal portion of abdomen strongly toothed (Fig. 127 D [vl]); ventrolateral papillae of sternite VIII with acuminate tubercles (Figs. 127 D-F [vp]) .............................................................................Simulium gorirossiae Accessory ventrolateral plates of distal portion of abdomen with serrated margin (Fig. 127 J [vl]); ventrolateral papillae of sternite VIII smooth ................................. .............................................................................................................. Simulium temascalense Rectal papillae with 60-78 lobules .................................................................................. 9 Rectal papillae with 45 or fewer lobules ..................................................................... 12 Rectal papillae with 72-78 lobules; length of mature larva < 7.0-7.7 mm .......... 10 Rectal papillae with 60-66 lobules; length of mature larva > 12.0 mm ................... ................................................................................................................. Simulium contrerense

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– 17(16) –

Anal ring with about 114 rows of hooks. Cephalic fan with 39-40 primary rays ................................................................................................................... Simulium delatorrei Anal ring with about 172-204 rows of hooks. Cephalic fan with 46-57 primary rays ...................................................................................................................................... 11 Anal ring with 172-178 rows of hooks; cephalic fan with 46-54 primary rays ...... ........................................................................................................................ Simulium burchi Anal ring with 194-206 rows of hooks; cephalic fan with 56-57 primary rays ...... ........................................................................................................ Simulium microbranchium Body length 8.1-10.0 mm; hypostomium with 11-17 lateral setae ........................ 13 Body length 7.0-8.0 mm; hypostomium with 10 or fewer lateral setae ................ 15 Cephalic fan with 42-44 primary rays; ratio of length of hypostomium/ hypostomial bridge = 0.8 .......................................................................Simulium dalmati Cephalic fan with 48-56 primary rays; ratio of length of hypostomium/ hypostomial bridge = 1.0 ............................................................................................... 14 Anal ring with 18-20 hooklets per row; rectal papillae with 15-24 lobules ............. .................................................................................................................. Simulium canadense Anal ring with 24-27 hooklets per row; rectal papillae with 36-45 lobules ............. ...................................................................................................................... Simulium ayrozai Body length 7.0 mm; cephalic fan with 36-40 primary rays ............ Simulium ethelae Body length 7.3-8.0 mm; cephalic fan with 42-50 primary rays ............................ 16 Body length 8.0 mm; anal ring with 98-104 rows of 24-30 hooks ...................... .....................................................................................................................Simulium estevezi Body length 7.5-7.7 mm; anal ring with 115-145 rows of 15-24 hooks .............. 17 Anal ring with 145 rows of about 17 hooks ....................................... Simulium juarezi Anal ring with 115-135 rows of 20-24 hooks ................................ Simulium capricorne

CLAVES

DE ESPECIES DE

– 11(10) – 12(8) – 13(12) – 14(13) – 15(12) – 16(15)

SIMULIUM (HEARLEA)

Hembras No incluidas Simulium nigricorne y S. temascalense por inseparables de otras especies, S. chiriquiense y S. paracarolinae por desconocida. 1 – 2(1) – 3(2) – 4(3)

Escudo con bandas plateadas submedianas uniéndose a 1+1 manchas plateadas anteriores (Figs. 120 A, B) ................................................................................................ 2 Escudo con bandas plateadas submedianas no uniéndose a 1+1 manchas plateadas anteriores (Figs. 120 K, L) .............................................................................. 7 Largo ala 2,5-2,7 mm .............................................................................. Simulium estevezi Largo ala >3,0 mm ............................................................................................................ 3 Cibario con 1+1 elevación en forma de uso en el espacio entre las ramas (Fig. 120 M) .................................................................................................................................. 4 Cibario sin 1+1 elevación en forma de uso en el espacio entre las ramas .............. ...................................................................................................................... Simulium ayrozai Escudo con 1+1 manchas plateadas anteriores submedianas subcuadradas (Figs. 120 A, B) .............................................................................................................................. 5

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–

Escudo con 1+1 manchas plateadas anteriores submedianas subtriangulares (Figs. 120 K, L) .............................................................................................................................. 6 5(4) Margen posterior de los tergitos II-V grisáceo .............................. Simulium canadense – Margen posterior de los tergitos II-V no grisáceo ............................. Simulium juarezi 6(4) Cibario con el borde anterior fino (Fig. 120 C); largo ala 3,1 mm ............................ ................................................................................................................... Simulium delatorrei – Cibario con el borde anterior grueso (Fig. 120 M); largo ala 3,4-3,6 mm ............... ................................................ Simulium dalmati, Simulium ethelae, Simulium larvispinosum 7(1) Cibario con espacios entrecortados suavemente aguzados entre las ramas (Fig. 120 G) ........................................................................................................................................... 8 – Cibario con espacios entrecortados bajos entre las ramas (Fig. 120 M) .............. 11 8(7) Escudo con 1+1 mancha plateada submediana subtrapezoidal a subcuadrada .......... 9 – Escudo con 1+1 mancha plateada submediana subtriangular ....... Simulium microbranchium 9(8) Largo ala 2,7-3,1 mm ...................................................................................................... 10 – Largo ala 3,4 mm ................................................................................. Simulium menchacai 10(9) Largo ala 2,7-2,8 mm; escudo con 1+1 manchas plateadas anteriores submedianas subtrapezoidales ................................................................................... Simulium capricorne – Largo ala 2,9-3,1 mm; escudo con 1+1 manchas plateadas anteriores submedianas subcuadradas ............................................................................................... Simulium burchi 11(7) Largo ala alrededor de 5,0 mm ........................................................ Simulium contrerense – Largo ala menos de 4,0 mm .......................................................................................... 12 12(11) Largo ala 3,9 mm; escudo con 1+1 manchas plateadas anteriores submedianas subtrapezoidales ..................................................................................... Simulium carolinae – Largo ala < 3,2 mm; escudo con 1+1 manchas plateadas anteriores submedianas subtriangulares ...................................................... Simulium gorirossiae, Simulium johnsoni Machos No se presenta clave por escasez de caracteres. Pupas Desconcida: Simulium paracarolinae. 1 – 2(1) – 3(2)

Ramas de la branquia subcilíndrica, lisa, con 4 o más ramas secundarias (Fig. 124); capullo típicamente en forma de chinela (Figs. 124 A, J) ......................................... 2 Ramas de la branquias achatadas o subcilíndricas con notoria rugosidad o anillación y sin pequeñas ramas secundarias (Figs. 122, 123); capullo generalmente en forma de zapato (Figs. 122 A, J, L; 123 A, G) ....................................................... 7 Rama secundaria ventral de las branquias más pequeñas que la rama primaria anterodorsal; rama ventral con 2 prolongaciones secundarias aguzadas; margen anterior del capullo con tejido compacto ..................................................................... 3 Ramas secundarias ventrales de la branquia tan larga como la primaria anterodorsal (Figs. 124 J, K); rama ventral con 6-7 prolongaciones secundarias aguzadas; margen anterior del capullo deshilachado ....................................................... Simulium menchacai Branquia con rama primaria anterodorsal aguzada (Figs. 124 F, G) ....................... 4

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Branquia con rama primaria anterodorsal roma (Figs. 124 A-E, H, I) .................. 5 Frontoclípeo y tórax con tubérculos esparcidos ...........................Simulium gorirossiae Frontoclípeo y tórax sin tubérculos ............................................. Simulium temascalense Frontoclípeo y tórax con abundantes tubérculos; capullo con el borde anterior reforzado .................................................................................................. Simulium carolinae – Frontoclípeo y tórax al máximo con algunos tubérculos esparcidos; capullo sin o con solo un ligero refuerzo en el borde anterior ........................................................ 6 6(5) Ramas secundarias de las branquias con fuertes espinas (Figs. 124 H, I); frontoclípeo sin tubérculos ................................................................... Simulium johnsoni – Ramas secundarias de las branquias romas o con débiles espinas (Figs. 124 A-D); frontoclípeo y tórax con pequeños tubérculos no elevados hasta aguzados especialmente en el dorso del tórax ........................................... Simulium larvispinosum 7(1) Branquia con 2 ramas principales: dorsal y ventral .................................................... 8 – Branquia con 3 ramas principales: dorsal, mediana y ventral ................................. 12 8(7) Capullo en forma de zapato (Figs. 122 A, J, L) ........................................................... 9 – Capullo en forma de chinela (Figs. 122 D, F) ............................................................ 11 9(8) Rama dorsal de la branquia 1,8 veces más larga que la ventral y fuertemente curvada (Figs. 122 J, K) ............................................................ Simulium microbranchium – Rama dorsal de la branquia 0,6-1,0 veces más larga que la ventral y débilmente curvada (Figs. 122 B, L, N); ápices de las ramas de la branquia romas ............... 10 10(9) Branquia sobresale bien más allá del borde anterior del capullo (Figs.122 L-N); ápice de las ramas romo ......................................................................Simulium nigricorne – Branquia bien cubierta por el capullo (Figs. 122 A, B); ápice de las ramas de las branquias aguzadas .................................................................................. Simulium ayrozai 11(8) Rama dorsal de la branquia aguzada, casi igual en grosor a la rama ventral y solo ligeramente curvada apicalmente (Figs. 122 F-I) .......................................................... .................................................................................. Simulium ethelae, Simulium chiriquiense – Rama dorsal de la branquia roma, el doble de gruesa que la rama ventral y fuertemente curvada apicalmente (Figs. 122 D, E) ...................... Simulium capricorne 12(7) Rama mediana de la branquia subglobosa, inflada, tuberculada, no anillada, excepto basalmente (Figs. 123 N, O) ........................................................................... 13 – Rama mediana de la branquia subglobosa, subcilíndrica, escasamente o no tuberculada o anillada ............................................................................................................................. 14 13(12) Ramas dorsal y media de la branquia infladas, la mediana membranosa ventralmente; rama ventral pequeña y subanillada ...................... Simulium contrerense – Rama mediana de la branquia inflada; ramas dorsal y ventral finas, subanilladas (Figs. 123 N, O) .........................................................................................Simulium juarezi 14(12) Rama mediana de la branquia redondeada o desde débil a fuertemente aguzada distalmente (Figs. 122 C; 123 C-I) ................................................................................ 15 – Rama mediana de la branquia engrosada y capitada distalmente (Figs. 123 A, B, J-M) ..................................................................................................................................... 17 15(14) Rama dorsal de la branquia con pequeña protuberancia ventral cerca de la mitad del largo (Figs. 123 C, G) ............................................................................................... 16 – Rama dorsal de la branquia sin protuberancia ventral (Fig. 122 C) ...... Simulium canadense

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16(15) Rama dorsal de la branquia con el ápice romo (Figs. 123 C-F); frontoclípeo con tubérculos lateralmente, medianamente solo con puntuaciones (Fig. 121 H) ........ ................................................................................................................... Simulium delatorrei – Rama dorsal de la branquia con ápice aguzado (Figs. 123 G-I); frontoclípeo con tubérculos en toda la superficie (Fig. 121 I) ....................................... Simulium estevezi 17(14) Rama dorsal de la branquia globosa apicalmente, con tuberosidades bien desarrolladas; capullo con forma de zapato (Fig. 123 A) .................. Simulium burchi – Rama dorsal de la branquia con 3 prominencias apicales faltando tuberosidades (Fig. 123 J-M; capullo con forma de chinela (Fig. 123 J) ................Simulium dalmati Larvas Simulium nigricorne y S. chiriquiense desconocidas. 1 – 2(1)

– 3(2) – 4(3) – 5(4) – 6(4)

Hipostomio con margen anterior convexo y con dientes intermedios arriba del nivel de los dientes laterales (Fig. 126 J); porción posterior del abdomen con placas accesorias (Figs. 127 B-E) .................................................................................... 2 Hipostomio con margen anterior recto, dientes intermedios más cortos que los dientes laterales (Fig. 126 B); porción posterior del abdomen sin placas accesorias ................................................................................................................................................ 8 Porción posterior del abdomen con solamente 1+1 placas accesorias dorsolaterales; sostenes ventrales del esclerito anal cortos no rodeando la base del anillo anal (Figs. 127 B, C); anillo anal con alrededor de 400 hileras de ganchos .............................................................................................Simulium paracarolinae Porción posterior del abdomen con 2+2 placas accesorias dorsolaterales (Figs. 127 D, E); sostenes ventrales del esclerito anal rodeando la base del anillo anal (Figs. 127 D, I); anillo anal con < 300 hileras de ganchos ........................................ 3 Porción posterior del abdomen sin placas accesorias ventrolaterales ...................... .................................................................................................................... Simulium carolinae Porción posterior del abdomen con robustas placas accesorias ventrolaterales (Figs. 127 D, H, J) .............................................................................................................. 4 Lóbulos de la placa accesoria ventrolateral con incisiones no profundas (Figs. 127 H, I [vl] ); segmento VIII sin papilas ventrolaterales ................................................. 5 Lóbulos de la placa accesoria ventrolateral con incisiones profundas (Figs. 127 D [vl] ); segmento VIII con bien definidas papilas esclerotizadas ventrolaterales (Figs. 127 D-F [vp] ) .......................................................................................................... 6 Abanico cefálico con 43-48 rayos; área anterior al esclerito anal con 1+1 placas accesorias dorsales, subrectangulares, achatadas (Figs. 127 H, I [df]) ...................... ............................................................................................................ Simulium larvispinosum Abanico cefálico con 56 rayos; área anterior al esclerito anal sin placas accesorias dorsales ...................................................................................................... Simulium johnsoni Placas accesorias de la porción posterior del abdomen con prolongaciones aguzadas anteriormente dirigidas; anillo anal con 310 hileras de ganchos .............. .................................................................................................................. Simulium menchacai

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– 7(6) – 8(1) – 9(8) – 10(9) – 11(10) – 12(8) – 13(12) – 14(13) – 15(12) – 16(15) – 17(16) –

Placas accesorias ventrolaterales de porción posterior del abdomen sin prolongaciones aguzadas anteriormente dirigidas; anillo anal con 242-282 hileras de ganchos ................................................................................................................................. 7 Placas accesorias de la porción posterior del abdomen fuertemente dentadas (Fig. 127 D [vl] ); papilas ventrolaterales del VIII esternito con tubérculos aguzados (Figs. 127 D-F [vp]) ........................................................................... Simulium gorirrossiae Placas accesorias de la porción posterior del abdomen con borde serrado (Fig. 127 J [vl] ); papilas ventrolaterales del VIII esternito lisas ............. Simulium temascalense Papilas rectales con 60-78 divertículos .......................................................................... 9 Papilas rectales con 45 o menos divertículos ............................................................. 12 Papilas rectales con 72-78 divertículos; largo de la larva madura < 7,0-7,7 mm .............................................................................................................................................. 10 Papilas rectales con 60-66 divertículos; largo de la larva madura > 12,0 mm ....... ................................................................................................................. Simulium contrerense Anillo anal con alrededor de 114 hileras de ganchos; abanico cefálico con 30-40 rayos ........................................................................................................ Simulium delatorrei Anillo anal con alrededor de 172-204 hileras de ganchos; abanico cefálico con 4657 rayos .............................................................................................................................. 11 Anillo anal con 172-178 hileras de ganchos; abanico cefálico con 46-54 rayos ..... ........................................................................................................................ Simulium burchi Anillo anal con 194-206 hileras de ganchos; abanico cefálico con 56-57 rayos ..... ........................................................................................................ Simulium microbranchium Largo del cuerpo 8,0-10,0 mm; hipostomio con 11-17 cerdas laterales .............. 13 Largo del cuerpo 7,0-8,0 mm; hipostomio con 10 o menos cerdas laterales ..... 15 Abanico cefálico con 42-44 rayos; relación largo hipostomio/puente postgenal = 0,8 ............................................................................................................Simulium dalmati Abanico cefálico con 48-56 rayos; relación largo hipostomio/puente postgenal = 1,0 .................................................................................................................................... 14 Anillo anal con 18-20 ganchos por hilera; papilas rectales con 15-24 divertículos .................................................................................................................. Simulium canadense Anillo anal con 24-27 ganchos por hilera; papilas rectales con 36-45 divertículos ...................................................................................................................... Simulium ayrozai Largo del cuerpo 7,0 mm; abanico cefálico con 36-40 rayos .......... Simulium ethelae Largo del cuerpo 7,3-8,0 mm; abanico cefálico con 42-50 rayos .......................... 16 Largo del cuerpo 8,0 mm; anillo anal con 98-104 hileras de 24-30 ganchos cada una ............................................................................................................... Simulium estevezi Largo del cuerpo 7,5-7,7 mm; anillo anal con 115-145 hileras de 15-24 ganchos cada una ............................................................................................................................. 17 Anillo anal con 145 hileras con 17 ganchos cada una ...................... Simulium juarezi Anillo anal con 115-135 hileras con 20-24 ganchos cada una ............................ .................................................................................................................. Simulium capricorne

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JUAREZI species group Simulium (Hearlea) ayrozai VARGAS (Figs. 122 A, B.) Simulium ayrozai VARGAS, 1945a: 74. Simulium (Hearlea) ayrozai: VARGAS et al., 1946: 106, 111, 164, 181, 184, 185; VARGAS & DÍAZ NÁJERA, 1957b: 166, 167, 170, 172, 173, 282; DÍAZ NÁJERA & VULCANO, 1962: 93, 108; COSCARÓN et al., 2004: 10-11.

Female: Wing length 3.4 mm. General coloration blackish. Scutum velvety black, with 1+1 silvery vittae joining anterior subtriangular spots with whitish pruinose prescutellar area. Fronto-ocular triangle longer than wide. Cibarium broadly U-shaped, with smooth edge, lacking ovoid sublateral prominences. Sc pilose, with 21 hairs. Hind basitarsus with length/width ratio of 6:1; claw with subbasal tooth. Genitalia as in S. capricorne: sternite VIII with median sclerotized area and 17-19 trichomes per side, gonapophysis with internal edge smooth and sclerotized along inner 1/2; cerci with distal margin straight; Paraprocts well sclerotized, with abundant hair distally. Male: Wing length 3.4 mm. Scutum velvety black. Sc with few hairs. Hind basitarsus with length/width ratio = 4.2. Genitalia as in S. capricorne, although ventral plate more elevated distally. Pupa: (Figs. 122 A, B). Cocoon shoe shaped, anterior margin reinforced, coarsely woven; basal length 3.5-4.7 mm. Frontoclypeus and anterodorsal portion of thorax smooth, with sparse tubercles; frontoclypeus with 2 small frontal and 1 large facial setae per side. Gill with 2 wide, curved, pseudo annulated branches (particularly along inner surface) whose apices meet along longitudinal midline, forming ring; gill typically not extended beyond anterior margin of cocoon; dorsal branch 0.9 mm, with small acute process distally; ventral branch 1.0 mm. Abdomen without terminal hooks. Larva: Body length 8.0-10.0 mm. General coloration grey. Cephalic apotome uniformly light brown, gradually darkened mediobasally and small anteromedian and submedian spots. Cephalic fan with 52-54 rays. Hypostomium with straight anterior margin, 14-17 lateral setae per side, and 4+4 discal setae. Mandible with 2 rows of 7-8 internal teeth, 2 thin marginal teeth, and 1 large latero-mandibular process. Ratio of proximal, medial and distal antennal articles = 1:1.5:1.0. Ratio of length of hypostomium/hypostomial bridge = 1:1. Postgenal cleft dome-shaped, with narrow anteromedian incision. Cervical sclerites joining postgenae posteriorly. Lateral sclerite of thoracic proleg with 46-52 teeth. Anal ring with about 150 rows of 24-27 hooks. Rectal papillae of 3 compound lobes, with 12-15 diverticula per lobe; dorsal lobe larger than ventral ones. Distribution: Guatemala: Solalá, Totonicapan; México: México, Distrito Federal. Bionomics: Unknown. Discussion: Simulium microbranchium is very close structurally to S. ayrozai, especially in features of the pupal gill, but the former species has the gill branches narrower and slightly longer than the ventral branch and a thickened anterior margin on the cocoon. Simulium microbranchium differs from S. ayrozai slightly in the larva, such as more greenish coloration, a posterior circlet with 194-206 hooklet rows, and rectal papillae with 25 lobules.

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Simulium (Hearlea) burchi DALMAT (Figs. 123 A, B.) Simulium (Hearlea) burchi DALMAT, 1951: 44 (1): 42-47; 1955: 72, 81, 76, 82, 239-244; COSCARÓN et al., 2004: 11-12.

Female: Wing length 2.9-3.1 mm. General coloration brown-blackish. Frons and clypeus light brown with silver pollinosity; scutum velvety brown-blackish, with 1+1 silvery submedian vitta not joining anteriorly with 1+1 subquadrangular silvery pruinose spots and posteriorly with whitish pruinose prescutellar area; lateral and anterolateral edges with silver bands; scutum pilosity dense, golden to silvery greenish. Pleurae light grey-brown, with silver pollinosity. Legs yellowish brown, with apices of femora, tibiae, and tarsomeres dark brown. Wing veins pale brown. Abdomen brownish, with tergite II silvery; tergites VI-IX waxy. Cibarium concave medially, slightly indented. Sc pilose. Genitalia structurally as in S. capricorne. Male: Wing length 3.0-3.1 mm. Scutum velvety black edged by silvery band along periphery with some light covering of anterior 1/4 area. Legs as in female, but apices of femora, tibiae, and tarsomeres black. Sc without setae. Length/width ratio of hind basitarsus = 3.9. Genitalia with gonocoxite having rounded internal protuberance; gonostylus elongate, ratio of gonostylus length/gonocoxite length = 2.1; ventral plate with small median carina similar to that of S. delatorrei; endoparameres with numerous, strong hooks. Pupa: (Figs. 123 A, B). Cocoon shoe-shaped, with weak texture and reinforced anterior edge. Length basally 3.8 mm, dorsal length 2.8 mm. Frontoclypeus smooth, lacking granules. Thorax with moderate number of tubercles. Gill strongly annulated, especially dorsally, comprised of 3 primary branches; anterodorsal branch largest (length = 1.4-1.5 mm), annulated, distally capitate, with small tubercles dorsally and apically; ventrolateral branch medium-sized (length = 1.0 mm), flattened, with peculiar longitudinal crest along internal margin; posterodorsal branch small, 1/5 or less length of other branches. Tergite X without terminal hooks. Larva: Length (mature) 7.3 mm. General coloration grey to tan. Cephalic apotome with narrow dark median stripe basally. Cephalic fan with 46-54 primary rays. Antenna surpassing stalk of cephalic fan; ratio of proximal, medial and distal antennal articles = 1.0:1.5:1.0. Hypostomium with median and corner teeth longer than others, 14-15 lateral setae per side, and 3+3 discal setae. Postgenal cleft dome shaped, with narrow median anterior incision; ratio of length of hypostomium/hypostomial bridge = 1.1:1.0. Lateral plate of thoracic proleg with 47-50 setae. Anal ring with 172-178 rows of 20-24 hooks. Rectal papillae with about 78 diverticula, 22 on dorsal (central) lobe and 28 diverticula on each lateral lobe. Anal sclerite simple. Distribution: Guatemala: El Quiché, Huehuetenango. Bionomics: Simulium burchi occurs in medium-sized torrential creeks on stones or leaves and twigs. Discussion: This species resembles S. dalmati based on the pupal gill, but the gill of S. dalmati has a more uniform, only slightly distally expanded, dorsal branch and a longer posterior branch. Furthermore, the cocoon of S. burchi is boot-shaped and more coarsely woven. In females of S. dalmati, the 1+1 silver anterior spots are subtriangular and join

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vittae that extend further posteriorly. The original description of S. burchi states that the vittae join the anterior 1+1 subquadrangular spots, but in the two pinned paratypes from the USNM, the vittae do not fully join the anterior silvery spots. Larvae of S. dalmati differ from those of S. burchi by their larger size, longer hypostomial bridge (index of 0.8), and smaller number of diverticula on the rectal papillae. Simulium (Hearlea) canadense HEARLE (Fig. 122 C.) Simulium canadense HEARLE, 1932: 14 (as Simulium virgatum ssp.); 1932: 15. Simulium (Hearlea) canadense: VARGAS & DÍAZ NÁJERA, 1957b: 262-284; DÍAZ NÁJERA & VULCANO, 1962: 110; PETERSON, 1981: 381; PETERSON & KONDRATIEFF, 1995: 33-34; COSCARÓN et al., 2004: 12-14. Simulium (Hemicnetha) canadense: ADLER et al., 2004: 369-370.

Female: Wing length 3.4-3.8 mm. General coloration dark grayish. Scutum blackish grey, clothed in closely adpressed silvery to greenish pilosity, with longitudinal submedian silvery vittae joining with 1+1 subtrapezoidal silvery anterior spots (changing to black with opposite light), lateral and posterior areas silvery grey pollinose; scutellum and metanotum brownish black; pleurae grayish pollinose. Abdomen grayish black, with lateral margin of tergite II and posterior 1/5 of tergites III-V silvery grey pollinose; tergites IIIV blackish, subquadrate. Frons convergent below; fronto-ocular triangle about as wide as high. Sensory vesicle of palpus about 1/2 of basal palpomere length, with abundant tubercles. Mandible with 12-27 teeth. Lacinia with 12-14 retrorse teeth. Basal portion of cibarium thick, smooth, medially concave, with 1+1 ovoid lateral prominences. Sc with 13 setae. Length/width ratio of hind basitarsus 5.7-6.4. Sternite VIII darkened medially, with about 14-20 hairs per side. Gonapophysis subtriangular, concave internally and blunt distally; cercus curved distally, paraprocts subrectangular, with relatively abundant hair internally and transverse sulcus subbasally; genital fork with median stem capitate basally and with lateral arms wide, bearing stout anteriorly directed apodemes; spermatheca ovoid, homogeneously sclerotized. Male: Wing length 3.1-3.3 mm. Scutum black, with anterior 1/3 light grayish pollinose, separated by black area and surrounding 1+1 subtriangular silvery spots. Length/width ratio of hind basitarsus = 3.8. Gonostylus about 1.8 times gonocoxite length, with small internally directed basal process; gonocoxite elevated laterally; ventral plate with median carina; endoparamere with strong hooks. Pupa: Cocoon slipper shaped, coarsely woven, reinforced along anterior edge. Gill membranous with 3 subsegmented branches; anterodorsal branch largest, blunt apically, ventral branch directed anteriorly, roughly 1/2 width of dorsal branch and digitiform distally (Fig. 122 C); posterior branch 1/4 length of other branches, curved posteromedially. Frontoclypeus and thorax with abundant small tubercles. Cephalic and thoracic trichomes single, stout. Abdomen with spines reduced. Larva: Length (mature) 8.0-9.0 mm. Cephalic apotome without ornamentation, head ventrally darkened basomedially. Cephalic fan with 48-56 primary rays. Ratio of length of hypostomium/hypostomial bridge = 1:1. Hypostomium with anterior margin straight, median tooth longer than corner teeth, 11-15 lateral setae per side, and 2-4 discal setae.

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Postgenal cleft dome shaped, with anterior incision. Mandible with internal teeth arranged in 2-3 rows of 5-8 teeth, 2 thickened marginal teeth, and 1 large latero-mandibular process. Ratio of proximal, medial and distal antennal articles = 1:1.3-1.4:1.0-1.2; medial article with 3-4 subdivisions. Lateral sclerite of thoracic proleg with 24 teeth. Anal sclerite with hair among struts. Anal ring with 102-158 rows of 18-20 hooks. Rectal papillae with 15-24 diverticula (5-8 per lobe), dorsal diverticulum of each lobe greatly enlarged. Distribution: Canada: British Columbia; USA: Arizona, California, Colorado, Florida, Idaho, Montana, Nevada, New Mexico, Oregon, South Dakota, Utah, Washington, Wyoming; Mexico: Distrito Federal, Durango, Mexico, Oaxaca, Puebla, Veracruz. Bionomics: Larvae are found on vegetation in cool, clear streams. Discussion: This species is structurally similar to S. burchi and S. dalmati but the former species has a shoe-shaped cocoon and a more strongly capitate, tuberculed, anterior gill branch; the latter species has both major gill branches multilobed apically. Simulium (Hearlea) capricorne DE LEÓN (Figs. 120 D-J; 121 B; 122 D, E; 126 A-C; 127 A.) Simulium capricornis DE LEÓN, 1945: 71-72. Simulium (Hearlea) capricornis: DALMAT, 1950: 72, 76, 82, 86; l955, 244-249; COSCARÓN, 1987: 34; COSCARÓN et al., 2004: l4-l6. Simulium (Simulium) deleoni VARGAS, 1945b: 72. Simulium (Hearlea) deleoni: VARGAS & DÍAZ NÁJERA, 1957b: 294; VARGAS et al., 1946: 182.

Female: Wing length 2.7-2.8 mm. General coloration brownish dark. Head blackish, eyes dark brown, frons and clypeus brown, with silver pruinosity; scape and pedicel light brown, flagellum dark grayish brown, palpus and proboscis dark brown to blackish; palpus with long blackish hair. Scutum dark brown to black, with faint grayish pollinosity, uniformly distributed recumbent hair, and 1+1 anterior submedial, subtrapezoidal black spots not joined to 1+1 silvery vittae; prescutellar area and lateral edges of scutellum with abundant, long golden hair, pleurae brown-grey pollinose; metanotum brown, silvery tomentose. Wing veins light brown. Legs light brown, with darkened apices on femur, tibiae and tarsomeres, tarsomeres of fore leg black. Abdomen dark brown; tergites II-V with dark brown median spots, tergite II with 1+1 silvery spots. Frons convergent below (Fig. 120 D); fronto-ocular triangle wider than high (Fig. 120 E). Basal portion of cibarium slightly wrinkled along median edge, with 1+1 submedian, subovoid prominences (Fig. 120 G). Mandible with 11+27 teeth. Lacinia with 13+17 retrorse teeth. Sensory vesicle of maxillary palpus slightly shorter than length of basal palpomere. Basal sector of R without hair; Sc with about 15 hairs. Length/width ratio of hind basitarsus = 5.2:1.0 (Fig. 120 F). Sternite VIII well sclerotized, with about 14 hairs per side; gonapophysis with distal edge externally curved and medially directed apices (Fig. 120 H). Cercus flattened distally. Paraproct subrectangular, with internal edge curved and with small longitudinally positioned crest (Fig. 120 I); genital fork (Fig. 120 J) with median stem long, distally capitate and lateral arms only moderately expanded apically. Male: Wing length 2.6-2.8 mm. General coloration same as for female but slightly darker. Scutum velvety black, with anterior 1+1 wide silver pruinose spot continuous with thin lateral silvery stripe that joins silver prescutellar area. Legs darkened, especially

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H P Figure 120. Female. A-B: S. (Hearlea) juarezi; A: scutum with front illumination; B: scutum with posterior illumination. C: S. (H.) delatorrei, cibarium. D-J: S. (H.) capricorne; D: frons; E: fronto-ocular triangle; F: hind basitarsus; G: cibarium; H: eighth sternite and gonapophysis; I: cercus and paraproct; J: genital fork. K-L: S. (H.) microbranchium; K: scutum with frontal illumination; L: scutum with posterior illumination. M-P: S. (H.) larvispinosum; M: cibarium; N: claw; O: genital fork; P: spermatheca.

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Figure 121. A-G: Male; A: S. (Hearlea) juarezi, scutum; B: S. (H.) capricorne, hind basitarsus; C-E: S. (H.) delatorrei; C: gonocoxite and gonostylus; D: ventral plate; E: endoparameres and median sclerite; F-G: S. (H.) larvispinosum; F: gonocoxite and gonostylus; G: ventral plate. H-I: Pupa, frontoclypeus of S. (H.) capricorne and S. (H.) delatorrei, respectively.

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Figure 122. Pupa and gill. A-B: S. (Hearlea) ayrozai; A: lateral view; B dorsal view. C: S. (H.) canadense. D-E: S. (H.) capricorne; D: lateral view; E: ventral view. F-I: S. (H.) ethelae; F: lateral view; G: lateral view; H: dorsal view; I: lateral internal view. J-K: S. (H.) microbranchium; J: lateral view; K: dorsal view. L-N: S. (H.) nigricorne; L: lateral view; M: latero internal view; N: antero dorsal view.

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prothoracic and mesothoracic coxae and most of metathoracic leg. Abdomen dark brown, with 1+1 silvery spots on tergite II. Sc bare. Hind basitarsus wide, length/width ratio = 3.03.2 (Fig. 121 B). Gonocoxite subquadrate, with posterior prominence. Gonostylus elongate, about twice as long as gonocoxite, with internal row of setae basally, and stout subapical spur; ventral plate with sinuous posterior margin and short longitudinal median carina. Pupa: (Figs. 122 D, E). Cocoon slipper shaped, coarsely woven, reinforced along anterior edge; basal length 3.5-4.3 mm; dorsal length 3.3-4.1 mm. Frontoclypeus with only lateral and ventral edges having few smooth apically rounded tubercles (Fig. 120 H) and 2+2 frontal and 1+1 facial simple trichomes. Anterodorsum of thorax with scarce to abundant acuminate granules and 5 stout, simple trichomes per side. Gill with 2 main branches, dorsal and ventral; branches thickened, subannulated, dorsoventrally flattened, strongly curved apically; dorsal branch largest, with strongly downturned apex; ventral branch curved along contour of cocoon. Maximum length of branch = 2.2 mm. Abdomen with spine-combs on tergites VII and VIII. Terminal spine absent. Larva: Length 7.3-7.7 mm. Coloration yellowish (in alcohol). Head light brown; cephalic apotome brownish and slightly darkened basally, without special ornamentation. Cephalic fan with 42-47 rays. Hypostomium with straight anterior margin, median tooth longer than corner teeth (Fig. 126 B), 8-10 lateral setae per side, and 1+1 to absent discal setae. Hypostomial bridge longer than hypostomium, ratio of length of hypostomium/ hypostomial bridge = 0.8-1. Postgenal cleft mildly curved, without apical incision (Fig. 126 A). Antenna longer than stalk of cephalic fan; medial article annulated; ratio of proximal, medial and distal articles = 1:1.1-1.5:1.0-1.4. Mandible with 2 rows of 4-5 internal teeth, 2 marginal teeth, and 1 curved lateral mandibular process (Fig. 126 C). Lateral sclerite of thoracic proleg with about 38 teeth arranged in 12-14 groups. Anal sclerite with sparse hair, ventral branches relatively elongate (Fig. 127 A). Anal ring with 115-135 rows of 20-24 hooks. Rectal papillae trilobate, and 4-6 diverticula per lobe. Distribution: Guatemala: Alta Verapaz, Chimaltenango, El Quiché, Guatemala, Huehuetenango, Sacatepequez, Sololá, Totonicapán; Mexico: Chiapas, Distrito Federal, Durango, Mexico, Morelos, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: The closest species to S. capricorne is S. estevezi, which can be differentiated by the absence of an anterior collar on the cocoon, presence of a process on the dorsal gill branch, and a cibarium with a thickened anterior edge; and S. burchi, which has a wide apically blunt dorsal branch, as does S. canadense. Pupal gill morphology, as used in the accompanying key, permits separation of S. capricorne from structurally similar species such as S. ethelae, S. dalmati and S. microbranchium. Simulium (Hearlea) chiriquiense FIELD Simulium (Hearlea) chiriquiense FIELD, 1967b: 194-199; VARGAS et al., 1977: 137; COSCARÓN et al., 2004: 16-17.

It was not possible to examine topotypical material of this species. Based on morphology of the pupal gill in the original description, it could be considered conspecific with S. ethelae DALMAT. Material from Costa Rica identified as S. chiriquiense by VARGAS et al. is similar to that

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of S. capricorne, based on illustrations of the pupa (Figs. 3, 4). Genitalia of adults cannot be reliably differentiated from those of several other Simulium (Hearlea) species. Distribution: Panama: Cerro Punta (Boquete), Chiriqui; Costa Rica: Vicinity of San José. Bionomics: Unknown. Simulium (Hearlea) contrerense DÍAZ NÁJERA & VULCANO Simulium (Hearlea) contrerense DÍAZ NÁJERA & VULCANO, 1962: 94-99; COSCARÓN et al., 2004: 17.

(Description based on DÍAZ NÁJERA & VULCANO, 1962). Female: General coloration blackish. Scutum velvety blackish, with 1+1 silvery vittae not joining anteriorly with 1+1 submedian silvery triangular spots; lateral and posterior edges silvery pruinose. Abdomen brownish, with posterior edge of tergites yellowish. Basal portion of cibarium concave medially, without wrinkled margin and thick rim. Genitalia similar to those of S. capricorne. Male: Wing length 5.0 mm. General coloration blackish. Scutum velvety black, bordered with whitish pruinosity and clothed in golden pile. Scutellum brown, with light brown posterior edge. Abdominal tergites black, with yellowish hair. Sc haired. Gonocoxite, gonostylus and endoparameres typical for subgenus (as in S. capricorne); ventral plate with prominent median carina. Pupa: Length (basal) 7.0 mm; dorsal 6.0 mm; gill 3.5 mm. Cocoon shoe shaped, gradually elevated, closed basally on anterior 1/4, compactly woven, not reinforced along anterior margin. Thorax with 5 simple trichomes per side. Gill inflated, with 2 major branches frontally and 1 small ventrolateral branch; dorsal branch stout, subglobulose, tapered distally, sclerotized and wrinkled dorsally, ventral branch membranous and translucent ventrally, with subconical lateral prominence, and subannulations; ventrolateral branch small, thin, and curved medially. Larva: Length 12.0 mm. Head brownish, body yellowish grey, without ventral tubercles. Cephalic fan with 42 primary rays. Hypostoma with 15-17 lateral setae per side and 4+4 discal setae. Mandible with 6-7 internal and 2 marginal teeth. Ratio of proximal, medial and distal antennal articles = 1:2.2:1.9. Postgenal cleft subtriangular. Anal ring with 185 rows of about 16 hooks. Rectal papillae with 60-66 diverticula, the dorsal lobe largest. Distribution: Mexico: Distrito Federal., Mexico (3000 m). Bionomics: Unknown. Discussion: The unique gill, with its inflated dorsal and ventral branches, membranous and translucent ventrally, distinguishes this species from all others. Simulium juarezi has a similar gill, but only its dorsal branch is markedly inflated. Simulium (Hearlea) dalmati VARGAS & DÍAZ NÁJERA (Figs. 123 J-M.) Simulium (Hearlea) dalmati VARGAS & DÍAZ NÁJERA, 1948b: 337-341; 1957b: 166, 169, 170, 171, 173, 290; DÍAZ NÁJERA & VULCANO, 1962: 93, 122; COSCARÓN et al., 2004: 18.

Female: (Diagnosis of female based on VARGAS & DÍAZ NÁJERA, 1948b). Wing length 3.5 mm. General coloration blackish, with golden pilosity. Scutum black with

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N O Figure 123. Pupa and gill. A-B: S. (Hearlea.) burchi; A: lateral view; B: dorsal view. C-F: S. (H.) delatorrei; C: lateral view; D: dorsolateral view; E: dorsal view; F: lateral internal view. G-I: S. (H.) estevezi; G: lateral view; H: dorsal view; I: ventral view. J-M: S. (H.) dalmati; J: lateral view; K: dorsal view; L: frontal view; M: ventral view. N-O: S. (H.) juarezi; N: lateral view; O: dorsal view.

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1+1 silvery vittae, joining anteriorly with 1+1 silvery subtriangular spots. Legs yellowish, with apices of femora, tibiae, and tarsomeres black. Basal portion of cibarium smooth, with thickened anterior margin gradually elevated sublaterally. Sc haired. Genitalia as in S. capricorne. Male: Wing length 3.0 mm. Coloration similar to that of female. Scutum black, with grayish pruinosity on anterior 1/3 interrupted medially. Sc with scarce hair. Length/width ratio of hind basitarsus = 3.6. Ventral plate with short, haired, median carina. Pupa: (Fig. 123 J). Cocoon slipper shaped coarsely woven, anterior edge reinforced. Length at base 5.0 mm. Gill (Figs. 123 K-M) inflated, pseudosegmented, comprised of 3 branches; dorsal and ventral branches subequal in size; posterior branch 1/4 or less length of other branches; dorsal branch 1.4 mm, subcylindrical, capitate, directed anterolaterally, with 3-4 small apical protuberances (Fig. 123 K); ventral branch 1.3 mm, laterally flattened, blunt apically, curved anteromedially, with 1 or more small apical protuberances (Fig. 123 L); posterior branch 0.4 mm, acuminate, directed posteromedially (Fig. 123 K). Frontoclypeus and thorax with sparse and dissimilarly sized tubercles. Larva: Length 9.0 mm. Coloration grayish green; cephalic apotome without head spots, slightly darkened mediobasally. Cephalic fan with 42-44 primary rays. Antenna as long as stalk of cephalic fan, median article with 2 whitish subannulations. Mandible with strong curvature on superior side, with 6 internal and 2 marginal teeth, the last well developed; ratio of proximal, medial and distal antennal articles = 1:1.3-1.4:1.0-1.1. Postgenal cleft gradually curved, with narrow apical incision. Hypostoma with straight anterior margin, 11-13 lateral setae per side, and 3+3 discal setae; median tooth longer than others; ratio of hypostomium/hypostomial bridge = 0.8. Anal ring with 144 rows of 21-23 hooks each. Rectal papillae with 18-24 diverticula, consisting of large dorsal and 57 smaller ventral diverticula per lobe. Distribution: Mexico: Veracruz. Bionomics: Unknown. Discussion: The female is very similar to those of S. delatorrei and S. larvispinosum. The pupa is structurally close to S. canadense and S. burchi, but S. canadense has a slippershaped cocoon and S. burchi has a more anteriorly erect cocoon with a finer textured weave. The larva is similar to that of S. ethelae, but has 36-39 lobules on the rectal papillae. The larva of S. delatorrei is also similar, but it has 114 rows of hooks on the posterior circlet and 7-9 lateral hypostomial setae per side. Simulium (Hearlea) delatorrei DALMAT (Figs. 120 C; 121 C-E, I; 123 C-F.) Simulium (Dyarella) delatorrei DALMAT, 1950: 137-147. Simulium (Hearlea) delatorrei: DALMAT, 1951: 53; 1955: 253-258; VARGAS & DÍAZ NÁJERA, 1957b: 170, 172, 174; DÍAZ NÁJERA & VULCANO, 1962: 93; COSCARÓN et al., 2004: 19-20.

Female: Wing length 3.1 mm. General coloration dark brown, clothed with yellow to greenish pile, very similar to S. capricorne; scutum with 1+1 submedian silvery vittae extending from subtriangular anterior silvery spots to silvery pollinose prescutellar area, changing from whitish silver to black depending on lighting. Frons convergent below, fronto-ocular triangle about as high as wide. Basal portion of cibarium smooth, with thin anterior edge and small

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median concavity (Fig. 120 C). Sensory vesicle of maxillary palpus with abundant tubercles, about 1/2 of basal article length. Genitalia similar to those of S. capricorne. Male: Wing length 3.0 mm. General coloration similar to that of female. Eyes dark brown; scutum dark brown, bordered by silver pruinosity with anterior and posterior areas wider than lateral areas, scutum with abundant closely appressed silvery pilosity; (in specimens without pilosity, light position enables vittae to be seen on grey tomentous pruinosity); pleurae brown with silver pollinosity. Legs brownish, with long yellowish brown hair. Abdomen brown blackish, with 1+1 silver spots laterally on tergites VI-VIII. Length/width ratio of hind basitarsus = 3.3. Genitalia as in figures 121 C-E. Pupa: Cocoon slipper shaped, closed anteriorly, compactly woven, threads not evident, reinforced on anterior edge (Figs. 123 C-F). Length (basal) 3.2-3.3 mm; (dorsal) 2.5-2.8 mm. Frontoclypeus with abundant, small tubercles laterally, continued internally by microscopic punctation, with 2+2 thin frontal and 1+1 stout and spine-like simple facial trichomes (Fig. 121 I). Thorax dorsally with abundant and minute tubercles more developed and conical along midline and with 5 spine-like trichomes per side. Gill 1.8 mm, comprised of 3 branches; filaments stout, gradually tapered, pseudosegmented; dorsal branch largest, subcylindrical, apically blunt, directed anteriorly, with noticeable ventrolateral protuberance submedially; ventral branch 1/2 width of dorsal branch, curved medially; posterior branch smallest, directed posteromedially. Abdomen lacking terminal hooks. Larva (description based on DALMAT, 1955): Body length 7.0 mm. General color grey, cephalic apotome darkened mediobasally. Cephalic fan with 39-40 primary rays. Postgenal cleft dome shaped, with deep anteromedian incision. Hypostomium with anterior margin straight and 7-9 lateral setae per side. Ratio of hypostomium/hypostomial bridge = 1.1. Mandible with 6 internal teeth disposed in 3 rows, 2 marginal teeth, second slightly longer than 1/2 length of first. Lateral sclerite of thoracic proleg with 40-42 comb teeth. Anal ring with 114 rows of 19-22 hooks each. Rectal papillae with 72-75 diverticula, 24-25 per lobe. Distribution: Guatemala: Chimaltenango, El Quiché, Guatemala, Huehuetenango, Quezaltenango, San Marcos, Sololá, Totonicapán; Mexico: Chiapas. Bionomics: Unknown. Discussion: The closest species is S. estevezi but differences in the pupa and larva enable their separation. The Díaz & Vulcano (1962) illustration of larvae from Mexico shows that the anal papillae have only 5 lobules per lobe and 8 lateral hypostomial setae, differing from Guatemalan populations. Simulium (Hearlea) estevezi VARGAS (Figs. 123 G-I.) Simulium (Hearlea) estevezi VARGAS, 1945b: 71; VARGAS & DÍAZ NÁJERA, 1948b: 341; DÍAZ NÁJERA & VULCANO, 1962: 92; COSCARÓN et al., 2004: l9-20.

(Diagnosis of female and male based on VARGAS, 1945b and VARGAS & DÍAZ NÁJERA, 1957b). Female: Wing length 2.5 mm. Scutum blackish, with 1+1 silvery vittae joined with anterior subtriangular silvery spots. Cibarium without ovoid sublateral prominences. Genitalia structure similar to that of S. capricorne.

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Male: Wing length 2.7 mm. Genitalia structurally similar to that of S. capricorne. Ventral plate with haired median carina. Pupa: Length (basal) 3.6 mm; (dorsal) 2.8 mm. Cocoon shoe shaped, compactly woven, threads not evident, slightly reinforced anterior edge. Frontoclypeus covered by platelets; thorax with abundant tubercles. Gill with 3 acuminate, pseudosegmented branches; dorsal branch largest, subcylindrical, directed anteroventrally, with median, internally directed prominence, and sharply upturned acuminate apex; ventral branch laterally flattened, curved ventromedially; posterior branch 1/3 or less length of other branches, directed posteromedially (Figs. 123 G-I). Larva: Length 8.0 mm. General coloration grayish. Cephalic apotome uniformly yellowish colored, or with slightly darkened spots forming cross. Antenna extended anteriorly beyond apex of cephalic fan stalk; ratio of proximal, medial and distal articles = 1:1.3-1.4:1.1-1.2; medial article appearing subsegmented into 3 pieces. Mandible with 2 rows of 7 internal teeth, and 2 internal teeth, the anterior one larger. Cephalic fan with 4450 primary rays. Postgenal cleft gradually narrowed apically, with small apical incision. Hypostoma with anterior margin straight. Ratio of hypostomium/hypostomial bridge = 0.9. Hypostomium with 8 lateral setae per side and 2+2 discal setae. Lateral plate of thoracic proleg sclerite with about 40 setae. Anal ring with 98-104 rows of 18-19 hooks each. Rectal papillae with 24-30 diverticula, 8-10 per lobe. Distribution: Mexico: Chiapas, Distrito Federal, Guerrero, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Externally, the female is very close to those of S. ayrozai, S. dalmati and S. ethelae. The pupa of these species, however, is quite distinct. The pupa of S. estevezi is closest to that of S. delatorrei, but it has the dorsal gill branch thicker and blunt apically, the medioventral process smaller, and a more basally flattened cocoon. The larva of S. estevezi is closest to that of S. delatorrei, but the anal ring has 114 rows of hooks, the rectal papillae have 24-25 diverticula per lobe, and the postgenal cleft is significantly shorter. Simulium (Hearlea) ethelae DALMAT (Figs. 122 F-I.) Simulium (Hearlea) ethelae DALMAT, 1950: 143-148; 1951: 52-53; 1955: 258-263; VARGAS & DÍAZ NÁJERA, 1957b: 167, 171, 174, 298 (as Simulium capricorne); DÍAZ NÁJERA & VULCANO, 1962: 93; COSCARÓN et al., 2004: 21-22.

Female: Wing length 3.6 mm. Scutum velvety black with 1+1 anterior silvery vittae joining 1+1 anterior silver pruinose subtriangular spots and light grayish prescutellar area. Basal portion of cibarium smooth, concave medially, with thickened margin bearing 1+1 lateral subovoidal prominences. Length/width ratio of hind basitarsus = 4.7:1.0. Genitalia as in S. capricorne. Male: Wing length 3.4 mm. Coloration of scutum velvety blackish with silvery pruinose spot on anterior 1/4; abdominal ornamentation and genitalia morphology similar to those of other S. (Hearlea) species, with ventral plate subovoidal with median carina. Length/width ratio of hind basitarsus = 3.3. Pupa: Length (basal) 2.4 mm. Cocoon slipper-shaped, closed anteriorly, compactly woven, threads not evident, with anterior reinforcement. Gill with 2, subequally sized,

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wrinkled, curved, gradually flattened, pseudosegmented, acuminate branches lacking granules or spicules (Figs. 122 F-I); dorsal branch slightly longer than ventral branch, with apex curled ventrally. Thorax with sparse acuminate tubercles; frontoclypeus mostly smooth, with rounded tubercles only along margins. Larva: Length 7.0 mm. General coloration grayish. Cephalic apotome with only small spots on midline and submedian area. Cephalic fan with 36-40 primary rays. Mandible with 5 or 6 internal teeth and 2 thin marginal teeth. Hypostoma with 12-16 lateral setae per side. Postgenal cleft narrow distally, deeply incised; ratio of hypostomium/hypostomial bridge = 0.9. Lateral sclerite of thoracic proleg with 36-42 teeth. Rectal papillae with 36-39 diverticula, 12-13 diverticula per lobe. Anal ring with 150-170 rows of 18-20 hooks each. Distribution: Guatemala: Chimaltenango, Guatemala, Huehuetenango, Quezaltenango, Solalá; Mexico: Chiapas. Bionomics: Unknown. Discussion: The closest species morphologically is S. capricorne, but the dorsal gill branch is more flattened and apically blunt and has a median crest and a small protuberance dorsobasally. The female differs in that the scutal vittae do not join the anterior subtrapezoidal spots. Simulium chiriquiense has an acuminate dorsal gill branch, frontoclypeus with numerous tubercles, and wing length (female 3.8 mm and male 3.0 mm) more similar to that of S. ethelae. Simulium (Hearlea) juarezi VARGAS & DÍAZ NÁJERA (Figs. 120 A, B; 121 A; 123 N, O.) Simulium (Hearlea) juarezi VARGAS & DÍAZ NÁJERA, 1957b: 205, 262, 304; DÍAZ NÁJERA & VULCANO, 1962: 100; COSCARÓN et al., 2004: 22-23.

Female: Wing length 3.3-3.8 mm. General coloration blackish brown. Head brown, with frons and clypeus grey pollinose. Scape and pedicel light brown; flagellum dark brown. Palpus and proboscis dark brown. Scutum brown, with grayish pruinosity, and yellowish-green to brown decumbent hair varying in color with light position. Scutum with 1+1 submedian vittae that are silver whitish with posterior lighting and blackish with anterior lighting, extending from 1+1 subquadrate anterior spots to prescutellar area; median blackish stripe about 3 times wider than silvery submedian stripes (Figs. 120 A, B); humeral spots light grey pollinose as are narrow lateral edges and prescutellar area; abdomen dark brown. Legs yellowish brown, with apices of femora, tibiae, and tarsomeres blackish. Basal portion of cibarium smooth, with thickened rim, and 1+1 subovoidal sublateral prominences. Genitalia similar to those of S. capricorne. Male: Wing length 3.2-3.6 mm. General coloration as in female but more darkened. Scutum velvety blackish, with anterior 1/3 grayish pollinose and separated medially by blackish area (Fig. 121 A). Legs mostly dark brownish, with closely adpressed yellowish brown short hair and blackish long hair. Length/width ratio of hind basitarsus = 4.0. Genitalia structurally similar to that of S. capricorne. Pupa: Cocoon slipper-shaped, closed anteroventrally, with reinforced anterior edge, compactly woven, frequently covered by sand particles. Length (basal) 3.4-4.0 mm; (dorsal) 3.2-3.5 mm. Frontoclypeus and anterodorsum of thorax with abundant, rounded,

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variably sized tubercles. Gill with 3 semi-membranous branches; dorsal branch largest, strongly clavate, with numerous tubercles dorsally and annulations basally; ventral branch laterally flattened, annulated, strongly curved medially; posterodorsal branch acuminate, subannulated, only slightly shorter than ventral branch, and directed dorsomedially. (Figs. 123 N, O). Larva (diagnosis based on DÍAZ NÁJERA & VULCANO, 1962): Length 7.5 mm. General coloration greenish dorsally and yellowish ventroapically. Cephalic apotome darkened basally and medially. Cephalic fan with 50 primary rays. Hypostoma with 8-10 lateral setae per side. Postgenal cleft dome shaped, narrowed distally. Anal ring with about 145 rows of 17 hooks. Rectal papillae with 5-7 diverticula per lobe, and median anterior diverticulum larger than others. Distribution: México: Chiapas, México, Michoacán, Oaxaca, Veracruz. Bionomics: Unknown. Discussion: The peculiar respiratory organ with its large, subglobose, superficially granulose dorsal branch distinguishes this species. The closest species with inflated gill branches is S. contrerense. Simulium (Hearlea) microbranchium DALMAT (Figs. 120 K, L; 122 J, K.) Simulium (Simulium) microbranchium DALMAT, 1949: 538-544. Simulium (Hearlea) microbranchium DALMAT, 1951: 50-52; 1955: 72, 76, 82, 87, 268-272; COSCARÓN et al., 2004: 23-24.

Female: Wing length 3.4 mm. Frons and clypeus brownish-grey silvery pollinose. Thorax brownish black. Scutum brownish dark, with silver submedian vittae not joining anterior 1+1 silvery subtrapezoidal spots, but joining silvery pollinose prescutellar area; area between these spots black, although changing light position changes this silver color to black (Figs. 120 K, L). Scutum with golden, closely adpressed pilosity. Shape and proportion of palpal sensory vesicle as in S. capricorne. Cibarium with median portion concave, hyaline, with serrated margins. Sc haired. Length/width ratio of hind basitarsus = 5.2. Genitalia structurally similar to those of S. capricorne. Male: Wing length 3.4 mm. Scutum velvety black with whitish pruinosity anteriorly and bordered with narrow silvery band on periphery. Scutum pilosity golden. Sc haired. Genitalia typical for subgenus; ventral plate with small median crest. Gonostylus about twice length of gonocoxite height at area of insertion; gonocoxite externally 1/3 longer than internal height. Pupa: Cocoon weakly shoe-shaped, closed and slightly enlarged anterobasally, with reinforced edge (Fig. 122 J). Length (basal) 3.5 mm. Frontoclypeus and anterodorsum of thorax with sparse tubercles laterally around base of frontoclypeus, and sparse on anterior area of thorax. Gill comprised of 2 laterally flattened, dorsally pseudosegmented, acuminate branches (Fig. 122 K); dorsal branch (length = 1.6 mm) directed anteriorly, forming about 90° angle with gill base; ventral branch (length = 0.9 mm) only slightly medially directed. Larva (diagnosis based on DALMAT, 1951): Body length 7.7 mm. General coloration grey. Cephalic apotome concolorous, with small black spots along midline and submedian

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anterior area. Cephalic fan with 56 or 57 primary rays. Hypostoma with median tooth and corner teeth longer than other teeth. Postgenal cleft curved with narrow anteromedian incision. Lateral sclerite of thoracic proleg with 37-39 teeth. Anal ring with 194-206 rows of hooks. Rectal papillae with about 75 diverticula, about 25 per lobe. Anal sclerite simple, lacking scales. Distribution: Guatemala: El Quiché, Huehuetenango, Quetzaltenango, Sololá, Totonicapán. Bionomics: Unknown. Discussion: This species is very similar to S. ayrozai, but its dorsal gill branch is projected anteriorly well beyond the anterior margin of the cocoon. Simulium (Hearlea) nigricorne DALMAT (Figs. 122 L-N.) Simulium (Hearlea) nigricornis DALMAT, 1950: 148-151; 1955: 76, 82, 273-275; COSCARÓN et al., 2004: 24-25.

(Diagnosis of female and male based on DALMAT, 1955). Female: General coloration unknown. Cibarium with definite indentation on internal edge. Cerci about 2 times wider than high with curved edge; paraproct with abundant hair distally; genital fork with median rod thin, elongate. Male: General coloration unknown. Gonocoxite with strong prominence basally along internal edge; ventral plate with small median carina. Pupa: (Figs. 122 L-N). Cocoon shoe-shaped, with anterior edge reinforced; length (basal) 2.9 mm; maximum 3.6 mm. Frontoclypeus with sparse tubercles. Thorax with tubercles. Gill extended well beyond anterior margin of cocoon, comprised of 2 subannulated, flattened branches; dorsal branch 0.9 mm, darkened on distal 3/4, distended apically, with blunt apex; ventral branch 1.5 mm, curved, elongate, thinning distally, with small projection distally. Abdominal terminal hooks absent. Larva: Unknown. Distribution: Guatemala: Chimaltenango. Bionomics: Unknown. Discussion: The presence of two annulated branches places S. ayrozai, S. ethelae and S. microbranchium close to S. nigricorne, but the short, wide, distally darkened dorsal branch permits separation. CAROLINAE species group Simulium (Hearlea) carolinae DE LEÓN Simulium carolinae DE LEÓN, 1945: 73-75. Simulium (Hearlea) carolinae: VARGAS et al., 1946: 122-124, 182; DALMAT, 1955: 72, 76, 82, 86, 249-253; DÍAZ NÁJERA & VULCANO, 1962: 93, 112; COSCARÓN et al., 2004: 25-26.

Female: Wing length 3.7-3.9 mm. Coloration blackish brown grayish with vittae not joining 1+1 anterior subquadrate silvery spots, pilosity adpressed, abundant, tin colored; pleurae light grayish, silvery pollinose; legs yellowish grayish with concolorous hair, with

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apices of femur, tibia, hind basitarsus, and tarsite blackish. Cibarium smooth, concave medially, with 1+1 sublateral ovoidal prominences. Sc haired. Length/width ratio of hind basitarsus = 4.8. Genitalia structurally similar to those of S. larvispinosum. Male: Wing length 3.2-3.6 mm. Scutum velvety black, homogeneously pigmented, or with light area on anterior 1/5, abundant pilosity tin to golden. Legs generally darkened. Length/width ratio of hind basitarsus = 2.8. Genitalia similar to those of S. larvispinosum. Gonocoxite slightly elevated laterally; ventral plate with large median carina. Pupa: Cocoon slipper shaped, reinforced along anterior edge. Length (basally) 3.0 mm. Thorax covered with numerous tubercles. Gill rigid, similar to that of S. larvispinosum, except main dorsal branch thicker and blunter apically, secondary branches relatively short, and ventral branch strongly curved. Larva: Length 7.8 mm. General coloration light grayish brown. Cephalic apotome darkened slightly along midline basally. Cephalic fan with 42-44 primary rays. Mandible with preapical teeth well developed, 5-6 internal teeth arrayed in 2 rows; marginal teeth with 2 denticles well developed and 1 accessory at half of internal teeth position. Ratio of proximal, medial and distal antennal articles = 1:1.6-1.8:1.1-1.2. Hypostomium with prominent median tooth and 11-15 lateral setae per side. Postgenal cleft dome shaped, only slightly acuminate distally, without deep anteromedian incision. Lateral sclerite of thoracic proleg with 24 teeth in 7-8 groups. Anal sclerite with ventral branches encircling posterior circlet. Dorsolaterally on posterior portion of abdomen there are 2+2 plates flattened and conical, very sclerotized and covered by acuminate tubercles and some spines with wide base. Anal ring with 190-200 rows of 25-27 hooks each. Rectal papillae with 24-33 diverticula, 8-11 diverticula per lobe. Distribution: Mexico: Chiapas, Oaxaca, Veracruz; Guatemala: Alta Verapaz, Chimaltenango, Guatemala, Quezaltenango, Sololá, Suchitepequez. Bionomics: Unknown. Discussion: The form of the pupal gill and absence of 1+1 ventrolateral papillae on the eighth segment of the larva is shared with S. larvispinosum, but the latter has 1+1 accessory ventrolateral accessory plates on the anal sclerite. Simulium (Hearlea) gorirossiae VARGAS & DÍAZ NÁJERA (Figs. 124 F, G; 126 D-G; 127 D-F.) Simulium (Hearlea) gorirossii VARGAS & DÍAZ NÁJERA, 1957b: 199-202; DÍAZ NÁJERA & VULCANO, 1962: 92; COSCARÓN et al., 2004: 26-28 (incorrect original spelling).

Female: Wing length 3.2 mm. Coloration blackish. Head dark brown, with frons and clypeus silvery pollinose. Scutum velvety black with 1+1 silvery vittae joined with submedian anterior subtriangular silvery spots; laterally bordered by narrow whitish grey pruinose band joining whitish prescutellar area. Abdomen blackish. Fronto-ocular triangle about as long as wide; cibarium with basal portion as in S. larvispinosum, with hyaline edge and 1+1 small rounded submedian processes. Sc bare. Genitalia typical for subgenus, with cercus rectangular, flattened distally; paraprocts with a transverse tegumentary fold and very haired apically; gonapophysis subtriangular, with blunt apex similar to that of S. larvispinosum. Male: Wing length 3.0 mm. Scutum velvety black with anterior 1/3 pruinose, continued by narrow band edging laterally and meeting with whitish grey pruinose

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A C

D

E

B

0.5 mm

H

I F

G

K

J

Figure 124. Pupa and gill. A-E: S. (Hearlea) larvispinosum; A: lateral view; B: dorsal view; C: lateral external view; D: latero internal view; E: dorsolateral view. F-G: S. (H.) gorirossiae; F: dorsal view; G: ventrolateral view. H-I: S. (H.) johnsoni; H: dorsal view; I: lateral external view. J-K: S. (H.) menchacai; J: lateral view; K: dorsal view.

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prescutellar area. Abdomen blackish. Sc bare. Genitalia similar to those of S. larvispinosum, except ventral plate more elevated medially. Pupa: Length: basally 2.4-2.7 mm; dorsally 1.5-1.7 mm. Cocoon slipper shaped, not closed anteriorly and thickly woven, compact with very narrow anterior rim, showing open weave. Clypeus and thoracic dorsum with moderately to abundant, small and acuminate tubercles; clypeus and thorax with typical strong single trichomes. Gill very similar to that of S. larvispinosum, but with large frontolateral branches narrower, acuminate apically, with spine at tip similar to other branches, first secondary branch more elongated and insertion area more sclerotized; in dorsal view, frontolateral branches more open (Figs. 124 F, G). Abdominal terminal hooks absent. Larva: Length 6.5-7.0 mm. General coloration light yellowish grey. Cephalic apotome uniformly colored, slightly darkened at base (Fig. 126 D); cephalic fan with 5052 primary rays. Antenna longer than stalk of cephalic fan; ratio of proximal, medial and distal antennal articles = 1:1.4:0.7; medial article with pseudosegmentation (Fig. 126 E). Mandible with strong curvature on superior side (Fig. 126 F); internal teeth arrayed in 2 rows with 4 teeth, with first and second teeth very curved, 2 marginal teeth, and 1 robust and long latero- mandibular process, 1/3 of which surpasses inferior edge of mandible (Fig. 126 G). Hypostoma with anterior margin as in S. larvispinosum, with prominent median tooth and intermediate teeth longer than corner teeth; 14-21 lateral setae per side.

I I

III

III

V

V

VII

VII

IX

Figure 125. S. (Hearlea) sp., abdominal chaetotaxy of pupa.

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Postgenal cleft deep, without anterior acuminate incision; ratio of hypostomium/ hypostomial bridge = 0.9-1.1. Comb of thoracic proleg lateral sclerite with 47-55 teeth. Abdominal segment VIII with 1+1 lateroventral, apically sclerotized papillae (Figs. 127 D, F [vp]). Anal sclerite with ventral branch projected distally forming ring around anal ring, laterally bordered by very sclerotized and spinous accessory plates of 2+2 conical dorsolateral projections [dd] and 1+1 flattened more expanded, ventrolateral, acuminate projections (Fig. 127 D [vl]), abundant acuminate tubercles and sparse minute trichomes; and 1+1 dorsolateral, irregularly bordered, elongate plates positioned diagonally to lateral lobes of rectal papillae (Fig. 127 E [df]) and 1+1 short and sclerotized ventrolateral papilla slightly anterior (Fig. 127 F [vp]). Anal ring with 272-282 rows of 30-35 hooks each. Rectal papillae with 18-30 diverticula, 6-10 per lobe. Distribution: México: Chiapas, Veracruz; Guatemala: Sololá. Bionomics: Unknown. Discussion: Simulium (Hearlea) larvispinosum, S. menchacai, S. carolinae, S. johnsoni and S. temascalense are very closely related to S. gorirossiae, but they can be distinguished by subtle differences in the pupal gills and accessory plates of the larval anal sclerite. Simulium larvispinosum has a pupal gill with the dorsofrontal branch more inflated and blunt apically, with the first secondary branch a little shorter (Figs. 124 A-D). The anal sclerite and accessory plates of the abdominal distal area of the larva are similar, but the ventrolateral plates lack hard spinous projections, and anterodorsal to the rectal papillae are 1+1 porous rectangular plates (Fig. 127 I) that in S. gorirossiae are more trapezoidal and obliquely positioned (Fig. 127 E). Simulium menchacai has a very similar larval anal sclerite, accessory plates and ventrolateral sclerotized papillae, but the pupal gill has stout secondary branches as long as the dorsal and ventral branches. Simulium temascalense larvae also have ventrolateral papillae on the eighth segment but they are less sclerotized, the postgenal cleft lacks an acuminate incision and the projections of the ventrolateral accessory sclerite anal plates are shorter. Also, the pupa has very reduced secondary gill branches. Simulium johnsoni, like S. temascalense, has very reduced secondary gill branches, and in the larva, the projections of the ventrolateral accessory plates are shorter, but S. johnsoni lacks ventrolateral papillae on segment VIII of the larva. Simulium carolinae has a very similar pupal gill, but the larva lacks the ventrolateral accessory plates on the distal portion of the abdomen, and the ventrolateral papillae of the eighth segment. Simulium paracarolinae has larvae with only 1+1 accessory dorsolateral plates on the distal portion of the abdomen, and the ventral branches of the anal sclerite are elongated, of subequal diameter, and enclose the anal ring. Simulium (Hearlea) johnsoni VARGAS & DÍAZ NÁJERA (Figs. 124 H, I.) Simulium (Hearlea) johnsoni VARGAS & DÍAZ NÁJERA, 1957b: 203-205; DÍAZ NÁJERA & VULCANO, 1962: 99-100; COSCARÓN et al., 2004: 28-29.

(Diagnosis of female and male based on descriptions from VARGAS & DÍAZ NÁJERA, 1957b and larva from DÍAZ NÁJERA & VULCANO, 1962). Female: Wing length 3.1 mm. General coloration blackish. Scutum black with 2 submedian silvery vittae not joining 1+1 anterior subtriangular silvery spots. Anterior

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portion of cibarium with thickened anterior edge, medially concave. Sc with 4-6 hair. Genitalia similar to those of S. larvispinosum. Male: Wing length 3.1 mm. Coloration blackish, scutum velvety black whitish on anterior edge. Abdominal tergites blackish. Genitalia as in S. larvispinosum. Pupa: Length basally 3.1 mm; dorsally 2.9 mm. Cocoon slipper shaped. Gill membranous with general shape similar to that of S. larvispinosum but secondary branches shorter and ventral branch more strongly curved; small branches acuminate, with well sclerotized spine-like apices (Figs. 124 H, I). Frontoclypeus smooth. Thorax dorsally with abundant small tubercles on median area, and sparse laterally. Posterodorsum of thorax with small, sparse, acuminate tubercles. Larva: Length 8.5 mm. Coloration grey. Cephalic apotome uniformly pigmented, with small thin darkened area mediobasally. Cephalic fan with 54-56 primary rays. Ratio of proximal, medial and distal antennal articles = 1:1.4:1.0. Mandible with preapical teeth robust, internal teeth in 2 rows of 5; 2 large marginal teeth and 3; lateral mandibular processes long and curved. Hypostoma with straight anterior margin and 10-12 lateral setae per side; ratio of hypostomium/hypostomial bridge = 0.7. Postgenal cleft dome shaped; abdominal segment VIII without ventrolateral tubercles. Anal ring with 242 rows of 26-30 hooks each. Anal sclerite with ventral branches encircling anal ring; 2+2 dorsolateral accessory plates with subconical processes, and 1+1 wide lateroventral plates bearing 4-5 short lateral processes. No ventrolateral papillae present. Rectal papillae with 21 diverticula, 7 per lobe, dorsalmost ones largest. Distribution: México: Oaxaca, Veracruz. Bionomics: Unknown. Discussion: Simulium gorirossiae is very similar to S. johnsoni but the pupal gill has the anterodorsal branch thinner and more acuminate, with a terminal spine, and the secondary and ventral branches longer. The larva has longer projections on the accessory ventrolateral plate, and 1+1 ventrolateral papillae on the eighth segment. Simulium temascalense has a very similar pupal gill but the dorsal anterior branch is spined apically, and the larva has ventrolateral papillae on segment VIII and the lateral projections of the accessory ventrolateral plate are shorter. Simulium larvispinosum has larger secondary branches and a more curved ventral branch on the pupal gill, and in the larva, 1+1 dorsolateral accessory rectangular plates before the anal papillae. Simulium (Hearlea) larvispinosum DE LEÓN (Figs. 120 M, P; 121 F, G; 124 A-E; 126 H; 127 G-I.) Simulium (Hearlea) larvispinosum DE LEÓN, 1948: 5-23; DALMAT, 1955: 72, 76, 81, 86, 263-268; DÍAZ NÁJERA & VULCANO, 1962: 93; COSCARÓN et al., 2004: 29-31.

Female: Wing length 3.4 mm. General coloration dark brown. Head blackish, frons brownish silvery pollinose; scutum velvety brown blackish bordered by narrow silver pruinose band, with 1+1 narrow (about 1/4 width of median black stripe) silvery vittae, not reaching to 1+1 anterior subtriangular silvery spots and joining posterior silvery prescutellar area. Frons with sides convergent below; fronto-ocular triangle about as long as wide. Cibarium with thickened, medially concave anterior margin bearing 1+1 short,

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smooth submedian processes (Fig. 120 M). Sc with 8-13 hair. Mandible with 10+25 serrations. Lacinia with 13-14 retrorse teeth. Claw with subbasal tooth (Fig. 120 N); length/width ratio of hind basitarsus = 4.6-4.7. Sternite VIII with 14-16 hairs per side; genital fork and spermatheca as in figures 120 O, P respectively; cercus subrectangular; paraproct with longitudinal fold, and thickly haired distally. Male: Wing length 3.4 mm. Scutum velvety black, bordered by silvery pruinosity, as are humeral angles. Scutellum black. Abdomen blackish. Sc bare. Length/width ratio of hind basitarsus = 3.5-3.6. Gonocoxite subquadrate, 1/2 of gonostylus length; gonostylus with abundant pilosity mediobasally, and subterminal accuminate spur (Fig. 121 F). Endoparameres with strong hooks; median sclerite wide basally; ventral plate subovoidal, without evident median carina (Fig. 121 G). Pupa: Length: basally 3.4 mm. Cocoon slipper shaped, compactly woven, threads not evident; anterior edge slightly reinforced. Frontoclypeus protruded basally without platelets but with impression of minute, very abundant tubercles; 1+1 facial and 2+2 frontal single, stout trichomes; thorax with small, acuminate, sparse tubercles on posterior area and impression of tubercles on remainder of integument, with 5 stout, single, spinelike trichomes per side. Gill rigid, granulose, cylindrical, with 2 principal branches: dorsal branch largest, directed anteriorly; ventral branch smaller, curved, and also directed anteriorly. Main dorsal branch subdivided basally into short secondary branches, the first of which is short and directed dorsally, the second of which is bifurcate, posteriorly directed, with 2 or 3 apical branches; the second laterointernal secondary branch also bifurcate distally; the last frontally directed branch short and apically subdivided (Figs. 124 A-E); except large frontal branch, other branches are sclerotized, and spined apically. Larva: Length 8.0 mm. Color yellowish to light brown (in alcohol). General aspect as in figure 127 G. Cephalic apotome homogeneous without special designs, only darkened on narrow basal stripe; cervical sclerites elongated, reaching edges; ratio of hypostomium/hypostomial bridge = 1.2-1.5. Cephalic fan with 43-48 primary rays. Hypostoma with median tooth stout, surpassing height of other teeth; corner teeth below intermediate teeth; 12-17 lateral setae per side, without discal setae, and few lateral serrations. Postgenal cleft dome shaped with deep projected anteriorly incision (Fig. 126 H). Antenna surpassing length of cephalic fan stem. Ratio of proximal, medial and distal antennal articles = 1:1.4-1.5:0.9. Mandible with 4-5 internal teeth; 2 marginal teeth, the second 1/2 length of anterior tooth. Lateral sclerite of thoracic proleg with 45-50 teeth. Anal sclerite with ventral branch projected basally, forming ring around anal ring. Accessory plates 2+2, subconical, well sclerotized, dorsolaterally and apically positioned (Figs. 127 H, I, [dd]), 1+1 ventrolateral subconical, subannulate, dentate on external edge (Fig. 127 I [vl]) and 1+1 dorsolateral subrectangular, flattened, anterior to rectal gill, (Figs. 127 H, I [df]). No ventrolateral papillae present. Anal ring with about 152 rows of 23-26 hooks each. Rectal papillae with 18-24 diverticula, 6-8 per lobe. Distribution: Guatemala: Chimaltenango, Sololá, Suchitepequez; Mexico: Chiapas. Bionomics: Unknown. Discussion: Simulium johnsoni is very similar to S. larvispinosum, but the pupal gills of these two species exhibit slight differences, as explained for the former species. Adults of S. carolinae do not differ externally. The closest species based on the pupa gill is S. gorirossiae, but

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Figure 126. Larva. A-C: S. (Hearlea) capricorne; A: postgenal cleft and hypostomium; B: hypostomial dental border; C: mandibular dental area. D-G: S. (H.) gorirossiae; D: cephalic apotome and neck sclerite; E: antenna; F: mandible, general aspect; G: mandibular dental area. H: S. (H.) larvispinosum, cephalic capsule, ventral view. I-L: S. (H.) paracarolinae; I: cephalic capsule, ventral view; J: hypostomial dental border; K: antenna; L: mandibular dental area.

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these species exhibit differences in the larval accessory plates, with the ventrolateral plate more subquadrate and having long, acuminate projections and the dorsolateral, flattened plates being positioned diagonally at the sides of the anal gill in S. gorirossiae. Simulium gorirossiae also differs in that it has sclerotized ventrolateral papillae on the eighth segment. Simulium (Hearlea) menchacai VARGAS & DÍAZ NÁJERA (Figs. 124 J, K.) Simulium (Hearlea) menchacai VARGAS & DÍAZ NÁJERA, 1957b: 208-210; DÍAZ NÁJERA & VULCANO, 1962: 101-102; COSCARÓN et al., 2004: 31-32.

Female: Wing length 3.5 mm. General coloration blackish. Thorax blackish brown; scutum with abundant adpressed silvery pilosity that somewhat obscure submedian silvery stripes; vittae not joining 1+1 anterior subtrapezoidal silvery spots; scutellum brownish, metanotum and pleurae blackish; legs yellowish brown. Abdomen brownish with tergites II-IV pale. Fronto-ocular triangle about as long as high. Metathoracic basitarsus length/ width ratio = 0.5-1.1. Cibarium wrinkled, with anterior edge thickened, translucent, concave medially and without lateral subovoidal prominences; Sc with 15-17 trichomes. Genitalia as in S. larvispinosum. Male: Wing length 3.0 mm. Coloration blackish; scutum velvety black with anterior area whitish. Sc with 2-3 trichomes. Metathoracic basitarsus length/width ratio = 3.0-3.1. Genitalia similar to those of S. larvispinosum. Pupa: Length: basally 3.5 mm; dorsally 3.0 mm. Cocoon slipper shaped, closed basally, with reinforced edge slightly projected laterodorsally; weave loose and coarse anteriorly. Gill well sclerotized, cylindrical, comprised of 9-10 terminal branches; 3 primary gill branches: dorsal one short, oriented posteriorly, and curved with rounded apex; median branch short, with 3 secondary branches of approximately equal length; dorsal secondary branch bifurcated basally, median secondary branch subdivided apically into 3 branches, third secondary branch also with 3 subdivided branches, although on different plane; primary ventral branch curved and longer than other three main branches (Figs. 124 J, K). All branches apically acuminated. Frontoclypeus smooth, and with robust spiniform setae, frontal setae separated and superior very long; thorax with abundant, small, rounded tubercles. Larva: (Diagnosis based on VARGAS & DÍAZ NÁJERA, 1957 and also DÍAZ NÁJERA & VULCANO, 1962). Length (mature) 7.5-8.0 mm. Coloration dark greenish. Cephalic apotome homogeneously pigmented, without special ornamentation. Cephalic fan with 50-55 primary rays. Hypostomium with anterior margin curved and 15-17 lateral setae per side. Postgenal cleft dome shaped, with apical incision. Anal sclerite, accessory plates and ventrolateral papillae similar to those of S. gorirossiae, but with ventrolateral plates having shorter accuminate projections and directed only anteriorly. Anal ring with about 310 rows of hooks. Rectal papillae with 18-36 diverticula. Distribution: Mexico: Oaxaca. Bionomics: Unknown. Discussion: The larva is very close to those of S. gorirossiae and S. temascalense, based on a similar anal sclerite and accessory plate and the presence of ventrolateral papillae on the larva. However, the peculiar form and arrangement of the pupal gill branches permits their separation.

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Simulium (Hearlea) paracarolinae COSCARÓN (Figs. 126 I-L; 127 B, C.) Simulium (Hearlea) paracarolinae COSCARÓN et al., 2004: 32-33.

Larva: Length (mature) 7.0-8.5 mm. Coloration light brown yellowish (in alcohol). Cephalic apotome light brown, with darkened base, and darkened medial longitudinal stripe on basal 1/3 and 1+1 small submedian spots near midline of sclerite. Cephalic fan with 44-48 primary rays. Medial antennal article with 4 annulations (Fig. 126 K); ratio of proximal, medial and distal antennal articles = 1:1.6:1.0. Postgenal cleft subtriangular, without anterior incision (Fig. 126 I); ratio of hypostomium/hypostomial bridge = 1.2. Hypostomium (Fig. 126 J) with median tooth longer than others; lateral setae 11-12 and 2 discal setae. Mandible with internal teeth arrayed in 3-4 rows with 6-8 teeth each; 2 marginal teeth, the second very small (Fig. 126 L); mandibular lateral process unique, elongated, stout and curved. Lateral sclerite of thoracic proleg with 42-46 teeth. Anal sclerite with ventral branches heavily sclerotized, relatively short, not encircling posterior circlet, although enclosing approximately 1/3 of it; accessory plates constituted of 1+1 well-sclerotized, dorsolateral conical plates covered with small setae (Figs. 127 B, C [ds]). No ventrolateral papillae. Anal ring with 398-400 rows of 42-48 hooks each. Anal papillae with 18-21 diverticula, 6-7 per lobe, median lobe largest. Distribution: Guatemala. Bionomics: Unknown. Discussion: Larval morphology permits differentiation of S. paracarolinae from all other species. The presence of accessory plates on the anal sclerite and a curved anterior hypostomial margin place this species into the CAROLINAE species group, for which the larva is known for all species. The larva cannot be distinguished from the unknown larva of other S. (Hearlea) species, as it could potentially be that of S. nigricorne, because pupal morphology suggests it corresponds to another group with different characters in the larva, such as absence of accessory plates on the distal portion of the abdomen, straight anterior edge of the hypostomium and distinct mandibular morphology and postgenal cleft. The closest species is S. carolinae, but it has the anal sclerite ventral branch forming a ring encircling the rows of hooks of the anal ring, and also has 2+2 dorsolateral accessory plates. The absence of a mature gill histoblast does not permit comparisons with other species. Simulium (Hearlea) temascalense DÍAZ NÁJERA & VULCANO (Fig. 127 J.) Simulium (Hearlea) temascalense DÍAZ NÁJERA & VULCANO, 1962: 102-105, 140, 142; COSCARÓN et al., 2004: 33.

(Diagnosis of female and pupa based on DÍAZ NÁJERA & VULCANO, 1962). Female: General coloration blackish. Scutum brown with 1+1 thin silvery vittae not joining anterior 1+1 subtriangular silvery spots. Pleurae black pollinose. Femora and tibiae yellowish, with black apices. Meso- and metathoracic coxae, trochanter, tarsomeres I-II, and apex of tarsomere III black. Cibarium with smooth, thickened, medially concave anterior margin. Genitalia similar to those of S. larvispinosum. Male: Unknown. Pupa: Cocoon (at base) 3.0 mm. Frontoclypeus and anterodorsum of thorax without tubercles. Gill with general facies of S. carolinae, with anterior branch elongate, relatively

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Figure 127. A-J: Larva. A-D, H-J: Terminal portion of abdomen showing anal sclerite and accessory plates (d = dorsal branch of anal sclerite, dd = dorsolateral conical accessory plates double, d f = dorsolateral flattened accessory plate, d s = dorsolateral conical accessory plate single, gd = diverticles of rectal papillae, v = ventral branch of anal sclerite, vl = ventrolateral accessory plate, vp = ventrolateral papillae). A: S. (Hearlea) capricorne. A: anal sclerite. B-C: S. (H.) paracarolinae; B: anal sclerite area in transparent slide view; C: abdomen posterodorsal view. D-F: S. (H.) gorirossiae; D: abdomen posteroventral view; E: abdomen posterodorsal view, with rectal papillae diverticles (gd); F: eighth segment ventrolateral papilla (vp) sclerotized. G-I: S. (H.) larvispinosum, general aspect; G: lateral view; H: abdomen posterodorsal view; I: same in transparent slide view. J: S. (H.) temascalense, abdomen posterolateral view.

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thin, with acuminate, spined apex, with 7 small subbasal branches and another curved and directed posteroventrally; apex of branches spined. Larva: Length (mature) 8.0 mm. Coloration dark greenish; head yellowish, cephalic apotome without noticeable ornamentation. Cephalic fan with 50 primary rays. Mandible with strong preapical tooth, 2 rows of internal teeth, and 2 large marginal teeth; lateromandibular process long and curved. Ratio of proximal, medial and distal antennal articles = 1:1.6:1.2. Comb of thoracic proleg sclerite with 38-39 teeth arranged in 10 groups. Hypostoma with anterior edge curved, median tooth longer than other teeth, and 15 lateral setae per side; ratio of hypostomium/hypostomial bridge = 0.8; postgenal cleft mildly curved apically, without acuminate anterior incision. Segment VIII with 1+1 sublateral, sclerotized papillae. Anal ring with 242 rows of 26-30 hooks. Distal portion of abdomen with 2+2 dorsolateral acuminate posteriorly directed processes, 1+1 wide ventrolateral positioned processes having small lateral projections (Fig. 127 J [dd and vl]); 1+1 dorsolateral flattened plates and 1+1 ventrolateral papillae; ventral arms of anal sclerite surrounding anal ring. Rectal papillae with 21-27 diverticula, 7-9 per lobe. Distribution: Mexico: Michoacan, Oaxaca. Bionomics: Unknown. Discussion: Simulium johnsoni and S. larvispinosum have very similar pupal gills, but the larvae lack ventrolateral papillae on segment VIII and the females have a blackish scutum in these species. Simulium (Trichodagmia) ENDERLEIN (Figs. 128-131; 153.) Trichodagmia ENDERLEIN, 1934b: 288 (as genus). Type-species: Simulium townsendi MALLOCH, 1912 = Trichodagmia latitarse ENDERLEIN, 1934b: 288. Grenieriella VARGAS & DÍAZ NÁJERA, 1951: 141. Type-species: Eusimulium lahillei PATERSON & SHANNON, 1927: 740.

Female: Big species (wing length 2.8-4.4 mm). Scutum brown-orange, dark brown to blackish, generally ornamented with 1+1 submedian longitudinal grayish stripes joined posteriorly, forming lyre-like pattern (Fig. 128 A); pilosity generally homogeneous but can be grouped simulating scales. Abdomen dark brown with blackish, generally with 1+1 grayish spots on tergite II. Frons relatively wide (Fig. 128 F) and fronto-ocular triangle about as long as wide (Fig. 128 G). Cibarium well sclerotized with small teeth generally like spiculae on median depressed area (Figs. 128 D, H). Claw with well developed subbasal tooth. Eighth sternite well sclerotized with median portion depressed; gonapophysis subovoidal to subtriangular with blunt apex and small anterior depression (Fig. 128 I). Genital fork with wide anterior branches and strong, well-sclerotized apodeme (Fig. 128 K). Male: Color brown-yellowish to black. Gonostylus nearly 1.5 times longer than gonocoxite, subcylindrical, acuminate distally and about 2 times longer than wide at base, generally with 1 apical spur. Ventral plate body wide, about 2.0 times wider than long (Figs. 129 G, E, K, M), with short median process. Endoparamere branches without hooks (Fig. 129 C). Pupa: Cocoon shoe shaped, generally with aperture anteriorly elevated protecting gill; weave thick. Frontoclypeus reinforced basally. Frontoclypeus and exposed portion of thorax

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generally with abundant tubercles; trichomes small, multibranched. Gill with 12-20 branches generally moderate thick and relatively short; abdomen with terminal spur very reduced. Larva: Body shape dorsally frequently straight on subterminal portion as in S. (Hemicnetha). Cephalic apotome dark brown, mostly homogeneous, darkened on basal margin and scarcely ornamented. Antenna relatively thick, with medial article much longer than proximal and distal; mandible with several internal teeth rows and thin marginal teeth. Postgenal cleft deep, with postgenal bridge shorter than hypostomium length; hypostomium with straight anterior margin. Anal sclerite with abundant hair, but no scales. Anal ring with 150-360 rows with 23-40 hooks each. Rectal papillae with elongated diverticula of 10-45 on each lobe. Distribution: The subgenus has 7 species in the South American temperate west sub-Andean and eastern mountains (Fig. 153). KEYS

TO

SIMULIUM (TRICHODAGMIA)

SPECIES

Females 1 – 2(1) 3 4(2) –

Scutum grayish brown, hair arranged in groups; paraproct relatively short (Fig. 128 J), ratio of length/width basally = 1.3; cibarium teeth well developed (Fig. 128 H) .......................................................................................................... Simulium nigrimanum Scutum yellowish brown or reddish to blackish; paraproct relatively long (Figs. 128 C, E), ratio of length/width = 1.9-2.0; cibarium teeth reduced, like spicules (Fig. 128 D) ......................................................................................................................... 2 Scutum blackish; paraproct well sclerotized; cercus with distal border straight (Fig. 128 E) ..................................................................................................... Simulium townsendi Scutum yellowish brown to grayish brown or blackish; paraproct medially sclerotized; cercus with distal border curved (Fig. 128 C) ........................................ 3 Scutum blackish golden to dark brown; wing length 2.8-3.8mm .............................. ................................................................ Simulium wgodzinskyorum, Simulium sumapazense Scutum blackish to reddish brown or grayish brown; wing length 3.9-4.4 mm ..... ............................................... Simulium huairayucu, Simulium lahillei, Simulium muiscorum

Males Not included: Simulium townsendi. 1 – 2(1) – 3(2)

Gonostylus with about 4 apical spurs (Fig. 129 L); apical process of ventral plate relatively long (Fig. 129 M) ............................................................. Simulium nigrimanum Gonostylus with 1 (or 2) apical spurs (Figs. 129 B, D, F, J); apical process of ventral plate short (Figs. 129 B, E, G) (except Simulium wygodzinskyorum) ............. 2 Scutum reddish brown to brownish orange; ratio of length/width of hind basitarsus = 4.5-5.0 ............................................................................................................ 3 Scutum black; ratio of length/width of hind basitarsus = 3.5 ................................ 4 Scutum reddish brown, showing with some illumination 1+1 subtriangular darker spots reaching lateral margin by thin anterior stripe (Fig. 129 A); ventral plate with median process short (Fig. 129 B) ......................................................... Simulium lahillei

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Scutum brownish orange, showing with some illumination 1+1 oblique subtriangular nacreous spots, reaching from antero-external angle to submedian darker stripes (Fig. 129 H); ventral plate with median process long (Fig. 129 K) ...................................................................................................... Simulium wygodzinskyorum Ratio of length/width of hind basitarsus = 3.5 ........................................................... ......................................................................... Simulium huairayacu, Simulium sumapazense Ratio of length/width of hind basitarsus = 2.5-2.7 ................... Simulium muiscorum

Pupae Not included: Simulium townsendi. 1 – 2(1) – 3(2) – 4(2) – 5(4) –

Gill with 18-20 branches (Fig. 130 L); frontoclypeus and thorax without tubercles ............................................................................................................. Simulium nigrimanum Gill with 12 branches; frontoclypeus and thorax generally with numerous tubercles (Fig. 130 F) ......................................................................................................... 2 Cocoon with anterior aperture border lower than posterior edge (Figs. 130 A, D) ................................................................................................................................................ 3 Cocoon with anterior aperture border higher than posterior edge (Figs. 130 B, C) ................................................................................................................................................ 4 Gill branches very thick, emerging from base without trunk, branches apically acute and with microscopic annulations only distally (Figs. 130 A, H) ......... Simulium lahillei Gill branches relatively thin, emerging from base with basal trunk, branches apically blunt and with microscopic annulations from base (Fig. 130 J) ................. ...................................................................................................... Simulium wygodzinskyorum Frontoclypeus and exposed portion of thorax without tubercles (Fig. 130 G); gill expanded (Fig. 130 K) .................................................................... Simulium sumapazense Frontoclypeus and exposed portion of thorax with numerous tubercles (Fig. 130 F); gill not expanded .......................................................................................................... 5 Ventral primary branches positioned far from other branches (Fig. 130 I) ............ ................................................................................................................ Simulium huairayucu Ventral primary branches positioned near other branches ........ Simulium muiscorum

Larvae No key to larvae is presented because of scarcity of material and characters. CLAVES

DE ESPECIES DE

SIMULIUM (TRICHODAGMIA)

Hembras 1 –

Escudo castaño grisáceo, pelos dispuestos en grupos; paraprocto relativamente corto (Fig. 128 J); relación largo/ancho en la base = 1,3; dientes del cibario bien desarrollados (Fig. 128 H) ............................................................... Simulium nigrimanum Escudo castaño amarillento o rojizo a negruzco; paraprocto relativamente largo (Figs. 128 C, E); relación largo/ancho = 1,9-2,0; dientes del cibario reducidos como espículas (Fig. 128 D) ............................................................................................ 2

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Escudo negruzco, paraprocto bien esclerotizado; borde distal del cerco recto (Fig. 128 E) ..................................................................................................... Simulium townsendi Escudo castaño amarillento a castaño grisáceo o negruzco, paraprocto medianamente esclerotizado; borde distal del cerco curvado (Fig. 128 C) ........... 3 Escudo negruzco dorado a castaño oscuro; largo ala 2,8-3,8 mm ............................ ............................................................... Simulium wygodzinskyorum, Simulium sumapazense Escudo negruzco a castaño rojizo o castaño grisáceo; largo ala 3,9-4,4 mm ......... ............................................... Simulium huairayacu, Simulium lahillei, Simulium muiscorum

Machos No incluida: Simulium townsendi. 1 – 2(1) – 3(2) –

4(2) –

Gonostilo con alrededor de 4 espolones apicales (Fig. 129 L); prolongación apical de la placa ventral relativamente larga (Fig. 129 M) .................. Simulium nigrimanum Gonostilo con 1 (ó 2) espolones apicales (Figs. 129 B, D, F, J); prolongación apical de la placa ventral corta (Figs. 129 B, E, G) (excepto Simulium wygodzinskyorum) ............. 2 Escudo castaño rojizo a castaño naranja; relación largo/ancho del basitarso posterior = 4,5-5,0 ............................................................................................................. 3 Escudo negro; relación largo/ancho del basitarso posterior = 3,5 ........................ 4 Escudo castaño rojizo mostrando con cierta iluminación 1+1 manchas oscuras subtriangulares, llegando al borde lateral por una fina banda anterior (Fig. 129 A); prolongación mediana de la placa ventral corta (Fig. 129 B) .......... Simulium lahillei Escudo castaño naranja mostrando con cierta iluminación 1+1 manchas nacaradas subtriangulares oblicuas, llegando desde el ángulo antero externo hasta las bandas submedianas más oscuras (Fig. 129 H); prolongación mediana de la placa ventral larga (Fig. 129 K) ............................................. Simulium wygodzinskyorum Relación largo/ancho del basitarso posterior = 3,5 ..................................................... ......................................................................... Simulium huairayacu, Simulium sumapazense Relación largo/ancho del basitarso posterior = 2,5-2,7 ............. Simulium muiscorum

Pupas No incluida: Simulium townsendi. 1 – 2(1)

3(2)

Branquia con 18-20 ramas (Fig. 130 L); frontoclípeo y tórax sin tubérculos ......... ............................................................................................................... Simulium nigrimanum Branquia con 12 ramas; frontoclípeo y tórax generalmente con numerosos tubérculos (Fig. 130 F) ...................................................................................................... 2 Capullo con borde frontal de la abertura más bajo que borde posterior (Figs. 130 A, D) ..................................................................................................................................... 3 Capullo con borde frontal de la abertura más alto que el borde posterior (Figs. 130 B, C) ...................................................................................................................................... 4 Ramas de las branquias muy gruesas, emergiendo desde la base sin tronco, ápice de las ramas aguzado y con anillaciones microscópicas solo distalmente (Figs. 130 A, H) ............................................................................................................ Simulium lahillei

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Ramas de las branquias relativamente finas, emergiendo desde la base con un tronco basal; ápice de las ramas romo y con anillaciones microscópicas desde la base (Figs. 130 J) ...................................................................... Simulium wygodzinskyorum Frontoclípeo y porción expuesta del tórax sin tubérculos (Fig. 130 G); branquia expandida (Fig. 130 K) ................................................................... Simulium sumapazense Frontoclípeo y porción expuesta del tórax con numerosos tubérculos (Fig. 130 F); branquia no expandida ...................................................................................................... 5 Rama primaria ventral de la branquia alejada de las otras ramas (Fig. 130 I) ......... ................................................................................................................ Simulium huairayacu Rama primaria ventral dispuesta junta con las otras ramas ....... Simulium muiscorum

Larvas No se presenta clave por escaso material y caracteres. Simulium (Trichodagmia) townsendi MALLOCH (Fig. 128 E.) Simulium townsendi MALLOCH, 1912: 651-652. Simulium (Trichodagmia) townsendi: HERNÁNDEZ & SHELLEY, 2005: 12-15. Simulium chalcocoma KNAB, 1914b: 85. Simulium (Trichodagmia) chalcocoma: VARGAS & DÍAZ NÁJERA, 1953b: 139; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429; COSCARÓN ARIAS, 2003: 191. Simulium (Grenieriella) chalcocoma: VARGAS & DÍAZ NÁJERA, 1951: 142; COSCARÓN, 1987: 38. Simulium (Hemicnetha) townsendi: CROSSKEY & HOWARD, 1997: 38; 2004: 74 Trichodagmia ecuadoriense ENDERLEIN, 1934a: 193. Trichodagmia latitarse ENDERLEIN, 1934b: 289. Trtichodagmia strigata ENDERLEIN, 1934b: 290.

Female: Wing length 3.5-3.7 mm. Scutum brown and grayish with 1 thin median and 1+1 wider submedian and lateral stripes; submedian stripes joined posteriorly, forming lyre pattern as in S. rubrithorax; with frontal illumination, vittae becoming blackish, and with posterior light, median and sublateral longitudinal areas grayish and black; pilosity recumbent, uniformly golden to silvery. Scutellum and metanotum brown and grayish. Legs blackish with distal portion of hind tibia and basitarsus with silver pruinosity. Abdomen blackish with 1+1 light grayish large lateral and posterior spots on tergite II, thinner on tergites III-V; tergal plates II-V dark brown. Cibarium as in S. huairayacu; fronto-ocular triangle slightly wider than high. Sensory vesicle of maxillary palp near 1/3 of palpomere length. Sc with hair all along vein. Claw with well-developed subbasal tooth. Eighth sternite and gonapophysis as in S. nigrimanum, but the first more sclerotized and with about 17 setae on each side; cerci and paraprocts well sclerotized; cerci with straight border (Fig. 128 E); paraprocts about 2 times longer than wide; genital fork with anterior apodemes relatively long and well sclerotized. Male, pupa and larva: Unknown. Discussion: The species is very close to S. huairayacu, S. muiscorum, S. sumapazense and S. wygodzinskyorum; small differences permit them to be separated, such as the curved distal border of the cercus, and the paraproct slightly sclerotized medially.

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B F

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E C Figure 128. Female. A-C: S. (Trichodagmia) lahillei; A: scutum, B: genitalia, ventral view showing eighth sternite and gonapophysis, cercus and paraproct, genital fork and spermatheca; C: cercus and paraproct, slide view. D: S. (T.) huairayacu, cibarium median portion. E: S. (T.) townsendi, cercus and paraproct. F-K: S. (T.) nigrimanum; F: frons; G: fronto-ocular triangle; H: cibarium; I: eighth sternite and gonapophysis; J: cercus and paraproct; K: genital fork.

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We accept the synonyms proposed by HERNÁNDEZ & SHELLEY, 2005, except S. muiscorum which shows differences such as the large size and short paraproct. The similarity among females is the principal element used to synonymize this species, but without knowledge of the other stages, it is doubtful if they are conspecific. Some species, such as S. muiscorum, S. huairayacu and S. sumapazense, have very similar females, but it is possible to separate them in the other stages, especially the immatures. Knowledge of more topotypical material, including all life stages, and cytological and molecular methods, will help to decide if S. muiscorum is a synonym of S. townsendi. Simulium (Trichodagmia) lahillei (PATERSON & SHANNON) (Figs. 128 A-C; 129 A-C; 130 A, H, N; 131 A-F.) Eusimulium lahillei PATERSON & SHANNON, 1927: 740; PINTO, 1931: 678. Simulium lahillei: VARGAS, 1945a: 153; WYGODZINSKY, 1949 : 306; 1953a: 317; STONE, 1963: 8. Simulium (Trichodagmia) lahillei: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429, 435. Simulium (Grenieriella) lahillei: VARGAS & DÍAZ NÁJERA, 1951: 141; COSCARÓN, 1987: 38; 1991: 276-278.

Female: Wing length 4.0-4.3 mm. Scutum brownish red with 1+1 submedian thin light gray pollinose stripe converging to posterior yellowish area, forming lyre pattern, with changing light position, thin stripes become blackish (Fig. 128 A). Scutum pile uniformly yellowish to silvery. Scutellum and metanotum dark brown, the latter with grayish pruinosity. Legs yellowish with silvery pilosity. Abdomen yellowish, tergite I darker with yellow posterior margin; II-V matte with 1 median and 1+1 submedian darker spots, and tergite II with 1+1 silvery anterior spots; VI-IX shinny. Gonapophysis subtriangular, about as long as wide at base (Fig. 125 B). Paraproct longer than wide at base, curved distally with slight anterior concavity (Fig. 128 C). Male: Wing length 3.6-4.0 mm. Scutum brownish red with 1+1 small light grayish anterior submedian subtriangular spots (Fig. 129 A); with changing light position, stripes become darker and reach lateral border by thin anterior stripe; pilosity uniformly grayish. Scutellum yellowish red; metanotum reddish to blackish; abdomen as in female. Ratio of length/width of hind basitarsus = 4.5-5.0. Gonostylus nearly half as long as gonocoxite, with curved apex and 1-2 apical spurs (Fig. 129 B); ventral plate with short median, relatively acute process (Fig. 129 B); endoparameres and aedeagal membrane as in figure 129 C. Pupa: Cocoon with aperture frontally projected, protecting gill branches (Fig. 130 A). Length basally 3.0-3.3 mm; dorsally 3.0-3.8 mm; maximum length 5.5-6.0 mm; gills 1.7-1.8 mm. Gills with 12 branches very thick, with acute apices; 4 primary branches emerging from base (Fig. 130 H). Frontoclypeus and exposed portion of thorax with small and abundant tubercles; trichomes with 5-8 branches and spiraled basal trunk as in S. huairayacu. Abdominal chaetotaxy as in figure 130 N with very reduced terminal spur. Larva: Maximum length 13.0 mm. Body general as in figure 131 A. Cephalic apotome dark brown, mostly uniform, darker on basal border and with negative spots (Figs. 131 B, C). Cephalic fan with 50-60 rays. Antenna thick (Fig. 131 D), ratio of proximal, medial and distal articles = 1:1.8:1.1. Mandible with much reduced second marginal tooth (Fig. 131 E). Postgenal cleft subtriangular (Fig. 131 F). Anal sclerite with

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K Figure 129. Male. A-C: S. (Trichodagmia) lahillei; A: scutum, B: genitalia, ventral view, gonocoxite and gonostylus, ventral plate; C: endoparameres and aedeagus. D-E: S. (T.) huairayacu; D: gonocoxite and gonostylus; E: ventral plate. F-G: S. (T.) muiscorum; F: gonocoxite and gonostylus, dorsal view; G: ventral plate. H-K: S. (T.) wygodzinskyorum; H: scutum; I: hind basitarsus; J: gonostylus; K: ventral plate. L-M: S. (T.) nigrimanum; L: gonocoxite and gonostylus, dorsal view; M: ventral plate.

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hair and scales, the last in 2-4 rows. Anal ring with 300-360 rows with 30-40 hooks each. Rectal papillae with about 10 diverticula on each lobe. Distribution: Argentina: Catamarca, Córdoba, Jujuy, Salta, Tucumán. Bolivia; Peru: Cuzco. Bionomics: This species breeds in creeks of median discharge, fast current and clear water, attached in groups on stones near the surface, frequently in winter and at the beginning of spring. Females were collected on horses and biting humans. Discussion: The closest species is S. huairayacu; differences are explained under that species. Probably S. lahillei is the most important vector of Mansonella ozzardi in northern Argentina (SHELLEY & COSCARÓN, 2001). Simulium (Trichodagmia) wygodzinskyorum COSCARÓN & PY-DANIEL (Figs. 129 H-K; 130 F, J.) Simulium (Grenieriella) wygodzinskyorum COSCARÓN & PY-DANIEL, 1989: 319. Simulium (Trichodagmia) wygodzinskyorum CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429-435; COSCARÓN ARIAS, 2003: 191, 206.

Female: Wing length 3.5 mm. General coloration brownish. Scutum dark reddish brown with 1+1 submedian stripes, with lyre pattern in some illumination; pilosity uniformly silvery to yellowish, in accordance with light incidence. Scutellum and metanotum dark brown, the latter with gray pruinosity. Legs yellowish with silvery pilosity. Abdomen brownish yellow with darker tergal plates. Cibarium smooth and sclerotized subbasally. Claw with short subbasal tooth. Genitalia as in S. lahillei; eighth sternite with about 10 setae on each side; gonapophysis apically curved. Male: Wing length 3.0-3.2 mm. Scutum yellowish brown with 1 median longitudinal and 1+1 thin submedian darker stripes converging posteriorly; with other illumination, 1+1 anterior oblique nacreous spots can be seen from outer anterior angle to submedian darker stripes (Fig. 129 H). Scutellum, metanotum and abdomen brown, changing to dark brown on the latter and with 1+1 lateral silvery spots on tergites II and IV. Legs as in female but darker; ratio of length/width of hind basitarsus = 4.8 (Fig. 129 I). Gonostylus about 2 times longer than gonocoxite, relatively acuminate apically (Fig. 129 J). Ventral plate with well-developed median process (Fig. 129 K). Pupa: Cocoon relatively low anteriorly, similar to that of S. sumapazense but lower than anterior portion of aperture; length basally 2.5-3.0 mm; dorsally 2.7-3.3 mm; maximum length 3.4-4.8 mm, gills 1.0-1.2mm. Frontoclypeus and thorax with numerous small and granulose tubercles (Fig. 130 F); basal portion of frontoclypeus reinforced. Cephalic and thoracic trichomes multibranched, frontal trichomes reduced. Gill with 12 medially thick branches, with microscopic annulations on surface and acute apically; 4 short primary branches emerging from short basal trunk (Fig. 130 J). Larva: Cephalic apotome dark brownish without much ornamentation and with negative spots. Cephalic fan with 30-32 rays. Ratio of proximal, medial and distal antennal articles = 1:2.5-2.8:1.2-1.5. Anal ring with 188-212 rows with 26-28 hooks each. Rectal papillae with 17-19 diverticula on each lobe. Distribution: Peru; Ecuador: Pichincha; Junín.

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Figure 130. Pupa. A-E: lateral view; A: S. (Trichodagmia) lahillei; B: S. (T.) huairayacu; C: S. (T.) muiscorum; D: S. (T.) sumapazense; E: S. (T.) nigrimanum. F-G: frontoclypeus; F: S. (T.) wygodzinskyorum; G: S. (T.) sumapazense. H-L: gills; H: S. (T.) lahillei; I: S. (T.) huarayacu; J: S. (T.) wygodzinskyorum; K: S. (T.) sumapazense; L: S. (T.) nigrimanum. M: S. (T.) huairayacu, thoracic trichome enlarged. N: S. (T.) lahillei, abdominal chaetotaxy.

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Bionomics: Unknown. Discussion: Simulium lahillei and S. huairayacu are very close, but they are bigger, have different male scutal ornamentation, darker color, especially in S. huairayacu, a ventral plate with a short median process, and a pupal gill without a basal trunk and with thicker branches in S. lahillei and a different arrangement in S. huairayacu. Simulium townsendi also is similar but can be differentiated by the wider, more sclerotized paraproct, distally flattened cerci, and general blackish coloration. Simulium (Trichodagmia) huairayacu WYGODZINSKY (Figs. 128 D; 129 D, E; 130 B, I, M.) Simulium huairayacu WYGODZINSKY, 1953a: 310; 1961: 460. Simulium (Grenieriella) huairayacu: COSCARÓN, 1987: 38; 1991: 278. Simulium (Trichodagmia) huairayacu CROSSKEY & HOWARD, 1997: 79; 2004: 74; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429, 435.

Female: Wing length 3.9-4.4 mm. Scutum brown-violaceous or reddish brown to brownish orange, with 1+1 submedian grayish stripes anteriorly starting as a subtriangle and converging posteriorly, forming lyre pattern. Scutellum yellowish brown and metanotum dark brown with gray pruinosity. Legs yellowish gray. Abdomen light yellowish gray with blackish velvety spots on tergal plates of segments II-V, and slightly lighter on VI-VII with dark grayish center; segments II-VI with 1+1 sublateral silvery spots, more developed on II. Cibarium concave medially and with abundant spiniform teeth (Fig. 128 D). Fronto-ocular triangle shorter than wide; claw with subbasal tooth moderately developed. Genital as in S. lahillei. Male: Wing length 3.5-4.0 mm. Scutum velvety black, showing with some illumination silvery lateral and posterior edges. Scutellum dark brown; metanotum blackish; some specimens lighter with reddish brown scutum. Legs brown to blackish. Abdomen velvety black; tergites II, V, VII with 1+1 submedian silvery spots, those of V smaller and in some specimens imperceptible. Ratio of length/width of hind basitarsus = 3.5. Genitalia very similar to those of S. lahillei, but gonostylus shorter, about 1.5 times longer than gonocoxite (Fig. 129 D). Ventral plate with short median process (Fig. 129 E). Pupa: Cocoon with aperture anteriorly elevated (Fig. 130 B), protecting gills; length basally 4.0-5.0 mm, on dorsum 2.6-3.0mm; maximum length 4.5-5.1mm; gills 1.4-1.6 mm. Frontoclypeus and exposed portion of thorax with abundant tubercles; trichomes frequently curled with thick basal trunk and 7-12 thin and elongated branches (Fig. 130 M). Gill with 12 thick branches with blunt apices, 4 relatively short primary branches emerging from common place, with 2 ventral branches directed down and forward and other directed dorsally, forming a bunch (Fig. 130 I). Larva: Maximum length 10-11mm. Very similar to S. lahillei. Ratio of proximal, medial and distal antennal articles = 1:2.3-2.7:1.3. Anal ring with about 250 rows with 30 hooks each. Rectal papillae with about 45 diverticula on each lobe. Distribution: Argentina: Jujuy, Salta, Tucumán; Bolivia: Cochabamba, Potosí. Bionomics: This species breeds in small to medium-sized torrential creeks in clear, cold water and is found frequently with S. lahillei.

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Discussion: This species is very close to S. lahillei but is easy to separate by the pupal gill shape. In the adults, it is possible to separate the male because S. lahillei is lighter reddish brown. Simulium (Trichodagmia) muiscorum BUENO, MONCADA & MUÑOZ DE HOYOS (Figs. 129 F, G; 130 C.) Simulium (Hemicnetha) muiscorum BUENO et al., 1979: 581-594. Simulium (Trichodagmia) muiscorum: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MUÑOZ DE HOYOS & COSCARÓN, 1999: 183; MIRANDA ESQUIVEL & COSCARÓN, 2001: 429, 435. Simulium (Grenieriella) muiscorum: COSCARÓN, 1987: 37.

Female: Wing length 3.9-4.1 mm. Coloration brown dark blackish. Scutum blackish with small golden to grayish hair and some darkened; with frontal illumination, 1+1 subtriangular light gray spots continued posteriorly by thin dark stripes. Scutellum and metanotum blackish, the latter shiny. Legs brown with apex of femora, tibiae and tarsomeres blackish. Cibarium and genitalia as in S. huairayacu, claw with well-developed subbasal tooth. Abdomen dark brown with 1+1 grayish spots laterally on tergites III-IV. Gonapophysis, genital fork and cercus as in S. lahillei, but the former more rounded apically; paraproctos as in S. townsendi but shorter and less sclerotized Male: Wing length 3.5-3.8 mm. Scutum blackish with golden to gray pilosity; scutellum, metanotum and legs as in female. Hind legs wide; ratio of length/width of hind basitarsus = 2.5-2.7. Abdomen dark brown with tergal plates darkened and 1+1 big silvery spots on tergite II. Gonostylus and ventral plate as in figures 129 F, G, respectively. Pupa: Cocoon with aperture very elevated frontally, covering mostly of gill base (Fig. 130 C). Length basally 3.5-4.0 mm, maximum length 4.8-5.0 mm; gills 1.2 mm. Frontoclypeus and exposed portion of thorax with numerous smooth tubercles. Facial trichomes with 7-12 branches, frontal with 2-5, thoracic with 3-5 branches. Gill with 12 branches emerging from short basal trunk with 4 primary branches; gill filaments with microscopic pseudoannulations and acute apex. Abdomen terminal spur absent. Larva: Maximum length 8.5-9.0 mm. Cephalic apotome brownish, homogeneously pigmented, only darkened on narrow basal margin and without definite spots. Cephalic fan with 36-39 rays. Antenna surpassing apex of cephalic fan stem; ratio of proximal, medial and distal antennal articles = 1:1.9-2:1.1-1.4. Mandible with thin marginal teeth, the second very small. Postgenal cleft dome shaped, postgenal bridge nearly half of hypostomium length. Thoracic proleg sclerite with about 47 teeth arranged in 15-17 groups. Anal sclerite without scales. Anal ring with about 196 rows of 30-34 hooks each. Rectal papillae with 24-35 diverticula on each lobe. Distribution: Colombia: Cundinamarca. Bionomics: This species breeds in torrential creeks with rocky bottoms and clear, cold water, attached to rocks or aquatic vegetation. Females bite cattle. Discussion: Adult morphology is similar to that of the other species in the subgenus. The closest species is S. huairayacu, with similar male blackish coloration. The adults of S. lahillei and S. wygodzinskyorum are lighter reddish brown. The anterior elevated

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aperture of the cocoon is very similar to that of S. huairayacu, but the gill branching, with more open basal primary branches, plus the larger larval size, greater number of anal ring rows and rectal papillae diverticula allow separation. It would be useful to do cytological and molecular studies to reaffirm the validity of these species. Simulium (Trichodagmia) sumapazense COSCARÓN & PY-DANIEL (Figs. 130 D, G, K.) Simulium (Grenieriella) sumapazense COSCARÓN & PY-DANIEL, 1989: 321. Simulium (Trichodagmia) sumapazense: CROSSKEY & HOWARD, 1997: 79; 2004: 74; MUÑOZ DE HOYOS & COSCARÓN, 1999: 183.

Female: Wing length 3.2-3.7 mm. Scutum blackish golden with 1+1 grayish submedian stripes forming lyre pattern, pilosity homogeneous. Paraprocts about as long as wide at base. Male: Wing length 2.8-3.8 mm. Scutum blackish. Ratio length/width of hind basitarsus = 3.5. Pupa: Cocoon anterior aperture border higher than posterior border (Fig. 130 D). Length basally 3.7 mm, on dorsum 3.3 mm, maximum length 5.5 mm; gills 0.9-1.2 mm. Frontoclypeus and exposed portion of thorax without tubercles except facial area with small tubercles (Fig. 130 G) and posterior area of thorax with numerous small acute tubercles; cephalic trichomes with 3-7 and thoracic with 8-9 branches. Frontoclypeus basal border reinforced. Gill with 12 branches arranged in 1 plane, emerging from short basal trunk, with acute apex (Fig. 130 K). Larva: Length over 7.0 mm. Antenna longer than cephalic fan stem. Anal ring with about 150 rows of hooks. Rectal papillae with 20-26 diverticula on each lobe. Distribution: Colombia: Cundinamarca. Bionomics: Unknown. Discussion: This species is near S. muiscorum but it can be separated by the numerous tubercles on the frontoclypeus and exposed portion of the pupal thorax, higher frontal aperture on the cocoon, and gill branches arranged very closely in a bunch. Simulium (Trichodagmia) nigrimanum MACQUART (Figs. 128 F-K; 129 L, M; 130 E, L; 131 H-M.) Simulium nigrimanum MACQUART, 1838: 88; SHELLEY et al., 1984: 146-147. Simulium (Trichodagmia) nigrimanum: CROSSKEY & HOWARD, 1997: 79; 2004: 74, MIRANDA ESQUIVEL & COSCARÓN, 2001: 429, 435; HERNÁNDEZ & SHELLEY, 2005: 11-12. Simulium (Grenieriella) nigrimanum: COSCARÓN, 1987: 37. Simulium pruinosum LUTZ, 1910: 250-251, 263, 265-266; PINTO, 1931: 704; PY-DANIEL, 1989a: 256. Simulium (Grenieriella) pruinosum: COSCARÓN, 1991: 280-281. Grenieriell pruinosa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 118. Eusimulium pruinosum: LANE & PORTO, 1939:169, 174-175; LANE & VULCANO, 1943: 436. Simulium spadicidorsum ENDERLEIN, 1934a: 194.

Female: Wing length 2.9-3.6 mm. General coloration grayish brown. Scutum and scutellum light brown with soft gray pollinosity; scutum pilosity tin to silvery, arranged in small groups. Metanotum velvety grayish brown; pleurae brown with gray pollinosity. Legs

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M Figure 131. Larva. A-G: S. (Trichodagmia) lahillei; A: larva, lateral view; B: cephalic apotome; C: basal portion of head dorsally enlarged, D: antenna; E: mandibular teeth; F: postgenal cleft and hypostomium; G: hypostomial teeth of anterior margin. H-M: S. (T.) nigrimanum; H: larva, in lateral view, I: cephalic apotome; J: antenna; K: mandibular teeth, L: postgenal cleft and hypostomium, M: rectal papillae diverticula.

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yellowish brown with distal portion of femora, distal 3/4 of tibia and tarsomeres blackish brown. Abdomen dark brown; tergites VI-IX cereus, shiny with grayish tones. Frons convergent (Fig. 128 F). Fronto-ocular triangle wider than high (Fig. 128 G). Basal portion of cibarium with well-sclerotized border and well-developed teeth on median depressed area, continued by slight serrations laterally (Fig. 128 H). Sensory vesicle of maxillary palp about 1/4 of palpomere length. Sc with 12-15 hairs; claw with large subbasal tooth. Eighth sternite with about 10 setae on each side. Gonapophysis subovoidal with rounded apex (Fig. 128 I). Paraproct longer than wide, rounded apically and with small anterior depression (Fig. 128 J). Genital fork with large posterior branches (Fig. 128 K). Male: Wing length 2.5-2.7 mm. Scutum brown-yellow to dark brown, showing with some light 1+1 submedian blackish stripes; pile golden. Abdomen dark brown with 1+1 silvery spots on tergite II and large spots on V-VIII. Gonocoxite subquadrate and nearly as long as gonostylus; thick gonostylus about 1.9 times longer than wide, with about 4 apical spurs (Fig. 129 L). Ventral plate with well-developed median apical process (Fig. 129 M). Pupa: Cocoon with aperture frontally elevated (Fig. 130 E); length at base = 3.0-3.8 mm, at dorsum 2.0-2.5 mm, maximum length 3.5-4.0 mm; gill 1.4-1.5 mm. Cocoon weave thick but translucent. Frontoclypeus and exposed portion of thorax without tubercles, small tubercles on facial area. Frontal trichomes single, facial bifid, thoracic short with 14 branches. Gill relatively short with 18-20 filamentous branches, emerging from short basal trunk and arranged in bunch (Fig. 130 L). Abdomen without terminal spur. Larva: Maximum length 8.0 mm. Shape as in figure 131 H. Cephalic apotome slightly darkened on posterior half and especially on basal border; median longitudinal and 1+1 submedian spots; the 1+1 basal spots negative (Fig. 131 I). Antenna relatively thin (Fig. 131 J), not surpassing apex of cephalic fan stem; ratio of proximal, medial and distal articles = 1:1.5-1.6:1:1.1. Mandible with several rows of internal teeth (Fig. 131 K), marginal teeth very thin. Hypostomium with median tooth slightly beyond level of lateral teeth; postgenal bridge shorter than hypostomium length (Fig. 131 L). Body integument with scarce and small filiform trichomes. Anal ring with 146 rows of 23-25 hooks each. Lateral sclerite of proleg with about 36 teeth arranged in 17-19 groups. Anal sclerite with abundant hair. Rectal papillae with 17-23 diverticula on each lobe (Fig. 131 M). Distribution: Brazil: Espírito Santo, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, São Paulo, Tocantins; Colombia: Vaupes; Paraguay; Venezuela. Bionomics: This species breeds in small creeks with fast-flowing, clear water 0.200.40 m deep. Females are very noxious to humans. Discussion: Simulium nigrimanum is easy to separate from similar species of the subgenus because the male has a gonostylus with one to four apical spurs and a longer median process on the ventral plate. SHELLEY et al. (1984: 146), based on a study of two females in the MNHN labeled as S. nigrimanum (that were designated as syntypes), decided that pruinosum LUTZ is a synonym. PY-DANIEL (1989: 256) did not agree with this designation because a male is mentioned in the original description of S. nigrimanum. PY-DANIEL (1989) prefers to maintain S. nigrimanum as species inquirenda.

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Simulium (Thyrsopelma) ENDERLEIN (Figs. 132-135; 154.) Thyrsopelma ENDERLEIN, 1934b: 284 (as genus). Type-species: Thyrsopelma brasiliense ENDERLEIN, 1934b: 284 = Simulium orbitale LUTZ, 1910: 231.

Female: Relatively big, wing length 2.0-3.6 mm. General coloration blackish. Scutum reddish brown with gray pollinosity and hair yellow to silvery, grouped, simulating scales. Abdomen black. Frons relatively wide. Fronto-ocular triangle about as high as wide to higher than wide. Basal portion of cibarium smooth and well sclerotized, as for cornuae and sometimes median area. Sensory vesicle of maxillary palp smaller than half of palpomere. Basal sector of R haired; Sc with or without hair. Claw without subbasal tooth. Eighth sternite with abundant hair; gonapophysis subovoidal, generally wider than long. Male: Coloration similar to that of female, showing in some cases black silvery lateral margin and 1+1 anterior spots as wide longitudinal stripes; abdomen blackish with 1+1 lateral silvery spots on segments II and smaller on V-VII. Gonostylus 1.5-2 times longer than gonocoxite, subtriangular, curved on apical third and becoming thinner distally with 1 apical spur. Ventral plate body wider than high with big median process. Median sclerite subrectangular with deep median incision. Endoparameres without teeth. Pupa: Cocoon shoe shaped projected ventrally, with reinforced border of aperture, protecting base of gill branches. Frontoclypeus and exposed portion of thorax with rounded or acuminate or spine-like tubercles on base of frontoclypeus and ventrolateral area of thorax. Cephalic trichomes (2+2 frontal, 1+1 facial) and thoracic (5-6) single and small. Gill branches 12-50, relatively thick, smooth, with apex acute and well sclerotized. Larva: Body cuticle with single hairs, generally with racket shape. Cephalic apotome weakly darkened basally and without special ornamentation. Antenna surpassing apex of cephalic fan stem, with medial article about 2 times longer proximal or distal articles. Cephalic fan rays with teeth of similar length and divergent at sides distally, appearing as 2 rows. Mandible with large preapical teeth nearly as long as apical tooth; marginal teeth large. Postgenal cleft subovoidal, postgenal bridge slightly shorter than hypostomium length. Hypostomium with median tooth lower than lateral teeth, and intermediate lateral teeth very wide. Anal sclerite without scales, anal ring with 120-300 rows with 17-35 hooks each. Rectal papillae with 5-30 diverticula on each lobe. Simulium (Thyrsopelma) is a Neotropical taxon distributed in tropical South America. It is composed of 5 species. It shares several characters with Simulium (Trichodagmia) (MIRANDA ESQUIVEL & COSCARÓN, 2001), and some authors, such as CROSSKEY & HOWARD (1997, 2004) and SHELLEY et al. (2000), include these species in Trichodagmia. Simulium (Hemicnetha) also shows similarities with Thyrsopelma and with Trichodagmia, including the morphology of the gonocoxite, gonostylus and paraproct; pupal cocoon and gill branches; and proportion of antennal articles and large number of hook rows and hooks in the anal ring of the larva. Simulium (Thyrsoplema) also has similarities with Ethiopian Anasolen, Xenosimulium and Freemaniellum (CROSSKEY, 1969; COSCARÓN, 1991; MIRANDA ESQUIVEL & COSCARÓN, 2001). The similarities are the smooth cibarium, haired basal sector of R, absence of a subbasal tooth on the claw, scutum not ornamented; cocoon with anterior projection, absence of spines on the abdominal terminal tergites of the pupa; and absence of ventral tubercles on the larval

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abdomen, elevated number of hook rows in the anal ring, presence of setae on the larval cuticle, and short and blunt teeth on the hypostomium, all showing a strong relationship with the Gondwanian fauna (COSCARÓN, 1991; MIRANDA ESQUIVEL & COSCARÓN, 2004). In the absence of a world revision, we maintain the subgeneric separation, based on morphological differences, as also indicated for the other Neotropical taxa that we have treated. KEYS

TO

SIMULIUM (THYRSOPELMA)

SPECIES

Females 1 – 2(1) – 3(2) – 4(3) –

Scutum dark reddish brown; gonapophysis wider than long (Fig. 132 M); paraproct subtriangular without anterior depression (Fig. 132 N) ............................ ..............................................................................................................Simulium scutistriatum Scutum blackish; paraproct with anterior deep depression, distally with tail-like process (Fig. 132 G); gonapophysis about as long as wide (Figs. 132 D, E, H) .......... 2 Paraproct continued distally, with relatively wide and curved membranous process (Fig. 132 G) ............................................................................................ Simulium hirtipupa Paraproct truncate at short distance from base, with short, thin and straight membranous process (Fig. 132 D) ................................................................................. 3 Claw with small subbasal tooth ........................................................ Simulium perplexum Claw without subbasal tooth (Fig. 132 C) .................................................................... 4 Wing length 1.8-2.5 mm; basal sector of R and Sc without hair .............................. ................................................................................................................... Simulium guianense Wing length 2.8-3.1 mm; basal sector of R with 20-40 hairs, Sc with 11-15 hairs ..................................................................................... Simulium orbitale, Simulium itaunense

Males 1 – 2(1) – 3(1) – 4(3)

Scutum dark brown to blackish, without silvery humeral spots; ventral plate body without lateral processes and with or without short median process (Fig. 133 K) ................................................................................................................................................ 2 Scutum blackish, generally with silvery humeral spots that reach posterior silvery area (Figs. 133 A, G); ventral plate body with lateral processes and long median process (Figs. 133 D, I) ..................................................................................................... 3 Ventral plate without median process; gonostylus with apical spur thin and pointed ................................................................................................... Simulium perplexum Ventral plate with short median process (Fig. 133 K); gonostylus with apical spur thick and blunt apex (Fig. 133 J) ...................................................Simulium scutistriatum Ventral plate with median process subpentagonal and wider at base than at apex (Fig. 130 I); gonostylus relatively thin basally (Fig. 133 H) .......... Simulium hirtipupa Ventral plate with median process pear shaped, wider at apex than at base (Fig. 133 D); gonostylus relatively wide basally (Figs. 133 B, C) ....................................... 4 Scutum with 1+1 silvery spots extended from humeral area to posterior silvery area as wide submedian stripes (Fig. 133 G); basal sector of R without hair ........ ................................................................................................................... Simulium guianense

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Scutum with 1+1 silvery spots reduced mostly to humeral area, not reaching posterior area (Fig. 133 A); basal sector of R with hair ............................................ 5 Wing length 2.5-2.7 mm; scutum with anterior 1+1 silvery spots elongated (Fig. 133 A) …. .................................................................................................. Simulium orbitale Wing length 2.8-3.0 mm; scutum with anterior 1+1 silvery spots rounded ........... ................................................................................................................... Simulium itaunense

Pupae 1 – 2(1) – 3(1) – 4(3) –

Gill with 20-50 branches .................................................................................................. 2 Gill with 12 branches ........................................................................................................ 3 Gill with 21-24 branches (Fig. 134 B); cocoon aperture relatively well elevated anteriorly (Fig. 134 A) ............................................................................. Simulium orbitale Gill with 46-52 branches; cocoon aperture weakly elevated anteriorly (similar to Simulium hitipupa, Fig. 134 F) .............................................................. Simulium itaunense Frontoclypeus and exposed portion of thorax covered by spines (Figs. 134 F, I); gill branches covered by strong spiculae (Fig. 134 H) ................ Simulium hirtipupa Frontoclypeus and exposed portion of thorax without spines; gill branches without spiculae or with very weak spiculae ................................................................ 4 Frontoclypeus and exposed portion of thorax with numerous apically acute tubercles (Figs. 134 L, M); cocoon length basally 4.0-4.5 mm .....................Simulium scutistriatum Frontoclypeus and exposed portion of thorax smooth or only with small rounded tubercles on frontoclypeus base (Fig. 134 C) or posterolaterally .............................. ............................................................................... Simulium guianense, Simulium perplexum

Larvae Not included: Simulium perplexum. 1 – 2(1) –

CLAVES

Maximum length 10.0-10.5 mm; anal ring with 280-300 rows with 28-35 hooks each; body cuticule with sparse trichomes filiform and racket shaped (Fig. 135 S) ..............................................................................................................Simulium scutistriatum Maximum length 6.8-8.8 mm; anal ring with 120-220 rows with 17-31 hooks each; body cuticle with abundant trichomes racket shaped (Figs. 135 G, L) .................. 2 Anal ring with about 220 rows with 28-31 hooks each ................ Simulium hirtipupa Anal ring with 133-170 rows with 17-30 hooks each ................................................... .................................................. Simulium guianense, Simulium itaunense, Simulium orbitale. DE ESPECIES DE

SIMULIUM (THYRSOPELMA)

Hembras 1 –

Escudo castaño rojizo oscuro, gonapófisis más anchas que largas (Fig. 132 M); paraprocto subtriangular sin depresión anterior (Fig. 132 N) ........Simulium scutistriatum Escudo negruzco; gonapófisis aproximadamente tan largas como anchas (Figs. 132 D, E, H); paraprocto con una depresión anterior profunda, generalmente mostrando como teniendo una cola (Fig. 132 G) ....................................................... 2

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Paraprocto continuado distalmente por una prolongación membranosa ancha y curva (Fig. 132 G) ................................................................................. Simulium hirtipupa Paraprocto truncado a corta distancia de la base con una prolongación corta, fina, membranosa (Fig. 132 D) ................................................................................................. 3 Uña con pequeño diente subbasal ................................................... Simulium perplexum Uña sin diente subbasal (Fig. 132 C) ............................................................................. 4 Largo ala 1,8-2,5 mm; sector basal de R y Sc sin pelos ................ Simulium guianense Largo ala 2,8-3,1 mm; sector basal de R con 20-40 pelos; Sc con 11-15 pelos ..... ..................................................................................... Simulium orbitale, Simulium itaunense

Machos 1 – 2(1) – 3(1) – 4(3) – 5(4) –

Escudo castaño oscuro a negruzco; sin manchas humerales plateadas; placa ventral sin prolongaciones laterales y con una corta prolongación mediana (Fig. 133 K) o ausente ................................................................................................................ 2 Escudo negruzco generalmente con manchas humerales plateadas (Figs. 133 A, G); placa ventral con prolongaciones laterales y prolongación mediana larga (Figs. 133 D, I) ............................................................................................................................... 3 Placa ventral sin prolongación mediana; gonostilo con espolón apical fino y aguzado .................................................................................................. Simulium perplexum Placa ventral con una corta prolongación mediana (Fig. 133 K); gonostilo con espolón apical grueso y con ápice romo (Fig. 133 J) ...............Simulium scutistriatum Prolongación mediana de la placa ventral subpentagonal y más ancha en la base que en el ápice (Fig. 130 I); gonostilo relativamente fino basalmente (Fig. 133 H) ................................................................................................................... Simulium hirtipupa Prolongación mediana de la placa ventral en forma de pera, con el ápice más ancho que la base (Fig. 133 D); gonostilo relativamente ancho basalmente (Figs. 133 B, C) .............................................................................................................................. 4 Escudo con 1+1 mancha plateada extendida desde el área humeral hasta el área plateada posterior, como una ancha banda submediana (Fig. 133 G); sector basal de R sin pelos ......................................................................................... Simulium guianense Escudo con 1+1 mancha plateada reducida mayormente al área humeral, no llegando al área posterior (Fig. 133 A); sección basal de R con pelos ................... 5 Largo ala 2,5-2,7 mm; escudo con 1+1 manchas plateadas anteriores, alargadas (Fig. 133 A) ................................................................................................ Simulium orbitale Largo ala 2,8-3,0 mm; escudo con 1+1 manchas plateadas redondeadas ............... ................................................................................................................... Simulium itaunense

Pupas 1 – 2(1) –

Branquias con 20-50 ramas .............................................................................................. 2 Branquias con 12 ramas .................................................................................................... 3 Branquias con 21-24 ramas (Fig. 134 B); abertura del capullo elevado frontalmente (Fig. 134 A) ....................................................................... Simulium orbitale Branquias con 46-52 ramas; abertura del capullo débilmente elevado frontalmente (similar a Simulium hirtipupa, Fig. 134 F) ........................................... Simulium itaunense

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Frontoclípeo y porción expuesta del tórax cubierta por espinas (Figs. 134 F, I); ramas de las branquias cubiertas por fuertes espículas (Fig. 134 H) ........................ ................................................................................................................... Simulium hirtipupa Frontoclípeo y porción expuesta del tórax sin espinas; ramas de las branquias sin espículas o muy débiles ..................................................................................................... 4 Frontoclípeo y porción expuesta del tórax con numerosos tubérculos con ápice aguzado (Figs. 134 L, M); largo capullo en la base 4,0-4,5 mm ................................. ..............................................................................................................Simulium scutistriatum Frontoclípeo y porción expuesta del tórax liso o solamente con pequeños tubérculos redondeados sobre la base del frontoclípeo (Fig. 134 C) o latero posteriormente ................................................... Simulium guianense, Simulium perplexum

Larvas No incluida: Simulium perplexum. 1 – 2(1) –

Largo máximo 10,0-10,5 mm; anillo anal con 280-300 hileras con 28-35 ganchos cada una; cutícula del cuerpo con escasos tricomas filiformes y en forma de raqueta (Fig. 135 S) ..........................................................................Simulium scutistriatum Largo máximo 6,8-8,8 mm; anillo anal con 120-220 hileras con 17-31 ganchos cada uno; cutícula del cuerpo con abundantes tricomas en forma de raqueta (Figs. 135 G, L) .............................................................................................................................. 2 Anillo anal con alrededor de 220 hileras con 28-31 ganchos cada una ................... ................................................................................................................... Simulium hirtipupa Anillo anal con 133-170 hileras con 17-30 ganchos cada una ................................... ................................................... Simulium guianense, Simulium itaunense, Simulium orbitale

Simulium (Thyrsopelma) orbitale LUTZ (Figs. 132 A-F; 133 A-F; 134 A-D; 135 A-G.) Simulium orbitale LUTZ, 1910: 231-234, 251; LUTZ & MACHADO, 1915: 14-46; NEIVA & PENNA, 1916: 93; LUTZ et al., 1918: 107, 109, 165; LUTZ & NUNES TOVAR, 1928: 47; PINTO, 1931: 706, 708, 741, 756; WYGODZINSKY, 1953a: 303. Simulium (Thyrsopelma) orbitale: COSCARÓN, 1987: 39; 1991: 287-288. Simulium nigrimanum (nec MACQUART, 1838: 38): LUTZ, 1909: 137-138, 145-146; LUTZ, 1910: 215; PINTO, 1931: 705-706, 726, 760. Simulium albimanum LUTZ, 1909: 127, 132. Simulium (Eusimulium) orbitale: D’ANDRETTA & D’ANDRETTA, 1946: 85-101. Thyrsopelma brasiliense ENDERLEIN, 1934b: 284; STONE, 1963: 19; VULCANO, 1967: 17. Simulium paraguayense (nec SCHROTTKY, 1909: 63): LANE & VULCANO, 1943: 437. Thyrsopelma orbitale: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 131-132. Simullium (Trichodagmia) orbitale: CROSSKEY & HOWARD, 2004: 74.

Female: Wing length 2.8-3.0 mm. Scutum blackish with whitish pruinosity and hair grouped like scales, silvery to yellowish, without special arrangement. Scutellum, pleurae and metanotum blackish, the latter one shiny nacreous. Legs brown with darker apical portion blackish. Abdomen velvety black on segments I-IV, black, cereus, shiny on V-VII;

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II with silvery pruinosity. Cibarium smooth with sclerotized border (Fig. 132 B). Sc with about 15-20 hairs. Fronto-ocular triangle slightly wider at base than high (Fig. 132 A). Claw without subbasal tooth (Fig. 132 C). Eighth sternite with 24-41 setae on each side, gonapophysis subovoidal, longer than wide and with dense hair (Fig. 132 E), paraproct relatively narrow and rounded apically, ventrally membranous with short tail-like projection (Fig. 132 D), genital fork with distal portion expanded (Fig. 132 F). Male: Wing length 2.5-2.7 mm. Basal portion of R with 20-40 hairs. Scutum blackish with 1+1 diagonal anterolateral silvery spots reaching anterior 1/3, and joined anteriority with silvery lateral and posterior margins (Fig. 133 A). Sc without hair. Ratio of length/ width of hind basitarsus = 6. Gonostylus narrow on distal half (Figs. 133 B, C). Ventral plate with peculiar elongated, subglobose and hairy distal process (Figs. 133 D, E). Endoparameres without teeth (Fig. 133 F). Pupa: Cocoon aperture relatively low anteriorly (Fig. 134 A); length basally: 3.1-3.7 mm, on dorsum 2.2-2.7 mm, gills 0.6-0.7 mm. Frontoclypeus with tubercles on basal 1/3 and laterally (Fig. 134 C); exposed portion of thorax smooth. Cephalic trichomes small and single. Gills with 21-24 branches, emerging without perceptible basal trunk; primary branches very short; branches medially thick, pseudoannulated and bordered by spiculae; apically acute with well-sclerotized terminal spine (Fig. 134 B). Abdominal chaetotaxy as in figure 134 D. Larva: Maximum length 6.7 mm. Shape as in figure 135 A. Cephalic apotome scarcely darkened basally, continued anteriorly by median line (Fig. 135 B). Antennae surpassing apex of cephalic fan stem; ratio of proximal, medial and distal articles = 1:2.2:1-1.1, shape as in figure 135 E. Cephalic fan with about 56 rays. Mandible with second marginal tooth about half of first (Fig. 135 F). Hypostomium with anterior margin straight (Figs. 135 C-D); postgenal bridge slightly longer than half of hypostomium length. Body with hair-like scales racket shaped; length 0.019 mm (Fig. 135 G), arranged in rows on anterior 5 segments and not ordered on following segments. Anal ring with 160166 rows with 25-30 hooks on each. Rectal papillae with 10-15 diverticula on each lobe. Distribution: Argentina: Buenos Aires, Corrientes, Entre Rios, Misiones; Brazil: Mato Grosso do Sul, Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: This species breeds in rivers with large discharge and fast flow attached to submerged tree branches at a depth of about 0.5 m. Female feeding habits are unknown. In Buenos Aires province, the highest prevalence is in autumn and winter (COSCARÓN et al., 2000). Discussion: The closest species is S. guianense, but it can be differentiated by the pupal gill with 12 branches and male with elongated silvery spots the length of the scutum. Simulium (Thyrsoplema) guianense WISE (Figs. 133 G; 134 E; 135 G 2.) Simulium guianense WISE, 1911: 248; 1912: 43; SMART, 1940: 4. Simulium (Trichodagmia) guianense: SHELLEY et al., 1997: 40-43; 2000: 183-184; CROSSKEY & HOWARD, 1997: 97; 2004: 74. Simulium (Thyrsopelma) guianense: COSCARÓN, 1987: 39. Thyrsopelma guianense PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

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Figure 132. Female. A-F: S. (Thyrsopelma) orbitale; A: fronto-ocular triangle; B: cibarium; C: claw; D: genitalia, ventral view showing eighth sternite and gonapophysis, cercus and paraproct (at left mounted on slide) and genital fork; E: eighth sternite and gonapophysis; F: genital fork. G-H: S. (T.) hirtipupa; G: cercus and paraproct; H: eighth sternite and gonapophysis. I-O: S. (T.) scutistriatum; I: hair of scutum; J: palpomere with sensory vesicle, K: claw; L: cibarium; M: eighth sternite and gonapophysis; N: cercus and paraproct; O: genital fork.

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Figure 133. Male. A-F: S. (Thyrsopelma) orbitale; A: scutum; B: genitalia in ventral view showing gonocoxite and gonostylus; C: gonocoxite and gonostylus; D-E: ventral plate, ventral view (D) and lateral view (E); F: endoparameres and aedeagal membrane. G: S. (T.) guianense, scutum. H-I: S. (T.) hirtipupa; H: gonocoxite and gonostylus; I: ventral plate. J-K: S. (T.) scutistriatum; J: gonocoxite and gonostylus; K: ventral plate.

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Simulium pintoi D’ANDRETTA & D’ANDRETTA, 1946: 101-103; SHELLEY et al., 1979: 41-43; LACEY & CHARLWOOD, 1980: 496; LACEY, 1981: 330. Simulium (Thyrsoplema) pintoi COSCARÓN, 1987, 39; 1991: 288-290. Simulium ortizi RAMÍREZ PÉREZ, 1971: 336-339. “A”, “B”, “C”,”D” CHARALAMBOUS et al.,1996: 111-120 (Simulium guianense cytoform).

Female: Wing length 1.8-2.5 mm. Coloration of scutum, scutellum, metanotum pleurae and abdomen as in S. orbitale. Sc without hair. Basal sector of R with single row of hairs, Cibarium, paraproct and cercus as in S. orbitale; eighth sternite with about 20 setae on each side. Male: Wing length 1.9-2.3 mm. Scutum velvety black with 1+1 silvery submedian stripes, extending from humeral callus on anterolateral area to posterior area; another silvery stripe on lateral and posterior margin (Fig. 133 G). Basal sector of R and Sc without hair. Ratio of length/width of hind basitarsus = 6. Genitalia as in S. orbitale. Pupa: Cocoon as in S. orbitale; length basally 2.5-3.0 mm; gills 1.0 mm. Frontoclypeus with small tubercles on basal 1/3, as in S. orbitale and on thorax only dorsally on midline. Gills with 12 branches arranged as in S. orbitale (Fig. 134 E). Larva: Maximum length 6.0 mm. Cephalic apotome, mandible and antenna as in S. orbitale. Cephalic fan with 54-58 rays. Hypostomium border with teeth arranged as in S. hirtipupa. Body cuticle with abundant trichomes racket shaped, bigger than in other species, length 0.028 mm (Fig. 135 G 2). Anal ring with about 140 rows with 23-25 hooks each. Rectal papillae with 5-12 diverticula on each lobe. Distribution: Brazil: Amapá, Amazonas, Espírito Santo, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Paraná, Rio de Janeiro, Roraima, Santa Catarina, São Paulo, Tocantins; Venezuela: Amazonas, Bolivar; French Guiana; Suriname. Bionomics: The larval stages were collected in moderately fast creeks 3-5 m wide and 0.4-0.5 m deep, but they can be found in larger streams (SHELLEY et al. 2000) with sandy bottoms in yellowish water and attached to aquatic vegetation especially Podostemaceae. Females are zoophilic and anthropophilic. They are important vectors of Onchocerca volvulus. Discussion: The closest species is S. orbitale, under which the differences were explained. Synonymy with S. pintoi was commented on by SHELLEY et al. (1997). Studies of CHARALAMBOUS et al. (1996) showed the presence of 4 cytospecies, which could explain the differences in feeding behavior. Simulium (Thyrsopelma) itaunense D’ANDRETTA & GONZÁLEZ Simulium itaunensis D’ANDRETTA & GONZÁLEZ, 1964: 106-108; VULCANO, 1967: 14; 1977: 292; PYDANIEL et al., 1985: 98-99. Simulium (Thyrsopelma) itaunense: COSCARÓN, 1987: 39; 1991: 290-291. Thyrsopelma itaunense: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119; STRIEDER, 2004: 131. Simulium (Trichodagmia) itaunense: CROSSKEY & HOWARD, 1997: 79; 2004: 74; HERNÁNDEZ et al., 2005: 3-12.

Female: Wing length 2.8-3.1 mm. Scutum blackish, humeri brown with grey pruinosity and recumbent grouped whitish hairs. Scutellum and metanotum black, the latter with silver pruinosity. Pleura blackish gray pruinose. Legs brown with darkened portion dark brown. Abdomen shiny blackish. Cibarium, wings, claws and genitalia as in S. orbitale.

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E Figure 134. Pupa. A-D: S. (Thyrsopelma) orbitale; A: Pupa lateral view; B: thorax, anterodorsal portion and gill; C: frontoclypeus; D; abdominal chaetotaxy. E: S. (T.) guianense, gill. F-I: S. (T.) hirtipupa; F: pupa; G: gill; H: gill apex enlarged view; I: frontoclypeus. J-M: S. (T.) scutistriatum; J: lateral view; K: gill; L: frontoclypeus; M: integument tubercles enlarged.

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Male: Wing length 2.8-3.0 mm. Scutum velvety black, nacreous shiny on lateral margins and prescutellum region, and 1+1 silvery rounded spots on anterior 1/3; humeri with silver pruinosity. Scutellum, metanotum and pleurae blackish. Abdomen velvety blackish with 1+1 lateral silvery spots on tergites II, VI-VIII, and anterior submedian are on segments IV-V. Basal sector of R haired and Sc with hair only on base. Genitalia similar to those of S. orbitale, but paraproct with distal membranous area more evident. Pupa: Cocoon aperture weakly elevated anteriorly, in lateral view similar to S. hirtipupa. Length basally 4.5-4.9 mm; dorsally 3.4-4.3 mm; gills 0.7-1.2 mm. Frontoclypeal and thoracic tubercles and trichomes as in S. orbitale, thoracic trichomes robust. Gill with 46-52 branches, relatively thick and short, emerging from thick and elongated basal trunk horizontally arranged; branches apically acute and well sclerotized, with spine-like aspect, similar to arrangement in S. orbitale but with more branches. Larva: Maximum length 6.8 mm. Antenna, hypostomial teeth and cuticular body hair as in S. orbitale. Cephalic fan with 48 rays. Postgenal bridge slightly shorter than hypostomium length. Lateral sclerite of thoracic proleg with about 65 teeth arranged in 14-16 groups. Anal ring with about 133 rows of 17-22 hooks each. Rectal papillae with diverticula on each lobe. Distribution: Brazil: Rio Grande do Sul, Santa Catarina, São Paulo. Bionomics: This species breeds in clean creeks with fast, semi-torrential flow, attached to stones and aquatic vegetation. Female feeding habits are unknown. Discussion: The closest species is S. guianense. Males of S. guianense can be separated by the elongated anterior silvery spots and the pupa by the 12 gill branches. Simulium (Thyrsopelma) hirtipupa LUTZ (Figs. 132 G, H; 133 H, I; 134 F-I; 135 H-L.) Simulium hirtipupa LUTZ, 1910: 200, 260, 263-264; PINTO, 1931: 702, 756; VARGAS, 1945a: 146; VULCANO, 1967: 13, 1971: 292; MAIA-HERZOG et al., 1985: 483-490. Simulium (Thyrsopelma) hirtipupa: COSCARÓN, 1987: 39; 1991: 292-293. Thyrsopelma hirtipupa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119. Simulium (Trichodagmia) hirtipupa: CROSSKEY & HOWARD, 1997: 79; 2004: 74.

Female: Wing length 2.7-2.9 mm. Scutum black with thin gray pruinosity and thin median and 1+1 sublateral stripes extending 2/3 of scutum length, scutum hair grouped like scales in longitudinal rows. Scutellum black, metanotum velvety blackish. Pleurae blackish with gray pollinosity. Abdomen dark grayish brown with tergal plates blackish. Cibarium as in S. orbitale. Fronto-ocular triangle short, higher than wide. Basal section of R and Sc without hair. Paraproct apically continued by relatively wide and curved membranous process with abundant hair (Fig. 132 G). Eighth sternite with 12-16 setae on each side, gonapophysis subtriangular, about as long as wide at base with blunt apex (Fig. 132 H). Male: Wing length 2.6-2.8 mm. Scutum blackish with much gray pollinosity and golden pile, with frontal illumination becoming leaden gray, and with posterior light blackish with 1+1 submedian grayish stripes on posterior 2/3. Scutellum and metanotum as in female. Abdomen black with 1+1 silvery spots on segments II, V-VIII. Gonostylus nearly 1.5 times longer than gonocoxite, becoming thinner distally (Fig. 133 H), basally narrower than in S. orbitale. Ventral plate with 1+1 lateral and 1 wider median processes (Fig. 133 I).

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Figure 135. Larva. A-G: S. (Thyrsopelma) orbitale; A: larva in lateral view; B: head, dorsal view; C: hypostomium; D: teeth of hypostomium, anterior margin; E: antenna; F: mandibular teeth; G: integument trichomes (1: S. (T.) orbitale; 2: S. (T.) guianense). H-L: S. (T.) hirtipupa; H: larva, lateral view; I: teeth of hypostomium, anterior margin; J: antenna; K: mandibular teeth; L: integument trichomes. M-S: S. (T.) scutistriatum; M: postgenal cleft and hypostomium; N: teeth of hypostomium, anterior margin; O: antenna; P: mandibular teeth; Q: dorsal posterior of head and cervical sclerites; R: anal sclerite (ventral branches up); S: integument trichomes.

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Pupa: Cocoon aperture lower anteriorly (Fig. 134 F), weave thick with wrinkled surface. Length basally 4.0-5.0 mm, dorsally 2.6-3.6 mm, gills 0.7 mm. Frontoclypeus and thorax with long spines (Fig. 134 I), these spines also in other places on head but smaller. Integument without tubercles. Gill with 12 branches relatively thick, with spine on apex and annulations covered with irregular shaped spiculae (Figs. 134 G, H). Larva: Maximum length 7.3-8.8 mm. General aspect as in figure 135 H. Cephalic apotome brown, darker at base. Postgenal bridge slightly shorter than hypostomium length; teeth with anterior margin of hypostomium very short (Fig. 135 I). Antennae surpassing length of cephalic fan stem; shape as in figure 135 J, ratio of proximal, medial and distal articles = 1:2-2.2:1-2. Mandible with long and thin first marginal tooth (Fig. 135 K). Cephalic fan with 55-60 rays. Body integument with petaloid hairs fewer in number than in S. orbitale and S. pintoi, length about 0.021 mm (Fig. 135 L). Lateral sclerite of thoracic proleg with 54-56 teeth arranged in 13-15 groups. Anal ring with 220-246 rows with 28-36 hooks each. Rectal papillae with 12-14 diverticula on each lobe. Distribution: Brazil: Bahia, Minas Gerais, São Paulo. Bionomics: The species breeds in fast-flowing, shallow creeks with rocky bottoms. Female feeding habits are unknown. Discussion: Adults are similar to those of several species of the subgenus, but the peculiar spines on the head and thorax of the pupa and the cocoon and pupa gill shape permit recognition of S. hirtipupa. Simulium (Thyrsopelma) perplexum SHELLEY, MAIA-HERZOG, LUNA DIAS & COUCH Simulium (Trichodagmia) perplexum SHELLEY et al., 1989: 343. CROSSKEY & HOWARD, 1997: 79; 2004: 74; SHELLEY et al., 2004: 446, 453, 469. Thyrsopelma perplexa: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

(Diagnosis based on original description). Female: Wing length 2.3-2.8 mm. Coloration and morphology similar to that for S. guianense, but with 1+1 lateral dark transverse spots on sternites II-VII, scutum hair arranged regularly in small groups, absence of hair on basal sector of R, presence of short subbasal tooth on hind leg claw and paraprocts without anterior tail-like projection. Male: Wing length 2.1-2.6 mm. Coloration and general morphology similar to that of S. guianense. Scutum with anterior illumination velvety black with dark longitudinal median line on anterior 2/3, changing light to posterior position shows light grey. Pilosity arranged in groups of hair-like scales, shiny golden to green intermixed with thin dark longer hairs on anterior and posterior edge. Gonostylus as long as gonocoxite; gonostylus with base thicker than in S. guianense and with apical spur pointed; ventral plate without median distal process. Pupa: Similar to S. guianense but gill branch annulations with border continued anteriorly as small processes instead of spiculae as in S. guianense. Larva: Unknown. Distribution: Guyana. Bionomics: Unknown. Discussion: This species is very close to S. guianense. In males of S. guianense, the biggest differences are in male coloration, the 1+1 silvery longitudinal stripes extended

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from the humeri to the posterior area, the gonostylus more elongated and the ventral plate with a median distal process. The female of S. guianense has a single row of setae on the basal section of R, a tail-like projection on the paraproct and the claw of the hind leg without a subbasal tooth. Simulium (Thyrsopelma) scutistriatum LUTZ (Figs. 132 I-O; 133 J, K; 134 J-M, 135 M-S.) Simulium scutistriatum LUTZ, 1909: 133-134; 1910: 225-226; LUTZ, 1922: 90; PINTO, 1931: 712, 725, 726, 762; VARGAS, 1945a: 195; VULCANO, 1967: 20; 1971: 293; MAIA-HERZOG et al., 1984: 350-351. Eusimulium scutistriatum: LANE & PORTO, 1939: 169, 173-174, LANE & VULCANO, 1943: 436. Simulium (Thyrsopelma) scutistriatum: COSCARÓN, 1987: 39; 1991: 294-296. Simulium (Trichodagmia) scutistriatum CROSSKEY & HOWARD, 1997: 79; 2004: 74. Thyrsopelma scutistriata: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

Female: Wing length 3.0-3.6 mm. Scutum dark reddish brown, without special ornamentation, with golden to tin, small, shiny pile, grouped, simulating scales (Fig. 132 I) and arranged in longitudinal rows, not continuous. Scutellum yellowish brown and metanotum velvety blackish brown. Pleurae dark grayish brown with gray pollinostity. Legs brown with darker areas dark brown and pale yellowish. Abdomen brown with tergal plates dark brown. Fronto-ocular triangle about as wide as high. Basal sector of R haired, Sc with about 40 hair. Sensory vesicle of maxillary palp shorter than half of palpomere length (Fig. 132 J). Claw without subbasal tooth (Fig. 132 K). Cibarium smooth with cornuae well sclerotized (Fig. 132 L). Eighth sternite with concavity and well-sclerotized median area and with about 26 setae on each side; gonapophysis subovoidal, wider than long (Fig. 132 M). Paraproct subtriangular, without depression on anterior border (Fig. 132 N). Genital fork with distal portion of anterior branches very wide (Fig. 132 O). Male: Wing length 2.9-3.3 mm. Coloration as in female but darker. Scutum dark brown with dark pollinosity increased on posterior region, with 1 median and 1+1 submedian longitudinal stripes, the last ones not reaching anterior margin; with changing illumination, scutum shows grayish pollinosity. Abdomen blackish with 1+1 silvery spots on segment I, VVII. Gonostylus relatively thick and shorter than in other species of subgenus (Fig. 133 J). Ventral plate body proportionately long and median process shorter (Fig. 133 K). Pupa: Cocoon with aperture inclined basally (Fig. 134 J). Length basally 4.0-4.5 mm, dorsally 3.8-4.0 mm, gill 1.2 mm. Weave resistant, threads not visible. Frontoclypeus and exposed portion of thorax with abundant, apically acute tubercles (Figs. 134 L-M); cephalic and thoracic trichomes single and relatively short. Gills with 12 branches, relatively short, with thick and acute apices (Fig. 134 K). Larva: Maximum length 10.0-10.5mm. Cephalic apotome darkened on median line and base, with positive spots. Antenna as long as or longer than cephalic fan stem; ratio of proximal, medial and distal articles 1:2-2.3:0.9-1. Postgenal bridge about same length as hypostomium (Fig. 135 M); hypostomial anterior border with large lateral teeth (Fig. 135 N); antenna as in figure 135 O; marginal teeth subequal in shape and relatively short and thick (Fig. 135 P). Cephalic fan with about 53 teeth. Cervical sclerite wide (Fig. 135 Q). Anal ring with 280-300 rows with 28-35 hooks each. Rectal papillae with 30 diverticula on each lobe. Anal

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sclerite with hair and scales (Fig. 135 R). Body integument with hair racket shaped, intermixed with filamentous hairs (Fig. 135 S). Lateral sclerite of thoracic proleg with 56 teeth. Distribution: Brazil: Goiás, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, São Paulo, Tocantins; Colombia; Paraguay; Venezuela. Bionomics: The larval stages can be found in fast-flowing creeks, attached to roots and stones. Females were collected on horses and around humans. Discussion: The similar species of the subgenus can be differentiated in the female by the blackish scutum and shorter paraprocts with the distal portion membranous; in the male by the blackish scutum with wide silvery stripes or big humeral spots and the ventral plate body with lateral and large median processes; and in the pupa by the absence of tubercles with acute apices on the frontoclypeus and thorax. Species inquirenda Simulium albopictum LANE & PORTO Simulium albopictum LANE & PORTO, 1940: 193-194. Simulium (Thyrsopelma) albopictum: COSCARÓN, 1987: 39; 1991: 296. Simulium (Trichodagmia) albopictum: CROSSKEY & HOWARRD, 2004: 74. Thyrsopelma albopicta: PY-DANIEL & MOREIRA SAMPAIO, 1995: 119.

Simulium albopictum was described based on 3 females deposited in Facultade de Saude Publica of São Paulo University. They are externally similar to those of S. guianense but without knowledge of the other stages, it is impossible to be sure if they are conspecific. We prefer to maintain S. albopictum as species inquirenda. Distribution: Brazil: Paraná. Species not placed to subgenus because of insufficient information Simulium costaricense SMART, 1944: 132 (n. nom. Acropogom rufidorsum ENDERLEIN, 1936: 119) (Costa Rica). Scarce information does not permit us to place S. costacricense in any supraspecific taxon; it probably falls into the S. (Ectemnaspis) DINELLII species group, based on the scutum coloration. Simulium diaznajerai VARGAS, 1943a: 136 (n. nom. Odagmia tenuifrons ENDERLEIN, 1936: 130) (Mexico). Based on the description, it is impossible to identify the species. Simulium nigrifemur (ENDERLEIN), 1936: 118 (Acropogon). (Chile: Cauquenes). With an insufficient original description and without the location of the type material, species identification is doubtful.

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Tlalocomyia revelata Mayacnephia aguirrei atzompensis fortunensis grenieri mixensis muzquicensis pachecolundi roblesi salasi tadai

Araucnephia montana iberaensis Araucnephioides schlingeri Cnesia dissimilis ornata pusilla Cnesiamima atroparva Paraustrosimulium anthracinum Lutzsimulium flavopubescens hirticosta pernigrum Kempfsimulium simplicicolor

Figure 136. Tlalocomyia, Mayacnephia, Araucnephia, Araucnephioides, Cnesia, Cnesiamima, Paraustrosimulium, Lutzsimulium, Kempfsimulium.

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Simulium (Pternaspatha)

“NIGRISTRIGATUM GROUP”

S. colopaxi S. deagostini S. diamantinum S. dureti S. horcochuspi S. limay S. nigristrigatum S. pichi S. pulchrum S. simile S. strigidorsum

Figure 137. Simulium (Pternaspatha), nigristrigatum species group.

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Simulium (Pternaspatha)

“NEMORALE GROUP” S. albicinctum S. albilineatum S. annulatum S. barbatipes S. bordai S. caprii S. hectorvargasi S. herreri S. luchoi S. nemorale S. philippii S. prodexargenteum S. quechuanum S. stelliferum S. yacuchuspi

Figure 138. Simulium (Pternaspatha), nemorale species group.

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Simulium (Notolepria)

S. blantoni S. cuasiexiguum S. exiguum S. llutense S. paraguayense S. subexiguum S. gonzalezi

Figure 139. Simulium (Notolepria).

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Simulium (Chirostilbia)

“PERTINAX GROUP” S. bifenestratum S. distinctum S. empascae S. friedlanderi S. laneportoi S. obesum S. pertinax

“SUBPALLIDUM GROUP”

S. riograndense

S. acarayense

S. serranum

S. papaveroi

S. spinibranchium

S. subpallidum

Figure 140. Simulium (Chirostilbia), pertinax and subpallidum species group.

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Simulium (Inaequialium)

“INAEQUALE GROUP” S. beaupertuyi S. clarki S. clavibranchium S. diversibranchium S. inaequale S. leapoldense S. mariavulcanoae S. nahimi S. nogueirai

“BOTULIBRANCHIUM GROUP”

S. parimaensis

S. botulibranchium

S. rappae

S. petropoliense

S. subclavibranchium

S. souzalopesi

S. subnigrum S. travassosi

Figure 141. Simulium (Inaequialium), inaequale and botulibranchium species group.

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Simulium (Psaroniocompsa)

“INCRUSTATUM GROUP” S. anamariae S. angrense S. auripellitum S. auristriatum S. bonaerense

“SIOLII GROUP”

S. brevifurcatum

S. damascenoi

S. fuliginis

S. quaporense

S. incrustatun

S. lourencoi

S. jujuyense

S. siolii

S. limbatum

S. tergospinosum

S. minuanum S. schmidtmummi

Figure 142. Simulium (Psaroniocompsa), incrustatum and siolii species group.

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Simulium (Cerqueirellum)

S. S. S. S. S. S. S. S. S. S. S.

amazonicum argentiscutum chaquense cuneatum delponteianum ganalesense minusculum oyapockense raraimense sanguineum venezuelense

Figure 143. Simulium (Cerqueirellum).

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Simulium (Coscaroniellum)

“QUADRIFIDUM GROUP” S. S. S. S. S. S.

cauchense cerradense dolianhanhani goeldii quadrifidum ulyssesi

“QUADRIVITTATUM GROUP” S. quadrivittatum

Figure 144. Simulium (Coscaroniellum), quadrifidum and quadrivittaum species group.

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Simulium (Ectemnaspis)

“BICOLORATUM GROUP” S. anaimense S. antonii S. arcabucense S. bicoloratum S. bicornutum S. cormonsi S. furcillatum S. ignescens S. jaimeramirezi S. pautense S. pifanoi S. roquemayu S. rubiginosum S. tolimaense S. tunja

Figure 145. Simulium (Ectemnaspis), bicoloratum species group.

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Simulium (Ectemnaspis)

“ROMANAI GROUP” S. albanense S. gabaldoni S. lutzianum S. mayuchuspi S. romanai S. wolffhuegeli

Figure 146. Simulium (Ectemnaspis), romanai species group.

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Simulium (Ectemnaspis)

“PERFLAVUM GROUP” S. kabanayense S. maroniense S. perflavum S. rorotaense S. suarezi S. trombetense

Figure 147. Simulium (Ectemnaspis), perflavum species group.

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Simulium (Ectemnaspis)

“DINELLII GROUP” S. antillarum S. dinellii S. nuneztovari S. ochraceum S. shewellianum

Figure 148. Simulium (Ectemnaspis), dinellii species group.

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Simulium (Psilopelmia)

“BLANCASI GROUP” S. blancasi S. penai S. tenuipes

“ESCOMELI GROUP” S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S.

bivittatum bobpetersoni callidum dandrettai downsi dugesi escomeli gonzalezherrejoni haematopotum iracouboense longithallum mangabeirai mediovittatum ochoai panamense pseudocallidum samboni travisi trivittatum veracruzanum zempoalense

Figure 149. Simulium (Psilopelmia), escomeli and blancasi species group.

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Simulium (Aspathia)

S. anduzei S. bustosi

S. metallicum S. morae

S. costalimai

S. parrai

S. covagarciai

S. piperi

S. hetchi

S. puigi

S. horacioi

S. putre

S. hunteri

S. racenisi

S. iriartei

S. ruizi

S. jacumbae

S. sandyi

S. jobbinsi

S. spilmani

S. kompi

S. tricorne S. wygoi

S. marquezi S. matteabranchium

Figure 150. Simulium (Aspathia).

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Simulium (Hemicnetha)

“PAYNEI GROUP” S. S. S. S.

bricenoi paynei rubrithorax virgatum

“MEXICANUM GROUP” S. S. S. S. S. S. S. S. S. S. S. S.

biuxinisa earlei freemani hinmani keenani mexicanum lobatai placidum seriatum smarti hieroglyphicum yepocapense

“BRACHYCLADUM GROUP” S. S. S. S. S.

brachycladum cristalinum guerrerense pulverulentum solarii

“OVIEDOI GROUP” S. oviedoi S. rivasi

Figure 151. Simulium (Hemicnetha), paynei, mexicanum, brachycladum and oviedoi species group.

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Simulium (Hearlea) S. S. S. S. S. S. S. S. S. S.

ayrozai burchi canadense capricome carolinae contrerense chiriquiense dalmati delatorrei estevezi

S. S. S. S. S. S. S. S. S. S.

ethelae gorirossiae johnsoni juarezi larvispinosum menchacai microbranchium nigricome paracarolinae temascalense

Figure 152. Simulium (Hearlea).

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Simulium (Trichodagmia)

S. huairayacu S. lahillei S. muiscorum S. nigrimanum S. sumapazense S. townsendi S. wygodzinskyorum

Figure 153. Simulium (Trichodagmia).

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Simulium (Thyrsopelma)

S. S. S. S. S. S.

quianense hirtipupa itaunense orbitale perplexum scutistriatum

Figure 154. Simulium (Thyrsopelma).

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7 REFERENCES ADLER, P.H., CURRIE, D.C. & D. M. WOOD (2004): The black flies (Diptera: Simuliidae) of North America. - Cornell University Press, Ithaca, New York, 941 pp. ALENCAR, Y.B., VEIGA LUDVIG, T.A., SOARES, C. & N. HAMADA (2001): Stomach content analysis of Simulium perflavum ROUBAUD 1905 (Diptera: Simuliidae) larvae from stream in Central Amazônia, Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 96: 571-596. ALVAN-AGUILAR, M.A., HAMADA, N., ADLER, P.H. & S. L.B. LUZ (2005): Cytotaxonomy of Simulium cauchense FLOCH & ABONNENC and Simulium quadrifidum LUTZ. (Diptera: Simuliidae) in Brazilian Amazonia. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 100: 249-257. ANDRADE, C.F. (1989): Manejo integrado de borrachudos. - Seminario sobre insectos e acaros. Actas XI Congreso Brasileiro de Entomologia, Campinhas, Fundação Cargill: 141-157. ANDUZE, P.J. (l947): Simulium matteabranchia. - Boletin de la Academia de Ciencias Físicas, Matemáticas y Naturales 10 (30): 374-376. ARTEAGA, L.T. & P. MUÑOZ DE HOYOS (1999): New cytotype in the Simulium metallicum complex (Diptera: Simuliidae) from Cundinamarca, Colombia. - Journal of Medical Entomology 36: 133-140. BARBOSA DE ALMEIDA, J.B. (1979): Simulium cerqueirai, nomen novum para Simulium septentrionale CERQUEIRA & ALMEIDA, 1970 (Diptera, Simuliidae). - Acta Amazônica 9 (4): 789. BARRETO, P. (1969): The species of black flies found in Colombia (Diptera: Simuliidae). - Journal of the New York Entomological Society 77 (1): 31-35. BECKERT, T. (1908): Dipteren der Kanarischen Inseln. - Mitteilungen aus dem Zoologischen Museum in Berlin 4:1-180 + 4 plates. BELLARDI, L. (1859): Saggio di ditterología messicana. Parte. I. - Memoire della Reale Accademia delle Scienze di Torino, Serie 2, 19: 1-80, 2 pls. BELLARDI, L. (1862): Saggio di ditterologia messicana. Appendice. - Memoire della Reale Accademia delle Scienze di Torino, Serie 2, 21: 1-18, 1 pl. BEQUAERT, J. (l934): Part III. Notes on the black-flies of Simuliidae, with special reference to those of the Onchocerca region of Guatemala. - In: STRONG, R.P., SOUNDGROUND, J.H., BEQUAERT, J.C. & M. MUNOZ OCHOA (eds): Onchocerciasis, with special reference to the Central American form of the disease: l75-224. Contributions from the Department of Tropical Medicine and the Institute for Tropical Biology and Medicine 6: 1-234. BEQUAERT, J. (1945): Dr. LUIS VARGAS on American black-flies - a review, with critical notes (Diptera). - Bulletin of the Brooklyn Entomological Society 40: 111-115. BIGOT, E. (1888): Diptères. - In: Mission Scientifique du Cap Horn 1882-1883. - Gauthier-Villars et Fils, Paris 6, Zoology: 1-45.

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BRICENO-IRAGORRY, L. (1941): Nota sobre Simuliidae (Diptera), con citación de una nueva especie para el país y lista de las conocidas en Venezuela. - Boletin del Laboratorio de la Clínica “Luis Razetti” 2 (6): 110-118. BUENO, M.L., MONCADA L. & P. MUÑOZ DE HOYOS (1979): Simuliidae (Insecta - Diptera) de Colombia I. Nueva especie de Simulium (Hemicnetha). - Caldasia 12 (60): 581-594. CAMPOS, J. & C.F.S. ANDRADE (2001): Considerações sobre os simulideos (Diptera, Nematocera) e seu controle. - Entomologia e Vectores 8 (1): 27-50. CERQUEIRA, N. & F. BARBOSA DE ALMEIDA (1970): Simuliidae de Amazônia. V. Descrição de Simulium septentrionale sp. n. (Diptera, Nematocera). - Boletim do INPA (Manaus), Patologia Tropical 2: 1-11. CERQUEIRA, N. & J.A. NUNES DE MELLO (1967): Simuliidae da Amazônia (II). Descrição de Simulium goeldii sp. n. (Diptera, Nematocera). - Amazoniana 1 (2): 125-130. CERQUEIRA, J. & J.A. NUNES DE MELLO (l968): Simuliidae de Amazônia (IV). Descrição de Simulium fulvinotum sp. n. (Diptera, Nematocera) - Amazoniana 1 (3): 205-210. CHARALAMBOUS, M., SHELLEY, A.J. & M. ARZUBE (1993a): The distribution and taxonomic status of chromosomal forms of the onchocerciasis vector Simulium exiguum (Diptera: Simuliidae). Medical and Veterinary Entomology 7: 299-305. CHARALAMBOUS, M., SHELLEY, A.J., MAIA-HERZOG, M. & A.P.A. LUNA DIAS (1996): Four new cytotypes of the onchocerciasis vector blackfly Simulium guianense in Brazil. - Medical and Veterinary Entomology 10: 111-120. CHARALAMBOUS, M., LOWRY, C.A., LOWELL, S., SHELLEY, A.J. & M. ARZUBE (l997): The value of larval head pattern for differentiating Simulium exiguum s. l. and S. gonzalezi (Diptera: Simuliidae) in the onchocerciasis focus of Ecuador. - Bulletin of Entomological Research 87: 19-24. CHARALAMBOUS, M., READY, P.D., SHELLEY, A.J., ARZUBE, M. & C.A. LOWRY (1993b): Cytological and isoenzyme analysis of the Bucay and Quevedo cytotypes of the onchocerciasis vector Simulium exiguum (Diptera: Nematocera) in Ecuador. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 88: 39-48. CONN, J., ROTHFELS, K.H., PROCUNIER, W.S. & H. HIRAI (1989): The Simulium metallicum species complex (Diptera: Simuliidae) in Latin America: a cytological study. - Canadian Journal of Zoology 67: 1217-1245. COQUILLETT, D.W. (1902): New Diptera from North America. - Proceedings of the United States National Museum 25 (1280): 83-l26. COSCARÓN, S. (1968): Ampliación del área de distribución de Simuliidae (Diptera) en Argentina. Revista de la Sociedad Entomológica Argentina 30: 65-67. COSCARÓN, S. (1971): Notas sobre Simúlidos Neotropicales I. Sobre una nueva especie del Norte Argentino (Diptera, Simuliidae). - Revista de la Sociedad Entomológica Argentina 33: 33-41. COSCARÓN, S. (1976a): Las especies de “jerjeles” (Simuliidae, Diptera, Insecta) de la zona de Arica. - Idesia 4: 25-34. COSCARÓN, S. (1976b): Notas sobre Simúlidos Neotropicales VI. Sobre dos especies nuevas de jejenes de la provincia de Misiones, Argentina (Diptera, Insecta). - Revista de la Sociedad Entomológica Argentina 35: 147-154. COSCARÓN, S. (1980): Notas sobre Simúlidos Neotropicales IX. Sobre un grupo de especies próximo al subgénero Simulium (Inaequalium) (Diptera - Insecta). - Revista de la Sociedad Entomológica Argentina 39 (3-4): 293-302.

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COSCARÓN, S. (1981a): Notas sobre Simúlidos Neotropicales XI. Sobre el subgénero Simulium (Chirostilbia) ENDERLEIN, con la descripción de dos especies nuevas del S.E. del Brasil (Diptera - Insecta). - Revista de la Sociedad Entomológica Argentina 40 (1-4): 157-164. COSCARÓN, S. (1981b): Insecta, Diptera, Simuliidae. - In: RINGUELET, R. (ed.): Fauna de Agua Dulce de la Republica Argentina. Vol. 38 (1). FECIC, Buenos Aires: 105 pp. COSCARÓN, S. (1982): Notas sobre Simúlidos Neotropicales X. Sobre un grupo peculiar de especies del género Simulium del S.E. de Brasil, Paraguay y N.O. de Argentina (Diptera - Insecta). Revista de la Sociedad Entomológica Argentina 41 (1-4): 65-76. COSCARÓN, S. (1983): Notas sobre Simúlidos Neotropicales XII. Las especies de Simulium “grupo amazonicum” de Argentina, Sur de Brasil y Paraguay (Simuliidae - Diptera). - Revista de la Sociedad Entomológica Argentina 42 (1-4): 89-99. COSCARÓN, S. (1984): Revisión del Subgénero Simulium (Ectemnaspis) ENDERLEIN (Simuliidae, Diptera, Insecta). - Revista de la Sociedad Entomológica Argentina 43 (1-4): 238-325. COSCARÓN, S. (1985): Sobre la larva de Cnesiamina atroparva EDWARDS y su significado para interpretar las relaciones filogenéticas de los simúlidos notogeicos (Diptera - Insecta). Boletín de la Sociedad de Biología, Concepción, Chile 56: 67-70. COSCARÓN, S. (1987): El género Simulium LATREILLE en la región Neotropical: Análisis de los grupos supraespecíficos, especies que los integran y distribución geográfica (Simuliidae, Diptera, Insecta). - Museu Paraense Emilio Goeldi; Col. E. Snethlage: 111 pp. COSCARÓN, S. (1989): Los estudios ecológicos en Simúlidos Neotropicales (Diptera-Insecta). - Actas XI Congreso Sociedade Entomológica Brasileira (Campinas 12-18 de julio de 1987). Fundação Cargill, Seminario Insetos e Acaros. Anais 5: 69-98. COSCARÓN, S. (1990): Taxonomia y Distribución del subgénero Simulium (Ectemnaspis) ENDERLEIN (Simulidae, Diptera, Insecta). - Iheringia, Sér. Zoologia (Porto Alegre) 70: 109-170. COSCARÓN, S. (1991): Insecta, Diptera, Simuliidae. - In: CASTELLANOS, Z.A. DE (ed.): Fauna de Agua Dulce de la República Argentina. Vol. 38 (2). FECIC, Buenos Aires: 304 pp. COSCARÓN, S. (1994): Los Simuliidae (Diptera) y su importancia sanitaria en Latinoamérica. Memorias XXI Congreso Sociedad Colombiana de Entomología (Medellín): 357-366. COSCARÓN, S. (2003): Biodiversidad de Simuliidae y Tabanidae del Iberá. - In: ALVAREZ, B. (ed.): Fauna del Iberá. Eudene (Corrientes): 3-6. COSCARÓN, S. & C.L. COSCARÓN ARIAS (1995): Distribution of Neotropical Simuliidae (Insecta, Diptera) and its areas of endemism. - Revista de la Academia Colombiana de Ciencias Exactas Físicas y Naturales (Bogotá) 19 (75): 717-732. COSCARÓN, S. & C.L. COSCARÓN ARIAS (1996a): On Simulium (Pternaspatha), with description of a new species (Diptera - Simuliidae). - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 91 (6): 703-710. COSCARÓN, S. & C.L. COSCARÓN ARIAS (1996b): Análisis cladístico de Simulium (Pternaspatha) ENDERLEIN con datos sobre su distribución geográfica (Diptera: Simuliidae). - Acta Entomológica Chilena 20: 71-82. COSCARÓN, S. & C.L. COSCARÓN ARIAS (1997): Cladistic analysis of the subgenera Inaequalium, Psaroniocompsa and Chirostilbia of the genus Simulium with comments on their distribution (Simuliidae: Diptera). - Revista de la Sociedad Entomológica Argentina 56 (1-4): 109-121. COSCARÓN, S. & C.L. COSCARÓN ARIAS (2000a): New species records for the blackflies (Diptera Simuliidae) fauna of Argentina with description of adults, pupa and larva of Simulium oyapockense s. l. and S. seriatum. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 95 (2): 179-187.

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COSCARÓN, S. & C.L. COSCARÓN ARIAS (2000b): Los simúlidos (Diptera - Insecta) del río Paraná en el área próxima a la ciudad de Corrientes (Argentina): Dinámica Poblacional de los Estados Inmaduros. - Facena 15: 25-37. COSCARÓN, S. & C.L. COSCARÓN ARIAS (2002): Araucnephia iberaensis n. sp., a Neotropical black fly with a peculiar distribution (Diptera - Simuliidae). - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 97 (1): 81-87. COSCARÓN, S. & S. IBÁÑEZ BERNAL (1993): Sobre una nueva especie de Simuliidae del sur de México: Simulium (Hemicnetha) biuxinisa n. sp. (Diptera). - Folia Entomológica Mexicana 88: 61-68. COSCARÓN, S. & S. IBÁÑEZ BERNAL (1994): Sobre la ubicación taxonómica de Simulium ganalesense VARGAS, MARTÍNEZ PALACIOS y DÍAZ NÁJERA (Diptera: Simuliidae). - Folia Entomológica Mexicana 90: 1-7. COSCARÓN, S. & A. MATTA (1982): Notas sobre Simúlidos Neotropicales VIII. Sobre dos especies nuevas de “jerjeles” del género Simulium Latreille del Norte de Chile (Simuliidae, Diptera, Insecta). - Idesia 6: 49-61. COSCARÓN, S. & D.R. MIRANDA ESQUIVEL (1998a): Pedrowygomyia a new neotropical Prosimuliini genus (Diptera: Simuliidae): Gigantodax s. lat. split of into two genera. - Entomologica Scandinavica 29 (2): 161-168. COSCARÓN, S. & D.R. MIRANDA ESQUIVEL (1998b): New synonym of Simulium (Inaequalium) inaequale (PATERSON Y SHANNON), 1927 (Diptera: Simuliidae). - Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 93 (3): 337. COSCARÓN, S. & P. MUÑOZ DE HOYOS (1995): Blackfly novelties from the area near the “Paramo de los Valles” in the Department of Tolima, Colombia (Diptera: Simuliidae). - Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 19 (74): 587-592. COSCARÓN, S. & V. PY-DANIEL (1989): Tres especies nuevas de Simulium LATREILLE de la región Neotropical: Simulium (Hemicnetha) cristalinun sp. n., (Grenieriella) wygodzinskyorum sp.n. y sumapazense sp. n. (Diptera, Simuliidae). - Revista de Saúde Pública (São Paulo) 23 (4): 313-321. COSCARÓN, S. & H. TAKAOKA (1989): Description of a new species of the black fly subgenus Simulium (Ectemnaspis) from the Andes of Ecuador (Diptera, Simuliidae). - Japan Journal of Tropical Medicine & Hygiene 17 (2): 191-196. COSCARÓN, S. & P. WYGODZINSKY (1960): Sobre la presencia de la familia Simuliidae (Diptera, Insecta) en la provincia de Buenos Aires. - Orientación Médica 432: 1124-1125. COSCARÓN, S. & P. WYGODZINSKY (1962): Simuliidae (Diptera - Insecta) de Tierra del Fuego, Patagonia e Isla de Juan Fernández. - Acta Zoológica Lilloana 18: 281-333. COSCARÓN, S. & P. WYGODZINSKY (1972a): Notas sobre Simúlidos Neotropicales III. Sobre tres especies de jejenes con hembras de color claro del S.E. del Paraguay y del N.E. de la Argentina (Simuliidae, Diptera). - Revista del Museo de La Plata (N.S.) Secc. Zoología 11: 209-231. COSCARÓN, S. & P. WYGODZINSKY (1972b): Taxonomy and distribution of the black fly genus Simulium (Pternaspatha) ENDERLEIN (Simuliidae, Diptera, Insecta). - Bulletin of the American Museum of Natural History 147: 199-240. COSCARÓN, S. & P. WYGODZINSKY (1973a): Notas sobre Simúlidos Neotropicales II. Sobre Simulium (Psaroniocompsa) opalinifrons (ENDERLEIN) y notas sobre el subgénero (Diptera, Insecta). - Physis 32: 161-172. COSCARÓN, S. & P. WYGODZINSKY (1973b): Notas sobre simúlidos Neotropicales IV. Sobre la presencia de Simulium inaequale (PATERSON Y SHANNON) en el N.E. de Argentina y S.E. del Paraguay y una nueva especie muy afín. - Revista de la Sociedad Entomológica Argentina 34: 141-149.

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COSCARÓN, S. & P. WYGODZINSKY (1975): Notas sobre simúlidos Neotropicales V. Aportes para el conocimiento del subgénero Simulium (Notolepria) ENDERLEIN (Diptera - Simuliidae). - Revista de la Sociedad Entomológica Argentina 34: 277-288. COSCARÓN, S. & P. WYGODZINSKY (1984): Notas sobre simúlidos Neotropicales VII. Sobre los subgéneros Simulium (Psaroniocompsa) ENDERLEIN y Simulium (Inaequalium) nov. - Arquivos de Zoología (São Paulo) 31 (2): 37-103. COSCARÓN, S., CERQUEIRA, A.L.P., SATO SCHUMAKER, T.T.V. & V.L.A. SALVIA (1992): Nuevos datos sobre la distribución de simulidos de Brasil y descripción de Simulium (Coscaroniellum) cerradense sp. n. (Diptera, Simuliidae). - Revista Brasilelira de Entomologia 36 (1): 111-119. COSCARÓN, S., COSCARÓN ARIAS, C.L. & G. PORCARO (2000): Simuliidae (Diptera) del río Quequén Grande en Las Cascadas, (Buenos Aires, Argentina): variación poblacional y relación con temperatura, caudal y precipitaciones. - Acta Entomológica Chilena 24: 29-36. COSCARÓN, S., IBÁÑEZ BERNAL, S. & C.L. COSCARÓN ARIAS (l996): Revisión de Simulium (Psilopelmia) ENDERLEIN en la región Neotropical y análisis cladistico de sus especies (Diptera; Simuliidae). - Acta Zoológica Mexicana (n.s.) 69: 37-l04. COSCARÓN, S., IBÁÑEZ BERNAL, S. & C.L. COSCARÓN ARIAS (1999): Revision of Simulium (Simulium) in the Neotropical realm (Insecta: Diptera: Simuliidae). - Memoirs of Entomology International 14: 543-604. COSCARÓN, S., MIRANDA-ESQUIVEL, D.R., MOULTON, K., COSCARÓN ARIAS C. L. & S. IBÁÑEZ BERNAL (2004): Simulium (Hearlea) VARGAS, MARTÍNEZ PALACIOS & DÍAZ NÁJERA (Diptera: Simuliidae): Taxonomic revision and cladistic analysis. - Zootaxa 396: 1-52. COSCARÓN, S., PY-DANIEL, V. & C.L. COSCARÓN ARIAS (1995): El subgénero Simulium (Psilopelmia) ENDERLEIN en Sudamérica. - Boletim do Museu Paraense Emílio Goeldi, Série Zoologia 1993. 9 (2): 283-311. COSCARÓN, S., SARANDÓN, R., COSCARÓN ARIAS, C.L. & E. DRAGO (1996): Análisis de factores ambientales que influyen en la distribución de los Simuliidae (Diptera: Insecta) en el Cono Austral de América del Sur. - Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 20 (78): 549-573. COSCARÓN ARIAS, C.L. (1998a): Los simúlidos y su control en el Valle del Río Negro y Neuquén, Argentina (Diptera: Culicoidea: Simuliidae). - Revista de la Sociedad Entomológica Argentina 58 (1-2): 276-281. COSCARÓN ARIAS, C.L. (1998b): The polytene chromosomes of Cnesia dissimilis (EDWARDS) and three species of Gigantodax ENDERLEIN (Diptera: Simuliidae) from Lanin National Park (Argentina). - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 93 (4): 445-458. COSCARÓN ARIAS, C.L. (2001): Insecta, Diptera, Simuliidae. - In: FERNÁNDEZ, H.R. & E. DOMÍNGUEZ (eds.): Guía para la determinación de los artrópodos bentónicos sudamericanos 195-219. Serie: Investigaciones de la UNT. Subserie: Ciencias Exactas y Naturales. Vol. 1. Editorial Universitaria de Tucumán, Argentina. COSCARÓN ARIAS, C.L. (2003): Clave de Simuliidae (Insecta, Diptera) de Ecuador. - Folia Entomológica Mexicana 43 (2): 191-219. COSTA I. DA CUNHA, M., COSCARÓN, S. & R.M. AZEVEDO DE BASSI (1998): Determinación de los estadios larvales de Simulium (Diptera, Simuliidae) de Paraná, Brasil. - Acta Biológica Paranaense 27: 57-66. CROSSKEY, R.W. (1969): A re-classification of the Simuliidae (Diptera) of Africa and its islands. Bulletin of the British Museum (Natural History), Entomology, Suppl. 14: 1-195.

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CROSSKEY, R.W. (1987): An annotated checklist of the world black flies (Diptera: Simuliidae). Part. VIII: Black fly species of the world. - In: KIM, K.C. & R.W. MERRITT (eds.): Black flies: ecology, population management and annoted word list: 425-520. Pennsylvania State University Press, University Park: 528 pp. CROSSKEY, R.W. (1990): The natural history of blackflies. - J. Wiley, Chichester, 711 pp. CROSSKEY, R.W. & T.M. HOWARD (1997): A new taxonomic and geographical inventory of world blackflies (Diptera: Simuliidae). - The Natural History Museum, London: 144 pp. CROSSKEY, R.W. & T.M. HOWARD (2004): A revised taxonomical and geographical inventory of world blackflies (Diptera - Nematocera). - Internetversion: http//www.nhm.ac.uk/entomology/projects/blackflies/Inventory-pdf (accessed 27 May 2004).Introl 8. Inventory 78. The Natural History Museum, London: 82 pp. DALMAT, H.T. (1949): Some species of Simuliidae (Diptera) from Guatemala 1. - Annals of the Entomological Society of America 42 (4): 538-553. DALMAT, H.T. (1950): New species of Simuliidae (Diptera) from Guatemala 2. - Annals of the Entomological Society of America 43 (1): 137-151. DALMAT, H.T. (1951): Notes on the Simuliidae (Diptera) of Guatemala, including descriptions of three new species. - Annals of the Entomological Society of America 44 (1): 31-58. DALMAT, H.T. (1952): Description of two new species of Simulium (Diptera, Simuliidae) from Guatemala. - Annals of the Entomological Society of America 45 (2): 339-347. DALMAT, H.T. (1953): Simuliidae (Diptera) of Guatemala. Description of Simulium (Dyarella) ardeni n. sp., and Simulium (Lanea) jacobsi n. sp. - Annals of the Entomological Society of America 46 (1): 35-42. DALMAT, H.T. (1955): The blackflies (Diptera, Simuliidae) of Guatemala and their role as vectors of onchocerciasis. - Smithsonian Miscellaneous Collections 125 (1): vii+ 425 pp. DAMPF, A. (1927): Un Simulium nuevo de México (Orden Diptera, Suborden Nematocera) procedente de Tiltepec, Estado de Oaxaca. - Revista Mexicana de Biologia 7 (6): 125-129. D’ANDRETTA, C. (1954): Sobre Simulium distinctum LUTZ (1910) (Diptera, Simuliidae). - Anais da Faculdade de Farmacia e Odontologia da Universidade de São Paulo 12: 63-66. D’ANDRETTA, C. (1956): Redescrição de Simulium perflavum ROUBAUD, 1908. - Anais da Faculdade de Farmacia e Odontologia da Universidade de São Paulo 13: 33-38 (1955). D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1947): As espécies Neotropicais da família Simuliidae SCHINER (Diptera, Nematocera). II. Lutzsimulium cruzi n. gen. e n. sp. e nova concepção de nervação das asas dos simulideos. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 44: 401-411, 35 figs. D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1948): Espécies Neotropicais da família Simuliidae SCHINER (Diptera, Nematocera). IV. Lutzsimulium cruzi ANDR. & ANDR., 1946: Descrição do alotipo macho e da larva. - Arquivos de Zoología (São Paulo) 5: 637-647. D’ANDRETTA, C. & M.A.V. D’ANDRETTA (1950): Espécies Neotropicais da família Simuliidae SCHINER (Diptera, Nematocera). IV. Redescrição de Simulium pertinax KOLLAR, 1832. - Papeis Avulsos do Departamento de Zoología (São Paulo) 9: 193-213. D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1946): As espécies Neotropicais da família Simuliidae SCHINER, 1864 (Diptera - Nematocera). Simulium (Eusimulium) orbitale LUTZ, 1910, Simulium (Eusimulium) pintoi n. sp. e Simulium nigrimanum MACQUART, 1837, sp. inquirendae [sic]. Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 45: 85-l52.

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D’ANDRETTA, M.A.V. & C.D. D’ANDRETTA (1947): Espécies Neotropicais da família Simuliidae (Diptera, Nematocera). III. Simulium botulibranchium LUTZ, 1910 e travassosi n. sp. - Papeis Avulsos do Departamento de Zoología (São Paulo) 8: 145-180. D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1949): Espécies Neotropicais da família “Simuliidae” SCHINER (Diptera, Nematocera). Redescrição de “Simulium auristriatum LUTZ, 1910 e “Simulium (Eusimulium) venustum infuscata” LUTZ, 1909 var. inquirendae [sic]. - Revista Brasileira de Biología 9 (1): 55-66. D’ANDRETTA, M.A.V. & C. D’ANDRETTA (1952): Espécies Neotropicais da família Simuliidae (Diptera, Nematocera). VII. - Redescrição de Lutzsimulium pernigrum (LUTZ, 1910), n. comb. e considerações sôbre o gênero. - Papeis Avulsos do Departamento de Zoología (São Paulo) 10: 307-324. D’ANDRETTA, C. & D. GONZÁLEZ (1964): Tres novas espécies Brasileiras de Simuliidae (Diptera, Nematocera). - Memorias do Instituto Butantan 30 (960-62): 103-116. DE ALMEIDA, F.B. (1979): Simulium cerqueirellum, nomen novum para Simulium septentrionale CERQUEIRA & ALMEIDA, 1970 (Diptera: Simuliidae). - Acta Amazônica 9 (4): 789. DE LEÓN, J.R. (1943): Preliminares para la descripción de cinco nuevas especies de la región occidental de Guatemala. - Boletín Sanitario de Guatemala 51: 94-101. DE LEÓN, J.R. (1945): Nuevas especies de simúlidos en la región occidental de Guatemala. - Boletín Sanitario de Guatemala 52: 66-77 (1944). DE LEÓN, J.R. (1948): Un nuevo simúlido de la región occidental de Guatemala. - Impreso no. 56, Instituto de Investigaciones Científicas, Universidad de San Carlos (Guatemala): 1-23 pp. DELLOME FILHO, J. (1983): Considerações sobre os fatores físico-químicos dos criadouros de Simulium goeldii CERQUEIRA & MELLO, 1967 (Diptera: Nematocera). - Revista Brasileira de Entomologia 27: 155-160. DÍAZ NÁJERA, A. (1961): Nota acerca de Simulium (Psilopelmia) antillarum JENNINGS, 1915. Nueva especie para México (Diptera, Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 21 (1-2): 79-92. DÍAZ NÁJERA, A. (1962): Claves para identificar las larvas del género Cnephia y descripción de dos nuevas especies. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 22: 271-287. DÍAZ NÁJERA, A. (1965): Acerca de un nuevo simúlido mexicano (Diptera: Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 15 (1): 25-45. DÍAZ NÁJERA, A. (1969): Simulidos de Aguascalientes y Zacatecas. Descripción de seis especies del subgénero Psilopelmia y datos de dimorfismo. - Revista de Investigaciones de Salud Pública, México 29 (1): 21-63. DÍAZ NÁJERA, A. (1971): Descripción de una nueva especie del género Cnephia del norte de México. - Revista de Investigaciones de Salud Pública, México 31 (4): 239-247. DÍAZ NÁJERA, A. (1979): Una nueva especie del género Simulium (Diptera: Simuliidae) del Estado de Coahuila, México. - Anales del Instituto Biológico, Universidad Autónoma (Mexico), ser. Zoologia 50 (1): 553-561. DÍAZ NÁJERA, A. & M.A. VULCANO (1961): Descripción de Simulium (Psilopelmia) longithallum n. sp. (Diptera - Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 21 (3-4): 221-235. DÍAZ NÁJERA, A. & M.A. VULCANO (1962): Claves para identificar las larvas de Simúlidos del subgénero Hearlea, con descripción de dos nuevas especies. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 22 (1-2): 91-144.

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DUMBLETON, L.J. (1960): Larval Simuliidae (Diptera) from Navarino Island, Tierra del Fuego. - New Zealand Journal of Science 3: 543-548. DYAR, H.G. & R.C. SHANNON (1927): The North American two-winged flies of the family Simuliidae. - Proceedings of the United States National Museum 69 (10): 1-54. EDMAN, J.D. & K.R. SIMMONS (1985a): Rearing and colonization of blackflies (Diptera: Simuliidae). - Journal of Medical Entomology 22: 1-17. EDMAN, J.D. & K.R. SIMMONS (1985b): Simuliids (mainly Simulium decorum WALKER): 145-152. - In: SINGH, P. & R.F. MOORE (eds.): Handbook of insect rearing Vol. 2, Elsevier, Amsterdam: 514 pp. EDWARDS, F.W. (1931): Simuliidae. - In: Diptera of Patagonia and South Chile. British Museum (Natural History), London 2 (4): 121-154. EDWARDS, F.W. (1933): Die Ausbeute der deutschen Chaco-Expedition.1925-26. XLIII. Simuliidae (Melusinidae). - Konowia l2 (3-4): 210-211. ELLIOT, M.J. & C. POTTER (1978): The future of pyrethroides in insect control. - Annual Review of Entomology 23: 443-469. ENDERLEIN, G. (1921a): Das System Kriebelmücken (Simuliidae). - Deutsche Tierärztliche Wochenschrift 29: 197-200. ENDERLEIN, G. (1921b): Neue aussereuropäische Simuliden. - Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 4-5: 77-81. ENDERLEIN, G. (1922): Weitere Beiträge zur Kenntnis der Simuliiden. - Konowia 1: 67-76. ENDERLEIN, G. (1925): Weitere Beiträge zur Kenntnis der Simuliiden und Verbreitung. - Zoologischer Anzeiger 62: 201-211. ENDERLEIN, G. (1929): Über einige neotropische Simuliiden des Genus Friesia (Dipt.). - Deutsche Entomologische Zeitschrift 4: 327-328. ENDERLEIN, G. (1930): Der heutige Stand der Klassifikation der Simuliiden. - Archiv für klassifikatorische und phylogenetische Entomologie 1: 77-97. ENDERLEIN, G. (l934a): Aussereuropäische Simuliiden aus dem Wiener Museum. - Sitzungsberichte der Gesellschaft Naturforschender Freunde (Berlin): 190-195. ENDERLEIN, G. (1934b): Weiterer Ausbau des Systems der Simuliiden. - Deutsche Entomologische Zeitschrift 2-3: 273-292 (l933). ENDERLEIN, G. (l935): Neue Simuliiden, besonders aus Afrika. - Sitzungsberichte der Gesellschaft Naturforschender Freunde (Berlin) 1934: 358-364. ENDERLEIN, G. (1936): Simuliologica. - Sitzungsberichte der Gesellschaft Naturforschender Freunde (Berlin): 113-130. FAIRCHILD, G.B. (1940): Notes on the Simuliidae of Panama (Diptera, Nematocera). - Annals of the Entomological Society of America 33 (4): 701-719. FAIRCHILD, G.B. & L. BRICEÑO IRAGORRY (1942): Nota sobre Simulium lutzianum. - Boletin del Laboratorio de la Clínica «Luis Razetti» 2:158-159 (3:187-188). FIELD, G. (1967a): Studies on black flies of Panama (Diptera: Simuliidae). I. Two new species of Simulium of the subgenus Notolepria. - Annals of the Entomological Society of America 60 (1): 188-194. FIELD, G. (1967b): Studies on black flies of Panama (Diptera: Simuliidae). II. A new species of Simulium of the subgenus Hearlea. - Annals of the Entomological Society of America 60 (1): l94-l99. FIELD, G. (1969a): Studies of black flies of Panama. III. Two new species of Simulium of the subgenus Hemicnetha. - Annals of the Entomological Society of America 62 (1): 157-163.

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HAMADA, N. & C.A. AZEVEDO (2002): Descrição do macho e fêmea de Simulium lourencoi e lista de espécies de Simuliidae (Diptera: Nematocera) de Carolina e municipios arredores, Maranhão, Brasil. - Livro de Resumos do 19° Congresso Brasileiro de Entomología, Manaus, AM: 232-233. HAMADA, N. & F. FOURQUE (2001): Black flies (Diptera: Simuliidae) of French Guiana: Cytotaxonomy and preliminary list of species. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 96 (7): 955-959. HAMADA, N. & M.E. GRILLET (2001): Black flies (Diptera: Simuliidae) of the Gran Sabana (Venezuela) and Pacaraima Region (Brazil): Distributional data and identification keys for larvae and pupae. - Entomotropica l6 (1): 29-49. HAMADA, N. & J.W. MCCREADIE (1999): Environmental factors associated with the distribution of Simulium perflavum group (Diptera: Simuliidae) among streams in Brazilian Amazonia. Hydrobiologia 397: 71-78. HAMADA, N. & M. PEPINELLI (2004): Simulium (Chirostilbia) bifenestratum (Diptera: Simuliidae), a new black-fly species from the Atlantic Forest, State of São Paulo, Brazil. - Memorias Instituto Oswaldo Cruz (Rio de Janeiro) 99 (1): 45-52. HAMADA, N., ALE-ROCHA, R. & S.J. BESSA-LUZ (2003): Description of Simulium damascenoi (Diptera: Simuliidae) male and the black-fly species from the state of Amapa, Brazil. - Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 98 (3): 353-360 HAMADA, N., BESSA-LUZ, S.J. & S. COSCARÓN (2004): New morphological information on Simulium cerradense adults and new records of black fly species (Diptera: Simuliidae) in the western region of Bahia state, Brazil. - Zootaxa 693: 1-8. HEARLE, E. (1932): The blackflies of British Columbia (Simuliidae: Diptera). - Proceedings of the Entomological Society of British Columbia 29: 5-19. HERNÁNDEZ, L.M. & A.J. SHELLEY (2005): New specific synonymic and taxonomic notes on Neotropical blackflies (Diptera: Simuliidae). - Zootaxa 853: l-46. HERNÁNDEZ, L.M., SHELLEY, A.J., LUNA DIAS, A.P.A. & M. MAIA-HERZOG (2005): Description of the female of Simulium itaunense D’ANDRETTA & GONZÁLEZ and redescription of the male and pupa (Diptera: Simuliidae). - Zootaxa 896: 1-14. HOFFMANN, C.C. (1930a): Un Simulium nuevo de la zona cafetera de Chiapas. - Anales del Instituto de Biología, México 1: 51-53. HOFFMANN, C.C. (1930b): Los simúlidos de la region onchocercosa de Chiapas (con descripción de nuevas especies). - Anales del Instituto de Biologia, México 1 (4): 293-306. HOFFMANN, C.C. (1931): Los simulidos de la region oncocercosica de Chiapas, segunda parte; los estados larvales. - Anales del Instituto de Biología, México 2: 207-218. IBÁÑEZ BERNAL, S. (l992): Las especies mexicanas de Simulium (Hemicnetha) y S. (Notolepria) (Diptera: Simuliidae). - M.Sc.-thesis. Universidad Autónoma de México. México D.F: 302 pp. IRIARTE, D.R. (1943): Contribución a la fauna Venezolana. - Boletin del Laboratorio de la Clínica “Luis Razetti” 3: 189-203. JENNINGS, A. (1915): Two new species of Simuliidae from tropical America. - Proceedings of the Entomological Society of Washington 17 (4): 199-200. JOAN, T. (1912): Nota sobre un díptero ponzoñoso. - Boletin del Ministerio de Agricultura, Buenos Aires 14: 363-385. KERTÉSZ, K. (1902a): Catalogus dipterorum hucusque descriptorum 1. - Leipzig: 339 pp. KERTÉSZ, K. (1902b): Catalogus dipterorum hucusque descriptorum 2. - Leipzig: 359 pp.

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KNAB, F. (1911): Dr. A. LUTZ‘s studies of Brazilian Simuliidae. - Proceedings of the Entomological Society of Washington 13: 172-179. KNAB, F. (1913): A note on some American Simuliidae. - Insecutor Inscitiae Menstruus 1 (12): 154-156. KNAB. F. (1914a): Simuliidae de Chile septentrional. - Anales de Zoologia Aplicada, Chile 1 (7): 17-22. KNAB. F. (1914b): Simuliidae of Peru. - Proceedings of the Biological Society of Washington 27: 81-86. KNAB, F. (1914c): Supplementary notes on Peruvian Simuliidae. - Proceedings of the Biological Society of Washington 27: 123-124. KNAB, F. (1915a): A new Simulium from Texas (Diptera: Simuliidae). - Insecutor Inscitiae Menstruus 3 : 77-78. KNAB, F. (1915b): New data and species in Simuliidae (Diptera). - Insecutor Inscitiae Menstruus 2 (12): 177-180 (19l4). KNAB, F. (1915c): Some new Neotropical Simuliidae. - Bulletin of Entomological Research 6 (3): 279-282. KOLLAR, V. (1832): Die vorzüglich lästigen Insecten Brasiliens. - In: POHL, J.E. (ed.): Reise im Innern von Brasilien, Vol. 1: 101-119. Wien: 448pp. LACEY, L.A. (1981): Simulideos antropofilicos no Parque Nacional da Amazônia (Tapajos), Brasil, com referencia aos efeitos no homen. - Boletin de la Oficina Sanitaria Panamericana 90 (4): 326-338. LACEY, L.A. & J.D. CHARLWOOD (1980): On the biting activities of some anthropophilic Amazonian Simuliidae (Diptera). - Bulletin of the Entomological Research 70: 495-509. LACEY, L.A. & A.H. UNDEEN (1988): The biological control potential of pathogens and parasite of black flies. - In: KIM, K.C. & R.W. MERRITT (eds.): Black flies: ecology, population management and annoted world list: 327-340 (1987). Pennsylvania State University, University Park: 528 pp. LANE, J. & C.E. PORTO (1939): Simulideos da região neotropical: o gênero Eusimulium. - Boletim Biológico (n. s.) (São Paulo) 4 (2): 168-176. LANE, J. & C.E. PORTO (1940): Simulideos da região Neotrópica III. Descrição de novas especies dos gêneros “Simulium “Eusimulium”. - Arquivos do Instituto Biológico (São Paulo) 11: 189-195. LANE, J. & M.A. VULCANO (1943): A armadura bucal dos simuliideos e seu valor taxonômico (Diptera: Simuliidae). - Revista de Entomologia (Rio de Janeiro) 14 (3): 430-440. LATREILLE, P.A. (1802): Histoire naturelle, générale et particulière des Crustacés et des Insectes. 3. Familles naturelles des genres. - F. Dufart, Paris: 467 pp. LEÓN L.A. & P. WYGODZINSKY (1953): Los simúlidos de Ecuador, su importancia en medicina humana. - Boletin de Informaciones Científicas Naciónales (Quito) 6 (57): 269-288. LEWIS, D.J. (1963): Simuliidae (Diptera) from the human onchocerciasis area of Venezuela. Proceedings of the Royal Entomological Society of London 32: 53-62. LEWIS, D.J. & R. IBÁÑEZ ALDECOA (1962): Simuliidae and their relation to human onchocerciasis in northern Venezuela. - Bulletin of World Health Organization 27: 449-464. LEWIS, D.J. & J.P. LEE POTTER (l964): Simuliidae (Diptera) from the Sierra Nevada de Santa Marta, Colombia. - Annals and Magazine of Natural History, Ser. 13, 7: 95-100. LOEW, H. (1862): Monographs of the Diptera of North America. Part I. - Smithsonian Miscellaneous Collections 6 (1) xxiv + 221 pp. LUNA DIAS, A.P.A., HERNÁNDEZ, L.M., MAIA-HERZOG, M. & A.J. SHELLEY (2004): Simulium lobatoi, new species of blackfly, (Diptera: Simuliidae) from the States of Mato Grosso and Goias, Central Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 99 (1): 37-43. LUTZ, A. (1909): Contribuição para o conhecimento das espécies Brasileiras do gênero “Simulium”. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 1: 124-146.

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MORENO RAMÍREZ, C. (1997): Estudio morfológico y citologico integrado de tres especies del Subgénero Simulium (Ectemnaspis) sensu COSCARÓN, 1997. - Magister-thesis in Biology (Sistematics), Facultad de Ciencias, Universidad Nacional de Colombia, Bogotá, Colombia: 146 pp. MOREIRA, G.R.P., PEGORARO, R.A. & G. SATO (1994): Influencia de fatores abioticos sobre o desenvolvimento de Simulium nogueirai D’ANDRETTA e GONZÁLEZ en um corrego da mata atlántica. - Anais da Sociedade Entomológica do Brasil 23 (3): 525-542. MOULTON, J.K. (1996): A new species of Gigantodax ENDERLEIN (Diptera: Simuliidae) from the United States. - Proceedings of the Entomological Society of Washington 98 (4): 741-751. MOULTON, J.K. (1998): Reexamination of Simulium (Psilopelmia) ENDERLEIN (Diptera: Simuliidae) of America North of Mexico. - Proceedings of the Entomological Society of Washington 100 (1): 50-71. MUHAMMAD, A. (1988): A cytological description of Simulium virgatum COQUILLET and related species. - M.Sc.-thesis, Sul Ross State University, Alpine, USA: 94 pp., 18 pls. MUÑOZ DE HOYOS, P. & S. COSCARÓN (1999): Claves para la identificación de simulidos (Diptera: Simuliidae) presentes entre las vertientes magdalenense y orinocense, en un sector al centro de Colombia. - Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 23 (Suplemento Especial): 181-214. NATHAN, M.B., TIKASINGH, E.S. & P. MUNROE (1982): Filariasis in Amerindian of Western Guyana with observations of Mansonella ozzardi by a Simulium species of the amazonicum group. Tropenmedizin und Parasitologie 33: 219-222. NEIVA, A. & B. PENNA (1916): Viagem cientifica pelo Norte de Bahia, Sudoeste de Pernambuco, Sul de Piauí e de Norte a Sul de Goiaz. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 8 (3): 1-224. NUNES DE MELLO, J.A.S. (1974): Simulideos (Diptera, Nematocera) do Territorio Federal de Roraima (Brasil). - Ph.D.-thesis, Doctoral Faculdade de Medicina, Universidade de Sorocaba, São Paulo: VII+56 pp. NUNES DE MELLO, J.A. & F. BARBOSA DE ALMEIDA (1974): Simuliidae da Amazônia VII. Descrição de Simulium pseudoexiguum sp. n. (Diptera, Nematocera). - Acta Amazônica 4 (1): 65-68. NUNES DE MELLO, J.A. & E. VIEIRA DA SILVA (1974): Simuliidae da Amazônia VIII. Descrição de Simulium mutucuna sp. n. (Diptera, Nematocera). - Acta Amazônica 4 (2): 69-71. NUNES DE MELLO, J.A., BARBOSA DE ALMEIDA, F. & J. DELLOME (1973): Simuliidae da Amazônia VI. Descrição do Simulium meruoca sp. n. (Diptera, Nematocera). - Acta Amazônica 3 (3): 59-62. OCHOA, J.O. & H. TAKAOKA (l993): A new blackfly species of Mayacnephia from Guatemala (Diptera: Simuliidae). - Japanese Journal of Tropical Medicine and Hygiene 21 (1): 5-7. OKAZAWA, T. & O. ONISHI (1980): Description of a new species of Simulium (Simulium) LATREILLE and redescription of Simulium (Simulium) metallicum BELLARDI from Guatemala (Diptera: Simuliidae). - Japanese Journal of Sanitary Zoology 32 (3): 167-179. ORFILA, R.N. (1939): Simuliidae (Dipt.) de la República Argentina. - Novena Reunión de la Sociedad Argentina de Patologia Regional 3: 1525-1534. ORTIZ, I. (1957): Nuevos representantes hematófagos de los géneros Culicoides (Diptera: Ceratopogonidae) y Simulium (Diptera: Simuliidae) de Venezuela. - Boletin Venezolano de Laboratorio Clínico 2 (3-4): 161-168. PATERSON, G. & R.C. SHANNON (1927): Los simulidos del noroeste argentino. - Revista del Instituto Bacteriológico del Departamento Nacional de Higiene 4 (7): 737-742.

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SHELLEY, A.J., LOWRY, C.A., MAIA-HERZOG, M., LUNA DIAS, A.P.A. & M.A.P. MORAES (1997): Biosystematics studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus. Bulletin of the Natural History Museum, London (Entomology) 66 (1): 1-121. SHELLEY, A.J., LUNA DIAS, A.P.A., MAIA-HERZOG, M., PROCUNIER, W.S. & M.A.P. MORAES (1987): Identification of vector species (Diptera: Simuliidae) of human onchocerciasis in the Amazonia focus of Brazil and Venezuela. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 82 (4): 461-465. SHELLEY, A.J., LUNA DIAS, A.P.A., MAIA-HERZOG, M., LOWRY, C.A., GARRITANO, P.R., PENN, M. & M. CAMARGO (2001): Simulium cuasiexiguum, a new blackfly species (Diptera: Simuliidae) from the Minaçu area in the State of Goias, Central Brazil. - Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 96 (4): 483-496. SHELLEY, A.J., MAIA-HERZOG, M., LUNA DIAS, A.P.A. & C.A. COUCH (1989): Description of the adults and pupa of Simulium (Trichodagmia) perplexum, new species (Diptera: Simuliidae). Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 84: 343-349 SHELLEY, A.J., MAIA-HERZOG, M., LOWRY, C.A., LUNA DIAS, A.P.A., GARRITANO, P., CAMARGO, M. & H.G. CARTER (2000): The Simuliidae (Diptera) of the secondary onchocerciasis focus at Minaçu in Central Brasil. - Bulletin of the Natural History Museum, London (Entomological Series) 69 (2): 171-221. SHELLEY, A.J., PINGER, R.R., MORAES, M.A.P., CHARLWOOD, J.D. & J. HAYES (1979): Vectors of Onchocerca volvulus at the river Toototobi, Brazil. - Journal of Helminthology 53: 41-43. SILVA FIGUEROA, C. (1917): Dos nuevos simulidos de Chile. - Boletin del Museo Nacional de Chile 10: 28-35. SMART, J. (1940): Simuliide (Diptera) from British Guiana and the Lesser Antilles. - Transactions of the Royal Entomological Society of London 90: 1-11. SMART, J. (1944): Notes on Simuliidae (Diptera) II. - Proceedings of the Royal Entomological Society of London, Ser. B, 13: 131-136. SMART, J. (1945): The classification of the Simuliidae (Diptera). - Transactions of the Royal Entomological Society of London 95: 463-528. SMITH, C.N. (1973): Pyrethrum for control of insects affecting man and animals. - In: CASIDA, J.E. (ed.): Pyrethrum, the natural insecticide. Academy Press, London: 329 pp. SPEISER, P. (1904): Zur Nomenclatur blutsaugender Dipteren Amerikas. - Insekten-Börse 21: 148 STAINS, G.S. & G.F. KNOWLTON (1943): A taxonomic and distributional study of Simuliidae of western United States. - Annals of the Entomological Society of America 36 (2): 259-280. STONE, A. (1948): Simulium virgatum COCQUILLET and a new related species (Diptera: Simuliidae). Journal of the Washington Academy of Sciences 38: 399-404. STONE, A. (1962): Notes on the types of some Simuliidae (Diptera) described by ENDERLEIN. Annals of the Entomological Society of America 55: 206-209. STONE, A. (1963): An annotated list of genus-group names in the family Simuliidae (Diptera). United States Department of Agriculture, Technical Bulletin 1284: 1-28. STONE, A. (1969): The black flies of Dominica (Diptera: Simuliidae). - Proceedings of the Entomological Society of Washington 61: 312-318. STONE, A. & M.M. BOREHAM (1965): A new species of Simulium from the southwestern United States (Diptera: Simuliidae). - Journal of Medical Entomology 2: 164-170. STRIEDER, M.N. (1998): Revisão e analise cladistica de espécies de Inaequalium (COSCARÓN & WYGODZINSKY, 1984): (Diptera: Simuliidae). - Ph.D.-thesis, Pontificia Universidade Catolica de Rio Grande do Sul, Porto Alegre, Brasil: 160 pp.

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VARGAS, L. & A. DÍAZ NÁJERA (1957a): Simúlidos de Durango y Simulium (Simulium) marquezi n. sp. (Diptera, Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 17 (1): 33-41. VARGAS, L. & A. DÍAZ NÁJERA (1957b): Simúlidos Mexicanos. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 17 (4): 143-399. VARGAS, L. & A. DÍAZ NÁJERA (1958): Simulium (Psilopelmia) bivittatum MALLOCH, 1914 (Diptera: Simuliidae). - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 18 (1): 13-30. VARGAS, L., DÍAZ NÁJERA, A. & A. MARTÍNEZ PALACIOS (1943): Tres simúlidos nuevos para México. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 4 (3): 287-291. VARGAS, L., MARTÍNEZ PALACIOS, A. & A. DÍAZ NÁJERA (1946): Simúlidos de México. Datos sobre sistemática y morfología. Descripción de nuevos subgéneros y especies. - Revista del Instituto de Salubridad y Enfermedades Tropicales, México 7 (3): 95-192. VARGAS, M. & J. RAMÍREZ PÉREZ (1988): Gigantodax bierigi & G. willei (Diptera: Simuliidae), two new black fly species from Costa Rica. - Revista de Biología Tropical 36 (2B): 457-469. VARGAS, M., VARGAS, J.V. & J. RAMÍREZ PÉREZ (1993): Simulium (Psilopelmia) travisi (Diptera: Simuliidae), a new black fly species from Costa Rica. - Proceedings of the Entomological Society of Washington 95 (2): 175-181. VARGAS, M., TRAVIS, B.V., DÍAZ NÁJERA, A. & F. FALLAS (1977): Bionomics of black flies (Diptera: Simuliidae) in Costa Rica. VII, genus Simulium, subgenus Hearlea. - Revista de Biología Tropical 25 (1): 137-149. VULCANO, M.A. (1958): Redescrição do Simulium rubrithorax LUTZ, 1909 e descrição do alotipo macho (Diptera, Simuliidae). - Papeis Avulsos do Departamento de Zoología (São Paulo) 13: 227-240. VULCANO, M.A. (1959): Descrição de Simulium obesum, sp. n. (Diptera, Simuliidae). - Papeis Avulsos do Departamento de Zoología (São Paulo) 13: 351-360. VULCANO, M.A. (1962): Simulideos da Serra de Bocaina. Observações bionômicas, revalidação de Simulium brevifurcatum LUTZ e descrição de uma espécie nova, S. anamariae (Diptera, Simuliidae). - Papeis Avulsos do Departamento de Zoología (São Paulo) 15: 239-272. VULCANO, M.A. (1967): Family Simuliidae. - A catalogue of the Diptera of the Americas South of the United States. Departamento de Zoologia, Secretaria de Agricultura (São Paulo) 16: 1-43. VULCANO, M.A. (1971): Simuliidae. - In: HURLBERT, S.H. (ed.): Biota Acuática de Sudamérica Austral: 285-293. San Diego State University: 342 pp. VULCANO, M.A. (1981): Simuliidae, Part I. - In: HURLBERT, S.H., RODRIGUEZ, G. & N. SANTOS (eds.) Aquatic biota of tropical South America: 275-285. San Diego State University: 342 pp. WALKER, F. (1861): Characters of undescribed Diptera in the collection of W.W. SAUNDERS. Transactions of the Entomological Society of London (n.s.) 5 (1858-1861): 297-334. W.H.O. (1982): Division of vector biology and control onchocerciasis control programme in the Volta river basin. A review of the literature concerning the feeding behaviour and general ecology of Simulium. - Technical Report: 40 pp. WILLISTON, S.W. (1893): List of Diptera of the Death Valley Expedition. - In: RILEY, C.V. (ed.): The Death Valley Expedition. A Biological survey of parts of California, Nevada, Arizona and Utah, Part II. 4. Report of a small collection of insects made during the Death Valley Expedition. North America Fauna 7: 235-268. WISE, K.S. (1911): The Simuliidae of British Guiana. - Timehri 1: 248-254. WISE, K.S. (1912): The Itanimi fly. A new Simulium in British Guiana. - Journal of Tropical Medicine and Hygiene 15: 43.

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WYGODZINSKY, P. (1949): Contribuciones al conocimiento de los Simuliidae Argentinos. I. Introducción. Redescripción de Simulium lahillei PATERSON & SHANNON, 1927 (Diptera). Anales del Instituto de Medicina Regional, Tucumán 2: 303-319. WYGODZINSKY, P. (1950): Contribución al conocimiento de los Simulidos Argentinos. III. Simulium dinellii (JOAN, 1912) y S. wolffhuegeli (ENDERLEIN, 1920). - Anales del Instituto de Medicina Regional, Tucumán 3 (1): 75-97. WYGODZINSKY, P. (1951a): Sobre Simulium jujuyense PATERSON & SHANNON, 1927, Simulium exiguum ROUBAUD, 1906 y Simulium opalinifrons (ENDERLEIN, 1934). - Anales del Instituto de Medicina Regional, Universidad Nacional deTucumán 3 (2): 207-220. WYGODZINSKY, P. (1951b): Sobre Simulium ignescens ROUBAUD, 1906, Simulium romanai sp. n. y Simulium adolfolutzi sp. n. (Diptera, Simuliidae). - Anales del Instituto de Medicina Regional, Tucumán 3: 221-236. WYGODZINSKY, P. (1953a): Sobre algunos simúlidos argentinos (Diptera). - Anales del Instituto de Medicina Regional, Tucumán 3 (3): 293-320. WYGODZINSKY, P. (1953b): Sobre algunos simúlidos de los países andinos (Diptera). - Anales del Instituto de Medicina Regional, Tucumán. 3 (3): 321-337. WYGODZINSKY, P. (1958): Notas y descripciones de Simuliidae patagónicos y andinos (Diptera). Acta Zoológica Lilloana 16: 121-148. WYGODZINSKY, P. (1961): Sobre una nueva especie de Simulium de la provincia de Buenos Aires (Simuliidae, Diptera). - Acta Kravsi, Cuadernos del Instituto Nacional de Microbiologia (Buenos Aires) 2: 30-36. WYGODZINSKY, P. (1971): Descriptions and redescriptions of species of the black-fly genus Simulium from northern Andes of Colombia (Simuliidae-Diptera). - American Museum Novitates 2447: 1-38. WYGODZINSKY, P. (1973): A black fly of the subgenus Simulium (Psaroniocompsa) (Simuliidae, Diptera) from the High Andes of Colombia. - American Museum Novitates 2513: 1-11. WYGODZINSKY, P. & S. COSCARÓN (1962): On the relationships and zoogeographical significance of Austrosimulium anthracinum (BIGOT), a black fly from southern South America (Diptera, Simuliidae). - Pacific Insects 44: 1-244. WYGODZINSKY, P. & S. COSCARÓN (1967): A review of Simulium (Pternaspatha) ENDERLEIN (Simuliidae, Diptera). - Bulletin of the Amertican Museum of Natural History 136: 42-116. WYGODZINSKY, P. & S. COSCARÓN (1970): A new species-group of the black fly genus Simulium from Western South America (Simuliidae - Diptera - Insecta). - American Museum Novitates 2433: 1-20. WYGODZINSKY, P. & S. COSCARÓN (1973a): A review of the Mesoamerican and South America black flies of the tribe Prosimuliini (Simuliinae - Simuliidae). - Bulletin of the American Museum of Natural History 151: 133-199. WYGODZINSKY, P. & S. COSCARÓN (1973b): Notes on South American black flies of the tribe Prosimuliini (Simuliinae, Simuliidae, Diptera). - American Museum Novitates 2529: 1-9. WYGODZINSKY, P. & S. COSCARÓN (1979): Description of a black fly of the subgenus Simulium (Pternaspatha) from the high Andes of Ecuador (Simuliida, Diptera). - American Museum Novitates 2670: 1-9. WYGODZINSKY, P. & S. COSCARÓN (1982): Description of three unusual species of black fly subgenus Simulium (Ectemnaspis) from the Andes of Colombia (Diptera, Simuliidae). - American Museum Novitates 2736: 1-13.

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WYGODZINSKY, P. & S. COSCARÓN (1989): A revision of the blackfly genus Gigantodax ENDERLEIN (Simuliidae, Diptera, Insecta). - Bulletin of the American Museum of Natural History 189: 269 pp. WYGODZINSKY, P. & A. DÍAZ NÁJERA (1970): Un nuevo género de simulido de la Republica Mexicana (Diptera: Simuliidae). - Revista de Investigación en Salud Pública, México 30: 83-110. ZETTERSTADT, J.W. (1838): Sectio tertia, Diptera. Dipterologis scandinaviae amicis et popularibus carissimis: 478-868. - In: Insecta Lapponica. - L.Voss, Leipzig: 1140 pp. (1840).

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8 SUBJECT INDEX Acropogon 139 barbatipes 191 edwardsi 159 Araucnephia 40, 41, 42, 44, 45, 47, 48, 49, 50, 51, 66, 76, 94, 629 iberaensis 67, 68, 69, 70, 71 montana 68, 69, 72, 73, 74 montanum 66, 72, 108 Araucnephioides 40, 41, 42, 44, 45, 47, 48, 49, 50, 51, 67, 76, 94, 629 schlingeri 76, 77, 79, 80 Archicnesia 90 Astega 96 Austrosimulium 99 Cnephia 56, 57, 59, 60, 61, 72 Cnephia (Stegopterna) 84 Cnesia 24, 40, 41, 42, 45, 47, 48, 50, 51, 81, 94, 629 dissimile 81 dissimilis 83, 84, 85, 86, 88, 90 gynandra 84 gynandrum 81,84 ornate 83, 84, 88, 89, 90 pusilla 83, 84, 85, 88 Cnesiamima 39, 41, 42, 43, 44, 45, 48, 49, 50, 51, 81, 93, 94, 97, 629 atroparva 95, 96, 98 atroparvum 93, 96 Dasypelmoza 139, 216 varipes 216 Eusimulium 109 Friesia 108, 216 *

Figures in bold

*

Gigantodax 24, 33, 39, 41, 42, 44, 45, 47, 48, 50, 67, 90 adleri 91 antarcticus 92 bierigi 91 bolivianus 90 brophyi 92 clandestinum 29 conviti 91 dryadicaudicis 92 femineum 22, 90 herreri 32 igniculus 30, 92 lazoi 91 marginalis 30, 92 minor 92 rufidulus 92 willei 91 Jalacingomyia 471 Kempfsimulium 40, 41, 42, 45, 47, 48, 50, 51, 110, 111, 629 hebeticolor 111 simplicicolor 110, 111, 112, 113 Lutzsimulium 39, 40, 41, 35, 45, 47, 48, 50, 51, 102, 110, 111, 629 cruzi 101, 106 flavopubescens 104, 105, 109 hirticosta 104, 105, 106, 107, 109 montanum 108 pernigrum 103, 104, 105, 108, 109 Mayacnephia 40, 41, 42, 44, 45, 47, 48, 49, 50, 51, 52, 54, 63, 67, 629

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alticola 65 aguirrei 53, 55, 56, 58 atzopensis 55, 56, 58 fortunensis 55, 56, 58 grenieri 55, 56, 58, 59 mixensis 55, 56, 58, 59 muzquicensis 55, 56, 58, 60 pachecolunai 52, 55, 56, 58, 60, 62 roblesi 53, 55, 56, 58, 61 salasi 55, 56, 58 tadai 55, 56, 58 Odagmia 626 Oestrus 114 columbacensis 114 Paraustrosimulium 39, 41, 42, 44, 45, 47, 49, 50, 51, 97, 99, 629 anthracinum 22, 94, 99, 100, 101 moorei 101 Pedrowygomyia 24, 33, 39, 41, 42, 44, 45, 47, 49, 50, 51 chacabamba 93 cortesi 91, 92, 93 jatunchuspi 92, 93 punapi 92, 93 Prosimulium 87 Simulium 39, 43, 45, 49, 51, 114 costaricense 626 diaznajerai 626 nigrifemur 626 spinifer 237 tenuifrons 626 Simulium (Acropogon) 626 Simulium (Anasolen) 518, 612 Simulium (Aspathia) 116, 117, 120, 122, 123, 124, 126, 127, 128, 131, 133, 134, 135, 471, 643 A-K (cytoforms) 495, 498 H (cytoform) 487 L (cytoform) 495 X (cytoform) 495 anduzei 471, 473, 474, 476, 477, 478, 480, 482, 483 avidum 495

bustosi 473, 474, 476, 477, 478, 480, 482, 483, 484, 496 costalimai 473, 474, 476, 477, 480, 481, 482, 483, 484, 485, 488, 489, 497, 499, 502 covagarciai 472, 474, 475, 477, 478, 479, 481, 482, 485, 503, 511 guatemalense 489 hechti 472, 474, 476, 477, 478, 479, 482, 483, 486, 499, 502, 506, 507, 517 hirtipes 488 horacioi 473, 474, 476, 477, 478, 480, 482, 483, 487, 490, 498 hunteri 471, 473, 474, 476, 477, 479, 480, 481, 483, 485, 487, 489, 496, 502, 507 iriartei 473, 474, 476, 477, 479, 480, 481, 483, 488, 500, 502, 507, 515 jacumbae 473, 474, 476, 477, 479, 480, 482, 485, 489, 496, 502, 504, 507 jobbinsi 473, 474, 476, 477, 479, 482, 483, 490, 498 knowltoni 503 kompi 472, 474, 475, 477, 478, 479, 480, 482, 491, 514 lassmanni 487 marquezi 473, 474, 475, 477, 478, 480, 481, 482, 492, 499, 500, 502, 506, 510, 512, 515, 516 matteabranchia 493 matteabranchium 473, 475, 477, 479, 480, 481, 482, 486, 493, 494, 496, 497, 500, 502, 503, 506, 510, 511 metallicum 366, 473, 474, 476, 477, 478, 480, 482, 483, 487, 490, 491, 494, 495, 496, 497, 499, 500, 501, 502, 506, 507, 508, 509, 511, 515, 516 morae 473, 474, 476, 477, 478, 480, 482, 501 nitidum 495 parnassum 517 parrai 473, 474, 476, 477, 479, 480, 481, 483, 495, 496, 499, 501, 502, 510, 511 patziciaense 483 piperi 473, 474, 476, 477, 479, 482, 485, 490, 494, 496, 499, 502, 503

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puigi 473, 474, 476, 477, 478, 479, 482, 483, 499, 502, 504, 507 putre 472, 475, 476, 477, 478, 480, 481, 482, 494, 496, 497, 499, 500, 502, 505, 506, 507, 515, 516 racenisi 473, 474, 476, 477, 478, 479, 482, 498, 501, 508, 509 riveti 495 ruizi 472, 475, 477, 480, 481, 483, 502, 509, 510 sandyi 472, 474, 475, 477, 478, 479, 481, 482, 496, 510, 512 sayi 486, 503 sp. G 508 spilmani 473, 474, 476, 477, 478, 480, 481, 483, 484, 500, 502, 507, 512 stonei 503 tescorum 484 tricorne 474, 478, 480, 481, 492, 506 tricornis 513 vandalicum 492 versicolor 495 violacescens 495 wygoi 472, 474, 476, 477, 478, 480, 482, 483, 487, 496, 497, 500, 502, 506, 507, 514, 515, 516 Simulium (Astega) 96 Simulium (Byssodon) 117, 120, 123, 125, 126, 127, 133, 134, 136, 138, 365 forbesi 138 meridionale 138 tamaulipense 138 Simulium (Cerqueirellum) 117, 120, 123, 124, 131, 133, 134, 135, 296, 325, 335, 360, 364, 366, 458, 636 amazonicum 325, 326, 327, 328, 329, 330, 331, 332, 333, 339, 343, 344, 346 argentiscutum 326, 327, 328, 329, 331, 332, 333, 334, 337, 346, 348, 361 chaquense 327, 328, 329, 330, 331, 332, 333, 334, 337, 339, 340, 341, 342, 343, 344, 345, 348 cuasisanguineum 335 cuneata 344

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cuneatum 327, 328, 329, 330, 331, 332, 333, 334, 335, 337, 339, 340, 342, 343, 344, 345, 346, 347, 348 delponteianum 326, 327, 328, 330, 331, 332, 333, 334, 337, 347, 348 exiguum 348 ganalesense 327, 328, 329, 330, 331, 332, 333, 334, 335, 339, 340 lutzi 348 minuscula 332 minusculum 326, 327, 328, 329, 330, 331, 332, 333, 334, 335, 337, 339, 340, 345, 348 minutum 348 oyapockense 327, 328, 329, 330, 331, 332, 333, 335, 337, 340, 341 pseudoamazonicum 336 pseudosanguineum 335 quadristrigata 348 quadristrigatum 348 roraimense 326, 327, 328, 329, 330, 33l, 332, 334, 335, 339, 340 sanchezi 335 sanguineum 327, 328, 329, 330, 331, 332, 333, 334, 337, 339, 340, 341, 343 tallaferroae 341 venezuelense 326, 327, 328, 330, 331, 339, 345 Simulium (Chirostilbia) 115, 118, 119, 120, 121, 122, 123, 124, 125, 126, 128, 130, 131, 132, 133, 134, 135, 136, 633 acarayense 240, 241, 242, 243, 244, 245, 246, 259, 260, 261, 262, 263, 264, 266 bifenestratum 239, 240, 241, 242, 243, 244, 245, 246, 257 cerqueirai 247 dekeyseri 249 distincta 255 distinctum 239, 240, 241, 242, 243, 244, 245, 246, 248, 250, 252, 254, 255, 256, 257, 259, 265 empascae 240, 242, 243, 244, 246, 250, 252, 254 flavifemur 238, 247 friedlanderi 239, 240, 241, 242, 243, 244, 245, 246, 252, 257

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guarani 259 inexorabile 247 infuscatum 247 laneportoi 239, 240, 241, 242, 243, 244, 245, 246, 248, 249, 250, 251, 253, 254, 256, 266 lutziana 247 obesa 258 obesum 239, 240, 241, 242, 243, 244, 245, 246, 248, 250, 252, 257, 258 papaveroi 240, 241, 242, 243, 244, 245, 246, 259, 260, 261, 262, 263, 264, 265 pertinax 239, 240, 241, 242, 243, 244, 245, 246, 247, 248, 249, 250, 251, 252, 253, 255, 265, 266 pilosum 249 prumirimense 255 riograndense 239, 240, 241, 242, 243, 244, 245, 246, 250, 252, 256, 259 septentrionale 247 serranus 239, 240, 241, 242, 243, 244, 245, 246, 248, 249, 250, 251, 252, 265 spadicidorsum 255 spinibranchia 253 spinibranchium 239, 240, 241, 242, 243, 244, 245, 246, 248, 249, 250, 251, 253, 254, 255, 265 striginotum 266 subpallidum 28, 239, 240, 241, 242, 243, 244, 245, 246, 259, 260, 262, 263, 264, 265 subviride 259 venustum 247 Simulium (Coscaroniellum) 117, 118, 120, 122, 123, 124, 128, 131, 133, 134, 135, 296, 326, 349, 364, 637 benjamini 365, 449 cauchense 349, 350, 351, 352, 353, 354, 355, 356, 361 cerradense 349, 350, 351, 352, 353, 355, 356, 357, 358, 359, 360, 361 daltanhani 349, 350, 351, 352, 353, 357, 363 fairchildi 365 goeldii 349, 350, 351, 352, 353, 354, 355, 357, 358, 359, 362, 363, 364

mallochi 365 metallicum 365 quadrifidum 349, 350, 351, 352, 353, 354, 355, 357, 358, 359, 360, 361, 362, 363, 365 quadrivittatum 326, 349, 350, 351, 352, 353, 354, 355, 357, 358, 364, 365, 449 rangeli 361 rasii 354 scorzai 362 sextobecium 361 sp. A 360 sp. B1 363 torrealbai 354 ulyssesi 349, 350, 351, 352, 353, 354, 358, 359, 361, 363, 364 Simulium (Cnephia) 81 gynandrum 81 Simulium (Dasypelmoza) 400, 401, 413 Simulium (Dyarella) 517, 531, 537, 539, 546, 552, 553, 554, 555, 581 Simulium (Ectemnaspis) 118, 120, 121, 122, 123, 125, 129, 131, 132, 133, 134, 136, 294, 366, 420, 461, 462, 463, 518, 626, 638, 639, 640, 641 adolfolutzi 403 albanense 28, 367, 371, 374, 375, 377, 379, 382, 384, 386, 389, 391, 393, 395, 399, 403, 406, 415 alirioi 404 anaimense 367, 370, 375, 378, 382, 385, 395, 396, 399, 402 antillarum 368, 370, 372, 374, 375, 378, 380, 383, 384, 386, 393, 417, 418, 419 antonii 367, 368, 370, 372, 375, 377, 378, 381, 383, 388, 392, 397 antunesi 409 arcabucense 367, 368, 371, 375, 377, 380, 382, 384, 386, 390, 391, 392, 393, 394, 395, 396, 399, 405 bicolorata 385, 400 bicoloratum 367, 371, 374, 375, 377, 379, 382, 384, 385, 386, 389, 394, 395, 400, 401, 402, 403, 404, 405

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bicornutum 367, 368, 371, 375, 377, 379, 381, 384, 388, 391, 395, 397, 399, 400 bipunctatum 416 cormonsi 367, 375, 377, 379, 382, 383, 401, 402 dinellii 368, 370, 372, 374, 375, 378, 380, 383, 384, 386, 389, 393, 395, 414, 416, 419 flavipictum 414, 415 fulvinotum 411 furcillatum 367, 368, 371, 375, 377, 379, 381, 384, 388, 389, 391, 394, 395, 397, 398 gabaldoni 367, 371, 375, 377, 380, 382, 383, 386, 390, 393, 405, 415 gaudeatum 400 haarupiana 407 ignacioi 411, 412 ignescens 367, 368, 371, 375, 377, 379, 381, 384, 385, 386, 387, 394 iguazuense 404 jaimeramirezi 367, 371, 375, 377, 379, 382, 383, 400, 402 kabanayense 367, 370, 372, 375, 378, 380, 381, 383, 386, 389, 391, 393, 394, 396, 410 kavanayense 410 lewisi 404 lutziana 404 lutzianum 367, 371, 374, 375, 377, 379, 382, 385, 386, 389, 395, 396, 400, 403, 404, 406, 415, 416 macca 400, 401 maroniense 367, 370, 372, 375, 378, 380, 381, 383, 386, 395, 396, 410, 411, 412, 413 martinezi 414 mauense 409 mayuchuspi 25, 367, 370, 374, 375, 377, 378, 383, 385, 386, 389, 391, 393, 395, 399, 408 miniata 414 molli 400 nilesi 409 nuneztovari 368, 370, 372, 374, 375, 378, 380, 383, 418

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ochraceum 368, 370, 372, 374, 375, 378, 380, 383, 384, 386, 389, 391, 393, 412, 413, 414, 416, 418 pautense 367, 368, 371, 375, 377, 380, 382, 384, 386, 389, 390, 391, 392, 393, 394, 395, 396, 399, 402, 406 perflava 409 perflavum 367, 372, 375, 378, 380, 382, 384, 386, 389, 391, 393, 396, 399, 409, 414 pifanoi 367, 368, 371, 374, 375, 377, 380, 382, 384, 388, 389, 390, 391, 394, 395, 396, 399 pseudoantillarum 418 romanai 25, 367, 371, 374, 375, 377, 379, 382, 385, 386, 391, 393, 395, 396, 399, 400, 403, 405, 406, 408 roquemayu 367, 370, 374, 375, 377, 378, 383, 387, 388, 389, 390, 396 rorotaense 367, 370, 372, 375, 378, 380, 381, 383, 386, 389, 391, 393, 396, 399, 410, 411, 412, 413 rubiginosum 367, 368, 371, 374, 375, 377, 379, 383, 384, 386, 387, 391, 393, 395, 396 scutellatum 416, 417 shewellianum 368, 370, 372, 374, 375, 378, 380, 383, 385, 386, 389, 391, 393, 395, 396, 399, 412, 415, 417, 418 simplex 397 sp. A 390 sp. B 390 suarezi 367, 370, 372, 375, 378, 380, 381, 383, 386, 389, 395, 396, 410, 411, 412 tolimaense 367, 368, 370, 374, 377, 378, 381, 383, 395, 397 trombetense 367, 370, 372, 375, 378, 380, 381, 383, 396, 413 tunja 367, 368, 371, 375, 377, 380, 381, 385, 389, 391, 393, 394, 395, 399 varians 419 wolcotti 416, 417 wolffhuegeli 367, 371, 374, 375, 377, 379, 383, 384, 386, 389, 391, 393, 395, 396, 399, 407 wuayaraka 411, 412

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Simulium (Eusimulium) 115, 119, 123, 124, 130, 133, 136, 137, 266, 461, 531, 533, 597, 616 aureum 137 diazi 137 donovani 137 Simulium (Freemaniellum) 518, 612 Simulium (Grenieriella) 597, 603, 605, 607, 609 Simulium (Hearlea) 33, 117, 121, 122, 123, 127, 131, 132, 134, 518, 560, 589, 645 ayrozai 562, 564, 566, 568, 570, 571, 577, 583, 586 burchi 562, 564, 566, 567, 569, 570, 572, 580, 581 canadense 562, 564, 566, 567, 568, 570, 573, 577, 578, 581 capricorne 562, 564, 566, 567, 568, 570, 572, 574, 575, 576, 577, 578, 579, 581, 582, 583, 584, 585, 593, 596 carolinae 562, 563, 565, 567, 568, 569, 586, 590, 592, 595 chiriquiense 562, 564, 568, 578, 584 contrerense 562, 564, 565, 567, 568, 570, 579, 585 dalmati 563, 564, 566, 567, 569, 570, 572, 574, 578, 579, 580, 583 delatorrei 562, 564, 566, 567, 569, 570, 572, 575, 576, 580, 581 deleonii 574 estevezi 562, 564, 566, 569, 570, 578, 580, 582 ethelae 563, 564, 566, 567, 568, 570, 577, 578, 581, 583, 584, 586 gorirossiae 563, 565, 567, 568, 570, 587, 588, 591, 592, 593, 594, 596 gorirossii 587 johnsoni 562, 564, 565, 567, 568, 569, 588, 590, 591, 592, 597 juarezi 562, 564, 566, 567, 568, 570, 575, 576, 580, 584 larvispinosum 563, 564, 565, 567, 568, 569, 575, 576, 581, 587, 588, 589, 590, 591, 593, 594, 595, 596, 597 menchacai 563, 565, 567, 569, 588, 590, 594

microbranchium 563, 564, 566, 567, 568, 570, 571, 575, 577, 578, 585, 586 nigricorne 562, 564, 565, 568, 569, 577, 586, 595 paracarolinae 562, 563, 567, 569, 590, 593, 595, 596 temascalense 562, 563, 565, 568, 570, 590, 591, 594, 596 Simulium (Hemicnetha) 116, 117, 119, 120, 121, 122, 124, 127, 129, 131, 133, 135, 517, 549, 573, 607, 612, 644 acatenangoensis 531 ardeni 555 aureopunctatum 548 bilimekae 531 biuxinisa 519, 521, 522, 524, 526, 528, 530, 532, 533, 535, 536, 538, 540, 542 brachycladum 519, 520, 521, 523, 524, 526, 527, 528, 529, 532, 534, 536, 538, 540, 542, 543, 544, 545, 547, 553, 554, 555, 556 brevibranchium 540 bricenoi 519, 521, 522, 524, 526, 527, 528, 530, 533, 535, 536, 537, 538, 539, 549, 551, 552, 555, 556, 560 chiapanense 537 cinereum 537 clavipes 560 coffeae 560 conviti 531 cristalinum 519, 520, 522, 523, 524, 526, 527, 528, 530, 532, 534, 536, 538, 541, 542, 544, 545, 546, 547, 556 dehnei 553, 554 earlei 519, 521, 522, 523, 524, 526, 528, 529, 534, 536, 538, 553, 554, 556, 560 falculatum 560 freemani 519, 520, 522, 523, 524, 525, 527, 528, 529, 549, 552 guerrerense 519, 521, 522, 523, 524, 526, 527, 529, 544, 546, 547, 553, 555 hieroglyphicum 519, 520, 522, 523, 524, 526, 527, 529, 530, 538, 540, 551, 552 hinmani 462, 519, 521, 522, 524, 526, 527, 530, 554, 557

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hippovorum 537, 539 keenani 519, 520, 521, 523, 524, 526, 529, 553, 554 laticalx 549 latidigitus 549 lobatoi 519, 521, 523, 524, 526, 527, 528, 529, 545 lugubre 548 magnum 533 major 533 mathesoni 531 mexicanum 517, 518, 519, 520, 522, 523, 525, 527, 529, 530, 531, 532, 536, 538, 547, 549, 550, 551, 552, 560 mutucuna 531 oviedoi 518, 519, 520, 521, 523, 524, 525, 526, 528, 530, 556, 557, 558, 559 paynei 517, 519, 521, 522, 524, 526, 527, 528, 530, 531, 532, 535, 536, 537, 539, 540, 560 pictipes 560 placidum 548 pulverulentum 519, 521, 522, 523, 524, 526, 527, 528, 529, 532, 534, 536, 541, 544, 546, 547, 554, 555, 556 ribasi 557 rivai 557 rivasi 518, 519, 520, 521, 523, 524, 525, 526, 528, 530, 557, 558, 559 rubicundulum 537 rubrithorax 519, 521, 522, 524, 526, 527, 528, 530, 531, 532, 533, 534, 536, 537, 538, 539, 540, 541, 542, 545, 601 seriatum 519, 520, 522, 523, 525, 527, 529, 530, 532, 534, 536, 538, 542, 549, 551, 552, 553, 555, 556 smarti 519, 520, 522, 523, 524, 525, 527, 529, 530, 532, 534, 536, 538, 549, 550, 551, 552 solarii 519, 521, 523, 525, 526, 527, 529, 530, 546, 553, 554 tarsale 560 tarsatum 549 tephrodes 537

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turgidum 548 virgatum 519, 521, 522, 525, 526, 527, 528, 530, 531, 534, 535, 536, 537, 538, 547, 560 virgatum chiapanense 537 yepocapense 519, 521, 522, 523, 524, 526, 528, 529, 530, 532, 553, 554, 555 Simulium (Inaequalium) 115, 120, 122, 123, 125, 126, 130, 132, 134, 136, 266, 294, 634 argentata 273 baiensis 273 beaupertuyi 267, 281, 282 botulibranchium 267, 268, 269, 270, 271, 273, 288, 289, 290, 291, 292 clarki 267, 268, 269, 271, 272, 275, 282, 283 clavibranchium 267, 268, 269, 270, 271, 272, 274, 276, 277, 278, 279, 281, 283, 284, 285, 287, 288 diversibranchium 267, 268, 269, 270, 271, 272, 274, 276, 277, 279, 280, 281, 285, 286, 287, 288, 289 diversifurcatum 278 hauseri 267, 268, 269, 271, 272, 275, 281, 283 inaequale 267, 268, 269, 271, 272, 273, 274, 275, 276, 277, 278, 279, 280, 281, 282, 283, 284, 285, 287, 291 inaequalis 273 jundiaiense 273 leopoldense 267, 268, 271, 272, 279, 281, 282 luribayae 294 manicata 273 mariavulcanoae 267, 268, 269, 270, 271, 272, 276, 277, 279, 286 mbarigui 278 missionum 287 nahimi 267, 268, 270, 271, 273, 276, 279, 285 nogueirai 267, 268, 269, 270, 271, 272, 275, 279, 281, 282 parimaensis 294

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petropoliense 267, 268, 270, 271, 273, 290, 292, 293 pseudoexiguum 294 rappae 267, 268, 269, 271, 276, 277, 280, 287 souzalopesi 267, 268, 269, 270, 271, 273, 290, 291, 292, 294 subclavibranchium 267, 268, 269, 271, 272, 274, 276, 277, 278, 279, 283, 284, 289 subnigrum 23, 267, 268, 269, 270, 271, 272, 274, 275, 276, 278, 279, 280, 281, 282, 283, 284, 285, 289 travassosi 267, 268, 269, 270, 271, 273, 274, 276, 277, 280, 281, 288 tulibranchium 289 Simulium (Lanea) 419, 437, 441, 444, 448, 451, 454, 456, 457, 458, 461 Simulium (Morops) 407 Simulium (Neosimulium) 138 Simulium (Nevermania) 115, 119, 123, 124, 130, 133, 136, 137 annulipes 137 carbunculum 138 pugetense 138 ruficorne 137 Simulium (Notolepria) 116, 120, 123, 126, 130, 134, 136, 317, 319, 360, 361, 632 blantoni 237 cuasiexiguum 225, 226, 227, 228, 233, 234, 313 delpontei 228 exigua 228 exiguum 29, 224, 226, 227, 228, 230, 231, 232, 233, 234, 313 glaucophthalmum 228 gonzalezi 225, 226, 227, 228, 229, 230, 232, 233, 234, 237 incertum 237 llutense 225, 226, 227, 228, 235, 236 paraguayense 226, 227, 228, 229, 230, 231, 232, 233, 234, 237, 313 paranense 237 sp. A 226, 227, 228, 229, 233, 234, 235 subexiguum 225, 226, 227, 228, 229, 233, 234, 237

urubambanum 228 Simulium (Odagmia) 400, 409, 626 Simulium (Pliodasina) 324 guttata 313, 324 Simulium (Psaroniocompsa) 117, 120, 122, 123, 125, 131, 133, 134, 136, 325, 335, 364, 635 aequifurcatum 314, 315 anamariae 296, 297, 298, 299, 301, 304, 308, 309, 313, 315, 316, 317, 318, 319 angrense 296, 297, 298, 300, 301, 305, 308, 309, 310 angustitarse 306 auripellita 302 auripellitum 296, 297, 298, 299, 300, 301, 302, 303, 304, 305, 307, 308, 309, 310, 311, 315, 320 auristriata 316 auristriatum 296, 297, 299, 300, 303, 304, 305, 308, 3l2, 313, 316, 318 bonaerense 296, 297, 298, 300, 301, 302, 303, 304, 305, 307, 308, 309, 310, 311, 312, 313, 314, 315, 318, 319 brevifurcata 317 brevifurcatum 296, 297, 298, 299, 301, 303, 308, 313, 316, 317, 319 catarinense 324 damascenoi 296, 297, 298, 300, 303, 305, 308, 309, 320, 322, 323 fuliginis 296, 299, 315 guaporense 296, 298, 301, 320, 322, 323 guttata 324 guttatum 324 hoffmanni 306 incaicum 306 incrustata 311 incrustatum 296, 297, 298, 299, 301, 302, 303, 305, 307, 308, 309, 310, 311, 312, 314, 315, 317 infuscatum 316 jujuyense 296, 297, 298, 300, 301, 302, 305, 306, 307, 308, 309, 310 limbata 306 limbatum 296, 297, 298, 299, 301, 302, 303, 304, 305, 308, 309, 313, 314, 315, 317

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lourencoi 296, 298, 301, 320, 321, 322 machadoallisoni 314 machadoi 314 mendozana 306 meruoca 314 minuana 305 minuanum 296, 297, 298, 299, 300, 301, 303, 305, 308, 310 opalinifrons 311 schmidtmummi 296, 297, 298, 299, 300, 304, 305, 308, 309, 318, 319, 321, 322 sicuani 306 siolii 296, 297, 298, 300, 301, 308, 312, 320, 321, 322, 323, 324 spinifer 306 stellatum 296, 297, 298, 299, 300, 319, 322 tergospinosum 296, 297, 298, 300, 304, 305, 308, 320, 321, 322 yarzabali 311, 313 Simulium (Psilopelmia) 116, 118, 120, 121, 122, 123, 124, 125, 127, 129, 131, 132, 133, 134, 135, 136, 360, 361, 365, 419, 642 bivittatum 422, 424, 428, 430, 432, 435, 437, 438, 440, 442, 443, 446, 447, 448, 449, 450 blancasi 420, 422, 425, 427, 428, 431, 433, 436, 464, 465, 467, 469, 470 bobpetersoni 422, 424, 426, 428, 430, 432, 435, 437, 440, 446, 447, 450, 456, 457, 458, 459 boydi 448 callidum 420, 421, 423, 425, 428, 429, 431, 437, 438, 440, 443, 446, 450, 452, 453, 461, 462 colvini 444 dandrettai 422, 424, 425, 427, 430, 432, 433, 436, 438, 440, 441, 446, 447, 450, 453, 454, 455 distinctum 455 downsi 422, 423, 426, 428, 430, 431, 435, 437, 440, 446, 447, 450, 451, 459 dugesi 421, 424, 426, 427, 429, 432, 435, 436, 437, 438, 439, 440, 441, 442, 443, 446, 447, 449, 450, 452

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escomeli 419, 422, 424, 428, 430, 432, 435, 436, 438, 440, 443, 444, 446, 447, 448, 450, 451, 452, 453, 454, 455, 457, 458 gonzalezherrejoni 422, 424, 426, 428, 430, 433, 434, 436, 440, 443, 446, 450, 456, 457, 458, 459 haematopotum 366, 419, 421, 424, 426, 428, 430, 432, 434, 436, 438, 439, 440, 441, 446, 447, 448, 470 idahoense 442 iracouboense 422, 423, 426, 428, 430, 431, 434, 436, 438, 440, 443, 445, 446, 447, 452 jacobsi 470 jerezense 459 letrasense 459 longithallum 422, 424, 426, 427, 430, 433, 434, 436, 450, 456, 457, 458, 459 mangabeirai 422, 424, 427, 430, 433, 435, 438, 443, 446, 447, 450, 452, 456, 457, 458 marathrumi 448, 449 mazzottii 455 mediovittatum 421, 423, 428, 429, 432, 435, 436, 439, 440, 441, 442, 447, 449, 452 mooseri 461 netteli 459 ochoai 421, 424, 426, 427, 429, 432, 435, 436, 439, 441, 442, 447, 449, 450, 452 olympicus 459 panamense 420, 421, 423, 425, 428, 429, 431, 433, 436, 438, 440, 446, 447, 450, 452, 460 penai 421, 423, 425, 427, 429, 431, 433, 435, 464, 467, 469, 470 pseudocallidum 420, 421, 423, 425, 428, 429, 431, 434, 437, 438, 450, 452, 461, 462 pseudohaematopotum 448 rufidorsum 419, 449 samboni 422, 423, 426, 428, 430, 431, 434, 437, 438, 440, 443, 444, 445, 447, 449, 452, 461, 470 santaelenae 445 sp. near dandrettai 425, 434, 447, 450 sucamense 445

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tenuipes 421, 423, 425, 427, 429, 431, 433, 435, 464, 467, 468, 470 travisi 462 trivittatum 422, 424, 425, 428, 430, 433, 434, 437, 438, 440, 441, 446, 449, 450, 452, 455, 456, 457, 458 veracruzanum 422, 424, 426, 428, 430, 432, 434, 437, 440, 443, 446, 447, 450, 456, 457, 458, 462 vulcanoae 459 zempoalense 422, 424, 425, 427, 430, 432, 434, 436, 443, 447, 450, 452, 454, 457 Simulium (Psilozia) 117, 120, 123, 127, 131, 134, 136, 138 argus 139 encisoi 139 groenlandica 138 kamloopsi 139 vittatum 139 Simulium (Pternaspatha) 115, 116, 119, 123, 125, 126, 127, 130, 134, 136, 630, 631 albicinctum 142, 144, 147, 151, 154, 157, 202, 210, 211, 213 albicinctus 210 albilineatum 143, 145, 147, 149, 152, 154, 157, 159, 199, 202, 204, 205, 206 annulatum 141, 143, 147, 149, 150, 153, 157, 159, 216, 217, 235 bachmanni 160, 163 barbatipes 142, 144, 147, 149, 152, 153, 156, 159, 174, 191, 193, 207 bordai 140, 143, 147, 150, 152, 157, 205 caprii 141, 144, 146, 149, 151, 153, 155, 159, 218, 220, 224 cotopaxi 142, 143, 147, 149, 151, 153, 157, 159, 185, 187 deagostinii 141, 143, 144, 146, 148, 150, 152, 153, 156, 158, 171, 172, 181, 183, 184 diamantinum 141, 144, 146, 148, 150, 153, 156, 158, 168, 171, 178, 180, 181, 183 dureti 143, 145, 146, 147, 149, 152, 154, 156, 159, 166, 168, 170, 172, 181, 183, 184, 214 edwardsi 160

figueroa 173 hectorvargasi 142, 144, 146, 149, 151, 154, 155, 158, 206, 207, 208, 209, 210, 212, 213 herreri 143, 144, 145, 147, 148, 152, 154, 155, 156, 158, 183, 196, 197, 200, 204, 206, 208, 210, 212, 214 horcochuspi 141, 143, 146, 149, 150, 152, 156, 159, 172, 173, 181 huemul 163 illiesi 173 limay 141, 144, 146, 148, 150, 153, 155, 158, 163, 164, 168, 169 luchoi 142, 144, 145, 147, 151, 154, 155, 157, 212 nemorale 142, 143, 144, 145, 148, 151, 152, 153, 155, 158, 166, 168, 169, 171, 172, 174, 176, 179, 188, 189, 192, 194, 208 nigristrigatum 141, 144, 146, 148, 150, 153, 155, 157, 159, 161, 165, 196 philippii 142, 144, 147, 148, 152, 153, 157, 158, 194, 195 pichi 143, 145, 146, 149, 152, 154, 156, 158, 165, 167, 171, 181, 183, 184 prodexargernteum 141, 142, 143, 144, 145, 148, 150, 151, 153, 154, 155, 157, 213, 215 pulchrum 142, 145, 146, 148, 152, 154, 155, 157, 174, 176, 177 punctativentris 173, 174 quechuanum 143, 144, 147, 149, 152, 154, 156, 158, 206, 207, 208, 213 schoenemanni 140, 143, 145, 147, 150, 152, 154, 157, 221, 224 simile 142, 145, 148, 152, 154, 155, 158, 173, 175, 176, 177 stelliferum 142, 144, 147, 149, 151, 153, 156, 158, 159, 194, 221, 223, 224 strigidorsum 29, 141, 143, 145, 147, 150, 152, 154, 157, 181, 182 varipes 139, 216 walterwittmeri 163, 165 yacuchuspi 143, 145, 147, 148, 152, 154, 156, 158, 199, 201, 204, 207, 210, 211 Simulium (Thyrsopelma) 116, 118, 121, 123, 126, 130, 131, 132, 134, 518, 612, 647

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A, B, C, D (cytoforms) 620 albimanum 616 albopicta 626 albopictum 626 brasiliense 612, 616 guianense 613, 614, 615, 616, 617, 619, 621, 622, 623, 624 hirtipupa 613, 614, 615, 616, 618, 619, 620, 622, 623 itaunense 613, 614, 615, 616, 620 itaunensis 614, 620 nigrimanum 616 orbitale 612, 613, 614, 615, 616, 618, 619, 620, 621, 622, 623, 624 ortizi 620 paraguayense 616 perplexa 624 perplexum 613, 614, 615, 616, 624 pinto 620, 624 scutistriata 625 scutistriatum 613, 614, 615, 616, 618, 619, 621, 623, 625 Simulium (Trichodagmiia) 116, 118, 119, 121, 122, 124, 126, 129, 130, 132, 133, 135, 518, 597, 612, 616, 617, 622, 624, 625, 626, 646

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chalcocoma 601 ecuadoriense 601 huairayacu 598, 599, 600, 601, 602, 603, 604, 605, 606, 607, 608, 609 lahillei 30, 597, 598, 599, 600, 602, 603, 604, 605, 606, 607, 608, 610 latitarse 597, 601 muiscorum 598, 599, 600, 601, 603, 604, 608 nigrimanum 598, 599, 600, 601, 602, 604, 606, 609, 610 pruinosa 609 pruinosum 609, 611 spadicidorsum 609 strigata 601 sumapazense 598, 599, 600, 601, 603, 605, 606, 609 townsendi 597, 598, 599, 600, 601, 602, 607 wygodzinskyorum 598, 599, 600, 601, 604, 605, 606, 608 Simulium (Wilhelmia) 228, 365 Simulium (Xenosimulium) 518, 612 Tlalocomyia 24, 39, 41, 42, 44, 45, 48, 49, 51, 54, 63, 67, 629 alticola 54, 65 revelata 63, 64, 65

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9 ADDENDUM Simulium (Chirostilbia) jefersoni HAMADA, HERNÁNDEZ, LUZ & PEPINELLI Simulium (Chirostilbia) jefersoni HAMADA et al., 2006: 23-35.

Diagnosis based on original description. Female: Wing length 1.3-1.6 mm. Scutum black with recumbent whitish setae, showing 1+1 submedian and 1+1 sublateral silver stripes, changing to blackish with different light position. Pleura dark brown. Legs yellowish with apical portion dark brown; Sc with hair, basal section of R bare. Claw with subbasal tooth. Paraproct subtriangular, about 1.5 times longer than cercus. Gonapophysis subovoidal, elongated, lightly sclerotized on internal margin. Male: Wing length 1.6-2.5 mm. Scutum black with golden hairs, showing with some light 1+1 submedian silver pruinose spots on anterior one third; scutellum and metanotum dark brown. Ventral plate very short with ventral margin mostly straight and well sclerotized. Gonostylus finger-like, slightly shorter than gonocoxite, with longitudinal ridge and 2 small apical spiniform setae. Pupa: Cocoon shoe shaped with loop-like fenestrations protecting gill branches; length ventrally 3.8-5.4 mm, and dorsally 2.8-3.8 mm. Gill length about 1.9 mm. Frontoclypeus and thorax dorsally with rounded tubercles and multiramous trichomes. Gill with 8 branches very open, with stout primary branches. Larva: Maximum length 5.8-6.6 mm. Head pale brown; cephalic apotome slightly darkened on median basal longitudinal area. Postgenal cleft subtriangular; postgenal bridge 0.7 times length of hypostomium. Antenna as long as cephalic fan stem; ratio of proximal, medial and distal articles = 1: 0.4-0.5: 1.5-1.6. Cephalic fan with 43-47 rays. Anal ring with about 100 rows with 13-14 hooks each. Rectal papillae with 7-9 diverticula on each lobe. Distribution: Brazil: Bahia. Bionomics: This species breeds in fast-flowing streams of tropical areas on trailing vegetation and rocks, at a median altitude of about 1600 m. Female feeding habits are unknown. Females have not been collected biting humans. Discussion: Adults are externally similar to other species of the S. (Chirostilbia) pertinax species group, but females can be differentiated by the relatively short paraproct, and males by the subovoidal finger-like gonostylus and short ventral plate. The pupa has 8 gill branches and a shoe-shaped cocoon, as in S. empascae, but can be differentiated by the gill primary and secondary branches grouped like a bundle, contrary to the isolated and open condition in S. jeffersoni. Larvae are similar to those of S. laneportoi.

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Simulium (Inaequalium) maranguapense COSTA PESSOA, RÍOS-VÉLASQUEZ & PY-DANIEL Inaequalium maranguapense COSTA PESSOA et al., 2005: 5-13. Diagnosis based on original description. Female (Pharate): Wing length 1.7 mm. General coloration dark brown to black; scutum without evident silver stripes; legs pale yellowish, with distal ¼ of fore leg tibia and tarsi blackish; coxa, trochanter and distal portion of femur and tibia of median and posterior leg blackish. Cibarium not elevated medially, similar to that of S. souzalopesi. Gonapophysis subtriangular, divergent, with internal border little sclerotized. Paraproct relatively short, subtriangular. Male: Unknown. Pupa: Cocoon slipper shaped. Length at base 2.6-3.1 mm, dorsally 2.1 mm. Frontoclypeus with abundant rounded tubercles and bifid trichomes. Gill with 4 thick branches, 2 dorsal and 2 ventral, anteriorly directed and becoming gradually thin distally; surface with small rounded tubercles. Thorax with rounded tubercles and filiform trichomes with 1-5 branches. Larva: Maximum length 4.1-4.8 mm. Cephalic apotome yellowish brown, pattern similar to that of S. diversibranchium. Antenna longer than cephalic fan stem; ratio of hypostomium/postgenal bridge = 0.6-0.7. Cephalic fan with 39-41 rays. Lateromandibular process simple, not bifurcated. Anal ring with 81-83 rows with 9-l3 hooks each. Rectal papillae with 0-3 diverticula on each lobe. Distribution: Brazil: Ceará. Bionomics: This species breeds on deciduous leaves and rocks in small, rocky streams at an altitude of about 600 m. Female feeding habits are unknown. Discussion: The female is similar to that of S. souzalopesi, with similar cibarium morphology, but it can be differentiated by the longer paraproct of S. souzalopesi. The pupa of S. travassosi is similar, but has only 3 gill branches. The larval rectal papillae, with 0-3 diverticula on each lobe, is a unique character of S. maranguapense among species of S. (Inaequalium). Simulium (Inaequalium) margaritatum PEPINELLI, HAMADA & LUZ Simulium (Inaequalium) margaritatum PEPINELLI et al., 2006: 48-58.

Female: Wing length 2.0 mm. General coloration blackish; Scutum uniformly black without silver stripes in any light, and golden green hairs. Cibarium with teeth arranged on median elevation. Sc with setae; basal section of R bare. Coxa, trochanter, distal portion of tibia and tarsomeres dark brown. Gonapophysis subtriangular with divergent internal margins sclerotized. Paraproct relatively short and with apical edge rounded. Male: Wing length 1.8 mm. Scutum black with 1+1 rounded silver spots, and golden hairs. Sc and basal section of R bare. Gonostylus subtrapezoidal and ventral plate relatively elongated, both similar to S. travassosi Pupa: Cocoon slipper shaped. Length at base 3.0-3.2 mm. Dorsally 2.4-2.7 mm. Frontoclypeus and thorax with abundant rounded tubercles, sometimes acuminate on thorax; cephalic trichomes with 2-3 and thoracic trichomes with 3-8 branches. Gill

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length 3.2-3.7 mm, with 4 filiform branches of same length, arising near base from 2 short primary branches. Larva: Maximum length 4.5-6.0 mm. Cephalic apotome pale dark brown, with pattern similar to that of S. diversibranchium. Postgenal cleft subtriangular with similar shape to that of S. diversibranchium. Postgenal bridge 0.7 times as long as hypostomium length. Subesophageal ganglion pigmented. Antenna as long as cephalic fan stem; ratio of proximal, medial and distal articles = 1: 0.7-08 : 1.3-1.4. Mandible with 1-2 lateromandibular processes surpassing teeth border. Cephalic fan with 44-48 rays. Rectal papillae with 5 or 6 diverticula on each lobe. Anal ring with about 80 rows with 12-13 hooks each. Distribution: Brazil: Bahia. Bionomics: The species breeds on deciduous leaves and rocks in small, rockybottomed creeks, with moderate temperature and slow flow, between 800 and 1700 m of altitude. Female feeding habits are unknown. Discussion: The adults of S. margaritatum are similar to those of other species of the subgenus S. (Inaequalium), particularly S. diversibranchium, which has the same female scutal coloration and male gonostylus morphology. The number of pupal gill branches separates S. margaritatum from other species of the subgenus. Simulium maranguapense, which also has 4 gill branches, has thicker branches. Another unique character of S. margaritatum, among species of S. (Inaequalium), is the bare basal section of R. Simulium (Thyrsopelma) duodenicornium PEPINELLI, HAMADA & TRIVINHO-STRIXINO Simulium (Thyrsopelma) duodenicornium PEPINELLI et al., 2005: 19-27.

Diagnosis based on original description. Female: Wing length 2.5 mm. Coloration and morphology as for S. guianense and S. orbitale, except paraproct internally concave in medial area, forming 2 short projections. Male: Wing length 2.4 mm. Scutum black without silver or gray spots. Genitalia similar to those of S. orbitale. Pupa: Similar to that of S. guianense, but gill branches shorter, thicker and without spicules in annular pattern. Larva: Maximum length 6.2-7.5 mm. Similar to other species of the subgenus; S. scutistriatum is most similar, but has more anal hook rows (280-300); S. duodenicornium has 174-184 rows with 26-31 hooks each. Distribution: Brazil: São Paulo. Bionomics: This species breeds on stems, leaves and roots of trailing vegetation in streams with fast flow and warm water at an altitude of about 900 m. Female feeding habits are unknown. Discussion: Simulium duodenicornium is similar to S. guianense, but can be differentiated by its thicker branches of the pupal gill, absence of silver spots on the male scutum, and medially concave region of the inner margin of the paraproct in the female.

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Simulium (Thyrsopelma) jeteri PY-DANIEL, DARWICH, MARDINI, STRIEDER & COSCARÓN Thyrsopelma jeteri PY-DANIEL et al., 2005: 465-471.

Diagnosis based on original description. Female and Male: Available material pharate, not showing characters that permit differentiation from other adults of Thyrsopelma species. Pupa: Cocoon shoe shaped, not completely protecting gill branches. Length at base 2.5-2.9 mm, dorsally 2.1-2.5 mm. Frontoclypeus and thorax with or without tubercles on different areas. Gill with 15 thick branches with rounded spicules on pseudoannulations. Larva: Maximum length 5.2-5.6 mm. Body cuticle with abundant racket-shaped scales. Cephalic apotome with uniform coloration. Antenna surpassing cephalic fan apex; ratio of proximal, medial and distal articles 1: 1.7-1.8: 1.1-1.5. Cephalic fan with 51-56 rays. Ratio of postgenal bridge/ hypostomium length = 1.0-1.2; postgenal cleft submiter shaped. Anal ring with 130-164 rows with 14-19 hooks each. Rectal papillae with 10-15 diverticula on each lobe. Distribution: Brazil: Rio Grande do Sul. Bionomics: The species breeds in rocky streams with crystalline water and marginal vegetation. Female feeding habits are unknown. Discussion: This species is difficult to differentiate from other Thyrsopelma species; only the pupal gill branches, numbering 15, permit identification. As in S. duodenicornium, it has thick gill branches.

REFERENCES COSTA PESSOA, F.A., RÍOS-VELÁSQUEZ, C.M. & V. PY-DANIEL (2005): First survey of Simuliidae (Diptera) from North Ceará State, Brazil, with descriptions of a new species and identification keys for the immature stages. - Zootaxa 1051: 1-18. HAMADA, N., HERNÁNDEZ, L.M., BESSA LUZ, S.L. & M. PEPINELLI (2006): Simulium (Chirostilbia) jeffersoni, a new species of black fly (Diptera: Simuliidae) from the state of Bahia, Brazil. Zootaxa: 1123: 21-37. PEPINELLI, M., HAMADA, N. & S. TRIVINO-STRIXINO (2005): Simulium (Thyrsopelma) duodenicornium, a new black fly species (Diptera: Simuliidae) from the southeast Region of Brazil. - Zootaxa: 1040: 19-27. PEPINELLI, M., HAMADA, N. & S.L. BESSA LUZ (2006): Simulium (Inaequalium) margaritatum, a new species of black fly (Diptera: Simuliidae) from the central region of Bahia State, Brazil. Zootaxa 1167: 47-60. PY-DANIEL, V., DARWICH, S.M., MARDINI, L.B., STRIEDER, M.N. & S. COSCARÓN (2005): Simuliidae (Diptera, Culicomorpha) no Brasil - XII sobre o gênero Thyrsopelma ENDERLEIN, 1934: descrição de T. jeteri sp.n, considerações sobre o cibário e chave para as pupas. - Acta Amazonica 35 (4): 463-472.