Encyclopedia of South American Aquatic Insects: Hemiptera - Heteroptera: Illustrated Keys to Known Families, Genera, and Species in South America

ENCYCLOPEDIA OF SOUTH AMERICAN AQUATIC INSECTS: HEMIPTERA - HETEROPTERA

Encyclopedia of South American Aquatic Insects: Hemiptera - Heteroptera Illustrated Keys to Known Families, Genera, and Species in South America

by Charles W. Heckman Olympia Washington, USA

Charles W. Heckman 93rd Ave. SW., 3625 98512-9193 Olympia Washington USA [email protected]

ISBN 978-94-007-0704-7 e-ISBN 978-94-007-0705-4 DOI 10.1007/978-94-007-0705-4 Springer Dordrecht Heidelberg London New York Library of Congress Control Number: 2011922502 © Springer Science+Business Media B.V. 2011 No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Cover photo: Gelastocoris angulatus Printed on acid-free paper

Springer is part of Springer Science+Business Media (www.springer.com)

This book is dedicated to my daughter, Sylvia Nickel

Contents Introduction…………………………………….………....……….………………1 An appeal for quality in taxonomic work……………..………….……...…………4 Scope of the work……………………………………...……...…………………....8 Acknowledgements………………………………………....……….……………10 Literature………………………………………………………....……….………10 Section 6: Hemiptera……...…..……………………………....…………………11 Key to the two major suborders of Hemiptera..………………....………………..12 Heteroptera……………………………………………....……………….13 Key to the infraorders of Heteroptera in South America…....................………… 15 6.1 Morphology……...….…...……………………………………………....……22 6.1.1. Morphology of the Gerromorpha…………...…………....………..34 6.1.2. Morphology of the Nepomorpha………………………....………..38 6.1.3. Morphology of the Leptopodomorpha………………....………….43 6.1.4. Changes during larval development…………………....……….…44 6.2 Ecology…………………………………………………...………....………...46 6.2.1. Ecology of the Gerromorpha…………...……………....…….……49 6.2.2. Ecology of the Nepomorpha………….......………....……………..52 6.2.3. Ecology of the Leptopodomorpha….......................…....………….54 6.3 Preservation and examination………………………...…………....……....….55 6.4 Zoogeography……………...………………………………………....……….59

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6.5 Taxonomic problems………...………………………………….………....….63 6.6 Suggestions for improvement………………………….………………....…...64 A note on the figures……….………………………………....…67 6.7 Keys to aquatic Heteroptera in South America….……………....……………68 6.7.1 Infraorder Gerromorpha…………………………………….....……68 Gerridae……………………………………………………....….72 Veliidae………………………...……………………….....……151 Hydrometridae……….……………………………....…………255 Hebridae…………………………………………………....….. 267 Mesoveliidae……………………………………….....………...275 6.7.2 Infraorder Nepomorpha………………………………....………...282 Corixidae…………………………………………………....….292 Nepidae………………………………………………....………356 Belostomatidae…………………………………………....……393 Gelastocoridae……………………………………....………….435 Ochteridae……………………………………….....………….. 457 Notonectidae……………………………………………........…464 Helotrephidae………………………………………....……….. 504 Pleidae…………………………………………………....…….509 Naucoridae…………………………………………....……….. 514 Potamocoridae………………………………………....……….574

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6.7.3 Infraorder Leptopodomorpha…………………………....………..578 Saldidae……………………………………………....………...579 Literature………………………………………………..………………....…….598 Index…………………………………………………..…………………....…....657

Introduction This work was begun to provide keys to the aquatic insect species known from Brazil. The original goal was to include all genera known from South America and all species from Brazil, but for most groups, the scope was expanded to encompass all species in South America, and, in some cases, to include terrestrial species of orders that include both terrestrial and aquatic taxa. In no case is a taxonomic revision of any group undertaken, although recommendations for such revisions are included where appropriate, and the probable synonymy of nominal species still treated as valid in the literature is noted. Two different approaches will be employed according to the taxon being treated. For phylogenetic groups encompassing overwhelmingly or exclusively aquatic species, such as the orders Plecoptera and Ephemeroptera or the families Dytiscidae and Culicidae, keys are provided to distinguish all genera and species known to occur in South America. An effort has been made to include every identifiable species so that the user of the key can determine with reasonable certainty whether or not his specimen belongs to a species that has already been described or whether it is one that is not yet known to science. Where feasible, complete keys will be prepared for groups containing both aquatic and terrestrial species, if they do not encompass an extraordinarily large number of species. This has already been done for the order Collembola. The second approach will be used for aquatic species belonging to predominantly terrestrial taxa, such as the order Lepidoptera or the families Curculionidae and Muscidae. In such cases, the number of terrestrial species involved is too great to deal with conveniently. For example, a work confined to aquatic insects cannot include a key to all of the approximately 45,000 weevil species (O'Brien and Wibmer, 1978) or even the portion of its enormous fauna inhabiting South America because only a small percentage of them live in or on aquatic plants. However, without such a comprehensive key, it becomes difficult for a person not specializing in the taxonomy of this group to be sure in which genus or subfamily his specimen belongs. For aquatic species in such groups, a descriptive approach is used. It is attempted to describe the morphology completely enough for the non-specialist to recognize his aquatic species. At the same time, descriptions or keys to higher taxa within these groups are provided as necessary so that the aquatic species can be distinguished from the terrestrial ones and phylogenetic affinities of the specimens can be recognized. The geographical limitations of this work are not strictly maintained throughout. For some genera, keys are provided that include mention or descriptions of species still known only from Central America or Caribbean islands, while for others, the keys are strictly limited to the South American fauna. The choice of comprehensiveness was dictated mainly by convenience. For genera encompassing few species and those that have been revised by competent taxonomists, keys to most or all Neotropical species could easily be C.W. Heckman, Encyclopedia of South American Aquatic Insects: Hemiptera - Heteroptera, DOI 10.1007/978-94-007-0705-4_1, © Springer Science+Business Media B.V. 2011

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provided. In the cases of other genera encompassing poorly described species, those which have not been reported south of Panama and the Antilles were not included in the keys. In general, the ranges of the aquatic insect species are very poorly known. The reported occurrences indicate more the locations at which entomologists have worked than the actual distribution of the species. The type specimens of a great many South American species were collected during expeditions to individual regions of the continent, and systematic surveys of the fauna have only been undertaken for a few groups. Among the prominent early collectors was Charles Darwin. Larger expeditions were undertaken during the late 19th and early 20th century. Considerable numbers of species have been described after examination of the specimens collected in Patagonia and South Chile during the early part of the 20th century. In Brazil, Rio de Janeiro and Santa Catarina appear very frequently in the distribution reports due to the extensive collection in those states by F. Plaumann, while many other species were described from Amazonas because of the presence of a large research station, the Instituto Nacional de Pesquisas da Amazônia, at Manaus. Naturally, a great many species were described from specimens collected at São Paulo and Rio de Janeiro, where most of the Brazilian institutes of research are located. Reports from other Brazilian states are generally much rarer, so the ranges of Brazilian species can generally be expected to be much more extensive than indicated by the published reports. Notable research efforts in other parts of South America have resulted in large numbers of species being recorded for Surinam and the Amazon region of Peru. Knowledge of the fauna in other regions varies considerably from taxon to taxon, depending upon the efforts of talented individuals specializing in individual orders or families. Generally, the insect taxa with the greatest impact on public health and agriculture are the best known. For some rarer species in neglected taxa, the ranges are scarcely defined at all. Specimens described during the 19th century were sometimes reported from "Brazil" or even "South America" without any more exact collecting data. The fact that the descriptions of such species are generally sketchy, at best, makes it doubtful whether they can be recognized again. There is mention in many of the review papers cited in this work of the confusion caused by falsely labelled specimens, incomplete collecting data, and doubt among the collectors about the names of the places from which their specimens originated. Names such as "Chapada" in Brazil frequently appear without mention of which Chapada is meant. This problem will continue to cause confusion in the future as it has in the past. The problems encountered in systematic biology worldwide are especially pronounced in the Neotropical Region. This is due to the dearth of detailed taxonomic studies on the regional fauna. Theoretically, the scientist who needs to know the correct name of a specimen can accomplish this by a procedure that assures a high probability of success, although a good deal of time and expense may be required. The first step in this procedure is the use of a comprehensive

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key treating the higher taxon to which the specimen belongs. This process is much easier for a specialist familiar with the group than for a person who needs to know the identity of species being used in physiological experiments or surveyed in synecological studies. In most cases, after the specimens have been identified using the keys, original or revised descriptions listed by the author of the key should be consulted to confirm the identifications. If the available descriptions are so poor that the identity of the species remains uncertain, a comprehensive taxonomic review of the taxon to which it belongs must be consulted. If such a review has not yet been published, the specimen to be identified must be compared with type specimens of each species belonging to the higher taxon to which the specimen belongs. These should be found in museum collections. According to taxonomic convention, the author of a published original description of a new species should designate one of his specimens as a holotype. This type then becomes the standard on which the identifications of all specimens collected in the future are based. The species to which the holotype belongs is then referred to by the name proposed by the author, at least when the same name has not previously been applied to another species. In addition to the holotype, other specimens believed to be of the same species by the author are designated as paratypes. If a holotype is not available, a paratype can be examined to confirm identifications. If the holotype has been lost or extensively damaged, a specialist revising the taxonomy of the group may designate a lectotype apparently belonging to the species in question. This then replaces the holotype as a standard for recognizing the species, at least until the lost holotype or paratype is found. Anyone wishing to take the trouble to identify a specimen beyond the shadow of a reasonable doubt should be able to follow this procedure. If his specimen is not the same species as any of the holotypes preserved in museums, he should prepare a description of his specimen and have it published with his proposed name for the new species or give it to a specialist who is interested in doing this. Unfortunately, the procedures described above often fail to work, especially when South American insects are involved. First of all, keys to identify the species are seldom available, and when they are, they are generally out of date or incomplete for the region being studied. This leaves the researcher with the chore of collecting a large number of original descriptions to match with his specimens. These descriptions are sometimes very sketchy, leaving the reader with no reliable way of identifying his specimen from the available literature. Comparisons with type specimens may not only be difficult in many cases because the types were deposited in museums on other continents, they are often impossible because some authors have failed to mention the name of the museum in which they intended to place their type specimens or because the specimen cannot be found again due to war damage or sloppiness by the curators of the museum in question. Even when the type can be found, it often proves to be in very poor condition, and relatively few paratypes of South

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American species have been designated that can be examined in the absence of a usable holotype. Using established procedures for identifying South American insects is made even more problematic by the practice of some authors of describing new species on the basis of only one life stage. Not only have many species been described by taxonomists who examined only adults, some are known only from larvae. Furthermore, countless species are known only from adults of one sex. While the description of an adult may provide a legitimate basis for establishing a new species, it is difficult to find justification for naming a species based only on larvae. Some authors have designated larvae as the types of species congeneric with others known only as adults. These individuals seemed to be in such a hurry to publish that they deliberately left the arduous task of matching larvae with adults to other researchers. In practice, convention should dictate which gender and life stage should be chosen as the holotype. For example, adult male chironomids are presently those on which species descriptions should be based. Earlier descriptions of female adults are generally useless for determining a species unless someone has taken the trouble to match the female to a described male. In an ideal system of nomenclature, descriptions of adults of both sexes and of the larval stages should be provided. However, the state of the art still requires specimens to be identified according to partial descriptions of one stage and often one sex. Therefore, before fully workable systems for identifying South American species will be possible, an enormous number of revisions and supplemental descriptions will be necessary. This work is meant as a first step in the process: providing keys as reliable as the available publications permit. Finally, keys to taxa higher than families are provided only to delimit groups of aquatic species from terrestrial ones. It must be noted that obsolete names of insect taxa are defined or noted in the keys whenever users of these books are likely to encounter the names in old publications. They are neither endorsed nor suppressed but merely defined. Keys to insect orders can be found in any number of textbooks on entomology or invertebrate zoology, and most specimens can be quickly assigned to the correct higher taxon by referring to the illustrations in this series. With a little experience, anyone can learn to recognize the order of most insects almost at a glance.

An appeal for quality in taxonomic work This appeal is addressed to two groups, the first consisting of those responsible for deciding who obtains what portion of the available research funds and the second being the taxonomists themselves. It has long been recognized by experienced ecologists that identification of the species present in a community is an absolute necessity for thorough ecological research. The biota of any water body, for example, is more than a quantity of "biomass" or a "pathway for energy". Many ecologists, especially those

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beginning their research careers, have the greatest difficulty in identifying the species present, and the quality of their work is limited by this difficulty. The ecologist may indeed investigate systems using methods very different from those engaged in descriptive biology, but he is nevertheless dependent on a sound basis of taxonomic information for the proper reporting of his results. Briefly stated, ten may be equal to ten, but ten oranges are not equal to ten cows, and ten of one dragonfly species are not equal to ten of another. The "emergency measure" of identifying a species only as far as the genus or family is not satisfactory because two congeneric species may have completely different habitat preferences, feeding habits, and seasonal activity cycles. In short, for one ecologist to properly compare his work with that of another, he must know the name of the species he has encountered and of those that have influenced the biotic communities he has been studying. In the past, ecologists usually relied on taxonomic specialists to identify their specimens. Since the specimens invariably belonged to a wide variety of taxa, they were distributed among a number of taxonomists, each of whom was an expert only for his own small phylogenetic group. This practice has ceased to function in recent years because of the failure of those in charge of distributing research funds to support scientists who wish to engage themselves in taxonomic work professionally. As the taxonomists reached retirement age, they left active research without being able to pass their knowledge on to a successor. As time went on, more and more personal knowledge and skill that is difficult or impossible to get from books was permanently lost. The disappearance of the best taxonomists was accompanied by a decline in the quality of ecological work. This may not be considered totally undesirable by persons in government and business who are constantly confronted by the demands of environmentalists. As ecology moves increasingly out of the laboratory and into the courtroom, the ground rules change, and where principles remain unclear, the lawyers have more room to maneuver, using their skills of persuasion rather than being forced to confront research results already proven beyond the shadow of a reasonable doubt. Furthermore, laws passed to protect endangered species can be better circumvented if no people can be found who are capable of identifying such species. Although there may well be incentives to suppress taxonomic research, the virtual demise of taxonomy as a field of biology has been instigated mainly by the scientific community itself. Ambition and the desire for quick success have motivated the staffs of many institutions to seek persons promising to achieve great breakthroughs, and no place was left for routine systematic work, which is by nature slow and methodical. Often, it was the number of publications rather than their quality or length that determined whether or not a person could succeed in finding gainful employment in science. Recently, science citation indices have become the criterion for judging success, prompting young researchers to investigate whatever everyone else is investigating in order to assure themselves of a maximum number of other scientists who would have reason to cite their work. Pressure was placed on the young scientist to come up

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with something clever very quickly, leaving no time to learn in depth about the organisms he was encountering. It is especially unfortunate that the worldwide elimination of taxonomist positions came just prior to the development of computer equipment that makes it very easy to establish data banks. Thus, science was robbed of the personnel competent to systematize and disseminate the great body of information that had accumulated during two centuries of intensive work by systematic biologists in all parts of the world. A long time went by without any crisis becoming evident because the taxonomists and systematic biologists already working in the museums and universities continued donating their time to help identify specimens collected or used in the laboratory by ecologists, physiologists, geneticists, and biochemists. Even after their retirements, many continued contributing time to help younger colleagues in other fields of biology. Moreover, a number of excellent specialists were employed in other fields of work rather than in the biological sciences and made their contributions to the knowledge of the world's flora and fauna as a hobby. With the gradual elimination of the competent taxonomists and systematic biologists through death or disability, however, the problems of defining community structure, determining species diversity, and identifying the organisms causing some problem in the field have been intensifying. For many taxa, there are no longer experts who can be consulted, or those experts that are still active are hopelessly overworked. The result is the loss of a vast body of information about the flora and fauna in all parts of the world. Without knowledge of the species that have been encountered, the ecologist is faced with a dilemma. He must learn to identify a vast array of species belonging to the widest variety of phylogenetic groups, or he must limit his research to superficial phenomena. Hence there has been an increasing tendency to use numbers instead of names and to try to equate quantities of unlike and undefined substances or objects, although this cannot yield any reliable results, as mentioned above. The second aspect of the problem must be dealt with by the taxonomists themselves, who have in the past often been guilty of laxity in quality control of their work. The ambition to gain status by naming new species has certainly been a motivating factor for many of these scientists, and for this purpose, it is easier to write short papers with sketchy descriptions of aberrant specimens than to produce comprehensive reviews compiling the available information on genera or families together with thorough, well illustrated descriptions of new taxa. As mentioned above, it is evident that the examination of type specimens for positive identification of South American insects is often impossible because none were designated, or if they were, they have been lost or greatly damaged. For better or worse, the published literature frequently provides the only criteria for identifying species. While compiling this key, the enormously broad spectrum of quality in the taxonomic publications became evident. Some papers are of very high quality, such as the works of Belle on the Odonata. These publications provide

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descriptions of all essential morphological characters, the exact locations of type specimens, and precise collecting data. Any morphological details lacking in the text can be seen in the detailed illustrations. They make it clear that the species described are actually distinct from all species previously described. In general, more recent publications tend to include better descriptions than older ones, but this is not the case with all taxa. Many works more than a century old are still useful for identifying species, and some of the information they provide cannot be found in any more recent publications. On the other hand, many recent publications are extremely poor, providing either very sketchy descriptions or showing a very poor comprehension of the concept of species as a closed phylogenetic grouping of individuals, which may possess a considerable degree of individual variability (Mayr, 1963). Many of the better taxonomic publications are rather long, but length is no guarantee of quality. There are some excellent concise works, which can be used with great reliability for the identification of species. Some very long works, including incredible amounts of detail, on the other hand, are very poorly organized and omit much essential information while including vast amounts of trivia. Authors who are more collectors than biologists have used very keen, practiced observation abilities to find the most minor differences among single populations to produce massive numbers of nominal species that are indistinguishable to less skilled observers. Among the European fauna, long lists of synonyms attest to the vast amount of printer's ink that has been squandered on the description and suppression of spurious taxa. Unfortunately, revisions of many South American taxa are still lacking, and many of the nominal species included in the keys will certainly prove to be junior synonyms when more is known about the variability of natural populations. In the following sections, minimum criteria will be suggested for describing new taxa. All of the features an author should describe to define a new species or subspecies within a particular taxonomic group will be mentioned. It is further suggested, as a general rule, to base descriptions on a holotype and several paratypes. New species described from only one or two specimens should be regarded as questionable, particularly when the morphological differences between them and specimens of other nominal species are very slight. The practice of choosing type specimens that are missing various structures, such as legs, abdomens, or antennae, should be discouraged. With modern means of mass transportation available, there should be no serious obstacle to collecting more material from the locations at which the specimens of the alleged new species were encountered, and funds should be provided for such collecting for the reasons outlined above. It would probably be justified to declare many names of species poorly described and without known type specimens as nomena inquerendae and omit them from the key. However, whenever distinctive features were found in the original description that can be used to distinguish a species from all other known species in the genus, the name is included in the keys. Names from the keys must therefore be used with caution because one or more undescribed

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species may also have the characteristics in question. Where such situations are apparent, it is hoped that specialists for the groups in question, if any are presently active, will designate lectotypes for the species most likely to have been referred to by the earlier authors. Holotypes, paratypes, and original descriptions should be provided for any similar, congeneric species that were hitherto undescribed.

Scope of the work The families recognized are mainly those found in the key of Brues et al. (1954), and some of the new families that have been described by various authors after the appearance of that key are omitted because they have not been generally accepted, are too poorly defined, are too similar to other families to recognize any general definitive characteristics for use in the key, or seem for another reason to be of doubtful validity.

Fig. 1: The geographical scope of this work is limited to continental South America and the offshore islands shown on this map.

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Fig. 2: The Brazilian states where the species were found are generally reported as shown on this map. Older reports, however, may not reflect the modern political boundaries, so it often requires knowledge of the travels of entomologists to determine the precise locations of known occurrence. D.F. designates the Federal District of Brasilia. Geographically, the work is limited to continental South America and islands very close to the coast, such as Tierra del Fuego (Fig. 1). Islands far enough offshore to be zoogeographically distinctive, such as the Galapagos Islands, are only included in some of the volumes, even though politically, they are parts of South American countries. Case by case decisions to include them depend upon the availability and reliability of published surveys of the local fauna. Except in

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the case of Brazil, the distribution reported in the keys usually refers only to the country in which the species have been found. When they have been reported in the literature, the Brazilian states are shown, as depicted in Figure 2. However, some inaccuracies may have been inadvertently introduced in the cases of states which have changed their boundaries since the insects were described. For example, Mato Grosso formerly encompassed the present states of Mato Grosso, Mato Grosso do Sul, and Rondônia, so species reported in older literature from Mato Grosso may actually be confined to any one of these three present states.

Acknowledgements The preparation of these keys required the search of a large amount of literature. A great deal of help was provided in obtaining many obscure or archaic papers by Frau G. Lechner of the Max-Planck-Institut für Limnologie Library in Plön, the staff of the Library of the Zoologisches Institut und Zoologisches Museum of the University of Hamburg, the New York Public Library, and the Timberland Public Library System in Washington State, U.S.A. I especially thank Dr. Silvia A. Mazzucconi of the Laboratorio de Entomologia, Universitad de Buenos Aires, for sending me a copy of a paper published in a journal that is especially difficult to obtain, thereby avoiding a delay in the completion of the manuscript. Other works were provided courtesy of Senhora Marilia Junqueira of the Fundação Centro Tecnológico de Minas Gerais in Belo Horizonte.

Literature Brues, C. T., A. L. Melander, and F. M. Cartenter, 1954: Classification of Insects. Keys to the living and extinct families of other terrestrial arthropods. - Bulletin of the Museum of Comparative Zoology at Harvard College 108:1-797. Mayr, E., 1963: Animal Species and Evolution. - Harvard Univ. Press, Cambridge, Mass. 797 pp. O'Brien, C. W., and G. J. Wibmer, 1978: Genera and species of Curculionidae (Coleoptera). - Entomological News 89(2-3):89-92.

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Section 6 Hemiptera According to the most widely accepted classification system, the order Hemiptera, originally named by Linnaeus (1758), includes two main groups (Latreille, 1810): Heteroptera, a suborder with a large number of truly aquatic species, and Homoptera, encompassing in South America only species which are fully terrestrial or which inhabit the emergent parts of aquatic plants. This nomenclature has long been controversial, and in the past, North American specialists called the order Heteroptera and the suborder herein given that name, Hemiptera, reversing the present meanings of these words. At other times, the name Rhyngota, later spelled Rhynchota, originating with Fabricius (1803), has been applied to the order or to the suborder. The taxonomy used in this volume is that currently accepted by the Zoological Record. The common name applied to all Heteroptera species is bug, often modified to true bug because bug has other colloquial meanings. Aquatic species are referred to as water bugs. Those few members of the Homoptera specialized for feeding on emergent aquatic plants have developed behavioral modifications for living on or near water bodies, where they colonize their hosts. Although they may be encountered in the littoral zone or in the pseudoterrestrium, or “floating islands” of vegetation covering large water bodies (Heckman, 1994), their habit of sucking juices from plants while resting on their emergent shoots would hardly differ in any way from that of related species living on the juices of fully terrestrial plants. However, personal observations indicate that they are better able to contend with falling onto the surface of the water than fully terrestrial species, especially during periods of heavy rainfall. Most homopteran species native to South America are poorly known and will not be considered further in this volume. Recent studies have called into question the traditional classification system separating the order Hemiptera into Heteroptera and Homoptera, but there is still no concensus on the validity of this or any other new system. The grounds for proposing changes include findings that a few groups now included in the Homoptera are more closely related to the Heteroptera than to the rest of their own suborder, as presently understood. The Homoptera itself is therefore thought to be an artificial assemblage of at least two groups that are not closely related. Cladistic studies suggest that members of Coleorrhyncha, now considered to be homopterans, are more similar to heteropterans than to any other group of Homoptera. The proposed taxon, Euhemiptera, would encompass these two groups, as well as the Cicadoidea, Cercopoidea, Cicadelloidea, and Fulgoroidea, now included in Homoptera. The rest of the Homoptera, as presently recognized, would be placed in a group called Sternorrhyncha, encompassing the groups Psylloidea, Aieyrodoidea. Coccoidea, and Aphidoidea.

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These considerations are beyond the scope of this book, and any recombination of the infraorders based on new phylogenetic discoveries would not affect the limitation of the scope of this volume to the three groups, usually regarded as infraorders, which encompass all of the families of aquatic true bugs in South America. For more information about recent studies on the higher classification of the Hemiptera, refer to Cranston and Gullan (2003).

Key to the two major suborders of Hemiptera 1. The piercing and sucking mouthparts arise at the anterior margin of the head, or, if they arise on the ventral side, they are wide, short, and have only one segment, or the fore-tarsus consists of a single segment, longer than the tibia and lacking a claw at the apex. The fore-wings are modified into covers for the hind wings called hemielytra, alternately spelled hemelytra, which have a characteristic morphological structure consisting of a corium, cunium, clavus, and membrane (Fig. 6.1). However, morphological forms of many aquatic species have reduced wings or lack them entirely. The adults of a few species shed their wings or parts thereof. ............................................................................................................Heteroptera - The piercing and sucking mouthparts arise on the ventral side of the head, often far posterior from the apex. The fore-tarsus is never modified to form a large, flattened structure longer than the tibia. The fore-wings are membranous, often differing little from the hind wing and not differentiated into distinct structures to cover the hind wing (Fig. 6.2). Adults of some species shed their wings. All known species live by sucking the juices of plants. ............................................................Homoptera….not included in this volume

Fig. 6.1 Habitus of Ambrysus scolius. Based on La Rivers (1970).

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Fig. 6.2 Peloridium hammoniorum Breddin, 1897, peloridiid from Chile. Peloridiidae is a family of homopterans more similar to heteropterans than any other members of the suborder. Based on China (1962a).

Heteroptera The suborder Heteroptera is well represented in South America, and some of its species have considerable impacts on agriculture and public health. Some terrestrial species are serious pests, but aquatic species are generally beneficial because of their roles in controlling harmful insect species, especially mosquitoes and blood-sucking gnats. The truly aquatic species are phylogenetically distinct from the primarily terrestrial ones, making it easy to delimit them for this volume. Thus, the systematic groups considered here can be characterized by their ecological requirements for aquatic or marine habitats. Entomologists formerly grouped most aquatic species in a separate taxon called Hydrocorisa, while terrestrial species were placed in the Geocorisa. Systematic entomologists no longer use the terms Hydrocorisa and Geocorisa, although they clearly delimit the water bugs phylogenetically and ecologically from the true bugs with terrestrial habits. This volume is devoted to all families that were formerly included in the Hydrocorisa plus the Leptopodomorpha, which includes the family Saldidae. A few species belonging to other families of Geocorisa may feed on the emergent parts of

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aquatic plants, but those are not considered in this volume, and little information about them is available. Current systematic divisions of the suborder Heteroptera that are considered in this volume are the infraorders Nepomorpha, Gerromorpha, and Leptopodomorpha, as defined by Schuh and Slater (1995). The other infraorders, encompassing predominantly terrestrial species, can be identified using the appropriate key in this section. They are not further considered. Nepomorpha includes the superfamilies Nepoidea, Corixoidea, Naucoroidea, and Notonectoidea. The Gerromorpha encompasses only the Gerroidea, and the Leptopodomorpha, the Leptopodoidea. The families belonging to each group are shown in the key in Section 6.7. While many of the terrestrial bugs are herbivorous, almost none of the aquatic species feed primarily on plants, and those few corixids that do seem to consume microscopic algae probably consume any organisms of suitable size and should be considered omnivores. The feeding habits of the individual South American species have seldom been studied, and in many older publications, the corixids were falsely classified as being primarily herbivorous. In fact, most seem to prey on a variety of microscopic metazoans, such as rotifers, as well as protozoans and algae. The South American water bugs belong to families with nearly cosmopolitan or, in the case of the Belostomatidae, circumtropical and subtropical distribution. The South American fauna is very rich, but it seems premature to make any conclusions concerning the total number of species in each family on that continent. Many regions and various kinds of habitat have not yet been adequately surveyed, and the intraspecific variability must be better understood to assure that the nominal species are valid. As already mentioned, Heteroptera was formerly divided into two groups that would be convenient to use in this work but no longer have any taxonomic significance: Geocorisa and Hydrocorisa. Geocorisa encompassed a large variety of terrestrial species and a few species associated marginally with aquatic habitats, notably all species of the Saldidae or shore bugs and three species that inhabit the pseudoterrestrium (Bachmann and Mazzucconi, 1995), the mats of floating vegetation that often cover large areas of the standing and slowly flowing water bodies. These names are introduced here because they are sometimes encountered in older publications with comprehensive keys to the major taxa, so anyone dealing seriously with taxonomy of the Heteroptera should be familiar with their meaning. The classification system used in this volume recognizes three major taxonomic groups, usually referred to as infraorders, encompassing species that are mainly or entirely aquatic. Most water bugs classified as Nepomorpha live under water and breathe atmospheric air, which they periodically refresh, either by actively swimming to the surface or by the use of a long or short breathing tube. The species of Gerromorpha typically seek their prey while moving across the surface tension layer of the water. Most species belonging to these two groups have hydrophobic layers of tiny setae protecting their spiracles from

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being blocked by water. The length and arrangement of these setae, which either bind air underwater or repel water from the bodies of insects that live on the surface, are distinctly different according to the habits of the species (Heckman, 1983). Members of Leptopodomorpha include the shore bugs, which live at the edges of water bodies. Within each of these groups, the genera of each family are also distinctive by possessing typical structures that make them immediately recognizable at a glance, with only a few exceptions. As already stated, almost all members of these three major groups are predators, with a few probable exceptions among the Corixidae. There are few reliable published reports on corixid feeding habits, and some older entomology texts suggest that they consume detritus or even aquatic plants. Many recent studies indicate that they prey on microscopic organisms, including “protists,” that are alternatively classified as plants, animals, or both. The nutrition of corixids is a rich field for entomological research since considerable speciesspecific differences in food preferences are suspected. Species in most other families of this group prey on relatively large animals, mainly arthropods but including some vertebrates. The size of the prey is roughly proportional to the size of the predator, with the smallest insects being captured by the microveliine veliids and relatively large fishes and amphibians falling prey to the largest belostomatids (Haddad and Bastos, 1997; MijaresUrrutia et al., 1997).

Key to the infraorders of Heteroptera in South America Information for the key was provided by Schuh and Slater (1995) and Schaefer (2003). 1. The head has a transverse constriction, dividing it distinctly into two lobes. Compound eyes are usually present, but they are absent or reduced in a few species. If ocelli are present, they are located on a lobe posterior to the compound eyes. The fore-tibiae are flattened and have distinct distal spines. The fore-tarsus has one or two segments, which form an opposing organ for grasping with the anterior part of the fore-tibia. If fore-wings are present, the texture of their surfaces is uniform. A claval commissure is absent, so a distinct corium and membrane are not evident (Fig. 6.3). ...............................................................................................Enicocephalomorpha Two families in this infraorder are known: Aenictopecheidae and Enicocephalidae. The species are terrestrial predators and are not further considered in this volume. - Either the head is not constricted and lacks distinct lobes, or there is a distinct sulcus in the prosternum to receive the apex of the labium, and the fore-tibia is not flattened or lacks distinct distal spines, which oppose the one or twosegmented tarsus (Fig. 6.4). Compound eyes are present, except in species parasitizing bats or inhabiting termite nests. .......................................................2

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Fig. 6.3 Enicocephalus schuhi Wygodzinsky and Schmidt, 1991: habitus without wings in dorsal view (middle left), antenna (upper left), head in lateral view (lower left), head with one antenna and thorax in ventral view (lower center), fore-wing (upper right), apex of hind wing (lower right), apex of the fore-leg (right of center) and details of the apex in ventral view (lower right center). This terrestrial species, found in Rio de Janeiro, belongs to the Enicocephalidae. Based on Wygodzinsky and Schmidt (1991).

Fig. 6.4 Habitus of a brachypterous Eurygerris fuscinervis in dorsal view (upper left), body of a macropterous individual (lower left), and the head and thorax in lateral view (right). Based on Alvarez and Roldán (1983), who used its synonym, E. kahli.

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2. (1) Three or four pairs of distinct trichobothria inserted in deep pits are present on the head near the inner margins of the compound eyes, which are always present. There is a coating of hair-like setae around the margin of each pit, which is deep enough to conceal the bothrium. If fore-wings are present, they lack a claval commissure, and a border separating the corium and clavus from the membrane is not evident (Fig. 6.4). A large part of the body is densely coated by tiny setae and many longer setae to repel water and maintain a coating of air on the surface of the body if the insect is forced below the surface of the water. The hind coxae are short and free-moving to permit rotary motion. They are typically found on the surface tension layer and occasionally along the shore of both freshwater and marine habitats. .................................................................................................Gerromorpha..p. 68 - Either the head lacks trichobothria, or, if present, they are not set in pits (Fig. 6.5). ......................................................................................................................3

Fig. 6.5 Ctenipocoris schadei (above, left to right): habitus of an adult and a fifth and a third instar larva, and (below, left to right): antenna, male genitalia, penis in ventral (above) and lateral view (below), paramere, and female genital capsule. Based on López Ruf (2004). 3. (2) The antennae are not as long as the head and are usually folded beneath it, in well-defined pits in all families except the Ochteridae. Compound eyes are always present and usually large; only the apices of the antennae are sometimes visible beyond the margins of these eyes. The poststernal sulcus is always absent. Most adults are macropterous, but a few have shortened hemielytra, resembling the wing covers of staphylinid beetles. Macropterous forms always have claval commissures (Fig. 6.5). The insects are typically found underwater, although flights may be undertaken during certain seasons. ................................................................................................Nepomorpha..p. 282

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- The antennae of most species are longer than the head and never concealed in well-defined grooves ventral to the compound eyes, or compound eyes are completely absent. If the antenna is shorter than the head, then a poststernal sulcus is present (Fig. 6.6). The insects do not live under water but are found on the surface tension layer or the shore, or they are terrestrial. .........................4

Fig. 6.6 Habitus (above) and lateral view of head and thorax (below) of Tropicoparapamphantus amazonicus Brailovsky, 1989, a species of Pentatomomorpha. Based on Brailovsky (1989). 4. (3) If compound eyes are present, either the fifth through seventh abdominal sternites each have at least one and usually two, three, or more trichobothria sublaterally, and usually the second and third sternites also have such trichobothria either submedially or sublaterally, or the first segment of the antenna is usually longer than the second; the third and fourth segments lack setae that are longer than the segment diameter; the body is flattened; the tarsi consist of two segments; an elongated pilvillus is always attached near the base of each claw. The pilvilli are elongated and attached to each claw only at the base. If compound eyes are completely absent, then the body is very much flattened and has marginal laminae (Fig. 6.6). Those lacking compound eyes and inhabiting the nests of termites belong to the family Termitaphididae, placed by some authors in this infraorder but in the Aradomorpha by others. See Couplet 6. ...................................................................................................Pentatomomorpha This infraorder encompasses many families and species, which are terrestrial and are not further considered in this volume.

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- If compound eyes are present, there is no more than one pair of setae resembling trichobothria along the midline of one or more abdominal sternites, or there is a single ventrolateral seta on the seventh sternite. If pulvilli are present, they are usually attached to the claws for most of their length. If compound eyes are absent, then the body is not flattened, and marginal laminae are absent (Fig. 6.7). Those lacking compound eyes are parasitic on bats. ......5

Fig. 6.7 Itagunannus itaguaiensis Wygodzinsky, 1947 male (above, left to right): habitus, 1.6 mm long, in dorsal view; outline of head and pronotum; claw on the fore-leg; and (below, left to right): head and thorax in lateral view, hemielytron; apex of the clypeus. The species belongs to the Schizoporidae, a family of the Dipsocoromorpha. Based on Wygodzinsky (1947). 5. (4) In all but the family Stemmocryptidae, the two basal segments of the antennae are short and subequal in length, and the terminal segments are long and narrow and bear setae much longer than the diameter of the segments. If hemielytra are present, they are tegminal or appear somewhat like the wing covers of a beetle (Fig. 6.7). Most species are shorter than 2.5 mm. The second segments of the antennae of members of the family Stemmocrypidae are longer than the first segments, and the fore-wings are membranous and undifferentiated with a few strong veins forming two cells, or, if more than two, they lack veins running from the posterior margins of the cells. Stemmocrypids also lack distinct pulvilli between the claws. Compound eyes are always present in all species. ....................................................................................................Dipsocoromorpha The species in the five families of this infraorder: Ceratocombidae, Dipsocoridae, Schizopteridae, Hypsipteryigdae, and Stemmocryptidae, are terrestrial and are not further considered in this volume.

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- The first segment of the antenna is usually longer than the second, or, if not, then the third and fourth segments lack setae longer than the diameter of the segment. Hemielytra are vestigial or absent, or, if developed, they are not tegminal and undifferentiated. The pronotum and hemielytra are covered with areoles (Fig. 6.8). Most species are longer than 2.5 mm. In a few species, compound eyes are absent. ...................................................................................6

Fig. 6.8 Emydocoris usingeri Wygodzinsky, 1948: habitus of a female in dorsal view (middle left to center), lateral outline of the body (lower right center), outline of the head of a male showing the location of one antennal segment (upper left), antenna (lower center), fourth or fifth instar larva (lower left), tarsal segments with claws and pulvilli (middle right), and rostrum showing the rostral groove (upper right). The species, known from Rio de Janeiro, belongs to the Aradidae, a family in the Aradomorpha. Based on Wygodzinsky (1948a). 6. (5) There are two tarsal segments bearing claws and usually elongate pulvilli, which are attached only at the bases of the claws. The body is greatly flattened and sometimes encrustate. If wings are present, they cover only the discal part of the abdomen, leaving the connexivum uncovered (Fig. 6.8). The compound eyes appear relatively small but have a number of ommatidia equal to those of most other species. Species without compound eyes and inhabiting the nests of termites belong to the family Termitaphididae, placed by some authors in this infraorder but in the Pentatomomorpha by others. ..........................................................................................................Aradomorpha This infraorder includes the family Aradidae, and some authors also assign the Termitaphididae here, as well. See Couplet 4.

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- The tarsi of most species consist of three segments, but if they consist of two, then either pulvilli are absent, or if present, they are not elongate and free but usually attached on one side to the claws. If wings are present, they usually extend to conceal the connexivum (Fig. 6.9). Compound eyes are usually present and appear relatively large, or, if reduced in size, relatively few ommatidia are present. If compound eyes are absent, the insects are parasitic on bats. .................................................................................................................7

Fig. 6.9 Saldolepta kistnerorum, a species in the Leptopodomorpha: habitus without legs in dorsal (upper left), ventral (lower left) and lateral view (upper right), dorsal view of the abdominal segments (center), fore-leg (middle right), and hind wing (lower right). Based on Schuh and Polhemus (1980). 7. (6) If compound eyes are present, the hemielytron of macropterous forms has a differentiated corium-clavus and a membrane, which sometimes has two closed cells and sometimes has more than two, each of which always has veins originating from their posterior margins. If the forewings are reduced or absent, they are not pad-like, and the body lacks a dense covering of minute setae for binding an air plastron (Fig. 6.10). If compound eyes are absent, then the body is not flattened and marginal laminae are absent. Those lacking compound eyes are parasitic on bats. .........................................................................................................Cimicomorpha The species in the 14 known families of this infraorder are terrestrial and not further considered in this volume.

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- Compound eyes are always present. If the fore-wings are tegminal, undifferentiated, or greatly reduced, they are pad-like, and part of the body is covered by a dense coat of minute setae for binding an air plastron (Fig. 6.9). The fore-wing may have a distinct membrane, may be reduced to small pads, or may be large and resemble the wing cases of a beetle. If the fore-wing has a membrane, even if it is reduced, its veins form three to five cells, none of which have veins originating from their posterior margins. If the fore-wings resemble those of a beetle, the compound eyes are large and cover almost the entire side of the head, and the length is about 1 mm. If the fore-wings are reduced to small pads, most of the body is covered by a dense coat of minute setae for binding an air plastron. The hind coxae are long and joined broadly to the thoracic pleurae. These are shore bugs, usually found at the water’s edge of both freshwater and marine habitats. .......................................................................................Leptopodomorpha..p. 578

Fig. 6.10 Habitus of Poicilocapsus citrinus Carvalho and Hussey, 1954, a species in the Cimicomorpha. Based on Carvalho and Hussey (1954).

6.1. Morphology Characteristic of the Heteroptera, correctly called bugs, are piercing and sucking mouthparts, as well as the modification of the mesothoracic wing pair to form structures covering the membranous posterior or metathoracic wings. These fore-wings are called hemielytra, sometimes spelled hemelytra, meaning “half wings” because their basal parts are modified to hardened wing covers, while their apical parts are membranous. Also typical is the wide thorax with its large triangular scutellum. Only a few other insects, notably several species of Coleoptera, can be mistaken at first glance for members of the Heteroptera,

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but an examination of the mouthparts quickly reveals whether or not the specimen is a true bug or a member of another order. Many species of Heteroptera have wingless or apterous forms (Fig. 6.11) or individuals with obviously reduced wings, called brachypterous forms (Fig. 6.12), in the adult stage. Adults with fully formed wings are referred to as macropterous (Fig. 6.13), which may be rare or even unknown among certain species. This causes some confusion and makes it more difficult to recognize adults at a glance. In some cases, separate keys or parts thereof have to be provided for macropterous, brachypterous, or apterous forms. Discussions of the aquatic species encompassed by this volume are usually limited to these three terms. However, Schuh and Slater (1995) recognized eight different degrees of wing modification, which will be discussed in more detail below.

Fig. 6.11 Habitus of an apterous male Rheumatobates klagei, a representative of the Gerromorpha, in dorsal view. Based on Hungerford (1954). In providing the keys to the higher taxa, characters of larval instars may be included with those of the adults because of their morphological similarities. The immature stages resemble the adults in general form and usually in habits, as well. However, they possess fewer body segments, are smaller, and always lack fully developed sex organs and wings. Because adequate descriptions of most larval instars are lacking, only a few keys exclusively for the larvae inhabiting individual parts of the world are available. Relatively few South American larvae have been described. However, in many cases, characteristics of adult species can also be recognized in the final larval instars, and it is possible to identify some of them with a fairly good degree of certainty. Early instars of congeneric species in South America are still indistinguishable and will remain so until considerably more is known about the developmental stages of the individual species.

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Fig. 6.12 Limnocoris espinolai, Nepomorpha: habitus of a brachypterous female (upper left) and a fifth larval instar (upper right), and (below, left to right): apex of the male abdomen in dorsal view, apex of the female abdomen in ventral view, left pseudoparamere, and the right pseudoparamere in medial (above) and dorsal view (below). Based on Nieser and López Ruf (2001).

Fig. 6.13 Habitus of a macropterous female Limnocoris aculabrum, a representative of the Nepomorpha. Based on La Rivers (1973).

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Fig. 6.14 The general morphological features of Nerthra ranina, a member of the Gelastocoridae, a family belonging to Nepomorpha. The legs are shown only in ventral view on the left side of the insect. Based on Todd (1955). Except for differences related to the various degrees of wing development, the general morphology of the water bugs usually remains relatively uniform within the individual families (Fig. 6.14). Usually, the general shape of the body and modifications facilitating the particular mode of life, such as the presence of a siphon for breathing underwater or modifications of the tarsi for life on the surface tension layer, characterize the members of the individual families. The pattern of the wing veins (Fig. 6.15), which is an important character for identifying species of many other insect orders, is seldom considered when examining water bugs belonging to the Gerromorpha and Nepomorpha, although the patterns are important taxonomic characters for some terrestrial taxa and for some members of the Leptopodomorpha. One exceptional study of gerrid wings, however, was completed by Mazzucconi and Bachmann (1993a). Instead of wing-vein morphology, specific features of the external morphology have been chosen by taxonomists in the past for distinguishing the individual species. These include the division of the fore-wings, called hemielytra, into distinctive structures. The hemielytra usually cover hind wings, although these may sometimes be greatly reduced or absent. The wings of many aquatic species are fully functional, but flights for seeking mates or moving to new habitats are typically seasonal and occur only when the weather conditions are conducive. The wings cannot be employed as the definitive characteristics for identifying most genera and species of water bug because apterous forms lack them completely, and the structure is incomplete in brachypterous forms. The morphological features most important for identifying the species include the shape, relative size, and color of a variety of body parts, including the beak, consisting of the rostrum at its base and usually

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apical segments modified for piercing and sucking out the partially digested insides of the prey; compound eyes and the ocelli, as well as the spaces separating them from one another; the antennae and the number and relative sizes of the individual segments; the pronotum, modified in many species into a broad dorsal plate covering the space between the head and anterior part of the mesothorax; legs, including their individual segments, the linear relationships between the individual segments, and special structures characteristic of individual families, including the pala of corixids and raptorial fore-legs of members of other families; pores for the release of secretions; abdominal segments; genital structures on the abdomen, and the presence and disposition of coverings of fine setae to bind and hold a “plastron” of air on the body while the insect is under water or to waterproof the bodies of species that live on the surface tension layer of the water.

Fig. 6.15 The fore (above) and hind wing (below) of Rhagovelia elegans, showing the nomenclature of the wing veins, according to Bacon (1956), who referred to the species by its synonym, Rhagovelia insularis Champion, 1898. The heads of most species are noticeably modified from the general insect plan. Some are elongate and have unusual shapes. Others are short and remarkably wide (Fig. 6.14). Compound eyes are generally well developed and functional. Some species also possess ocelli, but these may not be fully functional, and some individuals belonging to species normally possessing ocelli may even lack them. No ocelli are evident on fully aquatic members of Nepomorpha, and the dorsal surfaces of the heads of the benthic species are often covered by dense aufwuchs and debris, making it difficult to determine

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whether or not ocelli are present. In contrast, nepomorphs inhabiting the littoral zone, such as members of the Ochteridae and Gelastocoridae, typically possess ocelli. Morphometric features are frequently used for identifying species of some families. The relationships between the synthlipsis, defined as the minimum space between the compound eyes, and also called the interocular space, usually located between the inner posterior corners, and other distances on the head are frequently used by taxonomists specializing in heteropterans as diagnostic features for individual species. The synthlipsis is usually reported both in millimeters or hundredths thereof and as a decimal or percentage of the maximum width of a compound eye or, alternatively, as a percentage of the distance between the compound eyes measured across the anterior margin of the vertex (Fig. 6.16). Other commonly used features useful for species identification are the relative dimensions of the compound eyes and their relationships to the distance between their inner anterior corners measured along the vertex. In addition, the ocellocular space, that is, the distance between the inner margin of the compound eye and the nearest ocellus, is used in descriptions of those species possessing ocelli. The number of segments in the antenna is typically constant within a family; water bugs usually have four. Their relative lengths vary according to the genus or species. The antennae of Gerromorpha and Leptopodomorpha species are typically long and filiform. In contrast, those of Nepomorpha species are short and inconspicuous, and they are often clubbed at the apex. A beak, usually with three or four segments, depending on the family to which the insect belongs, is another characteristic. The base of the beak, called the rostum, is on the anterior surface of the head in all families of aquatic species, except Corixidae, which is characterized by an origin on the ventral side of the head. Using rostrum as a general term for beak may lead to confusion because rostrum is also used for the basal segment of the beak alone. The beak is formed through extreme modifications of the typical insect mouthparts to form a piercing organ with central hollow tubes through which toxins and digestive juices are injected into the prey and the partially digested emulsion on which the insects feed is sucked out. The modified labrum forms the basal part of the beak adjacent to the clypeus. The maxillae form a hollow tube surrounded by the mandible, which bears stylets for piercing the prey. The labium is also tubular and segmented. The substances injected into the prey pass through the salivary canal, while the liquified material passes into the insect through the food tube. Labial and maxillary palps are completely lacking. However, there is an epipharyngeal sense organ between the base of the labium and base of the labrum. In addition, notonectids typically have a pair of lateral processes, one on each side of the basal segment of the beak, called rostral prongs. A distinguishing characteristic of some species is the distance the beak extends posteriad relative to the coxae of the three leg pairs. Other characteristics of the head useful for identifying species include distinctive color patterns and relative dimensions of various structures. Such

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features as the distance between the compound eyes and between each compound eye and the median or lateral ocellus, if ocelli are present, are often recorded in species descriptions. The relative lengths of each segment of the beak, the width and height of the clypeus, and the width and length of the head are also morphometric features important for distinguishing species and genera.

Fig. 6.16 The head, prothorax, and scutellum of Tenagobia constricta, showing the measurements most frequently used to distinguish species. V = the interocular width across of the vertex; S = the synthlipsis; E = the width of an eye measured along its posterior margin; L = the length of the pronotal disc; W = the width of the scutellum. The length of the head and the scutellum are measured along the midline of each. According to Nieser (1975), the ocular index = 2V ÷ head width – V. Based on Hungerford (1948). The subdivisions of the thorax are usually distinct, although in some families, a streamlined body shape obliterates some of the typical lines of separation between the mesothorax and metathorax. The prothorax is usually distinctly separated from the mesothorax, and some families are characterized by strong modifications of the fore-legs, also called the prothoracic legs, for capturing prey. The dorsal plates of each section of the thorax are called the pronotum, mesonotum, and metanotum, respectively. The mesonotum and metanotum each bear one pair of wings. Those on the mesonotum are the modified hemielytra. In species with a dorsoventrally flattened body shape, the dorsal side of the prothorax is usually strongly modified to form a pronotum that protects the occipital part of the head, prothorax, anterior margins of the scutellum, and origins of the hemielytra (Fig.

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6.14). Some members of the Gerromorpha with narrowed and elongated body shapes have pronota that form tight collars merging with the posterior head margins. Like the general shape of the body, the legs of water and shore bugs are distinctively modified, allowing most families to be recognized at a glance. The fore-legs are usually modified in some way for catching and holding the prey. The species of most families have legs with unique modifications, and only a few families must be distinguished by more than a cursory examination. Because the mouthparts are not suited for grasping and, in most cases, must actually push against the surface of the prey to pierce the integument and suck material through a narrow tube, the fore-legs must firmly hold the prey while the rostrum penetrates it to inject enzymes, which first paralyze and then digest it. A similar process is often used in defense when the bug is grasped by a larger animal. The insertion of the rostrum into the skin and the sharp, burning pain caused by the substances injected are usually sufficient to convince the animal or entomologist grasping the bug to release it immediately. However, if a predator still persists in trying to consume the bug, it is usually deterred by acrid-smelling, toxic substances released through the pores of the scent glands. A few species are distinguished by the number and locations of these pore openings, but this character has been used far more frequently to distinguish terrestrial species than aquatic ones. Families of Nepomorpha have legs modified in various ways for life underwater. Some have characteristic flattened middle and hind legs with swimming hairs suitable for specific patterns of locomotion. For example, species of Notonectidae and Corixidae swim rapidly through open water, while members of Belostomatidae and Nepidae usually remain motionless or move slowly among plant masses or over detritus on the bottom and only occasionally swim rapidly for very short distances to seize prey or escape danger. The forelegs of belostomatids are modified for seizing and holding large prey, while those of notonectids are suitable for catching smaller, faster organisms. The one segment fore-tarsus of many corixids is modified into a large, flattened plate with rows of setae for filtering very small prey organisms out of the water. Such a plate-like tarsus is called a pala. The species that live underwater often have dense patches of tiny setae covering parts of the legs, which are contiguous with similar coats on other parts of the thorax and abdomen (Heckman, 1983). These coatings repel water and bind atmospheric air to the surface of the body to facilitate respiration under water. Spiracles, through which the air enters and leaves internal chambers for gas exchange, are usually covered by the thin, silvery layer of air held against the minute hydrophobic setae, which is called a plastron. The ultrastructure of the spiracles of a naucoroid species, Cryphocricos barozzii, was described and illustrated by Lopez Ruf et al. (2000). Some water bugs have the ability to make sounds underwater to communicate with each other. Chirping, clicking, or rasping noises are often heard from aquaria in which certain nepomorphs are kept. Some veliids and gerrids in the genus Metrobates also produce sound. Stridulatory structures,

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usually present on various parts of the legs or wings, produce these sounds when rubbed together. Specific names for the structures, given according to whether they are stationary or moved to produce the sound, include stridulitrum or pars stridens for the stationary structure and plectrum for the moveable structure (Schuh and Slater, 1995). More details will be provided in the discussions of the morphology of individual taxa. According to Polhemus (1994), stridulatory mechanisms have been observed in species belonging to the following genera known to inhabit South America: species of Metrobates in the family Gerridae possess fine ridges on the mesoacetabulae and a sharp ridge on the hind trochanter; all species of Stridulivelia in the family Veliidae possess ridges of denticles on the margin of the connexiva and patches of many spinules or pegs on the dorsal surfaces of the hind femora; Paravelia stenoptera in the family Veliidae has a row of pegs on the lateral parts of the third and fourth abdominal sternites and a patch of pegs on the hind trochanter; males of the family Corixidae, excluding Micronectinae, possess sharp ridges on the eighth abdominal segment and basal ridges on the right paramere; micronectine corixids in the genus Tenagobia possess ridges on the right lobe of the genital capsule; males in the genus Nerthra in the family Gelastocoridae have sclerotized ridges on a dorsally oriented part of the ninth abdominal segment and a ridge on the proctiger; males and females in the genus Limnocoris of the family Naucoridae have sclerotized ridges on the second and third abdominal sternites and a distal ridge on the hind femur; both adults and larvae in most New World species in the genus Ranatra of the nepid subfamily Ranatrinae have sclerotized ridges on the lateral edge of the anterior coxal cavity and ridges on the fore-coxa; males of the genus Buenoa in the family Notonectidae make noises using either the rostral prong and a comb on the foretibia or a file on the fore-femur and a peg on the fore-coxa; males and females of one species of Martarega in the family Notonectidae possess denticles on the costal margins of the hemielytra and a basal ridge on the hind femur. Stridulatory structures of saldids have also been described (Drake and Hottes, 1951a). Their structure is used in several keys to help distinguish species. Each fully-formed hemelytron, or “half wing,” consists of an anterior part, typically divided into three sections called the clavus, embolium, and corium, and a softer, more flexible membrane, located at the posterior end (Fig. 6.14). In some species, fractures, or indentations can be seen along the hemielytral margin at the point where two parts join. The degrees of wing modification within the Heteroptera vary from complete lack to full development, and Schuh and Slater (1995) were able to define the following eight distinct degrees of modification: 1. Aptery: no trace of wings; 2. Microptery: hind wings minute or absent and fore-wings reduced to minute pads; 3. Staphylinoidy: resembling staphylinid beetles with hind wings absent or appearing as flaps and the fore-wings covering no more than half of the

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abdomen; the clavus and corium appear fused, and the membrane forms only a thin posterior margin; 4. Brachyptery: the hind wings are short but do not resemble flaps; the hemielytra reach no farther posteriad than the fifth abdominal segment and have recognizable parts, which may be fused; 5. Coleoptery: the hemielytra resemble the shell-like wing coverings of beetles, while the hind wings are absent or reduced to small flaps; 6. Submacroptery: the hind wings are fully developed or only slightly reduced; the individual parts of the fore-wings are well differentiated, but their membranes are reduced to leave the posterior part of the abdomen uncovered; 7. Macroptery: both wing pairs are fully functional; the hind wings are long and the fore-wings fully differentiated into typical hemielytra covering the abdomen; 8. Caducous or deciduous: fully developed wings, entirely or partially shed. All individuals belonging to many terrestrial species always display one of these conditions. Among aquatic species, however, conspicific individuals with different degrees of wing development are frequently encountered (Andersen, 1993a). Among gerrids, for example, all adults of a generation may be wingless during periods of optimal weather, while macropterous individuals may appear in large numbers among generations that might be forced to migrate from habitats in which conditions are deteriorating due to desiccation. As in other insect taxa, the abdomens of heteropterans are protected by dorsal plates, called tergites, and ventral plates, known as sternites. In addition, marginal portions of the abdomen are separated from the median portions by sutures or grooves. The median plates are referred to as mediotergites and mediosternites, while the lateral plates forming the connexivum include dorsal laterotergites above and ventral laterotergites and laterosternites below. Ventral laterotergites are also called paratergites. Inner laterotergites refer to additional plates between the mediotergites and connexivum, which may also occur but are rare. The male genitalia of water bugs may include large and complex structures, often appearing to be shifted to one side of the abdomen by characteristic asymmetrical modifications of the apical segments. In a few families, they are rather simple, inconspicuous, symmetrical structures. Except in the Leptopodomorpha, they are almost always modified in ways typical for each species of a given genus and are the most definitive structures for the taxonomy of almost all groups. In the keys, the first and second genital segments are frequently considered definitive characters for the species of males. They are the last two abdominal segments, typically the ninth and tenth, which usually differ considerably from the eighth, which is the last pregenital segment of the male abdomen, recognized by its close resemblance to those anterior to it. The Heteroptera, and particularly the Nepomorpha, may show extreme modifications of the genital segments and those adjacent to them, so it is best to focus on the individual descriptions in the keys on a species by species basis. In counting the abdominal segments, it should always be kept in mind that the first and sometimes parts of the second are concealed by thoracic structures.

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Some authors have therefore numbered these segments from the “first visible abdominal segment.” Numbers determined in this way are usually one less than the actual number of the segment. Most females have internal genital chambers and therefore lack conspicuously modified external genitalia and often obvious secondary sexual modifications, as well. The last pregenital segment of the female is usually the seventh, while the eighth and ninth are modified as genital segments. In the cases of a few genera, the females of all known species cannot yet be distinguished with a reasonable degree of certainty. Unfortunately, however, there are also species that have been described only from female specimens, and descriptions of the males are lacking. For identifying species of many genera, differences in the genital structures are less distinctive than other features, some of which display conspicuous sexual dimorphism or dichromatism. Whenever possible, the individual keys include structures other than the genitalia as primary features for identifying the species, while the illustrations of most include depictions of primary or secondary sexual characters most important for distinguishing the genera and species. In this way, initial recognition of the species using general features can be followed by comparison of the male genitalia of the specimen to be identified with an illustration, providing an independent confirmation that the initial identification was correct. The individual parts of the male genitalia of a gelastocorid are illustrated in Figure 6.17. Each family shows characteristic differences in the visible genital structures, making it useful to provide illustrations with labeled parts for most of the families in addition to the general figure shown here.

Fig. 6.17 The male genitalia of Gelastocoris oculatus with the names of the individual structures, as employed by Todd (1955).

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The primary structure containing the individual parts of the male genitalia is often referred to as the male genital capsule, which is formed from the genital segments, that is, the ninth and tenth abdominal segments. Unfortunately, the nomenclature of the parts of the male genitalia is not uniform throughout the literature, making it confusing for those unfamiliar with the publications on individual families. To know which structures are being described or illustrated, knowledge of all names given to the individual body parts is often necessary. For example, the same structure may be called a paramere by one author, a clasper by another, and a harpagon by a third. For many years, entomological literature employed names other than penis for the male structure through which the sperm is transmitted. Taxonomists specializing in other insect orders now employ this term, but in literature on aquatic heteropterans, this structure in still often referred to as the phallus, which consists of a basal apparatus and the aedeagus, through which the sperm passes. Truxal (1952) defined “aedeagus” as the penis and a sheath surrounding it. In the keys, the term penis is used in a broad sense for the intromittent organ, and illustrations of the penis usually include the entire aedeagus and may also include parts of the basal apparatus of the phallus. Truxal (1952) discussed the nomenclature of the male genital apparatus of Notonectidae in great detail, and De Carlo (1963a) did the same for Lethocerus. Theoretical discussions of the correct terminology are avoided, but it is hoped that a consensus on general terminology will be reached as soon as possible. In species of some families, the penis is clearly part of a structure called the genital capsule, also called a pygophore or pygopher, which is formed mainly from the ninth abdominal sternite. The penis is accompanied by a pair of structures referred to by the names claspers or parameres in English publications, or harpagones in Spanish literature, depending upon the preference of the author. Especially among species of Nepomorpha, the right and left claspers are usually considerably different in size and shape, contributing to the asymmetry of the male abdomen. As already mentioned, some authors may call these structures dorsal and ventral parameres instead of right and left claspers, and the choice of these terms in the keys depends on what seems to be prevalent in the literature on the individual family. However, it should be remembered that these terms refer presumably to homologous morphological structures and, disregarding subtle differences envisioned by individual authors, the terms can be considered to be synonymous. The internal structure of the male genitalia has seldom been used as a diagnostic feature. The partially sclerotized phallotheca, also called the phallosoma or theca, is located in the basal portion of the penis, into which the apical structure, called the endosoma, can be withdrawn. A phallotheca may be absent in some species. The apical part of the ductus ejaculatorius in the phallotheca forms the ductus seminis, which may have an ejaculatory reservoir along its apical portion. The ductus seminis terminates through the secondary gonopore. De Carlo (1968a, b, c; 1983) provided an overview of the soft internal structures of the male genital apparatus. They are not further

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discussed here because their variability from individual to individual is too great for them to be used as reliable structures for identifying species (Viscarret and Bachmann, 1997). The structure of the female genitalia is not readily apparent during external examination, and it has only a limited value for identifying the species. The ovipositor, when well developed, is the most easily observed structure. It consists of an anterior and a posterior valvula, also known as the first and second valvulae. The valvulae are sometimes called gonopophyses or gonostyli. The gynatrial complex refers to the internal parts of the female genitalia. Many species do not have an ovipositor, so it is seldom used to distinguish species. It has not been well described in some of the South American aquatic heteropterans that possess them. The abdomen is distinctive in having plates separated by grooves or ridges, along which spiracles and sometimes pores for chemical secretions are located. The apical segments of the abdomen display various degrees of sexual dimorphism. The males of some species belonging to the Nepomorpha show extensive modifications of the abdomen involving most of the ventral surface. These modifications are asymmetrical, making it possible to recognize the males at a glance. The males and females of many species belonging to the Gerromorpha, on the other hand, appear nearly identical except for the different shapes and relative sizes of the eighth through tenth abdominal segments and sometimes also the average total length. Certain features of the color pattern are reliable characteristics for distinguishing many species. So are the locations of the coatings of fine hydrophobic setae, which form silvery coatings over various parts of the bodies and legs. 6.1.1. Morphology of the Gerromorpha Adult members of the Gerromorpha share many common features with almost all other true bugs, including their characteristic piercing and sucking mouthparts; modification of their mesothoracic wings, that is, the anterior pair, into modified wing covers called hemielytra, which are divided into distinct sections; and often scent glands for secreting noxious chemicals to discourage predators. Like species of the Leptopodomorpha, their antennae are typically longer than their heads. They differ from other groups of Heteroptera in bearing four pairs of trichobothria, which are hair-like sensory structures, arising from pits in the head. Most also have modifications of the tarsi to facilitate locomotion on the surface tension layer of the water, and many have dense coatings of setae on various parts of the body that repel water and are capable of holding a layer of air against the body for relatively short periods when the insect is forced beneath the surface of the water (Heckman, 1983). Because members of the Gerromorpha typically maintain contact with the atmosphere continuously, most cannot spend very long periods submerged.

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The spiracles, which are openings into the internal respiratory system, are typically large and easily observed. Their location relative to other structures is occasionally employed for distinguishing species or genera.

Fig. 6.18 Mesovelia mulsanti: habitus of a macropterous specimen with fully intact membranes on the hemielytra in dorsal view (left), lateral view of head and thorax (upper right), a hemelytron in dorsal view (lower right). Based on Alvarez and Roldán (1983). Few aquatic heteroptera are known to shed their wings, but caducous specimens are known. For example, macropterous individuals of Mesovelia mulsanti can lose the membranes of their hemielytra (Fig. 6.19) for reasons that are still not clear. Species of Gerromorpha typically have morphologically distinct apterous, brachypterous, and macropterous forms, although only one form has been described for many species. The different degrees of wing modification occurring among heteropteran species were already defined in Section 6.1, but all of them have not yet been encountered among South American gerromorphs. In most cases, macropterous forms appear to be fully capable of flight (Fig. 6.18). The significance the seasonal appearance of the distinct wing forms in the life history of almost all South American species remains to be determined. The wide-spread species, Mesovelia mulsanti, however, was found to have three developmental patterns leading to different degrees of wing development. Galbreath (1975) defined winged individuals capable of flying as those that had developed wing buds by the fourth instar and showed continued development of the buds in the fifth instar. Wingless adults never developed wing buds at all, while a form called “intermediate” developed wing buds by the fourth instar, but the development did not continue, and the adults were flightless. Which of the

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developmental patterns was followed was found to depend on the temperature. However, studies of the seasonal cycle of North American species suggest that the photoperiod seems to trigger the development of wings (Zera and Tiebel, 1991). Obviously, macropterous forms would be much better suited for colonizing new habitats than those with reduced wings, while flights and the wings necessary to undertake them would be superfluous during seasons when a complete life cycle could be completed in one stable aquatic habitat. This will be discussed more thoroughly in Section 6.2. Some families and genera of Gerromorpha have legs with characteristic modifications that facilitate certain kinds of movement across the surface tension of the water (Andersen, 1976). Others have legs that do not appear markedly different from those of terrestrial bugs but which nevertheless easily support the insect on the surface tension layer. Members of the Gerridae typically have very long and thin middle and hind legs, which hold the body well above the surface of the water and allow the insects to move rapidly with a jerky gait. Members of the Veliidae, on the other hand, usually have shorter legs held closer to the body, although there are a few exceptions. Many species appear less graceful on the water surface than gerrids, although species of several veliid genera are capable of rapid movement, and members of the genus Rhagovelia have greatly modified tarsi permitting the insects to race with remarkable speed across the surface of the water while frequently changing direction. For veliids, extreme modifications of the tarsi make it relatively easy to recognize some of the genera.

Fig. 6.19 Mesovelia mulsanti: dorsal view of the habitus of a caducous specimen, which has lost the membranes of the hemielytra. Based on Cobben (1960a).

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Size is another feature that can greatly simplify the identification of certain families. Several gerromorph families encompass species shorter than 5 mm and include many between 0.5 and 3 mm. The families Hebridae and Mesoveliidae are represented in South America mostly or entirely by such small species. These are among the insects that creep easily on the water surface without showing any elaborate leg modifications other than tarsi somewhat flattened on the ventral surface. Taxonomists have most frequently used the apical segments of the male abdomen, referred to as the genital segments, as definitive characters for distinguishing the species of most gerromorphs. The apices of female abdomens also display species-specific features useful in the identification of the species, but detailed descriptions of the females of all South American species have yet to be provided, limiting the usefulness of these features at the present time. In addition, there are other morphological characteristics that have been used to quickly recognize species in most families, such as the shape of the body and color of its markings; the extent and locations of silvery areas formed by dense coats of the fine setae that repel water from parts of the body; the relative sizes of the segments of the antennae and legs; certain extreme modifications of one or more of the leg pairs; shape and size of the claws and other structures at the apices of the tarsi; and the relative sizes and shapes of the individual parts of the mouth. Because gerromorphs do not live beneath the surface of the water, their hydrophobic setae are arranged primarily to keep the body surfaces near the spiracles dry. For example, the dense coats of setae covering much of the bodies of gerrids consist of both short, fine setae and longer, coarser setae (Heckman, 1983). While the coarse setae are typically curved to assist in supporting the insect on the surface tension layer, the fine setae hold a coat of air against the body surface for short periods during which the insect is forced beneath the surface because of turbulence or the actions of predators. These coats of setae, however, are not as effective as those on some of the fast-moving nepomorphs, and when forced to stay beneath the surface for extended periods of time, some gerromorphs become wet beneath the setose layers and drown. Pollution of water bodies with soaps, organic alcohols, or other substances that weaken the surface tension layer can largely eliminate gerromorphs from the habitat. With few exceptions, the family of a gerromorph can be determined at a glance from the general shape of the body and the size. The gerromorph family with the most extreme morphological modifications from the general heteropteran body plan is Hydrometridae. These small insects are extremely narrow, making them appear stick-like. Their heads are greatly elongated and account for at least ¼ of the body length. Mesoveliidae and Hebridae are distinguished easily from Gerridae and Veliidae by their small size, and they are readily distinguished from each other in the field with the assistance of a magnifying glass by the shapes of their bodies. From the relative length of the legs, almost all members of the families Gerridae and Veliidae can be

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distinguished, although exceptional veliids have legs long enough to confuse with those of gerrids. Among the Gerromorpha are groups of species, which are among the few insects adapted to life in marine habitats. Although some live near the shore, and others are common in mangrove forests, several gerrids have become fully adapted to life on the surface of the open ocean. Some marine species have bizarre modifications of the legs, and most of those encountered on the open ocean have a typically gray or silvery coloration (Polhemus and Manzano, 1992). Although the research on the genetics and molecular biology of South American heteropterans has been very limited, recent studies have revealed that the male diploid chromosome complement of Limnogonus aduncus, a gerrid, is 22 autosomes and 1 X chromosome, indicating that males have an XO set of sex chromosomes. Females would presumably have an XX set and a diploid number of 24. It was also found to have a nucleolar organizing region, which is rare among species of Heteroptera (Castanhole et al., 2008). Male Brachymetra albinerva and Halobatopsis platensis, both in the family Gerridae, have 24 autosomes + XO, while male Cylindrostethus palmaris has 28 autosomes + XO. The XO formula for males has also been found in the Veliidae. Rhagovelia whitei was found to have 38 autosomes + XO, while an unidentified South American species of Rhagovelia has 24 autosomes + XO (Castanhole, 2009). Some recent studies have focused on mitochondrial DNA to determine the extent of interbreeding between geographically distinct, conspecific populations. The results indicated that there are distinct populations of Halobates species in pelagic regions of the oceans, which apparently have very limited interaction (Andersen et al., 2000), and that individual populations of Potamobates williamsi and P. sumaco are somewhat isolated due to their distances from neighboring populations along river systems in Ecuador (Galactos et al., 2002). More studies are needed to determine the accuracy of the results obtained by this relatively new method. 6.1.2. Morphology of the Nepomorpha For meeting the requirements of the many ways of life adopted by members of the Nepomorpha, the morphology of the species in the different families displays a great variety of modifications. The representatives of most families live underwater, but a few, including the gelastocorids or toad bugs, leave the water and display habits at the water’s edge reminiscent of many amphibian species, which accounts for the common name of the group. Those that remain submerged include fast swimmers inhabiting the open water, including the corixids and notonectids, and slow-moving benthic species that breathe through long respiratory siphons, such as many of the nepids and belostomatids. Morphologically, the group is distinguished by the shortness of the antennae, which are typically concealed beneath the head (Fig. 6.20). The

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families of most species can be determined immediately by the size and general body shape, although close examination is necessary to distinguish a few species of Belostomatidae, Nepidae, and Naucoridae from one another. Generally, the habits can be deduced immediately from the body form. Those with streamlined bodies and at least one pair of legs somewhat resembling oars and lined with rows of setae modified for swimming are typically observed swimming rapidly in the water column, resting at the surface to refresh the air supply that they carry as a “plastron” on the body surface, or partially concealed among submerged plants or detritus. The general morphology is usually similar throughout the larval stages, and the family of first instar larvae can immediately be recognized after only a cursory examination in almost all cases (Fig. 6.20). The general morphology of corixids differs in several ways from that of all other groups of true bugs. The tarsi of the fore-legs each consist of only one segment lacking a claw and are enlarged, usually greatly, and flattened to form the pala. As a general rule, the pala displays strong sexual dimorphism, and its shape and armament in the male often provide reliable means for identifying the species. The palae of females are less distinctive, and some species cannot yet be distinguished when only female specimens are available for examination. In addition, the beak shows no distinct segmentation and is located on the ventral side of the head. Members of all other nepomorph families have forelegs with tarsi modified either for grasping or for locomotion, and their beaks are distinctly segmented and inserted on the anterior surface of the head.

Fig. 6.20 Belostoma bifoveolatum larval instars I through V with the antenna shown left of the habitus (above, left to right): instars I, II, III, and (below, left and right, respectively): instars IV and V. Based on Schnack (1971).

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Corixids also have distinctive color patterns on the disc of the pronotum, corium, and other parts of the hemielytra, making examination of these structures an additional useful means of species identification. It has also been noted that the shape of the metaxyphus, a structure located on the posterior margin of the sternum at the ventral midline, is distinctive for many species. Detailed information about the structure of the mesothorax and metathorax of notonectids and corixids was provided by Matsuda (1962), but taxonomists have usually confined their descriptions to the dorsal surfaces of the thorax, metaxyphus, and locations of the coxae. The male genital structures of most species are strongly modified, making the abdomens of males asymmetrical and usually strongly so. Females can be distinguished at a glance because of the bilateral symmetry of the abdominal segments. Respiratory structures include dense coats of fine, uniform setae, which bind the air supply to the surface of the abdomen while the insect is under water. Part of the air supply is also carried under the hemielytra. Individuals replenish their air supplies at the surface of the water. This kind of respiration from an air supply carried with the insect underwater is referred to as plastron breathing. The arrangement of spines on the legs and pegs on the palae have been used to distinguish species, but the swim hairs on the legs have seldom been used by taxonomists as distinguishing features.

Fig. 6.21 Buenoa cucunubensis: habitus in dorsal (left) and ventral view (right) and the beak in lateral view, showing how the length of a rostral prong (L) is measured (lower center). Based on Padilla-Gil (2002).

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The other nepomorph family noted for swimming strongly in the water column is Notonectidae. The morphology of the members of this family is distinctive in showing considerable streamlining (Fig. 6.21). The beak is generally similar to those of most other heteropteran species, although there are some distinctive structures used by taxonomists to characterize individual species, including the rostral prongs, which are inserted laterally in the rostrum. The length of these prongs relative to the length of the beak or its individual segments, the length of the segments relative to each other, and the total length of the beak are all features useful for species identification. In addition, the relative lengths of the synthlipsis, compound eye widths, and space between the eyes at the anterior margins of the vertex are useful for identifying the species. The name backswimmer used for members of the Notonectidae comes from the habits of the insects to swim with the dorsal surface down and replenish their air supply at the surface with the body oriented somewhat obliquely inverted. The other nepomorph species are modified either for benthic life and life concealed in plant masses or for life in the littoral zone. A detailed study of the male genitalia of a South American notonectid was provided by Angrisano (1982a).

Fig. 6.22 Ranatra unidentata female: habitus in dorsal view (middle), head in ventral view showing the location of the antennae (lower left), antenna (upper left), and fore-tibia and tarsus (lower left). Based on De Carlo (1946). Two families are noted for living close to or in contact with the surface tension layer of the water. However, unlike members of the Gerromorpha, species of the families Pleidae and Helotrephidae live on the underside of this interface. These insects are typically very small, so it is convenient for them to remain where they can easily replenish their supply of atmospheric air. The body shape of pleids and helotrephids facilitates their swimming in a typical

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position with their flattened ventral surfaces facing upward and in contact with the surface tension layer. This inverted position also allows the insects to scan the surface of the water for prey, especially terrestrial insects that have fallen on the surface of the water. Although the dome-like shape of the dorsal body surface and flat venter are common to species in both of these families, they are clearly distinct in the morphology of the head and thorax, which are clearly separated in pleids but fused internally in helotrephids. The cephalonotal suture, which externally marks the border between the head and pronotum, is straight and articulates in the Pleidae but is curved or sinuous and fused in the Helotrephidae. The general body shape of most species in the families noted for remaining submerged among the benthos or plant masses is usually widened and flattened, but a few, notably members of the genus Ranatra, are long and slender. Like members of the Corixidae and Notonectidae, the air supplies of these insects must be renewed periodically, but this is accomplished by members of the Nepidae and Belostomatidae without the need to swim to the surface. Members of the family Nepidae obtain atmospheric air with the aid of a nonretractible respiratory siphon arising at or near the posterior end of their abdomens (Fig. 6.22). Species of the family Belostomatidae also possess a respiratory siphon to refresh their air supplies, but it can be retracted. Respiration by members of the family Naucoridae, which also have widened and flattened bodies, requires the insect to make occasional trips to the surface to renew the air supply, which is carried with it underwater. Naucorids generally have a broad and flattened body shape similar to belostomatids and nepids. Few belostomatids are small enough to be mistaken for naucorids, but some nepids superficially resemble naucorids enough to be mistaken for them after a quick glance. On closer examination, however, several morphological differences even easier to detect than the presence of a breathing tube distinguish them immediately. The absence of raised veins on the hemielytra is one of the easiest characteristics of macropterous naucorids to see. Such veins are obvious on both belostomatids and nepids. Information about the chromosome complement of nepomorphs was summarized by Bressa and Papeschi (2007). Their data on South American species were augmented by information about corixids from other continents. Of the 28 species listed in their table on Corixoidea, only 2 had a 2n chromosome complement other than 24. One of these is Tenagobia fuscata, which was found to have 30: 14 pairs of autosomes, and the two sex chromosomes. All males listed have an XY complement of sex chromosomes, while the female complement is XX. The male of an unidentified South American species of Martarega was also found to have an X and a Y chromosome, in addition to 2 m chromosomes. Its formula was 2n=22A+2m+XY (Castanhole, 2009). Males of most of the other superfamilies which have been tested show that an XY complement is typical, although a few species in each higher taxon have been found to have an XO complement (Bressa and Papeschi, 2007). Special

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studies on the genus Belostoma, however, show that species of this genus seem to have developed considerable differences in the numbers of chromosomes through fusion and fragmentation (Papeschi, 1994). This hypothesis is supported by the presence of natural chromosome mutants within populations of Belostoma plebejum, a species which usually has a male diploid (2n) complement of 14 + XY. Two mutants were identified, one with 2n=13+XY, and the other with 2n=14+X1X2Y. Congeneric species have male karyotypes varying from 2n=6+XY in B. oxyurum to 2n=26+X1X2Y in B. bifoveolatum, B. dentatum, B. elegans, and B. martini. The male B. micantulum has a karyotype of 2n=14+XY (Papeschi, 1988, 1991). Each family of Nepomorpha has its own morphological peculiarities, which will be discussed in more detail in the individual keys. 6.1.3. Morphology of the Leptopodomorpha Most shore bugs are tiny heteropterans morphologically resembling many of the terrestrial species, but on closer examination, they can be seen to have parts of the body covered by the fine setae that repel water and bind layers of air to the body during periods of submersion. The South American leptopodomorph fauna encompasses members of only two families. The known fauna includes only one species in the family Leptopodidae, which has only been found once in South America and a second time in a seaport among vegetable material imported from Colombia. All other known South American species in this group belong to the family Saldidae. One additional saldid is known only from the Galapagos Islands.

Fig. 6.23 Saldula pallipes: head in dorsal (upper left) and anterolateral view (upper left center), the visible abdominal segments of a male in ventral view (upper right), habitus in dorsal view (lower left), and ventral view of a female (lower right). Based on Polhemus (1985).

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Apterous, brachypterous, and macropterous shore bugs are known, as are species with coleopteroid hemielytra. They differ from other aquatic heteropterans in having antennae longer than the head but lacking pairs of pits in which the trichobothria are set. The species are typically small, and some are among the tiniest heteropterans. The fauna is probably richer than it is now known to be because leptopodomorph specimens are easily overlooked. The morphology of the known shore bug species deviates less from the basic heteropteran body plan than that of members of the other water bug taxa, and on cursory examination, leptopodomorphs could easily be mistaken for species belonging to terrestrial groups (Fig. 6.23). The morphology and physiology of shore bugs require further study to determine the form and purposes of their minute sensory and glandular structures. 6.1.4. Changes during larval development Because the larvae so closely resemble the adults, separate keys to the larvae are seldom prepared. At times, it is not possible to distinguish between apterous or brachypterous adults and the late instar larvae at a glance. Closer examination is necessary to determine whether or not the full complement of abdominal segments is present and whether there is full development of the genitalia, indicating that the specimen is an adult.

Fig. 6.24 Morphological structures important for distinguishing the fifth larval instar of Heterocorixa braziliensis: habitus (upper left center), antenna (left), head in anterior view (lower left center), fore (lower right center) and hind leg (right). Based on Bachmann (1979a).

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The basic morphology of final instar larvae is almost identical with that of adults, except for the absence of wings in all specimens, fewer segments of the abdomen and sometimes of each antenna, and the absence of functional genitalia (Fig. 6.24). Earlier instars are recognizable by their small body size and often disproportionately long legs, lack of a complete complement of segments in most appendages, and undeveloped differentiation of the main body parts (Fig. 6.25). The larvae of some species can easily be confused because no reliable method has been developed to distinguish them. Some larvae, however, can be identified to species from the characteristics of the adults, at least during the final larval instars.

Fig. 6.25 The five larval stages of a corixid, Tenagobia fuscata, shown in dorsal view. The upper row (left to right) shows Stages I, II, and III, and below are Stages IV (left) and V (right). Based on Galliussi (1989). Based on sketchy information presently available about their larval development, most aquatic heteropterans seem to have five larval instars. The adult is considered the sixth instar. In a few cases, the adult seems to be the fifth instar. The final larval instars of many well-studied European species are similar enough to the adult to be identified from the description of the adult, often by the developing color pattern. However, relatively few South American water bugs have been studied well enough to permit at least a tentative identification from a larva. One of these is Belostoma oxyurum, for which Schnack (1971) provided descriptions and illustrations of each larval stage (Fig. 6.26).

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Because the larvae resemble adults in general shape and often in color, as well, treating them together in identification keys seems appropriate. In most cases, their preferred habitat and feeding habits are also similar to those of adults, with the prey being proportionately smaller, however. Future research will be required to eludidate the morphology, ecology, and behavior of the larvae of aquatic South American heteropterans, and until this is completed, knowledge of these early life stages will remain sketchy.

Fig. 6.26 Belostoma oxyurum larval instars I through V with the antenna shown to the left of the head, the claw on the middle tarsus below or to the lower right of the hind leg, and the apical segments of the hind leg above and right of the abdomen (above, left to right): instars I, II, III, and (below, left and right, respectively): instars IV and V. Based on Schnack (1971).

6.2. Ecology The piercing and sucking mouthparts of Heteroptera species may be used for sucking plant juices, or they may be employed by predatory species for piercing their prey and injecting toxic substances to paralyze or kill them and enzymes for digesting their inner organs and turning them to fluids that can be sucked out. A few terrestrial species are even adapted to sucking blood of vertebrates. The aquatic species considered in this volume are all carnivorous or scavengers on dead or dying animals, except for members of the Corixidae, some of which seem to be omnivores, consuming both algae and small aquatic animals, including protozoans, rotifers, and other organisms in the same general size range. In older literature, corixids are sometimes described as herbivorous because of unsubstantiated reports that they suck the juices of aquatic plants. It

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is certainly likely that they suck cellular material from filamentous algae that are too large to be consumed whole, but whether any South American corixids feed on vascular plants in this way is still unknown. Most other species in the three taxonomic groups now generally accepted as encompassing all of the water bugs, Nepomorpha, Gerromorpha, and Leptopodomorpha, have similar feeding habits, but each group is adapted for life in a different part of an aquatic or littoral habitat. All known South American larvae share the general habitat preferences with adults, although the smaller larvae must be somewhat more secretive to avoid predation. As far as known, larvae and adults show similar food preferences, with selectivity being based primarily on the size of the prey they can overpower. Hence, the final larval instars are frequently observed together with adults, sharing a common habitat and hunting similar food species in different size ranges. Several studies of the feeding habits of belostomatid species have been undertaken (Consoli et al., 1999; Pereira et al., 1993; Pereira and Melo, 1998; Pérez Goodwyn, 2001a), prompted by the potential of these species to control snail and mosquito vectors of serious tropical diseases (Ávila et al., 1996). In most cases, the species studied have been shown to have an impact on populations of mosquitoes, at least during the predators’ larval development. Looking at the other end of the size range of the prey, belostomatid adults, which are among the largest insects, have also been shown to capture and feed on vertebrates, especially fishes and amphibians (Haddad and Bastos, 1997; Mijares-Urrutia et al., 1997). Many general predators, which feed on almost any insect available, will avoid water bugs because of the chemical repellents they secrete. Some water bugs are also capable of inflicting a painful stab wound if grabbed by a potential predator or insect collector. The chemicals injected cause the wound to sting. While only large belostomatids can inject enough venom to produce a wound that might require the attention of a physician, even tiny gerrids are capable of inflicting a painful stab, which may cause pain for almost an hour. An early study of the venom was conducted by Pidado (1937), and an anatomical study of the salivary ducts was completed by De Carlo (1969a). Observations of heteropterans moving in plain sight of ravenous fishes without any fear of attack attest to the effectiveness of the chemical defenses of these insects. I have observed fishes steal prey from the fore-legs of gerrids and veliids without making any attempt to attack the water bugs themselves, which the fishes have apparently learned are highly unpalatable. Even corixids too small to break the skin on a human hand can leave bright yellow stains on the hand, produced by the acrid-smelling chemicals they secrete to make themselves unpalatable. Although the chemical repellants are very effective for keeping many predators away, in some localities, heteropterans are eaten by people because of the spicy taste these chemicals give the insects. In addition to some terrestrial species popular in Mexico, belostomatids are a favorite food in Southeast Asia. There are also a variety of arthropods that will prey upon heteropterans of suitable size, and studies have also shown that some water bugs are cannibalistic (Cárcamo and Spence, 1994).

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Nearly nothing is known about the micro-organisms that parasitize or live in commensal relationships with South American water bugs. Benthic nepomorph species or those inhabiting plant masses provide ideal substrates for sessile protozoans and rotifers. Although such commensal relationships are well known on other continents, epizoic species in South America have seldom been identified. In a pioneering study, Vianna and de Melo (2002) recorded the occurrence of flatworms in the family Temnocephalidae attached to naucorid, nepid, and belostomatid species in the Brazilian state of Minas Gerais. It was not determined with certainty whether the flatworms were living as parasites or commensals, but the observations suggested that the relationship was commensal, and the attachment to water bugs was only transitory. They found Temnocephala decarloi Moretto, 1978, attached to Belostoma testaceopallidum, and Moretto (1978) discovered it on Belostoma cummingsi. Temnocephala lanei Pereira and Cuocolo, 1941, was found on Limnocoris maculiceps, L. porphyros, L. saphis, L. submontandoni, and Ranatra montei (Vianna and de Melo, 2002). Macropterous water bugs are sometimes observed flying around street lamps, especially during the driest periods of the year, just before the freshet. Whether any fly in search of mates is uncertain since mating pairs are frequently observed on the surface of the water during seasons when flights do not seem to occur. Some species apparently display distinct seasonal behavior. The appearance of macropterous individuals of some species seems to be related to seasonal patterns. In the Pantanal, the collection of aquatic heteropterans under street lights at the end of the rainless season often yielded specimens of many species not otherwise observed in the region, indicating they prefer aquatic habitats that are not easily found or sampled (Heckman, 1998). Occasionally, massive numbers of one species may be encountered in flight, as reported for Lethocerus annulipes by Orfila (1932). Although Polhemus (1993a) noted no immediate danger to South American water bug species, uninhibited habitat destruction is certain to endanger those with highly specialized habitat requirements. Whether mating takes place before, during, or after the flight period, the seasonal flights certainly serve to distribute the species and facilitate the colonization of new habitats. Traces of dried sediment observed on specimens personally found around streetlights near the Pantanal in Mato Grosso, Brazil, demonstrate that the insects were aestivating in desiccated sediments shortly before the weather conditions immediately preceding the freshet induced them to take to the air after sundown. The flying insects usually anticipate the first heavy rains by a few hours, giving some of them habitats to settle in immediately after the flights. The tropical thunderstorms are typically limited to relatively small areas, into which large amounts of rain fall. When a particular area is missed by the storms on the first night, the mass flights of insects repeat themselves night for night until the area is finally inundated. Aestivation of adult heteropterans during dry summer periods is also known on

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other continents, but most attention has been given to hibernation because most studies have been conducted in temperate zones (Spence and Andersen, 1994). In temperate regions, the flights of aquatic heteropterans seem to be most frequent during the spring, although the flight habits of most South American species remain to be learned. Studies that have been completed in the temperate zone indicate that more than one generation can be completed during the course of a single year, but a review of those studies already completed indicate that reproductive habits and life history are species specific but can be modified according to habitat and climatic differences. Cases of hibernation as adults, larvae, and eggs have been reported (Spence and Andersen, 1994). As the distribution of the South American climatic zones suggests, diapause of most species is induced by dryness rather then cold, but the species inhabiting temperate South America must adapt themselves to summer and winter conditions similar to those known from the North Temperate Zone. In some locations, it is likely that no diapause occurs. However, year round activity is known with certainty for few South American species, among which those of Halobates inhabiting the open ocean are included. As already mentioned, each of the three taxonomic groups, Nepomorpha, Gerromorpha, and Leptopodomorpha, has a particular niche in the aquatic habitat, for which it is ideally specialized. It is therefore necessary to discuss the ecology of each group separately. It must be emphasized, however, that each species displays ecological specializations for its own individual niche, and generalizations made about genera, families, and higher taxa do not adequately represent the many adaptations displayed by the individual species. Studies of the relationships between the aquatic Heteroptera species and their physical, chemical, and biological environments are still in their early stages, and almost nothing is known about the ecology of the majority of South American species beyond the generalizations reported below. Water bug ecology will provide an extremely rich field of study and almost certainly yield a considerable number of surprises. 6.2.1. Ecology of the Gerromorpha Typical habitats for species belonging to the Gerromorpha are the surfaces of all kinds of water bodies. Individual species show specializations for lentic or lotic freshwater bodies or even for life at sea. The legs are typically constructed to support the body on the surface tension layer of water; the tarsi must be modified to spread the weight on the water surface and not penetrate the surface tension layer that must support them. Therefore, many of these insects die when certain pollutants that weaken the tension, such as soaps and detergents, are discharged into the water. In the United States, heavy alcohols have been poured on the surfaces of reservoirs in arid regions to reduce evaporation, and these resulted in decimation of the gerrid populations by

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weakening the surface tension enough to cause these insects to sink into the water and drown. A few gerromorphs are adapted to less typical habitats, such as the water’s edge, waterlogged soils, or on the water trapped in bromeliads. Little has been reported on the habits of species that inhabit moist soils at the water’s edge or in rainforests, and mention of this will be made as remarks about the individual species in the keys. Polhemus and Polhemus (1991) reviewed the information on veliids that live in bromeliads. Unfortunately, in most cases, the bromeliad hosting the insect has not been identified, but one found to host Paravelia recens is Guzmania brasiliensis Ule (Pereira, Neiss, and Ferreira, 2007). Veliids have developed a considerable number of morphological modifications for survival in lotic habitats flowing at different velocities. Highly modified structures on the tarsi permit members of the genus Rhagovelia to move rapidly enough to maintain their positions along relatively fast-flowing streams, while other veliids can move rapidly enough only to inhabit waters with only moderate flow (Andersen, 1976). One South American gerrid, Brachymetra furva, and two veliids, Rhagovelia sinuata and Microvelia cavernula, have been found to inhabit caves (Galan and Herrera, 2006). The bodies of gerromorphs are typically protected by dense layers of fine setae, which repel water from the body surfaces bordering the spiracles. When the insects are forced below the surface of the water or deliberately conceal themselves below the surface to escape from predators, the fine, silvery coats of setae can bind a layer of air to the surfaces for a limited period of time. Gerrids were found to have setae of two distinct lengths, the longer of the two assisting the insects to resist being forced beneath the surface of the water, and the shorter and denser coat binding the layer of air against the body whenever the insects are forced beneath the water (Heckman, 1983). The legs of the individual species show various modifications for supporting the insects on the surface tension layer. The apices of the tarsi often appear as narrow pads with simple preapical claws. Some vellids, however, have elaborate modifications to permit movement rapid enough for the insects to colonize the surface of fast-flowing streams. Species of Rhagovelia are capable of such rapid movement thanks to a fan-like arrangement of long setae around the claws. They show little difficulty in maintaining position on the surface of rapidly flowing streams, and they are skilled at avoiding capture by natural enemies and entomologists by suddenly allowing themselves to be carried away from the danger by the current and regrouping downstream. Naturally, standing water allows the insects resting on the surface tension layer to conserve their energy by remaining motionless unless attracted to prey or disturbed by a large predator. Those species that prefer lentic habitats often congregate in relatively sheltered parts of lakes when winds become strong. As already mentioned, however, a few highly specialized marine species are capable of living among waves on the open ocean. The life cycles of these species deserve considerably more attention than they have been given. They are most frequently collected in sheltered areas along the coasts, but it should be

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determined whether the individuals captured from ships far from any land were simply carried out to sea from their preferred neritic habitats or whether they are primarily pelagic in their habits and congregate near the shore only as part of a complex migration pattern. It seems that at least the species of the gerrid genus Halobates are primarily adapted to life on the open ocean. Analyses of their mitochondrial DNA suggest that separate populations of Halobates micans have inhabited the Atlantic and Pacific Oceans for at least a million years with little contact or exchange of genetic material. Similarly, the separate populations of H. sericeus inhabiting the northern and southern parts of the Pacific Ocean seem to have had little contact with one another (Andersen et al., 2000). Almost all species in this group utilize terrestrial insects that fall into the water as a convenient source of food, and some are also skilled at capturing small flies and other insects that mate just above the surface of the water and deposit their eggs on the surface. Many species are known to be very adept at detecting slight vibrations that pass across the surface of the water, and some gerrids reportedly communicate with each other by causing and detecting such vibrations. In addition, most members of the gerromorpha seem to have excellent vision, by which they can quickly locate their prey and also observe potential predators in time to flee. Some gerromorph species reportedly scavenge dead insects that they find on the surface of the water, but it is not certain how widespread this habit is among the individual species. Like most other predatory bugs, their food is primarily ingested in liquid form. Upon capture, animals seized as prey are injected with toxins and enzymes that not only immobilize them but also begin their digestion into a fluid mass that can be sucked through the rostrum. This external digestion process may take some time to complete, so the predatory bug may either carry its food or leave it on the surface and return to it to complete feeding as digestion continues. More extensive observations are needed to reveal how much of the diet of each species consists of carrion encountered randomly and how much consists of the remains of insects killed earlier by the predators and left to digest to a form that can be ingested through the rostrum. An interesting aspect of water strider ecology is the appearance of macropteous, brachypterous, or apterous adults during different seasons and perhaps under certain meteorological conditions. Unfortunately, most studies of this phenomenon have been undertaken on other continents (Järvinen and Vepsäläinen, 1976; Selvanayagam and Rao, 1986, Zera and Tiebel, 1991), where environmental conditions, including season, photoperiod, and rainfall, have sometimes been found to influence the frequency of wing development within populations. One of the only investigations in the Neotropical Region focused on populations of Limnogonus franciscanus on the island of Barbados. Although a peak in reproduction was reached during rainy periods, no correlation could be found between the development of wings and season, weather, or photoperiod. Throughout the year, about 3.5% of the females had wings and were capable of flight (Nummelin, 1997). More information on the

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conditions that induce the occurrence of the individual wing forms among South American populations would be greatly appreciated. 6.2.2. Ecology of the Nepomorpha Most species of Nepomorpha typically live below the surface of the water, employ special morphological structures to obtain and store an atmospheric air supply, and feed on organisms they are able to overpower. Exceptions to this are members of the Gelastocoridae and Ochteridae. Gelastocorids are most commonly observed hopping over sandy or muddy areas at the edge of water bodies, while ochterids are usually found near the water’s edge, but some species are encountered far from water. The members of four superfamilies grasp relatively large prey with powerful fore-legs and stab them with their mouthparts to inject toxic substances containing digestive enzymes to paralyze, kill, and externally digest the prey. Members of one of the superfamilies, Corixoidea, typically feed on minute organisms, which they gather using enlarged fore-tarsi and consume through the relatively short and broad mouthparts. The species belonging to two superfamilies of the Nepomorpha, Corixoidea and Notonectoidea, swim rapidly in the water column, actively gathering or hunting their food. Their air supply is carried with them as a thin coating of air, referred to as a plastron, which is renewed periodically, as necessary, at the surface of the water. The plastron itself can also participate in gas exchange with the surrounding water, which can remove carbon dioxide from the air layer, allowing the insect to remain under water longer before having to renew the air at the surface.

Fig. 6.27 Eggs placed on a male Belostoma micantulum by the female. Based on Nieser (1975).

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Many members of the superfamily Nepoidea typically remain motionless in masses of plants, accumulations of detritus, or beneath the sediment, waiting for suitable prey to approach. Most match the color of their background and are additionally camouflaged by dense growths of aufwuchs. Some species avoid having to renew their air supply at the surface by extending a long breathing tube called a siphon and refreshing the air supply through it. When an organism that can be overpowered approaches within grasping range, the nepoid water bug seizes it with a rapid movement of the fore-legs. It is immediately injected with the toxic secretions of the salivary glands and held firmly until paralyzed. Nepoidea encompasses two families which are not always immediately distinguishable: Nepidae and Belostomatidae. These families include some of the largest species of Hemiptera, and several belostomatids are among the largest of insects and protein-rich components of the human diet in some parts of the world. Most belostomatids are also noted for protecting their eggs in an unusual way. The female pastes them on the dorsal surface of the thorax and hemielytra of the male while she lays them (Fig. 6.27). Detailed descriptions and illustrations of the mating and egg deposition of Belostoma oxyurum were provided by Schnack et al. (1990). Apparently, one male may carry the eggs of several different females at the same time. Females may also deposit eggs on several different males. They are carried on the dorsum of the adult until they hatch. In a few cases, females have also been found with eggs pasted to their body surfaces, where they were able to develop and hatch successfully (Estévez et al., 2006). Evidently, the embryos in eggs not attached to an adult die prior to hatching. Even belostomatids that do not carry eggs attached to their bodies have been found to care for them (De Carlo, 1962a). Members of the Lethocerinae do not carry eggs attached to either parent. However, males shade and defend the eggs deposited above the water’s surface, as observed during a detailed study of the parental behavior of Lethocerus medius (Smith and Larsen, 1993). Schnack et al. (1982) found that the mortality among first instar larvae of several Belostoma species is usually far higher than that of any other developmental stage. Survival of the second instar in the laboratory, however, may be little better than that of the first. Species of the superfamily Naucoroidea are small to medium-sized insects, which hunt actively while creeping through plant masses or on other objects in the water. They are more active than members of the Nepoidea but move much slower than members of the Corixoidea and Notonectoidea. The family Naucoridae includes some species with breathing tubes and others that renew the air supply in a plastron at the surface. Seasonal periods of flight seem to be characteristic of the behavior of most species of Nepomorpha. Although a casual observer might be skeptical about the flying abilities of a large belostomatid heavily coated with aufwuchs or a tiny, fast-swimming corixid, both are abundant around street lights at night in

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the tropics during certain times of year, most typically late in the dry season, shortly before the first heavy rainstorms of the year. In addition to the Belostomatidae, several other groups belonging to Nepomorpha are noted for various kinds of parental care of the eggs. After hatching, many nepid and belostomatid larvae may swim freely in the water before settling to the bottom or on substrates to take up the way of life of the adults. The habits of the early larvae of few of the many species in these families have been adequately studied, and this would be a fertile field for ecological research. Eggs of some nepomorph species are deposited in groups on objects in the water. Some appear to be capable of passing the winter in a dormant state and then resuming development and hatching after the water warms in the spring. Much has to be learned about the eggs of species inhabiting tropical and subtropical climates. The production of drought-resistant eggs would be an obvious strategy for surviving dry seasons, but the appearance of adults of many species around street lights just before the freshet indicates that at least some water bugs survive the dry season as adults, which mate and spawn at the time of the first rains. Obviously, studies of the life cycles of individual species will be required to elucidate the species-specific differences that permit each to survive in its own special habitat. Few such studies have been completed to date, although several publications have provided valuable information on the activities of Belostoma species reared in the laboratory (Pereira et al., 1991; Pereira and Melo, 1998). Because their predatory feeding habits have raised hopes that these species can contribute to the control of disease vectors and intermediate hosts of parasites, several studies have been undertaken in the laboratory, where species of Belostoma have actually been shown to consume both the larvae and pupae of mosquitoes (Consoli et al., 1989) and snails that serve as the intermediate hosts of blood flukes (Pereira et al., 1991, 1993). The larvae of at least one South American Belostoma species were shown to select prey according to size (Pérez Goodwyn, 2001a), and one instar was particularly well suited for consuming larvae of the mosquito, Aedes aegypti, an important vector for yellow and dengue fever. 6.2.3. Ecology of the Leptopodomorpha Morphologically, species of Leptopodomorpha appear more similar to terrestrial bugs than do the members of the other two infraorders, but most of them are actually specialized for life in littoral habitats, either at the edge of freshwater bodies or along the seashore. This infraorder includes relatively few species, most of which are in the family Saldidae, the family best represented in South America. One species in the family Leptopodidae was found in an Ecuadorian termite nest and among vegetables shipped from Colombia to a North American seaport (Schuh and Slater, 1995). Aside from

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that species, no leptopodids or members of the other two families in this group are presently included in the leptopodomorph fauna of South America. Two leptopodomorph families, Aepophilidae and Omaniidae, include only species that inhabit the marine intertidal zone of the Palaearctic and Indo-Pacific Region, respectively. There are no known representatives of either family in South America (Cobben, 1970). Little is known about the habits of the saldids in South America, other than whether they inhabit marine or freshwater habitats. Members of those species already described seem to hunt small organisms along the shores of water bodies. Few species are longer than 2 mm, indicating that their prey encompasses the tiniest of insects or other invertebrates. It can be assumed that these insects are adapted to periodic flooding and employ their coats of fine setae to bind atmospheric air to the body surface while submerged. The habits of Holarctic species are better known, but there are too few of them to permit generalizations to be made concerning the life cycles of congeneric species in South America. Ecological studies of saldids were delayed until recent decades because the insects are small, inconspicuous, and lacking in obvious economic importance. Cobben (1960b) included much of what had been learned about saldid life cycles in his taxonomic publication on Caribbean shore bugs, and Polhemus (1985) summarized most of what has been learned about Central American species. Both publications include information about species also known to inhabit South America.

6.3. Preservation and examination Each of the three aquatic heteropteran infraorders requires a different collecting strategy. Most members of the Nepomorpha live below the surface of the water and can be captured using sieves, large hand nets, or dredges. Variations on these methods have proven effective for members of the individual families. Corixids and notonectids are strong swimmers and are best captured using fast sweeps of a large, long-handle net. Corixids are often captured near the bottom among detritus or among dense beds of submerged plants. Some notonectids are captured in such places, as well, but many prefer open water. Many more of these insects are usually captured on the second of two successive net sweeps at the same location because the disturbance of the water carries prey into the water column and strongly attracts the predatory bugs. The relatively small species in the families Pleidae and Helotrephidae are most often encountered in masses of submerged vegetation. Some are small enough to pass through coarse netting and are best captured using sieves with holes in the screening finer than 1 mm2. They can sometimes be found by placing a white laboratory tray just below the surface of the water at the top of plant masses and raising the tray with some water after several of the insects are seen swimming inside.

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Many families of Nepomorpha include mainly slow-moving species, which are not capable of escaping rather slow raking or scraping across the sediment surface. Belostomatids are particularly slow-moving, but they are effective at camouflaging themselves and must be searched for carefully among detritus or sediment samples. Nepids are generally smaller but capable of concealing themselves in the same way. Naucorids are somewhat faster and may escape if the sample material is collected too slowly. A different strategy is necessary for capturing gelastocorids, which frequent the littoral zone. They can sometimes be seen hopping out of the way when a person walks along the sandy shore of a water body, which explains their common name, toad bugs. They can be captured with the fingers either on the shore or in very shallow water. Adults seem to be most easily found and captured beside water bodies in tropical regions with alternating rainy and dry seasons while the water is receding during the dry season. Members of the Gerromorpha can be collected effectively using a large and fairly deep dip net, preferably at the end of a long pole. They are easily spotted on the surface of the water and can be captured by swift movements of the net. An effective trick to catch the larger, faster-moving species is to submerge the net beneath a location at which the insects congregate and remain motionless until some individuals return and position themselves above the net. If the net is raised quickly, some of the insects are caught before they have detected the net moving up from below. Because most species remain close to the shore, the use of a boat is rarely necessary to capture freshwater species. Exceptional groups include the marine species, some of which may be encountered on the surface of the open ocean. Most salt-water species, however, are most abundant in mangrove swamps, bays and harbors, and other relatively sheltered locations near the shore. Collecting these insects in tropical and subtropical habitats often requires the use of a boat. Members of the Leptopodomorpha are relatively uncommon in insect collections not only because they are actually less common than members of the other two aquatic infraorders but also because they are easier to overlook. Collectors of terrestrial bugs do not often visit the littoral habitat, and limnologists are generally distracted from the habitat along the shore by the more abundant members of other families on and in the water. Furthermore, casual collectors may not immediately distinguish these small insects from typical terrestrial heteropterans. It remains to be seen whether marine species belonging to the Leptopodomorpha somewhat similar those known from other continents will be found among seaweeds and debris along the seashores of South America. Studies of this group will probably yield some interesting results and prove to be a fertile field of research. At many localities in South America, heteropterans of many species undertake seasonal mating flights and may be collected under street lamps at night. The flight of massive numbers of these and other insects usually occurs in tropical and subtropical regions at the end of the dry season just before the

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freshet. At this time, the ground beneath street lights is often covered with aquatic heteropterans ranging in size from tiny corixids to large belostomatids. In the temperate zones, the flights of different species occur at different times, but a great many aquatic species undertake their mating flights in late spring or early summer on hot, muggy evenings shortly before heavy rainstorms. Care should be taken when collecting heteropterans because some of them can break the skin with their beaks and inject poisonous substances. Even relatively small gerrids can inflict painful bites. Other species secrete noxious chemicals from certain glands, which stain the skin yellow and have strong, unpleasant odors. After collection, most heteropterans can be brought back to the laboratory alive. Those species that depend on a so-called plastron of air trapped within a fine coat of minute setae will not survive if subjected to rubbing against other objects in the container. Gerrids and other surface-dwelling species will drown if agitated in a water-filled container. They are best carried is containers filled with moist, absorbent paper without folds or excessive water that might form small pools in the container during transport. The containers for the insects must be kept fairly cool and out of direct sunlight during transport. Because of their size and the hardness of their exoskeletons, aquatic heteropterans have usually been pinned and dried for preservation in collections. Large specimens are less likely to become distorted during the drying process than most other insects. In general, only members of the Coleoptera and Hymenoptera are less likely than heteropterans to be distorted by pinning and drying. Prior to preservation, the insects should be killed in special jars containing a toxic substance, such as chloroform, which does not change the colors or damage the surface of the chitin. Cyanide has sometimes been used, but if the specimens are left too long in the killing jar, their colors may change. Cyanide is also dangerous to work with. Specimens with dense coats of fine setae, which hold the plastron of air against the body, may be cleaned of fine debris that accumulates on the setae through an ultrasonic treatment. Specimens of large heteropterans are pinned by passing an insect pin through a stable part of the thorax just to the right of the dorsal midline so that it exits the thorax along the ventral midline. Insect pins should be made of a metal that does not rust or corrode, especially when the insects are to be kept for an extended period of time in moist, tropical climates. Such needles can be purchased from dealers specializing in scientific or laboratory supplies. After pinning, the insects should not be moved until they are completely dry so that the body will adhere to the pin. Many aquatic heteropterans are too small to be pierced with conventional insect pins. For these, several methods are used for preserving them dry in insect cases. The most common of these is affixing the insects to small pieces of stiff paper, such as that used for index cards. In some collections, the paper is cut in the form of tiny rectangles, and in others, small triangles are used, often referred to as points. The insect may then be glued to one end of the rectangle

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or to the acute point of the triangle, preferably using a paste that can be dissolved away easily using water. The insect pin is passed through the point and through labels, which should show the name of the species, the date and location of collection, and the collector’s identity. Alternatively, the insect can be affixed to the stiff paper using tiny pieces of very fine stainless steel wire, which is narrow enough to pass through the thorax of the specimen without crushing or otherwise damaging it. In recent decades, preservation in a liquid has become more popular, even for relatively large specimens. Ethanol, also called ethyl alcohol, is a fully satisfactory preservative for heteropteran specimens not collected for special analyses. This alcohol hardly reacts at all with chitin or proteins, and it should be avoided only if the lipids of the insects or fat-soluble organic substances, such as various kinds of pesticide, are to be analyzed. Solutions of 70% to 96% should prove effective for long-term preservation, but it is recommended to add up to 5% glycerine to keep the specimens soft and protect them somewhat from accidental desiccation. Other liquid preservative solutions may be used as long as they do not damage the chitin or setae, but ethanol is by far the most economical liquid preservative that does little damage to the specimens preserved. Most aquatic species of Heteroptera are characterized by large areas of the body covered with fine hydrophobic setae, which serve either to repel water from parts of the body where spiracles are located or to bind a layer of air, called a plastron, while the insect is underwater. Although most preservatives do not damage the tiny setae, fine suspended matter contaminating the solutions often become entrapped among the setae, making them difficult to examine. This material is often hard to clear away before preparation for electron microscopy and other special examinations. It is therefore recommended that preservatives be kept free from such suspended matter and to transfer the specimens as soon after fixation as possible into clean fluids after washing them gently with a flow of the preservative to remove debris that might have been placed in the fixative along with the insect. This washing serves the same purpose as the ultrasonic treatment of specimens to be pinned and dried. Acetone has sometimes been used for preserving various insects because it preserves the colors, but if it is used, the specimens will become useless for DNA studies and determinations of their lipid content. It may also result in loss of the tiny setae that hold the layer of air to the body while the insect is submerged. It is therefore not recommended for aquatic heteropterans. Although heteropterans resist damage from formaldehyde better than most insects, solutions of this preservative are not recommended because they discolor specimens, attack the surface structure of the chitin, and sometimes cause swelling, which prevents accurate measurement of the length and width of the insects. Working with formalin required a glove-box and elaborate exhaust systems to prevent harm to the health. The problem of preserving the colors present in life has never been fully solved. Some colors persist on preserved specimens for years, while others

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change rapidly. The patterns of insects preserved for long periods of time may also fade, making it difficult to match type material with fresh specimens. Color photography is therefore recommended to minimize later confusion resulting from color changes. Of course, it is necessary to make prints using photographic paper that does not change color too much with time. Electronic files of color images may prove superior to negatives for storage of images, but they also may deteriorate with time. Needless to say, the photography should be completed while the insect is alive. An important advantage of keeping specimens in a preservative solution is the exclusion of small destructive insects, which can unexpectedly appear in collections of dry arthropods and rapidly destroy them before their presence is even noted. Dermestid beetles are famous consumers of museum collections, requiring the construction of special storage cases to keep them out. In tropical countries, tiny ants are able to gain access to specimens in seemingly air-tight boxes and eliminate entire collections in a matter of hours. Poisons are often placed in the cases to kill the insects, creating a hazard for entomologists who work many hours with the collection. These problems have prompted increasing numbers of entomologists to opt for a liquid preservative, even in the case of relatively large insects. The size and shape of heteropterans and the natural position of their wings beneath robust hemielytra permit many of them to be stored in relatively little space. Many collectors have their own preferred methods of fixing and storing their specimens. Those judged to be best preserve specimens for the longest time with the least change. Unlike many other kinds of insect, no special methods are needed for fixing and storing specimens of water bug larvae. The larvae of almost all heteropterans share the habitat with adults and have body structures that react similarly to preservative solutions. Because larvae are smaller and somewhat more delicate, killing and fixation times may be slightly shorter. As in the case of other insects, rearing larvae in aquaria can provide useful information on the morphology and ecology of poorly known species (Pereira et al., 1991). It also permits larval stages to be identified positively to species. In recent years, several such studies have provided very useful information. The exuvia of each instar should be preserved for later study in 70% to 96% ethanol with up to 5% glycerine added or in another suitable preservative. Ethanol not only keeps the exuviae pliable, it protects them from the mold and bacteria that may attack dried specimens if the humidity becomes too high. A reducing agent can also be added to protect the color pattern.

6.4. Zoogeography Almost all South American aquatic species of Heteroptera are endemic to the Neotropical Region, and the ranges of most are limited to the South American continent. Some of those found in northern South America are also known

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from islands not far from the coast or the southern part of Central America. Notable exceptions are the marine water striders, some of which have the ability to live on the open ocean long enough to cross significant oceanic barriers, which limit the ranges of almost all other insects. Relatively few taxa are known only from the colder parts of the Andes Range, Patagonia, and the islands at the southern tip of South America. This pattern of distribution seems plainly evident only in the case of the Leptopodomorpha (Schuh and Polhemus, 2009). Among the groups most interesting for zoogeographers are the water striders that have succeeded in colonizing the oceans. The Halobatinae, a subfamily of the family Gerridae, have long been known from marine coastal regions in various parts of the world (Aguilar and Méndez, 1987). This subfamily is one of the few groups of Heteroptera with unique distribution patterns among the insects, mainly due to their ecological specialization. They were found on the open sea at times, and recent studies indicate that the pelagic habitat is the primary one for Halobates species (Andersen et al., 2000). Others seem more abundant in neritic zones. Marine veliids are often encountered in mangrove swamps and estuaries, where they find food, mates, and shelter from rough seas. In contrast, the migration of most water bugs to new habitats is limited, not so much by any weakness in their flying ability but rather by the limitation of their flight periods to specific seasons and weather conditions, which usually prevail for only a few days each year. Individuals of many species may reproduce for many generations without the adults developing wings at all. The adults of some species have never been found with functional wings, although that does not preclude their ability to produce macropterous generations under certain seasonal and climatic conditions without their having been discovered yet by entomologists. Judging from the apparent random appearance of water bugs far from water during the flight periods, many species seem to be distributed more by the winds than by any form of purposeful navigation. However, in many parts of South America, such random dispersal of adults would tend to guarantee that each species with occasional flight capability would eventually reach every water body within vast regions not interrupted by formidable natural barriers. Nevertheless, individual families show some interesting zoogeographical affinities, some being primarily associated with the African, Indian, and Indomalayian regions and others represented in South America by apparent migrants from North America. The family Belostomatidae is widespread in tropical and subtropical climatic zones and rare or absent elsewhere. This, of course, has more significance for the physiology and ecology of the family than for its zoogeography since the inability to adapt adequately to cold winter weather easily accounts for its absence north and south of subtropical regions. Belostomatidae is well represented in both the Old and New World, indicating that during some period of geological history, it has been able to overcome the oceanic barriers presently separating the continents. In spite of their failure to develop physiological adaptations to cold periods, many belostomatid species have been successful in colonizing habitats which undergo periodic desiccation.

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Other families of water bug known from South America are almost cosmopolitan, and some of their species have been successful in adapting to relatively severe winter weather. None of the families has a distribution limited enough to suggest that the species are relicts of ancient fauna or typical of identifiable geographical regions, such as the lands surrounding Antarctica. It should be noted that this applies only to the families of the aquatic species considered in this volume. Among terrestrial Hemiptera, relicts and families with ranges encompassing the Australasian Region and South America are also represented in the South American. More thorough studies are certain to reveal some interesting affinities, such as the apparent close relationship between one Chilean gelastocorid species, Nerthra praecipua, and congeneric Australian species. N. praecipua is obviously distinct morphologically from all other South American species (Polhemus and Cekalovic, 1998). In contrast to the cosmopolitan families, many genera are endemic to South America and closely adjacent parts of Central America. Others are confined to the North and Central America, the West Indies, and the northern parts of South America. However, entomologists making revisions of certain genera must be cautious of bias shown by their predecessors in grouping species from the geographic region in which they primarily work into new genera. Other insect species are known to have benefited from shipping traffic to colonize new continents, so a certain augmentation of the South American fauna with heteropterans introduced by man from other continents probably occurred, as well. Based on published records, most South American water bug species seem confined to very limited ranges, although many of these belong to genera common on other continents. Few species have been reported from many parts of the continent, but some of those with a limited distribution there are also encountered in Central or North America or in the West Indies. This may be due largely to the limited number of studies on the ranges of the species, which certainly results from the difficulties most researchers have in identifying the species they collect. Water bugs have not attracted as much attention as some terrestrial heteropterans, which have considerable economic impact on agriculture or present dangers to public health. They are not as attractive as dragonflies nor as useful for sport fishing as mayflies, so they do not seem to be as interesting for amateur entomologists as some of the other aquatic insect taxa. In addition, many species look enough alike to have made their identification too difficult for persons without the facilities to assemble a library of many publications from different countries, which were written during the past two centuries and account for our current knowledge of the fauna. On the South American continent, several distinct geographic and climatic regions can be characterized by their water bug fauna. The high Andes and cool southern parts of the continent harbor a relatively small number of species, while the tropical and subtropical lowlands provide habitats for a particularly rich heteropteran fauna. A few studies have provided records of the species that occur at different terrain elevation in mountainous regions (Alvarez and

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Roldán, 1983). Distributional studies have been conducted in various parts of South America, but most have provided lists of species for relatively small areas, such as national parks and nature reserves (Bachmann, 1962a; López Ruf et al., 2003; Torres et al., 2007). Some have been performed to survey the fauna in specific water bodies or drainage areas, such as the Retiro das Pedras in the State of Minas Gerais, Brazil (Vianna and de Melo, 2003), and the Paraná Delta (Bachmann, 1962b); special ecological regions, such as the llanos of Colombia (Roback and Nieser, 1974), the altiplano of Peru and Bolivia (Hungerford, 1927a; Roback and Nieser, 1980), a subtropical pampasic ecotone in Argentina (von Ellenrieder and Perez Goodwyn, 2000), and the Canastra Mountains in the State of Minas Gerais, Brazil (Pelli et al., 2006); surveys of individual families within political divisions of countries, such as Belostomatidae in Corrientes Province in Argentina (Reyes and Estévez, 1999) and Gelastocoridae at the sites in central Goiás, Brazil, visited by the Machris Expedition (Todd, 1957a); or local districts, such as the Município of Mariana (De Souza et al., 2006) in Minas Gerais; Pitinga in Amazonas, Brazil (Pereira and de Melo, 2007); the region of the Mar del Plata (Bachmann and Farina, 1989) and Martín García Island (Fernández and López Ruf, 1999) in the province of Buenos Aires, the mouth of the Iberá in Argentina (Estéves et al., 2003), the Meseta del Somuncura of Patagonia, Argentina (Muzón et al., 2005), and the reef at Maricá in the state of Rio de Janeiro, Brazil (Ribeiro et al., 1998). A study limited to aquatic heteropterans active during the season preceeding the annual period of flooding in a district of Rio Grande do Sul, Brazil, was completed by Neri et al., (2005). A few faunistic studies of larger regions, such as Surinam (Nieser, 1968, 1970a, b, c), the entire Guiana region (Nieser, 1975), Colombia (Aristizabal García, 2002), Ecuador (Froeschner, 1981), Uruguay (Rufinelli and Pirán, 1959; Mazzucconi and Bachmann, 2004), the Amazonian region of Peru (Roback, 1966), Patagonia (Bachmann, 1963a), southern Chile (China, 1962b), and Argentina (Contartese and Bachmann, 1987; Bachmann, 1962b, c; 1966a; 1979a, b; 1981) have yielded much longer lists of species in the groups encompassed by their surveys, but because of the size of the regions they cover, they are certainly less complete that those produced by intensive studies conducted at only one site. One of the few studies designed specifically to determine the ranges of individual species focused on the gerrid species of Argentina (Mazzucconi and Bachmann, 1995a). Froeschner (1985) provided an illustrated list of heteropterans inhabiting the Galápagos Islands, which are more affiliated politically than zoogeographically with continental South America. Since most studies include new distribution records for at least several species, it can be concluded that the known ranges of many species underestimate the real extent of their distribution. Furthermore, because of their inaccessibility, vast areas of South America have never been subjected to exhaustive entomological surveys, and some regions have never even been visited by zoologists. As a result, many species certainly remain to be discovered, and it is not yet possible to provide a reasonable estimate of the

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percentage of South American species that remain to be discovered. Naturally, the Amazon Basin comes to mind as the probable range of the greatest number of species that have thus far escaped the attention of taxonomists. In the case of aquatic heteropterans, there are very few examples of species confined to isolated habitats because they are unable to disperse to other regions, either via large drainage systems of through the air. Therefore, the distribution is determined primarily by the climate and by the ecological requirements of the individual species. There are many habitats in which several congeneric species occur together, indicating that many individual niches exist within relatively small areas. Occupation of discrete niches may depend upon the sizes of the insects, feeding specialization, or seasonal factors. Taking species of Belostoma as examples, it is easy to see how size determines which species of prey can be captured and eaten. Because individual Neotropical lakes often harbor several species of Belostoma, each with a different size range, it is easy to understand how each species could feed on its own group of prey species without competing with any congeneric species. Food competition could also be prevented by specialization in the feeding habits of species of similar size, such as certain corixids, which may feed on different groups of microorganisms and therefore not compete. Little is known about such specialization, and knowledge about the habits of the South American species is especially poor. Therefore, it is premature to conclude that niches created by specific selection of prey are anything more than possibilities. Seasonal activity patterns certainly govern the life cycles of most South American water bugs, but there is less evidence of niches being established based on seasonal activity differences among water bugs than among members of other insect orders. The large numbers of adult heteropterans belonging to a great variety of species observed flying around street lamps just at the time of the freshet in the Pantanal indicates that similarities rather than differences in the seasonal activity patterns in the tropical wet-and-dry climatic region are the rule (Heckman, 1998). Since many water bug species range throughout more than one distinct climatic zone, it is clear that much still must be learned about the zoogeographical factors limiting the ranges of the individual water bug species. Another rich field for future research is the determination of what constitutes the geographical barriers that limit the northward movement of the South American species and the southward movement of Nearctic species into South America. Because both larval and adult water bugs usually share common habitats, their ranges would logically depend more upon the nature of the water bodies in which they live than upon surrounding terrestrial habitats.

6.5. Taxonomic problems Conventional taxonomy of water bugs has focused almost entirely on adult specimens. The final instar larvae of almost all species closely resemble

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apterous adults and, for that reason, they are easy to confuse with adults during cursory examination, especially while collecting in the field. There are relatively few publications on interspecific differences among the larvae, and fewer still concern South American species. It is therefore wise to plan field collections only of adults for surveys and to collect larvae only when they are to be the subjects of special studies, which are needed to provide better descriptions, elucidate their ecological requirements, and for various other purposes. Special taxonomic studies of females and larvae will eventually be necessary to provide comprehensive descriptions of the species. The females of many genera have already been adequately described, thanks to the comprehensive monographs by some very prolific 20th century authors, whose works were thorough, accurate, and well illustrated. Problems have arisen with only a few taxa, mainly because several authors disagreed about the validity of many described species. The synonymy of the names of species in the genus Gelastocoris produced considerable disagreement in the past, and there has also been some controversy over the generic classification of the family Belostomatidae. More recent changes have involved the family Gerridae, in which several names used for many years as subgenera are being elevated to the rank of genus. Most of the disagreements in the past seem to have been settled, or they may not be debated at the present time due to a lack of active specialists to disagree. A major problem has resulted from the failure of earlier authors to provide illustrations of the species they described. In the case of some genera, more recent revisions have provided excellent figures of the morphological features useful for species identification. However, there are no existing illustrations of a few species, and some authors who provided reviews reported that the type specimens of some of these species could not be located. In conclusion, there are few problems in the general taxonomy of aquatic Heteroptera in South America. These include the need for major revisions of several genera, such as Paravelia and Limnocoris. In addition, still unknown sexes, wing morphs, and larvae urgently require descriptions, and most certainly, undescribed species remain to be discovered.

6.6. Suggestions for improvement Thanks to many extensive monographs, some dating back more than half a century, the problems encountered working with the aquatic Heteroptera are relatively easy to overcome. Most publications have been richly illustrated, making work with the South American water bugs easier and more pleasant than studies of most other insect groups. Future problems can be avoided if taxonomists continue describing the same morphological features that have been used in the past to distinguish members of the same families and genera. Descriptions of new taxa without illustrations

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should be avoided, and drawings should be provided of the most important taxonomic features. Photographs of water bugs appearing with some recent publications have not been of the best quality, probably due, in part, to the methods used for printing the journals in which they appeared. The most urgent problems which taxonomic and systematic entomologists will face in the immediate future is finding and providing descriptions of the various morphological forms of both sexes of each species. Apterous, brachypterous, and macropterous specimens of one species may appear according to seasonal cycles or randomly. In some cases, wing development of only one sex is known to vary, and many observations of populations in the field may be necessary to determine whether and under what circumstances other forms of the wings occur and the influence seasonal breeding patterns exert on the occurrence of winged and wingless forms. To facilitate faunistic and ecological studies, adequate descriptions of all forms that occur will eventually be needed. In addition, more information on the larval instars will be necessary to expand the keys to encompass immature specimens. It should be a goal to provide researchers in the field with sufficient information to identify all phases of each species and to elucidate their seasonal activity cycles. The methods used by the authors who have aleady provided accounts of the entire life cycles of a few South American species should be adopted for studies of other species, and raising the insects in the laboratory should be encouraged. As a general rule, each description of a heteropteran species should provide the following information: 1. A description of its general shape and dimensions, including the total length, width, and lengths of each antennal segment and the segments of each leg; 2. Dimensions of the head, including the space between the compound eyes measured along the vertex, the synthlipsis, the head length along the midline, and the width of the head between its posterolateral corners; 3. The number of segments of the beak and their absolute and relative lengths, as well as which of the coxae the beak reached when folded under the thorax; 4. Relative lengths of any prominent structures on the beak, such as rostral prongs; 5. The shape and dimensions of the pronotum with descriptions of its surface, including the presence and location of projections, depressions, or other obvious features; 6. The shape, size, and color of the scutellum; 7. The dimensions of the metaxyphus; 8. Obvious veins and the location of fractures in the hemielytra, if present, and the extent of their development; 9. The posterior extension of each leg relative to other parts of the body; 10. Descriptions, preferably with detailed illustrations, of all body parts modified to produce the male genital apparatus and whether the modifications are predominantly on the left side or right side of the body;

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11. External and, when possible, internal structures that form part of the female genitalia; 12. All structures that display sexual dimorphism, including, where appropriate, the appendages, wings, stridulatory apparatus, and setae; 13. The general color pattern, including any obvious markings or bright colors, and the differences, if any, between the color patterns of the males and females; 14. Any morphological character selected by earlier authors as a means for distinguishing species within any genus or confamilial genera; 15. Descriptions of the larval instars, whenever they are available for examination; 16. Illustrations of the eggs, whenever they are found with the adults. In many cases, size and color can provide the quickest means of identifying a water bug species. Once familiar with the local fauna, a specialist can immediately identify many living insects in the field from the general form, size, and color pattern. However, few South American species are known well enough to be identified so easily, and careful comparisons with type specimens are often necessary. Natural variability of the individual species has seldom been determined, and many species have been described based on only one sex or wing morph. A few nominal species are still known from only one specimen. Certainly, various species names still regarded as valid will be found to be synonymous with others after more information becomes available. Most species of certain genera, such as Belostoma, are very similar in coloration, making rapid identification without careful examination impossible in many cases. Total length and the dimensions of specific structures are often helpful to narrow down the number of possible species to which a specimen might belong. In most cases, however, examination of the genitalia or parts of the body displaying secondary sexual characteristics is necessary for a reasonably reliable identification. In many cases, only males or rarely only females can be identified, and considerable additional taxonomic work will be required to provide descriptions of both sexes and the larvae of all known species. It must always be remembered that absolutely reliable identifications of specimens from South America are still difficult at this time. First, there are many species which have not yet been described. Many parts of South America have still not been thoroughly searched for species. Regions within the Amazon River drainage area are thought to support the most diverse aggregations of species on earth, and these have not been systematically surveyed for insects. Other parts of the continent, including much of the Andes and Patagonia, may not be nearly as rich in species, but they have still to be explored thoroughly by entomologists, and it is certain that new species will be found. In addition to the problem of undescribed species that will occasionally be encountered, serious difficulties will occur due to the failure to have sufficiently accounted for intraspecific variability. After more specimens of each species become available, it will almost certainly be determined that two or more

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nominal species have actually been created for minor morphological varieties of single species. Until the natural variability of each species is better known, many hours of researchers’ time will be spent in puzzling over specimens with features intermediate between those of the type specimens of what are actually synonyms of one variable species. Even in parts of the world in which the insect fauna is far better known, problems with undescribed species and unrecognized synonyms occur occasionally. The extreme richness in species, for which South America is noted, results in a much greater magnitude of difficulty in working with its biota. Clearly, improvements will be made only by intensifying research, which will require adequate funding for taxonomic, systematic, and ecological studies and for maintaining collections of type specimens. An immediate goal should be the preparation of photographic catalogs of South American water bugs together with accurate drawings of the morphological structures, which can be be used to produce reliable field guides for use by researchers and amateur entomologists. Field studies could yield considerably more information about the habits of heteropteran species. There is relatively little information on the behavior of water bugs in South America. Studies on other continents have yielded valuable information about their communication with one another, their tracking of prey, and their reproductive behavior. Similar studies in South America have been undertaken on relatively few species, and more such studies will certainly result in the discovery of hitherto unknown survival strategies. New methods must be developed to study some of the smallest heteropterans, which interrupt their normal behavior to flee when approached close enough for observation in the field. Those that live on the surface tension layer of the water could easily be observed by mounting electronic cameras above locations at which they congregate and recording their interactions with prey, predators, and each other remotely. More elaborate methods will be necessary to observe the activity of nepomorphs beneath the surface of the water. Artificial habitats produced in the laboratory are also proving effective for raising and observing some of these species (Consoli et al., 1989; Pereira et al., 1991, 1993). A note on the figures Most illustrations were drawn based on figures appearing with the original descriptions of the species or in later review papers. In a few cases, they are based on figures prepared to justify taxonomic revisions or with descriptions of taxa later found to be synonyms of species already described. When using illustrations of specimens other than a holotype, there is a risk of perpetuating earlier published errors in identification. Original figures on which those in this book are based should be consulted as primary sources whenever they exist. Minor discrepancies between illustrations and specimens being examined will exist because individual variability of these insects can be considerable, and few

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taxonomists have worked with sufficient numbers of specimens from South America to appreciate the degree of this variability. In descriptions of water bugs, the relative dimensions of the body are frequently used to distinguish the species. Length to width ratios of various parts of the body are particularly useful for distinguishing many nepomorphs. It should be remembered, however, that individual species are known to have forms with wings developed to different degrees, and that wingless and winged individuals also display various other modifications of the thorax, which affect the relative dimensions of the body. The fact that only apterous or only macropterous individuals have been found and described by taxonomists does not mean that other forms do not exist. It is therefore well to remember that the illustrations of the habitus do not always show the general appearance of all adults belonging to any given species. Macropterous specimens should only be compared with illustrations of other macropterous individuals, and apterous forms with other apterous forms. The fact that specimens of all wing forms are not yet available for many species is a warning against hasty identification and an invitation to perform further research to better diagnose the species. As in the case of species belonging to most insect orders, the genitalia are usually the most definitive structures. After a specimen is tentatively identified based on other features described in the keys, the genitalia or secondary sexual structures, such as modifications of the appendages, patterns of setae on the pala or legs, and color patterns, should be compared with the illustrations provided. If no illustration is provided, the original species description should be consulted. Unfortunately, in some cases, there are neither good illustrations nor adequate descriptions of the genitalia, and it will be necessary to compare a specimen with type specimens designated in the original description to make a positive identification. In such cases, taxonomists are encouraged to undertake revisions of the group to which the species belongs.

6.7. Keys to aquatic Heteroptera in South America Information is based mainly on Schuh and Slater (1995). Only the three infraorders of species predominantly aquatic in their habits, Gerromorpha, Nepomorpha, and Leptopodomorpha, are included.

6.7.1 Infraorder Gerromorpha Key to families in South America 1. At least the fore-tarsi bear claws inserted dorsally and proximal to the apex. The coxae are inserted ventrolaterally with much more than their own length separating them across the ventral midline (Fig. 6.28). .......................................2

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- All claws are inserted at the apices of the tarsi. Exceptionally, members of the Mesoveliidae have claws inserted slightly proximal to the apex, but these have coxae inserted ventrally with less than their own length separating them across the ventral midline (Fig. 6.29). ..................................................................3 2. (1) The middle and hind coxae are clearly closer together than the fore and middle coxae. The hind femora are long and usually reach well beyond the apex of the abdomen. Adults have median metasternal scent gland openings dorsally (Figs. 6.4, 6.11, 6.28). .........................................................................................................Gerridae..p. 72 - The distances between the fore, middle, and hind coxae are subequal. The hind femora of the females and most males usually reach no farther than slightly beyond the apex of the abdomen. Adults have lateral metathoracic scent gland openings dorsally (Figs. 6.15, 6.30). ........................................................................................................Veliidae..p. 151

Fig. 6.28 Habitus of an apterous male Ovatametra thaumana. Based on Drake (1959).

Fig. 6.29 Habitus of Hydrometra caraiba with a lateral profile of the head and thorax at lower left. Based on Alvarez and Roldán (1983).

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3. (1) The head is as long as the entire thorax and longer than ¼ the total length. The body is long and cylindrical, and the legs are very narrow throughout their entire length. Nymphs lack dorsal abdominal scent gland openings (Fig. 6.29). .............................................................................................Hydrometridae..p. 255 - The head length does not exceed the combined length of the pronotum and scutellum and is less than ¼ the total length. The legs are not narrow for their entire length. Nymphs lack a single dorsal scent gland opening (Fig. 6.31). ...4

Fig. 6.30 Habitus of Stridulivelia cinctipes (left), its head and thorax in lateral view (upper right), and the right hemelytron in dorsal view (middle right). Based on Alvarez and Roldán (1983).

Fig. 6.31 Hebrus major: habitus in dorsal view (left), fore-wing (upper right), and habitus in ventral view without the legs (lower right). Based on Alvarez and Roldán (1983).

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4. (3) The body is less than 3 mm long and covered by small setae. The color pattern typically consists of blackish and reddish colors. Adults usually have two tarsal segments. The rostral sulcus is a groove on the ventral side of the head (Fig. 6.31). ..........................................................................Hebridae..p. 267 - Adults usually have three tarsal segments. No rostral groove is present, so the base of rostrum is clearly visible dorsally (Fig. 6.32). ..................................5

Fig. 6.32 Chepuvelia usingeri male: habitus of an apterous form (left), genital capsule with anal tube removed to reveal the parameres and penis (upper right), and the anal tube in dorsal view (lower right). Based on China (1962b).

Fig. 6.33 Mesovelia zeteki male (left to right): fore, middle, and hind legs, clasper in dorsal view (below), apical abdominal segments in ventral (above) and dorsal view (below). Based on Moreira et al. (2008). 5 (4) There are scattered stiff black bristles on the legs. On winged adults, the scutellum is always visible, and ocellli are present. Forms without wings lack a scutellum and ocelli (Fig. 6.18, 6.19, 6.33). ................................................................................................Mesoveliidae..p. 275

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- The legs do not bear stiff black bristles. The scutellum of macropterous and brachypterous adults is concealed by a plate of the pronotum. Ocelli are distinct dorsally (Figs. 6.32). ..........................................................................................................Macroveliidae The only South American species in this small family, Chepuvelia usingeri China, 1962, seems to be confined to Chile, where it was found in wet forest litter.

Gerridae Supplemental key to the subfamilies of adults Information for the key was provided by Matsuda (1960) and Andersen (1982). Matsuda (1960) recognized five groups as subfamilies and considered Cylindrostethini and Charmatometrini to be tribes in the subfamily Gerrinae. The classification in the key follows the system of Andersen (1982), although Charmometrinae might prove to be the correct spelling, based on priority. 1. In apterous specimens, the metacetabular suture continues dorsally to the posterior margin of the mesonotum and forms a secondary, definitive suture separating the mesonotum and metanotum, called the intersegmental suture. The intersegmental suture does not end at the posterior margin of the mesonotum but reaches the anterior border of the first abdominal tergite. The primary intersegmental suture is not produced anteriad to form lateral branches passing anterior to the metathoracic spiracle (Fig. 6.34). .....................2 - In apterous specimens, the metacetabular suture does not continue dorsally to the posterior margin of the mesonotum and does not form a secondary, intersegmental suture separating the mesonotum and metanotum, or, if it does, it ends at the posterior margin of the mesonotum. The intersegmental suture reaches the anterior border of the first abdominal tergite and does not end at the posterior margin of the mesonotum. The primary intersegmental suture is produced anteriad, forming lateral branches passing anterior to the metathoracic spiracle, or, rarely, a dorsolateral branch is absent (Fig. 6.35). ......3 2. (1) The first abdominal segment appears very prominent ventrally. The genae are well developed and lobate. The ovipositor is well formed. Vein R+M+Cu in the fore-wing is simply branched into two apical veins beyond the middle of the wing. There is always a pale line of weakness horizontally at the middle of the wing. A definitive intersegmental suture is represented by the posterior margin of the mesothoracic postnotum and metacetabular suture. A lateral suture separating the mesonotum from the mesopleuron is always present. The omphalium is absent (Fig. 6.34). ............................................................................................Rhagadotarsinae..p. 90

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- The first abdominal sternite is not visible. The definitive intersegmental suture is dorsally represented by the posterior margin of the mesothoracic postnotum and laterally by the metacetabular suture. There is no lateral longitudinal suture separating the mesonotum from the mesopleuron. The omphalium is usually absent. Vein R+M+Cu in the fore-wing is simply branched into two apical veins. There is always a horizontal line of weakness in the middle of the wing (Fig. 6.36). ................................................................................................Trepobatinae..p. 102

Fig. 6.34 Rheumatobates imitator male: habitus in dorsal view (center), outside view of the left antenna (upper left), and the right hind leg (below). Based on Hungerford (1954). 3. (1) The metacetabular suture is dorsally connected to the anterolateral angle of the first abdominal tergite. The primary intersegmental suture is always distinct laterally. A lateral longitudinal suture separating the mesonotum from the mesopleuron is absent, except in the genus Potamometra. An omphalium is present. Veins R+M and Cu of the forewing are either separated completely or fused basally. Sc is not connected to R+M by an oblique vein (Fig. 6.35). .............................................................................................................Ptilomerinae No species in this Old World subfamily has yet been found in South America. - The metacetabular suture is not connected to the anterolateral angle of the first abdominal tergite. The genae are not lobately developed. An ovipositor is not developed. The definitive intersegmental suture is not dorsally represented by the posterior margin of the mesothoracic postnotum or laterally, by the metacetabular suture. Veins R+M and Cu of the fore-wing are fused basally. The first abdominal segment is not evident ventrally (Fig. 6.37). .......4

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Fig. 6.35 Potamometra berezowskii Bianchi, 1896: habitus without appendages of a male (upper left) and a female in dorsal view (middle left), head in lateral view (lower left), fore and hind wing (upper right and right center, above and below, respectively), ventral view of the metasternum and abdomen of a male (below wings) and a female (middle right), antenna (lower left center), fore-leg of a male (lower right center), middle tarsal segment (space below fore-leg), and apical tarsal segments of the hind leg (lower right). This species is known only from China. Based on Matsuda (1960).

Fig. 6.36 Trepobates trepidus: dorsal view of the macropterous form (left) and the body of apterous form (right). Based on Alvarez and Roldán (1983).

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Fig. 6.37 Limnogonus aduncus: habitus of an apterous male in dorsal view (upper left), head with the beak and the basal antenna segments in anterior view (upper right center), enlargement of the joint between the basal and second segments of the antenna (middle right), apex of the male abdomen in lateral view (lower left) and its enlarged outline lacking details (lower right), and outline of the apex of the female abdomen in lateral view (upper right). The outlines of the apices of the male and female abdomens are based on Nieser (1970a), and the remaining illustrations are based on Jurberg and Gomes (1983).

Fig. 6.38 Halobates sericeus habitus (left) and apex of the male abdomen in dorsal (upper right), ventral (middle right), and lateral view (lower right). Based on Herring (1961) and Schuh and Slater (1995).

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4. (3) Vein R+M+Cu forks to form R+M and Cu. Sometimes vein R+M is not connected to the embolium by an oblique vein. Veins Cu and A join apically. The definitive intersegmental suture is dorsally represented by the posterior margin of the mesothoracic scutoscutellum and is obliterated laterally in most genera. There is no lateral longitudinal suture separating the mesonotum from the mesopleuron (Fig. 6.38). Marine species. ..................................................................................................Halobatinae..p. 114 - Vein R+M in the fore-wing is connected to Sc by Sc2. The primary intersegmental suture between the mesonotum and metanotum is either distinct or obliterated laterally. The lateral longitudinal suture between the mesonotum and mesopleuron is usually absent. The omphalium is always present (Fig. 6.37). The insects are usually confined to fresh or brackish water. ...............................................................................................................................5 5. (4) The metacetabular groove and posterior margin of the mesopostnotum join dorsally. The fourth segment of the antenna is short and curved. The short beak does not extend past the border of the prosternum. The second segment of the fore-tarsus is more than twice as long as the first (Fig. 6.39). .........................................................................................Cylindrostethinae..p. 118 - Dorsally, the metacetabular groove is indistinct. The fourth segment of the antenna is straight. The beak clearly passes the border of the prosternum (Fig. 6.40). .....................................................................................................................6

Fig. 6.39 Cylindrostethus palmaris: habitus of an apterous female in dorsal view (below) and (above, left to right): apex of the male abdomen in ventral view and apex of the female abdomen in ventral and lateral view. Based on Matsuda (1960) and Nieser and Alkins-Koo (1991).

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6. (5) On the metasternum, there are lateral omphalial grooves, i.e. scent channels, connecting the opening to the scent organ to the metacetabular region. There is a straight transverse line across the metanotum. Connexival spines are absent (Fig. 6.40). .......................................................................................Charmatometrinae..p. 131 - The metasternum lacks lateral omphalial grooves, or, if they are present, long connexival spines are formed, and the length of the insect is at least 30 mm. There is no transverse line on the metanotum (Fig. 6.37). ...................................7 7. (6) In the apterous form, the pronotal lobe is always reduced. There are pretarsal areolae. There is a distinct lateral suture between the mesothorax and metathorax (Fig. 6.41). ..............................................................................................................Eotrechinae This subfamily seems confined to the Oriental Region and is apparently not represented in South America. - In the apterous form, the pronotal lobe is usually not reduced, but if it is, then there are no pretarsal areolae, or the lateral suture between the mesothorax and metathorax is indistinct (Fig. 6.37). .......................................................................................................Gerrinae..p. 135

Fig. 6.40 Charmatometra bakeri: habitus of an apterous female in dorsal view (upper left), abdomen in ventral view (upper right), and (middle row, left to right): head in lateral view, apical segments of a male abdomen in ventral view, male genital segment in lateral view, female genitalia with the apical segment of the endosome shown at upper middle right, and (below, left to right): antenna, the joint between the male fore-femur and tibia, apical part of the fore-leg, hind tarsus, and the apical segment of the middle tarsus. Based on Matsuda (1960).

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Fig. 6.41 Eotreches kalidasa Kirkaldy, 1902: habitus of a macropterous specimen without appendages in dorsal view (upper left), head in lateral view (upper right), thorax and anterior part of the abdomen in lateral view (center), habitus of a macropterous male without limbs in ventral view (lower left), apical segments of the abdomen of a male in ventral (lower center) and lateral view (lower right), male fore-leg (middle right), and the generalized depiction of the thorax and anterior abdomen of an apterous member of the Eotrechinae in dorsal view showing the typical morphology of the subfamily (middle left). Based on Matsuda (1960) and Andersen (1982). Key to the genera of adults in South America Information for the key was provided by Drake and Harris (1934), Kenaga (1941), Kuitert (1942), Hungerford and Matsuda (1958a, b, c), Matsuda (1960), Nieser (1970a), Nieser and Alkins-Koo (1991), Polhemus (1991a), Bachmann and Mazzucconi (1995), García (2002), and Pereira et al. (2007). 1. The metacetabular suture of apterous specimens continues dorsally to the posterior margin of the mesonotum and forms a secondary, definitive suture separating the mesonotum and metanotum, called the intersegmental suture. The intersegmental suture does not end at the posterior margin of the mesonotum but reaches the anterior border of the first abdominal tergite. The primary intersegmental suture is not produced anteriad to form lateral branches passing anterior to the metathoracic spiracle (Fig. 6.34). .....................2 - In apterous specimens, the metacetabular suture does not continue dorsally to the posterior margin of the mesonotum or form a secondary intersegmental suture separating the mesonotum and metanotum, or, if it does, it ends at the posterior margin of the mesonotum. The intersegmental suture reaches the anterior border of the first abdominal tergite and does not end at the posterior

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margin of the mesonotum. The primary intersegmental suture is produced anteriad, forming lateral branches passing anterior to the metathoracic spiracle, or, rarely, a dorsolateral branch is absent. The genae are not lobately developed (Fig. 6.38). An ovipositor is not developed. ....................................................10 2. (1) The first abdominal sternite is plainly visible. The genae are well developed and lobate. The female has a well-developed ovipositor. Vein R+M+Cu in the fore-wing is simply branched into two apical veins beyond the middle of the wing. There is always a pale line of weakness horizontally at the middle of the wing. A definitive intersegmental suture is represented by the posterior margin of the mesothoracic postnotum and metacetabular suture. A lateral suture separating the mesonotum from the mesopleuron is always present. The omphalium is absent. The first abdominal segment appears very prominent ventrally. The third antennal segment and posterior margin of the eyes bear long, hair-like setae (Figs. 6.11, 6.34). .....................................................................Rheumatobates Bergroth, 1892..p. 90 - The first abdominal sternite is not visible (Fig. 6.42). .....................................3 3. (2) The hind femur is obviously longer than the body length. The distal ends of the second and third antennal segments are swollen, and those of the male sometimes bear conspicuous groups of spines. The second hind tarsal segment bears long, knife-like claws at or distal to the middle. The middle tibia is the longest leg segment (Fig. 6.42). ...............................................................................Metrobates Uhler, 1871..p. 102 - The hind femur is equal to or shorter than the body length (Fig. 6.43). ..........4 4. (3) The apical segment of the fore-tarsus more than twice as long as the basal segment (Fig. 6.43). ..............................................................................................5 - The apical segment of the fore-tarsus is not more than twice as long as the basal segment (Fig. 6.28). The omphalium is distinctly evident. ......................8

Fig. 6.42 Metrobates amblydonti male (left to right): antenna; one side of the thorax and abdomen in ventral view, showing the spine on the mesothoracic acetabulum; thorax and abdomen in ventral view showing the parameres; a paramere showing the narrowing apex. Based on Posso and González (2008).

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Fig. 6.43 Trepobatoides boliviensis (above, left to right): habitus of an apterous female in dorsal view, female abdomen in ventral view, and female genitalia, and head in lateral view (middle left), antenna (lower left), apical segment of the endosoma (upper middle right), male abdomen in ventral view (lower left center), male genital segment in lateral view (lower right center), fore-leg of a female (lower right center to lower right), and the middle (above fore-leg) and hind tarsus (above middle tarsus). Based on Matsuda (1960). 5. (4) The first antennal segment is much longer than the combined length of the second and third (Fig. 6.43). The color is mainly yellow with a few black markings, generally longitudinally oriented. .......................................................Trepobatoides Hungerford and Matsuda, 1958 This genus is represented by a single species, Trepobatoides boliviensis Hungerford and Matsuda, 1958, known only from Bolivia. - The first antenna segment is no more than about equal in length to the second and third together (Fig. 6.44). ...............................................................................6 6. (5) Along the inner margin of the first tarsal segment on the hind leg, there is a row of 5 or 6 short, stout, black spines. The mesosternum has prominent black and yellowish brown markings, and there is an irregular black marking along the posterior margin of the pronotum, which forms a stripe to the anterior margin in the middle (Fig. 6.44). The apterous male of the only known species is about 3.15 mm long, and the length of the apterous female is about 3.7 mm. .........................................................................Telmatometroides Polhemus, 1991 The only species yet described in this genus is Telmatometroides rozeboomi (Drake and Harris, 1937) from Central America, Colombia, and Ecuador. Syn: Telmatometra rozeboomi Drake and Harris, 1937.

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- The first tarsal segment on the hind leg lacks a row of 5 or 6 short, stout, black spines. The body is dark yellow with black and white markings, reddish brown and black, or uniform in color (Fig. 6.45). ................................................7 7. (6) The third segment of the antenna is more than twice as long as the second and distinctly longer than the first (Fig. 6.45). .....................................................................Telmatometra Bergroth, 1908..p. 104 - The third antenna segment is less than 1.5 times as long as the second and clearly shorter than the first. The body is yellowish brown with indistinct darker markings (Fig. 6.46). The only species in this genus is 2.6 to 3.2 mm long. ..............................................................................Cryptobatoides Polhemus, 1991 The only species yet described in this genus is Cryptobatoides brunneus Polhemus, 1991, from Amazonas.

Fig. 6.44 Habitus of Telmatometroides rozeboomi in dorsal view without appendages. Based on Kenaga (1941).

Fig. 6.45 Telmatometra whitei: habitus of an apterous specimen (left) and male clasper (right). Based on Drake and Harris (1937) and Kenaga (1941).

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8. (4) The middle tibia is distinctly shorter than the body (Fig. 6.28). ...........................................................................Ovatametra Kenaga, 1942..p. 107 - The tibia on the middle leg is about as long as or slightly longer than the body (Fig. 6.47). ............................................................................................................9

Fig. 6.46 Cryptobatoides brunneus: habitus of a female without appendages in dorsal view (left), habitus of a male without appendages in lateral view (upper right), and paramere of the male genitalia (lower right). Based on García (2002).

Fig. 6.47 Halobatopsis spiniventris: habitus of an apterous male in dorsal view (upper left), apical segments of the male abdomen in ventral view (upper right center), head in lateral view (lower left), male hind tarsus (lower center), fore (center) and hind wing (lower left center), male genital segment in lateral view (middle right), and female genitalia (lower right). Based on Matsuda (1960).

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9. (8) Viewed laterally, the compound eyes do not extend posterior to the middle of the propleurae. The hind tibia is clearly less than twice as long as the tarsus (Fig. 6.36). ...............................................................................Trepobates Uhler, 1894..p. 111 - The compound eyes extend posterior to the middle of the propleurae. The hind tibia is more than twice as long as the tarsus (Fig. 6.47). ........................................................................Halobatopsis Bianchi, 1896..p. 112 10. (1) The metasternum is reduced to a very short, transverse, nearly triangular plate, which usually does not reach laterad as far as the metacetabular region, or it is evident only as the omphalium. The basal margin of the clypeus is welldefined. The anterior margin of the head is not evenly rounded. There is a fringe of long erect hairs on the tibia and first tarsal segments of the middle leg (Fig. 6.38). All species are marine. .......................................................................Halobates Eschscholtz, 1822..p. 114 - The metasternum is plainly evident and at least about 1/10 as long as the mesosternum (Fig. 6.37). ....................................................................................11 11. (10) The metacetabular suture extends dorsally as far as the suture between the mesonotum and metanotum (Fig. 6.48). .......................................................12 - The metacetabular suture does not extend dorsally as far as the suture between the mesonotum and metanotum (Fig. 6.37). ........................................14

Fig. 6.48 Platygerris depressus: habitus of an apterous male (left) and the apical abdominal segments of a female in ventral view (right). Based on Matsuda (1960). 12. (11) The body is greatly flattened and short, and it lacks an omphalial groove (Fig. 6.48). The only South American species is about 7 to 8 mm long. ..........................................................................................Platygerris White, 1883 Only one species has been reported from South America, Platygerris depressus White, 1883, known from Mexico and Colombia. It is distinguished from two congeneric species known only from Costa Rica by the eighth abdominal segment of the male, which is four times longer than wide, while in the Central American species, it is not more than twice as long as wide.

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- The body is cylindrical and much shorter than wide. The rostrum does not reach the base of the prosternum (Fig. 6.39). .....................................................13 13. (12) The body is long and cylindrical, and its length is at least four times its width. The middle and hind femora extend only slightly beyond the apex of the abdomen. The antennae are relatively short. The abdominal spiracles are closer to the anterior margins than the posterior margins of the segments. The male genital segments are symmetrical (Fig. 6.39). .......................................................................Cylindrostethus Fieber, 1861..p. 118 - The body is less than four times as long as broad (Fig. 6.49). The body is not shiny. In most species, the abdominal spiracles are closer to the anterior margins than the posterior margins of the segments. The male genital segments are rotated and asymmetrical. .....................................................................Potamobates Champion, 1901..p. 122

Fig. 6.49 Potamobates peruvianus: habitus of an apterous female in dorsal view (upper middle left), antenna (upper left), head in lateral view (lower left), apex of a female abdomen in lateral view (upper right), fore-leg (lower left center), middle tarsus (lower center), and hind tarsus (lower right center), and the female genitalia (lower right). Based on Matsuda (1960). 14. (11) An omphalial groove is present (Fig. 6.50). .........................................15 - An omphalial groove is absent (Fig. 6.51). ....................................................17 15. (14) The middle tibia is longer than the middle femur (Fig. 6.50). ............................................................................Brachymetra Mayr, 1865..p. 131 - The middle femur is longer than the middle tibia (Fig. 6.52). .......................16 16. (15) There are four distinct black longitudinal stripes on the pronotum, which has a ground color of pale yellow in the only known South American

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species. The first tarsal segment on the fore-leg is shorter than the second (Fig. 6.52). Apterous and macropterous forms are encountered. ..............................................................................Eobates Drake and Harris, 1934 The only species in this genus known from South America is Eobates vittatus (Shaw, 1933) from Haiti and Peru, also referred to by its synonym, Brachymetra vittata Shaw, 1933. - The pronotum is uniform brown and lacks black longitudinal stripes. The first tarsal segment on the fore-leg is longer than the second (Fig. 6.40). Members of this genus are large, reaching a length greater than 12 mm. ..............................................................................Charmatometra Kirkaldy, 1899 The only South American species in this genus is Charmatometra bakeri (Kirkaldy, 1898), described under the synonym Brachymetra bakeri Kirkaldy, 1898. It is known from Venezuela, Colombia, and Ecuador. This correct generic name may be Charmometra, but the spelling Charmatometra persists in the literature.

Fig. 6.50 Brachymetra albinerva: habitus of an apterous specimen in dorsal view and the head and pronotum in lateral view (lower left). Based on Alvarez and Roldán (1983).

Fig. 6.51 Tachygerris adamsoni: macropterous male in dorsal view (upper left), ventral view of the male abdomen (upper right), and apex of a female abdomen in ventral view (lower center). Based on García (2002).

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Fig. 6.52 Eobates vittatus: a macropterous (upper left) and an apterous form (lower left) in dorsal view without appendages or markings shown, head and pronotum of the macropterous form showing the dark pattern (center), apex of the male abdomen in ventral view (upper right center) and enlarged as shown with the original species description (right), and apex of the female abdomen in ventral view (lower right center). Based on Shaw (1933) and Matsuda (1960). 17. (14) The second segment of the antenna is much shorter than the third. The antenna is about as long as or longer than the entire body. The fourth antennal segment is distinctly the longest (Fig. 6.51). Only macropterous specimens are known. .............................................................................Tachygerris Drake, 1957..p. 135 Species now placed in this genus were often found in older literature in the genus Tenagogonus Stål, 1853, a genus presently with no known representatives in South America. - The third segment of the antenna is, at most, slightly longer than the second and usually shorter (Fig. 6.53). ..........................................................................18 18. (17) The pronotum is usually shiny and has a yellow stripe along the dorsal midline, short lateral yellow stripes, or a single large yellow spot on the anterior lobe (Fig. 6.53). The rostrum reaches the base of the mesonotum. .................19 - The pronotum is matt with a yellow stripe along the dorsal midline, which fades out on the posterior lobe. A pair of large black spots is sometimes present lateral to the yellow stripe along the midline (Fig. 6.54). ......................20

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Fig. 6.53 Habitus of a macropterus Limnogonus hyalinus. Based on Andersen (1975).

Fig. 6.54 Eurygerris flavolineatus male: habitus in dorsal view (lower left), forefemur (upper left), and abdomen in ventral view (right). Based on Matsuda (1960) and Drake (1963). 19. (18) The pronotum reaches the metanotum and almost completely covers the mesonotum, and its anterior lobe has only one pale mark in the middle. The length of the first antenna segment is usually less than but sometimes subequal to the width of the head through the eyes. Claws are absent on the middle and hind legs (Fig. 6.55). ........................................................................Neogerris Matsumura, 1913..p. 139

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- The pronotum reaches at least to the middle of the mesonotum, and its anterior lobe has a pair of wide, pale lines usually reaching its anterior margin, but if they become obsolete before reaching the margin, then the first antenna segment is longer than the width of the head through the eyes (Fig. 6.53). ................................................................................Limnogonus Stål, 1868..p. 144 20. (18) The first segment of the antenna is much shorter than the second and third combined. The hind and middle femur are approximately equal in length. The pronotum does not cover the entire mesonotum. The metathoracic spiracle is separated from the pronotum by more than its own length. The posteriormost sternite of the male is doubly emarginate at the apex. The apices of the connexivum lack spines (Figs. 6.4, 6.54). .................................................Eurygerris Hungerford and Matsuda, 1958..p. 148 - The first segment of the antenna is equal to or longer than the second and third combined (Fig. 6.56). .................................................................................21

Fig. 6.55 Neogerris hesione: habitus of an apterous female without appendages in dorsal view (upper left), antenna (lower left), head in lateral view (left center), fore-leg (right center), and apical segments of a female abdomen in lateral view (right). Based on Matsuda (1960). 21. (20) The hind tibia is at least four times the length of the first segment of the hind tarsus. The ventral emargination along the midline of the apical abdominal segment is simply rounded (Fig. 6.56). The spine at each posterolateral corner of the connexivum is very short and inconspicuous in the only South American species but long in other species of the genus. Length: about 10 to 11 mm. ...................................................................................Aquarius Schellenberg, 1800 Aquarius was long recognized as a subgenus of Gerris with worldwide distribution, represented by several species in North and Central America as far south as Guatamala and by one South American species, Aquarius chilensis

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(Berg, 1881), known from Chile and Peru and reported questionably from Venezuela. Syn: Limnotrechus chilensis Berg, 1881; Gerris chilensis (Berg, 1881). Its morphological characteristics were compared with those of congeneric species by Andersen (1993b). Aquarius is now treated as a genus by most authors. - The hind tibia is never longer than 3.2 times the length of the first segment of the hind tarsus. The ventral emargination along the midline of the apical abdominal segment is sinuous and forms a doubled emargination (Fig. 6.57). The connexiva of the only known South American species is distinctly produced at the apex. The fore-femora are not strongly curved. The length of the only species reported from South America is 8 to 11 mm. .............................................................................................Gerris Fabricius, 1794 Since some of its former subgenera have been given generic status, only one species of Gerris appears in species lists from South America: Gerris marginatus Say, 1832. This North American species was previously thought to be very widespread from Canada to Brazil, possibly due to errors in identification or because individuals were introduced from North America at seaports. Reports of its presence in South America are few and questionable, and Andersen (1993b) stated positively that no South American species of Gerris are known.

Fig. 6.56 Aquarius chilensis: habitus of an apterous male (middle left), dorsal view of a macropterous individual without appendages (middle right), apex of the male abdomen in ventral (upper left) and lateral view (upper right), and apex of a female abdomen in ventral (lower left) and lateral view (lower right). Based on Andersen (1990).

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Fig. 6.57 Gerris marginatus: apex of the female abdomen in ventral view (upper left), apical segment of the endosoma (upper right), and female genitalia (lower center to right). Based on Matsuda (1960). Subfamily Rhagadotarsinae Key to the species of adult male Rheumatobates found in South America Information for the key was provided by Esaki (1926a), Drake and Harris (1932a, 1942a), Hungerford (1954), Drake (1958a), Nieser (1970a), Polhemus and Manzano (1992), Westlake et al. (2000), and García (2002). Several species found only in North America and the northernmost West Indies are excluded from the key. Because species in this genus are usually associated with marine habitats, all of the species known only from southern Central America and Trinidad could and probably will be encountered along the northern coast of South America and are therefore included in the key. 1. The hind femur is straight, and the segments of the legs are unmodified (Fig. 6.58). .....................................................................................................................2 - The hind femur is not straight, and other leg segments may be incrassate, curved, twisted, or spurred (Fig. 6.59). ................................................................9

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Fig. 6.58 Rheumatobates trinitatis male: habitus in dorsal view (left), outer surface of the left antenna (lower right), and a posteroventral view of the forefemur (upper right). Based on Hungerford (1954).

Fig. 6.59 Rheumatobates probolicornis male: antenna in dorsal view (upper left), fore-leg (middle left), claws on fore-leg (upper center), femur and trochanter in oblique view (lower left), and apex of the abdomen in dorsal (upper right) and lateral view (lower right). Based on Polhemus and Manzano (1992).

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2. (1) The entire ventral surface of the first abdominal segment is deeply excavated and appears cup-like. There are thick, hair-like setae scattered on the ventral margin of the fore-femur (Fig. 6.60). Longitudinal furrows are not present on the sixth or seventh abdominal sternites. The color of the mesosternum is light, and the mesocoxae are dark with a light spot. Length of apterous male: c. 2.4 mm; apterous female: c. 2.9 mm. .......................................Rheumatobates peculiaris Polhemus and Spangler, 1989 (Ecuador, Colombia). - The ventral surface of the first abdominal segment is not deeply excavated, although a small excavation localized on part of the surface may be present (Fig. 6.61). .....................................................................................................................3

Fig. 6.60 Rheumatobates peculiaris male: antenna (upper left), fore-leg (upper center), apex of the abdomen in ventral view (lower left), genital segments (upper right), and hind leg (lower right). Based on Polhemus and Spangler (1989). 3. (2) The fore-femur is robust and has a fringe of coarse setae along the anterior margin (Fig. 6.58). The length is greater than 2.5 mm. .....................................4 - Either the fore-femur is not notably robust or it lacks a fringe of coarse setae along the anterior margin (Fig. 6.61). The length is less than 2.5 mm. ...............5 4. (3) There is a dense patch of long, curved setae near the apex of the forefemur of the male. The anterolateral margin of the head of the female is yellow (Fig. 6.58). The length of the apterous male is 2.8 to 3.2 mm. The apterous female is 3.2 to 3.8 mm long. ..................................................................Rheumatobates trinitatis (China, 1943) (West Indies, Surinam). Syn: Hynesia trinitatis China, 1943.

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- There is no dense patch of long, curved setae near the apex of the fore-femur of the male (Fig. 6.62). The anterolateral margin of the head of the female is black. Length of the male: c. 2.8 mm; female: c. 3.3 mm. ..........................................................Rheumatobates mangrovensis (China, 1943) (Trinidad).

Fig. 6.61 Habitus of a male Rheumatobates minutus in dorsal view. Based on García (2002).

Fig. 6.62 Rheumatobates mangrovensis: antenna of a female (left); fore-leg of a male (right). Based on Matsuda (1960). 5. (3) The longest segment of the antenna is the third (Fig. 6.61). The anterior femur is slightly thicker than the middle one. The first genital segment is not hairy. Length of male: 1.6 to 1.9 mm; female: 1.7 to 2.2 mm. .............................................................Rheumatobates minutus Hungerford, 1936 (North and Central America, West Indies, Colombia, Peru, Argentina, Minas Gerais, Rondônia, Amazonas?). Syn: Rheumatobates curracis Drake and Carvalho, 1954 is considered a synonym of Rheumatobates minutus flavidus Drake and Harris, 1942. The other subspecies presently recognized is Rheumatobates minutus minutus Hungerford, 1936, which occupies the part of the range as far south as Panama. R. m. flavidus Drake and Harris, 1942, is

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found exclusively in South America. It is distinguished by the mainly yellowish dorsal part of the thorax, while that part of R. m. minutus is mainly black. The validity of the subspecies has not been confirmed by recent studies, but they have been maintained as such (Westlake et al., 2000). - The longest segment of the antenna is the first (Fig. 6.63). .............................6

Fig. 6.63 Rheumatobates clanis (left to right): antenna of a male and a female; fore-leg of a male. Based on Matsuda (1960). 6. (5) The fore-femur of the male is thickened and has a conspicuous curving row of long setae on its ventral side, which has a parallel row of setae on the basal third (Fig. 6.63). Length of male: c. 2.1 mm; female: c. 2.3 to 2.4 mm. .....................................................Rheumatobates clanis (Drake and Harris, 1932) (Belize, Cuba, suspected to occur in Colombia). - The fore-femur of the male lacks a conspicuous curving row of long setae on the ventral side and lacks the parallel basal row, as well, although five or six long, thickened setae may be present (Fig. 6.64). ................................................7 7. (6) The first antennal segment is the longest, and it is yellow (Fig. 6.65). There is a row of small spines in the middle of the internal surface of the middle femur. The hind femur has a row of small setae running its whole length on the internal surface. Length of the apterous male: 2.0 to 2.1 mm. Length of apterous female: 2.5 to 2.6 mm. .............................................................Rheumatobates drakei (Hungerford, 1954) (Peru, Ecuador, Surinam, Guyana). - The first antennal segment is the longest, and it is black or blackish fuscous, sometimes with a whitish testaceous basal band and whitish ventral surface (Fig. 6.64). .....................................................................................................................8 8. (7) The fore-femur does not bear thickened, hair-like setae. The first antennal segment is black (Fig. 6.64). There is a shallow longitudinal groove running along the last abdominal and basal genital segment. Length of male: 1.89 to 2.1 mm. Length of female: 2.25 to 2.52 mm. ...............................................................Rheumatobates bonariensis (Berg, 1898) (Argentina, São Paulo, Santa Catarina, Rio Grande do Sul). Syn: Rheumatobates wrighti Drake and Harris, 1937.

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- The fore-femur bears a row of five or six thickened, hair-like setae. The first antennal segment is blackish fuscous with a narrow white fuscous basal ring and ventral surface. Length: c. 1.75 mm. Maximum width: c. 0.6 mm. Brief descriptions of an apterous and a macropterous male have been provided, but further study is required to determine that the two species in this couplet are really distinct. .....................................................................Rheumatobates minimus Drake, 1958 (Peru, Santa Catarina).

Fig. 6.64 Antenna of a male Rheumatobates bonariensis. The black first segment is not shown in full length or color. Based on Matsuda (1960).

Fig. 6.65 Antenna of a male Rheumatobates drakei.

Based on Matsuda (1960).

Fig. 6.66 Rheumatobates ornatus male: antenna (left) and the apical abdominal segments in ventral view (right). Based on Westlake et al. (2000). 9. (1) The first segment of the antenna is greatly enlarged and widest at the distal end (Fig. 6.59). Length of the apterous form: c. 2.3 to 2.5 mm. It is known from estuaries. Macropterous forms are unknown. ................................Rheumatobates probolicornis Polhemus and Manzano, 1992 (Colombia).

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- The first segment of the antenna is not clearly widest at the distal end (Fig. 6.66). ...................................................................................................................10 10. (9) The middle femur is excavated near its midlength and bears claw-like structures formed by rigid setae. The third segment of the antenna bears a row of long setae on its basal half (Fig. 6.66). ..............................................Rheumatobates ornatus Polhemus and Cheng, 1977 (Costa Rica). - The middle femur is not excavated and bears, at most, a row of rigid setae (Fig. 6.67). ..........................................................................................................11 11. (10) Some of the abdominal segments have a fringe of long setae inserted along the margins, folded and directed ventrad and inward. Some of these setae may reach or almost reach the center of the segment (Fig. 6.67). .............12 - There are no fringes of setae on the abdomen. The hind trochanter is incrassate and as thick as the coxa (Fig. 6.68). ..................................................14

Fig. 6.67 Rheumatobates prostatus male: tubercle on the fourth abdominal segment (left) and the abdomen in ventral view (right). Based on Polhemus and Manzano (1992).

Fig. 6.68 Rheumatobates spinosus: remains of a type specimen in dorsal view (below) and enlarged antenna (above). Based on Hungerford (1954).

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12. (11) The fourth through seventh abdominal segments bear long setae. The fourth segment has a median tubercle directed toward the anterior margin, and the second through sixth segments converge in the middle from this tubercle (Fig. 6.67). ..............................................................Rheumatobates prostatus Polhemus, 1975 (Costa Rica, Panama). - There are long setae on all abdominal segments, or only on the fourth. There is no central tubercle on the fourth abdominal segment (Fig. 6.69). .......13

Fig. 6.69 Rheumatobates carvalhoi male: habitus in dorsal view (left) and a ventral view of the abdomen showing the location of the long setae (right). Based on Hungerford (1954).

Fig. 6.70 Rheumatobates longisetosus male (left to right): antenna, fore-leg, and posterolateral view of the abdomen. Based on Polhemus and Manzano (1992). 13. (12) Only the fourth abdominal segment bears long setae (Fig. 6.70). Total length: 2 to 3 mm. Only the apterous form has been described. ..................................Rheumatobates longisetosus Polhemus and Manzano, 1992 (Colombia).

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- The ventral part of the abdomen is concave with very long, curved hairs arising from the edges of the segments (Fig. 6.69). The hind trochanter is smaller than the hind coxa. There is a large, prominent tubercle slightly beyond the base of the fore-femur. Length of apterous form: 2.9 to 3.0 mm. ..................................................Rheumatobates carvalhoi Drake and Harris, 1944 (Colombia, Ecuador, Minas Gerais?). 14. (11) The hind trochanter has a very long basal spur (Fig. 6.68). .................15 - The hind trochanter is not armed with a long spur (Fig. 6.71). .....................16

Fig. 6.71 Habitus of Rheumatobates crassifemur crassifemur. Based on Drake and Hottes (1951b). 15. (14) Length of apterous male: 1.7 to 1.8 mm; macropterous specimens: c. 3.25 mm. The hemielytra are about 2.25 mm long, much longer than the abdomen. Color of apterous specimens: mainly black with a pale spot on the pronotum, reddish eyes, and a frosted dorsal surface of the abdomen (Fig. 6.68). Color of hemielytra: pale brownish fuscous with pale and prominent sutures and dark fuscous veins. The wings are fumose and opaque. ............................................................Rheumatobates spinosus Hungerford, 1954 (Bolivia, Amazonas, Pará). - Length of apterous male: c. 1.55 mm. Color: velvety brown with a large testaceous area in the middle of the pronotum. The eyes are large and fuscous brown. The ventral surface is dark brown with testaceous areas on the prosternum, acetabulum, and last two abdominal segments. The two nominal species in this couplet may very well belong to a single species. Better descriptions are urgently needed. ..............................................Rheumatobates hamatus Drake and Chapman, 1954 (Pará).

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16. (14) The distal end of the hind trochanter joins the basal end of the femur. The femur has a T-shaped or anchor-like structure on its inner surface. The middle tibia is sinuate (Fig. 6.72). ......................................................................17 - The hind trochanter is inserted beyond the base of the femur. The femur lacks a T-shaped or anchor-like structure on its inner surface (Fig. 6.71). .......18

Fig. 6.72 Rheumatobates bergrothi male: habitus in dorsal view (center), outside view of the left antenna (upper left), and the right hind leg (below). Based on Hungerford (1954). 17. (16) The hind trochanter is at least 0.8 of the femur length. The middle tibia is sinuate (Fig. 6.72). Length of apterous male: 2.3 to 3.0 mm. Length of female: 2.6 to 3.5 mm. Length of winged female: 3.36 mm. ................................................................Rheumatobates bergrothi Meinert, 1895 (Central America, West Indies, Colombia, Venezuela). - The hind trochanter is slightly less than half the length of the femur (Fig. 6.34). Length of apterous male: 2.0 to 2.6 mm, apterous female: 2.5 to 3.0 mm. ....................................................................Rheumatobates imitator (Uhler, 1894) (West Indies, Venezuela, Colombia). Syn: Hymenobator imitator Uhler, 1894. 18. (16) There is a dense tuft of bristles extending forward from the base extremity of the hind femur (Fig. 6.11). There is also a tuft of bristles on the trochanter. The first segment of the male antenna is swollen, and the second and third are subequal in length. Length of apterous male: 2.2 to 2.6 mm;

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macropterous male: c. 3.6 mm. Length of apterous female: 2.7 to 3.0 mm; macropterous female: 3.3 to 3.4 mm. The color is brownish to black, and there is a fulvous brown, U-shaped band on the posterior part of the head. The bases of the legs and genital segments are often yellow. There is more than a usual discrepancy in the sizes given by different authors, so the size ranges given for this species are subject to doubt, and other features should be given more weight for the identification of specimens. ..................................................................Rheumatobates klagei Schroeder, 1931 (Peru, Amazonas, Pará). - The basal extremity of the hind femur is bare (Fig. 6.71). Length of apterous male: 2.0 to 2.4 mm; apterous female: 2.5 to 3.2 mm; winged female: 3.2 to 4.0 mm. ................................................................Rheumatobates crassifemur Esaki, 1926 (Guyana to Argentina). Three subspecies have been described; the males can be distinguished using the following supplemental key. Supplemental key to the subspecies of male Rheumatobates crassifemur in South America Information for the key was provided by Hungerford (1954). taxonomic information above.

See range and

1. The terminal segment of the antenna has some very long, stout setae forming a dorsolateral, marginal row, which extend as a small group far beyond the other setae (Fig. 6.73). Length of apterous male: 2.2 to 2.4 mm; macropterous male: 2.4 to 2.5 mm; apterous female: 2.6 to 2.8 mm; macropterous female: 2.8 to 3.0 mm. ...............................................Rheumatobates crassifemur esakii Schroeder, 1931 (West Indies, Ecuador, Colombia, Guyana, French Guiana, Surinam, Peru, Amazonas, Pará). - The terminal segment of the antenna has a fairly uniform row of strong setae on the dorsolateral margin, but none are remarkably longer than the others (Fig. 6.74). ....................................................................................................................2 2. (1) The basal lobe of the hind trochanter is densely margined ventrally with long hair (Fig. 6.71). .............................................Rheumatobates crassifemur crassifemur Esaki, 1926 (Central America, Colombia, Bolivia, Paraguay, Uruguay, Argentina, Rio de Janeiro, São Paulo, Minas Gerais). - The basal lobe of the hind trochanter is not densely margined ventrally with long hair (Fig. 6.74). Length of apterous male: 2.1 to 2.3 mm; apterous female: 2.5 to 2.8 mm; macropterous female: c. 2.7 mm. .....................................Rheumatobates crassifemur schroederi Hungerford, 1954 (Ceará, Rio Grande do Norte, Pernambuco, Pará, Amazonas, Goiás, Mato Grosso).

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Fig. 6.73 Habitus of a male Rheumatobates crassifemur esakii. Hungerford (1954).

Based on

Fig. 6.74 Habitus of a male Rheumatobates crassifemur schroederi. Hungerford (1954).

Based on

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Subfamily Trepobatinae Key to the adult males of Metrobates in South America Information for the key was provided by Drake and Harris (1933a, 1945a), Hungerford (1951), Drake and Roze (1954), Nieser (1993a), Polhemus and Polhemus (1993a), Mazzucconi and Bachmann (1995b), and Posso and González (2008). Apterous and winged forms occur in South America. 1. There is a ventral spine on the fore-femur (Fig. 6.75). ....................................2 - The fore-femur lacks a ventral spine (Fig. 6.76). ............................................4

Fig. 6.75 Paramere of a male Metrobates laetus. Based on Nieser (1993a).

Fig. 6.76 Outline of the right paramere of a male Metrobates curracis. Based on Drake and Roze (1954). 2. (1) There is a spine on the ventral side of the base of the acetabulum of the middle leg. The parameres do not become widest at the apex. .........................................................Metrobates fugientis Drake and Harris, 1945 (Peru, Bolivia). - The base of the middle acetabulum lacks a spine. ...........................................3 3. (2) There is a ventral spine on the acetabulum of the hind leg. The paramere is narrow and ends in an acute point (Fig. 6.75). ...............................................................................Metrobates laetus Drake, 1954 (Brazil). - There is no ventral spine on the acetabulum of the hind leg. The paramere is relatively wide and has three short processes near the apex (Fig. 6.77). Length: c. 4 mm. Macropterous specimens have not been described. ...............................................................................Metrobates vigilis Drake, 1958 (Santa Catarina, Minas Gerais).

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Fig. 6.77 Paramere of a male Metrobates vigilis. Based on Nieser (1993a). 4. (1) The combined length of the second and third antennal segments is longer than the first antennal segment. The paramere is funnel-shaped and widest at the apex (Fig. 6.76). Length: 3.2 to 3.7 mm. Color: blackish with a bluish pruinose coat on the median part of the pronotum, most of the metanotum, and all of the ventral part of the body, and there is a wide bluish pruinose median stripe and a narrower lateral stripe on each side on the mesonotum. ............................................................Metrobates curracis Drake and Roze, 1954 (Venezuela). - The first antennal segment is longer than the second and third combined. The parameres do not become widest at the apex (Fig. 6.42). .............................5

Fig. 6.78 Metrobates plaumanni male: tarsus of the middle leg (above) and apex of the abdomen in ventral view (below). Based on Matsuda (1960). 5. (4) The fourth antennal segment is longer than the third. There is a midventral spine at the base of the middle acetabulum (Fig. 6.42). Length of male: 3.3 to 3.6 mm; female: 3.6 to 3.9 mm. The color is mainly blackish with recumbent yellow pilosity of the body, a brown V-shaped mark between the compound eyes, and patches and bands of bluish gray pilosity on the thorax and abdomen. ......................................................................Metrobates amblydonti Nieser, 1993 (Trinidad, Colombia).

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- The fourth segment of the antenna is shorter than the third. The parameres of the male narrow to an acutely pointed apex (Fig. 6.78). ...............................................................Metrobates plaumanni Hungerford, 1951 (Argentina, Uruguay, Santa Catarina, Minas Gerais). Nieser (1993a) recognized two sub-species of Metrobates plaumanni: M. p. plaumanni Hungerford, 1951, from Argentina, Uruguay, and Santa Catarina, characterized by a curve in the paramere just proximal to a distinct widening, and M. p. genikos Nieser, 1993, from Minas Gerais, which lacks a widening of the paramere distal to the curve (Fig. 6.78). Key to the species of adult Telmatometra in South America Information for the key is from Kenaga (1941) and Polhemus (1992). 1. On the mesonotum, narrow black stripes end before reaching the posterior margin. The clasper (paramere) is stout and nearly straight for most of its length but curves dorsad about 90º near its apex (Fig. 6.45). Length of apterous male: 4.6 to 4.8 mm; macropterous male: c. 5.0 mm; apterous female: 5.0 to 5.5 mm; macropterous female: 5.0 to 5.6 mm. Color: dark yellow with black and white markings. .......................................................................Telmatometra whitei Bergroth, 1908 (Mexico, Central America, Colombia). - All black stripes on the mesonotum, if any, reach the posterior margin (Fig. 6.79). .....................................................................................................................2

Fig. 6.79 Telmatometra panamensis: habitus in dorsal view without appendages (upper left), male clasper (lower center), and apex of the abdomen (lower right). Based on an illustration labeled Telmatometra acuta by Kenaga (1941).

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Fig. 6.80 Telmatometra indentata (left to right): habitus in dorsal view without appendages, male clasper, and apex of the abdomen. Based on Kenaga (1941). 2. (1) The anterior margin of the mesonotum is black (Fig. 6.79). ......................3 - The anterior margin of the mesonotum is light colored (Fig. 6.80). ................4 3. (2) The posterior femur is slightly more than twice the length of the tibia (Fig. 6.79). Length of the apterous form, males: 3.6 to 3.8 mm; females: 4.1 to 4.6 mm. The macropterous form has not been described. ................................................Telmatometra panamensis Drake and Harris, 1941 (Central America, Colombia, Peru). Syn: Telmatometra acuta Kenaga, 1941. - The posterior femur is less than twice the length of the tibia (Fig. 6.81). Length of apterous form, males: 4.0 mm; females: 4.5 to 4.7 mm. Length of macropterous form disregarding the wings, males: 4.8 to 5.1 mm; females: 5.9 to 6.1 mm. ..........................................................................Telmatometra ujhelyii Esaki, 1926 (Central America, Colombia).

Fig. 6.81 Telmatometra ujhelyii: habitus in dorsal view without appendages (upper left), male clasper (lower left), and apex of the abdomen (right). Based on Kenaga (1941).

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4. (2) The mesonotum has two black longitudinal stripes on the sides with prominent silvery sheen or a white stripe between them (Fig. 6.82). Length of the apterous form, males: 3.5 to 3.9 mm; females: 3.9 to 4.6 mm. The macropterous form has not been described. ........................................................................Telmatometra retusa Kenaga, 1941 (Bolivia). - There is no silvery sheen or white stripe separating the two black stripes on the sides of the mesonotum (Fig. 6.80). ...............................................................5

Fig. 6.82 Telmatometra retusa male: habitus in dorsal view without appendages (left) and male clasper (right). Based on Kenaga (1941).

Fig. 6.83 Telmatometra parva (left to right): habitus in dorsal view without appendages, male clasper, and apex of the abdomen. Based on Kenaga (1941). 5. (4) The first genital segment of the male is emarginated ventrally and reaches the posterior margin of the last abdominal segment, making it acutely V-shaped. The markings on the head are very dark and distinct (Fig. 6.80). Length of the apterous form, males: 3.3 to 3.5 mm; females: 3.6 to 4.1 mm. The macropterous form has not been described. ...................................................................Telmatometra indentata Kenaga, 1941 (Bolivia).

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- The first genital segment of the male is emarginate, but it does not nearly reach the posterior margin of the last abdominal segment, making it broadly Vshaped. The markings on the head are light and indistinct (Fig. 6.83). ............6 6. (5) There is a median dark line on the posterior half of the mesonotum. The anterior femur is black ventrally. The male has a median longitudinal groove on the ventral side of the first genital segment (Fig. 6.83). Length of the apterous form, males: 3.2 to 3.4 mm; females: 3.7 to 3.8 mm. The macropterous form has not been described. .........................................................................Telmatometra parva Kenaga, 1941 (Amazonas). - There is no median dark line on the posterior half of the mesonotum. The anterior femur is dark brown ventrally. The male has a median saucer-shaped depression on the ventral side of the first genital segment (Fig. 6.84). Length of apterous male: 3.2 to 3.7 mm; females: 3.9 to 4.2 mm. Length of macropterous females: c 4.0.mm. Color: reddish brown dorsally with black margins to the pronotum and mesonotum and black stripes on the mesopleurae. ..........................................................................Telmatometra fusca Kenaga, 1941 (Surinam, Amazonas).

Fig. 6.84 Telmatometra fusca male: habitus in dorsal view without appendages (left) and male clasper (right). Based on Kenaga (1941). Key to the species of adult Ovatametra found in South America Information for the key was provided by Drake and Harris (1935), Kenaga (1942), and Drake (1958b). 1. The apterous male is velvety black with a yellowish marking running along the anterior margin of the head and curving posteriad about midway between the central dark marking and compound eyes. There are two pairs of pale stripes on the dorsal surface of the thorax, which has an orange tinge, and silvery pleurae (Fig. 6.28). The margins of the three or four posteriormost abdominal tergites are yellowish brown. Length of male: c. 2.5 mm; female: c. 2.85 mm. ........................................................................Ovatametra thaumana Drake, 1959 (Paraná).

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- The color on the thorax is predominantly yellow or chestnut brown with dark markings, or there are multiple dark markings on a lighter background, or one or more of the posteriormost abdominal segments is almost entirely pale (Fig. 6.85). .....................................................................................................................2 2. (1) The length is less than 2.5 mm. ..................................................................3 - The length is 2.5 mm or more. .........................................................................4

Fig. 6.85 Dorsal surface of the mesothorax of Ovatametra minima. Kenaga (1942).

Based on

Fig. 6.86 Dorsal surface of the mesothorax of Ovatametra parvula. Kenaga (1942).

Based on

Fig. 6.87 Dorsal surface of the mesothorax of Ovatametra obesa. Kenaga (1942).

Based on

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3. (2) The dorsal part of the mesothorax has dark longitudinal bands on a chestnut brown ground color. The female abdomen lacks dark markings (Fig. 6.85). ..........................................................................Ovatametra minima Kenaga, 1942 (Peru, Rondônia). - The dorsal part of the mesothorax has dark longitudinal bands on a yellow ground color. The female has dark markings on the ventral part of the abdomen (Fig. 6.86). .......................................................Ovatametra parvula (Drake and Harris, 1935) (Amazonas). 4. (2) The pronotum of apterous males and females is brownish ochraceous with a black basal spot and a broad fuscous black inverted U-shaped marking, which fails to reach the base of the pronotum. The mesonotum is brownish ochraceous and depressed along the midline with the following fuscous black markings: a transverse, median, anterior spot; a triangular median spot in the middle; lateral stripes, which are widely interrupted in the male but uninterrupted in the female. On the outer face of the fore-femur, there is a longitudinal black stripe. The fore-tibia of the male has an ovate patch of hairlike setae. Length: c. 2.7 mm. Only apterous specimens have been found. The species was poorly described and not illustrated, so its identification should be considered tentative without access to type specimens. ................................................................................Ovatametra bella Drake, 1957 (Bolivia). - The pronotum is not brownish ochraceous and marked with a black basal spot, U-shaped marking and lateral stripes, or, there is no longitudinal black stripe on the fore-femur or ovate patch of hair-like setae on the fore-tibia of the male (Fig. 6.87). .....................................................................................................................5 5. (4) The posteriormost abdominal tergite has a tuft of dark hair-like setae on the median posterior margin. Dark median marks of the mesonotum are continuous from the anterior to the posterior margins. Most dark longitudinal bands on the dorsum of the thorax are connected by a transverse stripe along the posterior margin of the mesonotum (Fig. 6.87). Length of male: 1.7 to 2.2 mm; female: 2.2 to 2.6 mm. Color: reddish yellow and black, with a variable pattern. .............................................................................Ovatametra obesa Kanega, 1942 (Amazonas, Pará). - The posteriormost abdominal tergite lacks a tuft of dark hairs on the median posterior margin. Dark median marks on the mesonotum are not continuous from the anterior to the posterior margins. Most dark longitudinal bands on the dorsum are not connected by a transverse stripe along the posterior margin (Fig. 6.88). .....................................................................................................................6 6. (5) The mesonotum of apterous individuals has a thick, black median stripe, a pair of thick, black dorsolateral stripes, and a pair of thin, black lateral stripes, all of which end shortly before reaching the posterior margin of the mesonotum (Fig. 6.88). The contrasting color is bright yellow. The pronotum of the winged form is large and darkly colored with a pair of short, bright yellow

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stripes bordering the dorsal midline just posterior to the anterior border. Length of apterous and winged forms: 2.7 to 3.0 mm. ................................................................Ovatametra gualeguay Bachmann, 1966 (Argentina, São Paulo, Minas Gerais). - The mesonotum of apterous individuals has short, dark markings only along the anterior and posterior margins, leaving the central area uniformly light in color (Fig. 6.89). Only apterous individuals have been described. .............................................................................Ovatametra fusca Kanega, 1942 (Guyana, Amazonas).

Fig. 6.88 Habitus of an apterous (left) and a winged female (right) Ovatametra gualeguay without appendages in dorsal view. The light color is bright yellow, and the dark color is black. Based on Bachmann (1966b).

Fig. 6.89 Dorsal surface of the mesothorax of Ovatametra fusca. Based on Kenaga (1942).

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Key to the species of adult Trepobates found in South America Information for the key was provided by Drake and Harris (1932b) and García (2002). 1. The first genital segment of the male is covered by long hair-like setae. The apical abdominal segment of the female has a ventral prolongation that partially conceals the genital segments, which also bear long, hair-like setae, which are coarse and dark along the posterior margin, best seen in lateral view (Fig. 6.36). Total length: 3.0 to 4.5 mm. ...........................................................Trepobates trepidus Drake and Harris, 1928 (North and Central America, Venezuela, Colombia). - The first genital segment of the male lacks long hair-like setae, although it may be covered by a short pubescence (Fig. 6.90). The apical abdominal segment of the female lacks long, hair-like setae or the fringe may consist of very delicate setae. ................................................................................................2

Fig. 6.90 Trepobates taylori male: apex of the abdomen in ventral (left) and right lateral view (right). Based on García (2002). 2. (1) The parameres are usually short, narrow, and curved. The fore-femur of the male has a ring-like constriction near the apex. The third antennal segment of the female is distinctly longer than the second. The seventh abdominal segment of the female is not elongated ventrally and does not partially cover the genital segments (Fig. 6.90). Total length of apterous male: c. 3.5 mm; macropterous male: 3.2 to 3.7 mm; macropterous female: 3.8 to 3.9 mm. Color: black and yellow with a highly variable pattern. ........................................................................Trepobates taylori (Kirkaldy, 1899) (North and Central America, West Indies, Venezuela, Colombia, Peru, Ecuador, Argentina, Pará, Amazonas). Syn: Kallistometra taylori Kirkaldy, 1899; Trepobates comitialis Drake and Harris, 1928; Trepobates commitialis Hynes, 1948.

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- The parameres are usually wide and curve abruptly before the apex, sometimes far enough to form a right angle. The fore-femur of the male is constricted apically. The seventh abdominal segment of the female has short and delicate setae, which are sometimes difficult to see. Total length of apterous male: 3 to 4 mm. ....................................................Trepobates panamensis Drake and Hottes, 1952 (Mexico, Central America, Colombia). Key to the adult male Halobatopsis in South America Information for the key was provided by Drake and Harris (1938), Bachmann (1962d), Nieser and Melo (1999a), and García (2002). 1. There are no black longitudinal markings on the mesonotum (Fig. 6.91). Length of male: 3.4 to 3.6 mm; female: 4.0 to 4.2 mm. Color: mainly golden brown with dark brown or black markings. .............................................Halobatopsis chrysocastanis Nieser and Melo, 1999 (Minas Gerais). - There are black longitudinal markings on the mesonotum (Fig. 6.92). ...........2

Fig. 6.91 Halobatopsis chrysocastanis: dorsal view of the habitus of a female (left) and the male paramere (right). Based on Nieser and Melo (1999a). 2. (1) The first genital segment of the male lacks a ventral spine. The connexiva of the female is oriented horizontally or slanted only slightly upwards (Fig. 6.92). Total length of apterous specimens: 3.5 to 4.5 mm. Color:

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mainly ochraceous with dark markings, a reddish and black mesonotum, and black on the anterior part of the abdomen. Macropterous forms have not been found. .......................................................................Halobatopsis platensis (Berg, 1879) (Argentina, Uruguay, Mato Grosso, Goiás, Distrito Federal, Minas Gerais, Rio de Janeiro, Rio Grande do Sul). Syn: Halobates platensis Berg, 1879. - There is a black or dark brown ventral spine on the first genital segment of the male (Fig. 6.93). The connexiva of the female is oriented vertically or bent over the abdomen. ................................................................................................3

Fig. 6.92 Habitus of Halobatopsis platensis. (1938).

Based on Drake and Harris

Fig. 6.93 Halobatopsis delectus: habitus of a female in dorsal view without appendages (upper left), the color pattern on the head and pronotum of a macropterous specimen (lower left), fore (upper right center) and hind wing (lower right center), apex of a male abdomen in lateral view (lower left center), and the trochanter and fore-femur of a male (right). Based on Nieser and Melo (1999a).

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3. (2) There is no tubercle on the basal ¼ of the male fore-femur (Fig. 6.47). ..................................................Halobatopsis spiniventris Drake and Harris, 1936 (Uruguay, Argentina, Santa Catarina). - There is a tubercle on the basal ¼ of the male fore-femur (Fig. 6.93). Macropterous forms occur. .......................................................Halobatopsis delectus Drake and Harris, 1941 (Minas Gerais) Subfamily Halobatinae Key to the species of adult male Halobates found in South America Information for the key provided by Drake (1957a), Cheng and Schulenberger (1980), Froeschner (1985), and García (2002). Males are recognized by the obviously modified genital structures at the apex of the abdomen. 1. The styliform left genital process is much shorter than the right process and is twisted near the base to project away from the body axis (Fig. 6.94). .................2 - The left and right genital processes are about equal in length, and the left process is nearly straight and not strongly twisted (Fig. 6.95). ............................3

Fig. 6.94 Halobates splendens male: habitus in dorsal view (lower left); head with antenna (lower right); fore-tarsus (middle right); apical segments of the

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male abdomen in dorsal, ventral, and lateral view (above, left to right). Based on Herring (1961) and Aguilar and Mendez (1987).

Fig. 6.95 Halobates sobrinus: habitus of a larva without appendages (left) and a male abdomen in ventral view (right). Based on Matsuda (1960) and García (2002). 2. (1) The right styliform genital process is nearly straight. The styliform left genital process is twisted nearly 90° from the longitudinal axis. The ninth abdominal segment has a lateral process (Fig. 6.96). Total length: 4.0 to 4.5 mm. ......................................................................Halobates micans Eschscholtz, 1822 (Circumtropical oceanic species, reported from the coast of Colombia). Syn: Halobates Wuellersdorffi Uhler, 1890. - The right genital process curves strongly inward. The narrow left genital process is twisted only about 45° from the longitudinal axis. The ninth abdominal segment does not have a lateral process (Fig. 6.94). Total length: c. 5 mm. ......................................................................Halobates splendens Witlaczil, 1886 (Eastern Pacific Ocean, reaching the coasts of Peru, Colombia, and Ecuador, including the Galapagos Islands).

Fig. 6.96 Halobates micans male (left to right): apical segments of the male abdomen in dorsal, ventral, and lateral view. Based on Herring (1961).

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3. (1) The styliform genital processes are asymmetrical with the left process somewhat twisted near the base so that it projects away from the body; the right process extends nearly straight posteriad. The ninth abdominal segment has well developed lateral processes (Fig. 6.95). This species is less frequently encountered in the pelagic zone than the other species of the genus. Total length: c. 4.1 to 4.4 mm. .............................................................................Halobates sobrinus White, 1883 (Eastern Pacific Ocean from California to Colombia, Peru, and the Galapagos Islands). - The narrow genital processes are symmetrical or almost so, and neither is twisted to point away from the body. The ninth abdominal segment lacks lateral processes (Fig. 6.97). Length of male: c. 3.5 mm. .................................4 4. (3) The styliform genital processes are about 2/3 as long as the subgenital plate, so no part of them is visible beyond the edges of that plate (Fig. 6.97). ............................................................................Halobates robustus Barber, 1925 (Pacific Ocean, known only near the Galapagos Islands). - The styliform genital processes are subequal in length to the genital capsule so that their apices extend slightly beyond the lateral angle of the dorsal genital plate (Fig. 6.38). ....................................................................Halobates sericeus Eschscholtz, 1822 (Tropical and subtropical Pacific Ocean, mainly along the coasts of Peru, Ecuador, and Colombia).

Fig. 6.97 Halobates robustus male (above, left to right): apical segments of the male abdomen in dorsal, ventral, and lateral view, and (below) habitus. Based on Herring (1961) and Froeschner (1985).

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Key to the species of adult female Halobates found in South America Information for the key was provided by Cheng and Schulenberger (1980), Froeschner (1985), and García (2002). Females are recognized by the simple, unmodified apex of the abdomen. 1. In posterior view, short, suberect, black bristles can be seen on the mesonotum and metanotum (Fig. 6.97). Length of female: c. 4.2 mm. ............................................................................Halobates robustus Barber, 1925 (Pacific Ocean, found only near the Galapagos Islands). - There are no short, suberect black bristles on the mesonotum or metanotum (Fig. 6.95). ............................................................................................................2 2. (1) The two fore-tarsal segments are approximately the same length. The basal tarsal segment of the middle legs is approximately six times longer than the second segment (Fig. 6.94). Length of female: less than 4.5 mm. .......................................................................Halobates splendens Witlaczi, 1886 (Eastern Pacific Ocean, including the coasts of Peru, Ecuador, and Colombia). - The first segment of the fore-tarsus is clearly shorter than the second and about half as long as the fourth (Fig. 6.38). Length: c. 3.4 mm. .......................3 3. (2) The second antennal segment is very slightly longer than the third and approximately half as long as the fourth. The first tarsal segment of the foreleg is half the length of the second segment (Fig. 6.38). ....................................................................Halobates sericeus Eschscholtz, 1822 (Subtropical Pacific Ocean, mainly along the coasts of Peru, Ecuador, and Colombia). - The second antennal segment is 1 1/3 the length of the third and 2/3 as long as the fourth. Total length of female: c. 4.0 mm. ...................................................4 4. (3) With the legs extended posteriad, the hind femur is 3/4 as long as the middle femur. Color: gray. ......................................................................Halobates micans Eschscholtz, 1822 (Circumtropical oceanic species). - The hind femur is almost 8/9 as long as the middle femur. Color: nearly uniform blackish red (Fig. 6.95). .............................................................................Halobates sobrinus White, 1883 (Eastern Pacific Ocean from California to Peru, including the Galapagos Islands).

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Subfamily Cylindrostethinae Key to the species of adult Cylindrostethus in South America Information for the key was provided by Drake and Harris (1934), Kuitert (1942), Drake (1952a, 1954, 1958a), and Matsuda (1960). 1. The mesonotum is uniform brownish or brownish black, longitudinally impressed along the sides. The sixth sternite of the male is doubly emarginated and has a deep notch in the middle (Fig. 6.98). ....................................................2 - The mesonotum is brownish with conspicuous fuscous or black longitudinal stripes or black with longitudinal brown or fuscous stripes. The sixth sternite of the male has a broad, simple emargination (Fig. 6.99). ...................................5

Fig. 6.98 Cylindrostethus linearis: apical segments of the male abdomen in ventral view (left) and of a female abdomen in right lateral view (right). Based on Matsuda (1960).

Fig. 6.99 Cylindrostethus hungerfordi: apical segments of the male abdomen in ventral view. Based on Matsuda (1960). 2. (1) The ventral surfaces of the fifth and sixth abdominal segments of the male show no evidence of median impressions. The dorsum or posterior border of the mesothorax of the female has two median lobes directed dorsoposteriad (Fig. 6.100). The relative lengths of the male antennal segments from the first to the

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fourth, respectively, are 122:52:33:64; relative lengths of female antennal segments: 151:55:33:66. Length of male: 14 to 15 mm; female: 16 to 17 mm. Maximum width: 2.0 to 2.5 mm. ..................................................................Cylindrostethus bilobatus Kuitert, 1942 (Bolivia, Brazil). - The ventral surface of the sixth or of both fifth and sixth abdominal segments of the male display distinct impressions near the middle. The dorsum and posterior border of the female mesothorax lack median lobes (Fig. 6.101). ........3

Fig. 6.100 Cylindrostethus bilobatus: apical segments of the male abdomen in ventral view. Based on Matsuda (1960).

Fig. 6.101 Cylindrostethus erythropus male: body without appendages in dorsal view (left) and apex of the abdomen in ventral view (right). Based on Matsuda (1960). 3. (2) The mesonotum is fuscous or black. There are medial impressions on the ventral surfaces of both the fifth and sixth abdominal segments. The lateral spines of the connexiva are very stout, and the abdomen is widened toward the apex. The first genital segment of the female is as long as broad above, narrowed distally, and broadly rounded at the apex (Fig. 6.101). Length of apterous male: 13.9 to 15.0 mm; apterous female: 15.2 to 17.6 mm. The general color is grayish with yellowish brown connexiva and legs. ...........................................Cylindrostethus erythropus (Herrich-Schaeffer, 1850) (Colombia, Ecuador, Peru, Pará, Amazonas, Rondônia). Syn: Hydrometra erythropus Herrich-Schaeffer, 1850. - The mesonotum is uniformly brownish, testaceous, or ferrugineous (Fig. 6.98). .....................................................................................................................4

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4. (3) The general color is dark fuscous or black. The fourth segment of the antenna is 1.9 to 2.0 times as long as the third (Fig. 6.98). The relative lengths of the first through fourth antennal segments are roughly 16:7:5:10, respectively. Length of apterous male: 12.8 to 16.2 mm; apterous female: 15.4 to 16.3 mm. ...............................................................Cylindrostethus linearis (Erichson, 1848) (Guyana, Peru, Bolivia, Pará, Amazonas, Rondônia). - The general color is dark ferrugineous brown (Fig. 6.102). The fourth antennal segment is about 2.3 times as long as the third. The relative lengths of the antennal segments from the first to the fourth, respectively, are 41:15:10:23. There is no midline tooth at the base of the U-shaped posterior emargination of the last abdominal segment. Length of apterous specimens: 16 to 17 mm. .....................................................................Cylindrostethus bassleri Drake, 1952 (Peru, Amazonas).

Fig. 6.102 Cylindrostethus bassleri: apical segments of the male abdomen in ventral view. Based on Matsuda (1960). 5. (1) The color is mainly black, but the body surface is clothed in a short grayish or bluish white pubescence. There is a pair of very narrow longitudinal stripes on the dorsal surface of the thorax or abdomen. .......................................6 - The color is mainly fuscous brownish with fuscous or black stripes. ..............7 6. (5) The body surface is clothed in a short grayish pubescence. There is a pair of very narrow longitudinal stripes on the dorsal surface of the abdomen. The antennae and legs are brownish fuscous with the antennae somewhat darker than the legs. Macropterous specimens have not been reported. The connexiva are fairly wide and nearly erect, and each ends in a spine extending posteriad as far as the middle of the first genital segment of the male and slightly farther in the female. Length: c. 18 mm. Width at the base of the metanotum: c. 2 mm. Lengths of the four segments of the antenna of the type specimen: 2.6, 0.8, 0.5, and 1.28 mm, respectively. .......................................................................Cylindrostethus stygius Drake, 1961 (Peru). The two species in this couplet were not illustrated and only briefly described. Their apparent similarity warrants a thorough examination of the specimens to assure that they are distinct species. - The body is black with recumbent grayish white pubescence dorsally and bluish pubescence ventrally. There is a chocolate brown stripe on each side of the mesonotum and the metanotum. The exterior edges of the connexiva are

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also chocolate brown. The fore-legs are yellowish brown dorsally and mainly fuscous ventrally. The middle and hind legs are yellowish brown with the apices of the segments dark testaceous. Length: 18.5 to 19.5 mm. Width: 2.1 mm. Lengths of the four segments of the antenna of the type specimen: 2.5, 0.8, 0.25, and 1.30 mm, respectively. ...................................................................Cylindrostethus podargus Drake, 1958 (Peru). 7. (5) The length of the first antennal segment is subequal to the width of the head through the eyes (Fig. 6.99). The fore-femora have many black setae near the base of the dorsal side and dark, bristly hairs on the ventral surface. The fore-tibiae and tarsi are blackish. Length: 17 to 22 mm. The general color is shiny light brown with some blackish markings. .............................................Cylindrostethus hungerfordi Drake and Harris, 1934 (Guyana, Surinam, Bolivia, Pará). - The length of the first antennal segment is obviously greater than the width of the head through the eyes (Fig. 6.103). ................................................................8

Fig. 6.103 Cylindrostethus regulus: apical segments of the male abdomen in ventral view. Based on Matsuda (1960). 8. (7) The first two segments of the antenna are black. The convexial spines of the male reach to the middle of the first genital segment. The female has long, reflexed convexial spines, which are contiguous dorsal to the genital segments. The first genital segment is longer than broad dorsally and ends in an acute point (Fig. 6.39). Length of apterous specimens: 16 to 20 mm. ..................................................Cylindrostethus palmaris Drake and Harris, 1934 (Trinidad, Guyana, French Guiana, Surinam, Venezuela, Colombia, Ecuador, Bolivia, Argentina, Rondônia, Amazonas, Pará, Minas Gerais, Mato Grosso). Syn: Cylindrostethus linearis Drake and Harris, 1930 (nec Erichson). - The first two segments of the antenna are rather pale, mainly testaceous. The convexial spines of the males do not reach the middle of the first genital segment. The female does not have long, reflexed convexial spines that are contiguous dorsal to the genital segments (Fig. 6.103). Length: 18.5 to 20 mm. ...................................................................Cylindrostethus regulus (White, 1879) (Amazonas). Syn: Halobates regulus White, 1879.

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Key to the adults of Potamobates in South America Information for the key was provided by Drake and Harris (1934), Polhemus and Polhemus (1995), Cognato (1998), García (2002), and Buzzetti (2006). 1. In the middle of the pronotum, there are only narrow yellow longitudinal lines on a dark background. ..........................................................................................2 - In the middle of the pronotum, there is a yellow triangular marking. The eighth abdominal segment of the male is asymmetrical, and the ninth and tenth segments are rotated between about 10° and 90° (Fig. 6.104). ............................3

Fig. 6.104 Potamobates sumaco: habitus of three males with varying color patterns in dorsal view without appendages (above, left and right, and below left), apical segments of the male abdomen in oblique (center) and lateral view (middle right), and the apex of a female abdomen in dorsal view (lower right center). Based on Cognato (1998).

Fig. 6.105 The abdomen of a male Potamobates bidentatus in dorsal view. Based on Polhemus and Polhemus (1995). 2. (1) The mesonotum is unmarked except for lines or blotches formed by a gold or silver pubescence. The vertex is black with a yellow fleck between the compound eyes. The apical segments of the male abdomen are elongate, and there is a lobe bearing two blunt teeth on the eighth abdominal segment (Fig.

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6.105). The eighth abdominal segment of the female bears a fairly long dorsal process. Length: c. 11 mm. ...............................................................Potamobates bidentatus Champion, 1898 (Mexico, Colombia, Ecuador). - The mesonotum of the apterous adult has a median yellow line, or rarely, it is entirely dark. The eighth abdominal segment of the male is elongate, not expanded at the apex, and has only one small tooth on the apical margin. The genital segments of the male bear two teeth. In dorsal view, the eighth segment of the female appears triangular, and ventrally, it displays a short finger-like protuberance. The second segment of the antenna is distinctly longer than the third. There is a short subapical tuft of setae on the ventral side of the first antennal segment (Fig. 6.106). Length: 7.9 to 8.2 mm. ..........................................................................Potamobates horvathi Esaki, 1926 (Central America, Colombia). Two species known only from Mexico also lack a yellow triangular marking on the pronotum. The males of Potamobates thomasi Hungerford, 1937, show no modification at all on the eighth abdominal segment and no rotation of the ninth and tenth segments. Potamobates osborni Drake and Harris, 1928, has no light markings on the mesonotum except for lines or blotches formed by a gold or silver pubescence and has a yellow vertex with a central black marking. 3. (1) The ninth and tenth abdominal segments of the male are rotated a maximum of 45°. Females do not have an extension of the connexiva (Fig. 6.107). ...................................................................................................................4 - The ninth and tenth abdominal segments of the male are rotated about 90°. Females have an extension of the connexiva (Fig. 6.108). ..................................5

Fig. 6.106 Potamobates horvathi (left to right): first segment of the antenna and base of the second in ventral view and ventral views of the apex of a male and a female abdomen. Based on García (2002). 4. (3) The distal part of the ninth abdominal segment of the male is pale in color. The male proctiger is spatulate and usually concealed. The ninth and tenth segments are rotated between 5° and 10° from the longitudinal axis. The

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eighth abdominal segment of the female is triangular in dorsal view and has a triangular structure ventrally (Fig. 6.109). Length: 8 to 10 mm. .............................................................Potamobates unidentatus Champion, 1898 (Central America, Colombia). - The distal part of the ninth abdominal segment of the male is dark. The male proctiger has an acuminate process, which is not concealed. The ninth and tenth segments are usually rotated between 35° and 45° from the longitudinal axis. The eighth abdominal segment of the female is produced to form a large spiniform structure dorsally and has a spoon-like structure ventrally (Fig. 6.107). Total length: 9 to 10.5 mm, with apterous forms usually shorter than 10 mm, and macropterous forms, longer. ......................................Potamobates manzanoae J. T. and D. A. Polhemus, 1995 (Colombia).

Fig. 6.107 Potamobates manzanoae (upper row, left to right): apex of the male abdomen in dorsal, ventral, and lateral view, and (below, left and right): apex of the female abdomen in ventral and lateral view. Based on Polhemus and Polhemus (1995) and García (2002). 5. (3) The eighth abdominal segment of the male is short, narrow, and only slightly asymmetrical. The terminal segments of the female abdomen are truncate (Fig. 6.108). Length: c. 10 to 10.5 mm. Color: black with yellowish markings. .........................................Potamobates carvalhoi J. T. and D. A. Polhemus, 1995 (Colombia, Venezuela). - The eighth abdominal segment of the male is wide and forms two denticulate or spiny lobes on the right side. The eighth abdominal segment of the female has either a short, finger-like process or an extension in the form of a spine, in which case the segment is truncate (Fig. 6.110). ..................................................6 6. (5) The distal lobe on the right side of the eighth abdominal segment of the male forms a single angle and points posteroventrad. The eighth abdominal

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segment of the female is produced to form a robust spine directed posterodorsad. Total length of apterous form: 9 to 11 mm. ...........................................................Potamobates vivatus Drake and Roze, 1954 (Central America, Colombia, Venezuela). - The distal lobe on the right side of the eighth abdominal segment of the male does not form only one angle or is not directed posteroventrally. The process on the eighth abdominal segment of the female is never directed posterodorsad (Fig. 6.110). ..........................................................................................................7

Fig. 6.108 Potamobates carvalhoi: dorsal view of habitus without appendages of a male (upper left) and a female (lower left), apical segments of a male abdomen in lateral (upper right) and oblique ventral view (center), and apical segments of a female abdomen in dorsal (middle right) and ventral view (lower right center). Based on J. and D. Polhemus (1995).

Fig. 6.109 Potamobates unidentatus: apex of the male abdomen in ventral view (left), and apex of the female abdomen in dorsal view (right). Based on García (2002). 7. (6) The distal lobe on the right side of the eighth abdominal segment of the male forms an acute angle directed posteriad and a twisted spatulate lobe directed posteroventrad. The process on the eighth abdominal segment of the female is short and triangular on the dorsal side and has a robust ventral process that extends posteriad much farther than the dorsal triangular plate (Fig. 6.110). .........................................................Potamobates woytkowskyi Hungerford, 1937 (Peru).

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- The distal lobe on the right side of the eighth abdominal segment of the male does not form only one acute angle directed posteroventrad. The ventral process on the eighth abdominal segment of the female does not extend considerably farther posteriad than the dorsal process on the same segment (Fig. 6.111). ...................................................................................................................8 8. (7) The distal lobe on the right side of the eighth abdominal segment of the male forms a short spine directed posteriad and an acute angle directed posteroventrad. The process on the dorsal side of the eighth abdominal segment of the female is robust, finger-like, and directed posteriad, while on the ventral side of the segment, there is a robust process twisted to the left side and extending half as far as the dorsal process (Fig. 6.111). ...........................................Potamobates spiculus J. T. and D. A. Polhemus, 1983 (Peru). - The distal lobe on the right side of the eighth abdominal segment of the male does not form both a short spine and an acute angle directed posteroventrad. The process on the ventral side of the eighth abdominal segment of the female is plate-like and does not extend very far posteriad (Fig. 6.112). ............................9

Fig. 6.110 Potamobates woytkowskyi male (left to right): apical segments of the abdomen in lateral, ventral, and dorsolateral view. Based on Polhemus and Polhemus (1983a).

Fig. 6.111 Potamobates spiculus (above, left to right): apex of the male abdomen in lateral and dorsolateral view, apex of the female abdomen in dorsal view, and (below) apex of the male abdomen in ventral view. Based on Polhemus and Polhemus (1983a).

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9. (8) The distal lobe on the right side of the eighth abdominal segment of the male forms a twisted, spatulate protuberance directed posteriad. The dorsal side of the eighth abdominal segment of the female is triangular, and on the ventral side, there is a flat process with a lobe on the left side (Fig. 6.49). Wingless specimens lack a median brown stripe on the mesonotum. Length of wingless male: c. 10.5 mm; winged male: c. 12 mm; wingless female: c. 9.3 mm; winged female: c. 12 mm. ............................................................Potamobates peruvianus Hungerford, 1936 (Peru, Colombia). - The distal lobe on the right side of the eighth abdominal segment of the male has one or two spines or denticles directed ventrad or posteroventrad. The ventral side of the eighth abdominal segment of the female has a flat process that does not extend far posteriad, if at all (Fig. 6.112). ...........................................10

Fig. 6.112 Potamobates variabilis (left to right): female abdomen in dorsal view, the apex of the male abdomen in ventral view, and the apices of the abdomens of two different male specimens in right lateral view. Based on Hungerford (1938a). 10. (9) The distal lobe on the right side of the eighth abdominal segment of the male has only one spine on the right side. The posterior apices of the female connexiva are drawn out into long processes, which entwine and are in contact dorsally over the two posteriormost abdominal segments, concealing the apical segment from view from above (Fig. 6.113). Length of apterous male: c. 8.1 to 8.2 mm; apterous female: c. 8.5 mm. Winged specimens have not been found. Color: mainly black with creamy white markings. The legs are mainly black and dark brown. ..........................................................................Potamobates shuar Buzzetti, 2006 (Ecuador). - The distal lobe on the right side of the eighth abdominal segment of the male has two spines or denticles directed ventrad or posteroventrad. If the apices of the female connexiva are drawn out, they are not entwined or in contact, and they do not conceal the posteriormost abdominal segment from dorsal view (Fig. 6.112). .................................................................................................................11

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Fig. 6.113 Potamobates shuar male: habitus without appendages in dorsal view (upper left), apex of the abdomen showing the extensions of the connexiva (upper right), and the male genitalia in ventral (lower left) and lateral view (lower right). Based on Buzzetti (2006). 11. (10) The distal lobe on the right side of the eighth abdominal segment of the male has two denticles directed posteroventrad, of which the ventral denticle is longer. The male proctiger has a forked distal process. The eighth abdominal segment of the female has a short and robust dorsal process directed posteroventrad, and on the ventral side, it has a short, flat process covering the left side of the dorsal process (Fig. 6.112). ..............................................................Potamobates variabilis Hungerford, 1938 (Peru). - The distal lobe on the right side of the eighth abdominal segment of the male has two spines of approximately the same length directed ventrad or posteroventrad. The process on the male proctiger is not forked distally. The eighth abdominal segment of the female has a short, wide process dorsally and a short, flat process on the ventral side (Fig. 6.114). ............................................12 12. (11) The distal lobe on the right side of the male eighth abdominal segment has two spines directed ventrad, which are about of equal length and are close to one another. The eighth abdominal segment of the female has a triangular process on the dorsal side and a short, flat ventral process (Fig. 6.114). ...........13 - The distal lobe on the right side of the eighth abdominal segment of the male has two spines directed posteroventrad, which are relatively far from one another and have a process of the proctiger between them that extends slightly farther. The eighth abdominal segment of the female has a short and wide process on the dorsal side and a short, flat ventral process (Fig. 6.115). ...........14

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Fig. 6.114 Potamobates anchicaya (upper row, left to right): color pattern on the dorsal surface of the thorax, the apex of a male abdomen in dorsal and oblique view, and (below left to right): the apex of the male abdomen in oblique view showing the ventrad direction of the spines on the distal lobe located on the right side of the eighth abdominal segment, and the apex of a female abdomen in dorsal and ventral view. Based on Polhemus and Polhemus (1995).

Fig. 6.115 Potamobates williamsi (left to right): apex of the male abdomen in ventral view and apex of the female abdomen in dorsal and lateral view. Based on Polhemus and Polhemus (1995).

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13. (12) The distal lobe on the right side of the eighth abdominal segment of the male has two strong spines directed ventrad and a deep, rounded incision between them. The eighth abdominal segment of the female has a short, wide process that is never doubled over to the ventral side; on the flat ventral process, only the left lobe is developed (Fig. 6.116). Length: 8.5 to 11.0 mm. ......................................................................Potamobates tridentatus Esaki, 1926 (Costa Rica, Panama, Colombia) - The distal lobe on the right side of the eighth abdominal segment of the male has two strong spines directed ventrad and a narrow incision separating them. The eighth abdominal segment of the female has a short, wide process that is doubled over to the ventral side; on the flat ventral process, both the left and right lobes are developed (Fig. 6.114). Total length: c. 9 to 10 mm. .......................................Potamobates anchicaya J. T. and D. A. Polhemus, 1995 (Central America, Colombia).

Fig. 6.116 Potamobates tridentatus: habitus of an apterous male without appendages in dorsal view (upper left), head and thorax of a specimen with a large pronotum in dorsal view (lower left), antenna (upper right center), fore-leg of a male showing the color pattern (upper right), male genital segments in ventral (lower center) and left (lower right) and right lateral view (right of center), and apex of the female abdomen in ventral view (middle right). Based on Esaki (1926b). 14. (12) The distance between the stout, posteroventrally directed spines and the right lobe on the eighth abdominal segment of the male is 0.3 to 0.4 mm. The proctiger has a flat, curved projection. The abdomen of the female has a flat process that touches the short triangular process on the eighth segment. The connexiva of the female ends in long, tapering processes (Fig. 6.115). Total length: 9.0 to 11.9 mm. ...............................................................Potamobates williamsi Hungerford, 1932 (Ecuador, Colombia).

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- The distance between the posteroventral spines and the right lobe on the eighth abdominal segment of the male is 0.4 to 0.5 mm. The proctiger has a projection that is not curved but rotated 90°. The abdomen of the female has a short, flat process that is produced asymmetrically. The left side is extended parallel as far as the middle of the digitate connexival lobe and never touches the short triangular process on the eighth segment (Fig. 6.104). Total length of the apterous form: 10 to 11 mm. ......................................................................Potamobates sumaco Cognato, 1998 (Ecuador, Colombia). Subfamily Charmatometrinae Key to the species of adult Brachymetra found in South America Information for the key was provided by Shaw (1933), Drake and Harris (1942b, 1945b), Harris and Drake (1945), Drake (1957b, 1958b), and Nieser (1975). Some species are too poorly described to distinguish with confidence based on the original descriptions, and a revision of the genus would be helpful. 1. The pronotum of the winged male is very long, about 7/15 of the total body length. It almost almost reaches the base of the abdomen and is obtusely rounded at the apex. The abdomen, including the tergites and connexiva, are rectangular in outline. The fore-femur is much shorter than the other femora. The body of the male is rufous brown dorsally and brownish stramineous ventrally. The paramere is short, flattened, concave on its inner margin, and tapers to an apical point. Length of winged male: c. 7.5 mm. Females and apterous forms of this poorly described species are unknown. ............................................................................Brachymetra accola Drake, 1957 (Found only at an undisclosed location in Brazil). - Both winged and apterous specimens are usually not between 7.0 and 8.0 mm, long, or, if a specimen is in this size range, the pronotum of the male is shorter than 7/15 of the total length (Fig. 6.117), or the color pattern includes blackish markings, or the outline of the abdomen is not quadrangular, or the fore-femur is longer than the middle femur. ...........................................................................2 2. (1) The first antennal segment is as long or longer than the next two together. The male clasper bends sharply just beyond mid-length in one species, and according to the sketchy description of the other, it is similar but shorter (Fig. 6.117). ...................................................................................................................3 - The first antenna segment is shorter than the next two segments together. The pronotum is rather uniform in coloration, except sometimes at the margins (Fig. 6.118). ..........................................................................................................4

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3. (2) Dorsally, the color is reddish brown with black markings, including a black median line on the pronotum (Fig. 6.117). Macropterous and apterous forms are known. Length: 6.3 to 7.0 mm. ................................................................................Brachymetra unca Shaw, 1933 (Trinidad, Surinam, French Guiana, Colombia, Bolivia, Brazil). - The color on the dorsum is brownish with dark fuscous markings around the margins of the pronotum and connexivum, on the ventral surfaces of the forefemora, and on the entire fore-tibiae and tarsi. The ventral surface is mainly yellowish. The first genital segment and claspers are described as being shorter than those of Brachymetra unca, but there are no illustrations, and the general description is too sketchy to permit a reliable identification of this species, if it is valid. Only apterous specimens have been described. Length: c. 8 mm. ........................................................Brachymetra anduzei Drake and Harris, 1942 (Venezuela).

Fig. 6.117 Brachymetra unca male (above, left to right): head and pronotum in dorsal view and apex of the abdomen in dorsal and left lateral view, and (below) an enlarged apex of the abdomen in right lateral view (left) and a clasper (right). Based on Shaw (1933) and García (2002). 4. (2) The length of the brachypterous form is about 8.8 mm or longer, and the winged form is nearly 10 mm in length. The pronotum is broadly rounded at the apex. The general color is brown dorsally and pale ventrally. There are usually silvery spots on the middle and hind acetabula. The ratios of the antennal segment lengths from I to IV are 90:45:68:50. There is a deep, broad emargination in the first abdominal segment. The pronotum is long and deeply impressed on the disc of the anterior lobe; its posterior lobe is about three times as long as the anterior lobe. The length of the middle femur equals the body length, and the hind femur is longer. The legs are dark fuscous but slightly

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paler on the ventral surface and yellow on the coxae and trochanters. The basal segments of the beak are yellowish brown, and the apical segment is blackish and reaches just posterior to the prosternum. The paramere is long and narrow with a curved inner margin. While minor size differences are poor criteria for recognizing rare species, the cursory original description provides no better characters than those clearly shared with other species. .............................................................Brachymetra mera Harris and Drake, 1945 (Peru). - The wingless and short-winged forms do not exceed 8.6 mm in length, and the apterous form reaches only c. 9.2 mm. If it exceeds 7 mm, the apex of its pronotum forms an acute angle. ...........................................................................5

Fig. 6.118 Brachymetra kleopatra: head and pronotum (upper left), apex of the male (lower right center) and female abdomen (upper right center) in ventral view, male paramere (upper right), male genital capsule (lower left), and apex of the penis showing the sclerotized plates (lower right). Based on Shaw (1933), Hungerford and Matsuda (1957), and Matsuda (1960).

Fig. 6.119 Brachymetra lata male (left to right): head and pronotum in dorsal view, clasper, and apex of the abdomen in ventral view. Based on Shaw (1933).

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5. (4) The apex of the pronotum forms an acute angle and extends well beyond the apices of the mesothoracic acetabula (Fig. 6.118). ........................................6 - The apex of the pronotum is bluntly angulate, or it does not extend beyond the apices of the mesothoracic acetabula (Fig. 6.119). ........................................7 6. (5) The wings are reddish brown, contrasting with the lighter orange yellow pronotum, mesonotum, and metanotum. The widest part of the pronotum of winged specimens is anterior to its mid-length. The posterolateral angles of the seventh abdominal segment of the male are broadly rounded and somewhat recurved anteriad. The inner sclerotized plates of the penis are only slightly convergent, and the narrow outer plates are angled (Fig. 6.120). The relative lengths of the antennal segments from I to IV are 40:25:32:29, respectively. Length of winged male: c. 9.2 mm; wingless male: 8.3 to 8.4 mm; winged female: 9.0 mm; wingless female: 8.0 to 8.6 mm. ...............................................Brachymetra shawi Hungerford and Matsuda, 1957 (West Indies, Guyana, Surinam, French Guiana, Bolivia, Pará, Amazonas). Syn: Brachymetra kleopatra Shaw, 1933 (nec Kirkaldy, 1899). - The wings and pronotum are fairly uniform reddish brown in color. The widest part of the pronotum of winged specimens is at its mid-length. The posterolateral angles of the seventh abdominal segment of the male are evenly rounded and not recurved. The inner sclerotized plates of the penis are narrow and parallel, and the outer plates are slightly convergent (Fig. 6.118). The relative lengths of the antennal segments from I to IV of the male are 104:55:76:62, and those of the female, 50:25:37:29, respectively. Length: 7.4 to 8.1 mm. ................................................................Brachymetra kleopatra (Kirkaldy, 1899) (Ecuador). Syn: Gerris kleopatra Kirkaldy, 1899.

Fig. 6.120 Brachymetra shawi: paramere (upper left) apical segments of the abdomen of a male (upper center) and a female (upper right) in ventral view, genital capsule of a male (lower left), and the apical part of the penis (lower right). Based on Hungerford and Matsuda (1957) and Matsuda (1960).

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7. (5) The fore-femur is considerably shorter than the other femora. The color is dark chocolate brown dorsally and brownish yellow ventrally, and there is a patch of long, silvery white setae on the dorsal surface of each acetabulum of the middle and hind legs. Length of winged specimens: 6.0 to 6.5 mm. Apterous forms are unknown. ..............................................................................Brachymetra furva Drake, 1957 (Argentina, Minas Gerais, Rio de Janeiro). - The fore-femur is longer than the middle femur. .............................................8 8. (7) The fore-femur is only slightly larger than the middle femur. The rear margin of the eye is definitely longer than the interocular space; the eyes are directed posteriad. There is a dense field of small pegs on the posterior surface of the thin anterior femur (Fig. 6.119). Macropterous and apterous forms are known. Length: 6.2 to 6.9 mm. .................................................................................Brachymetra lata Shaw, 1933 (Ecuador, Surinam, Colombia, Pará, Amazonas). - The eyes are not directed posteriad. The fore-femur lacks a dense field of small pegs on the posterior surface, and it is at least 1.25 times as long as the middle femur (Fig. 6.50). Length of apterous adults: 5.4 to 6.9; length of macropterous adults: 6.0 to 6.4 mm. ...............................................Brachymetra albinerva (Amyot and Serville, 1843) (Central America, Trinidad, Lesser Antilles, Surinam, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia, Paraguay, Argentina, Ceará, Bahia, Pará, Amazonas, Mato Grosso, Minas Gerais, São Paulo, Rio de Janeiro). Syn: Helobates albinervus Amyot and Serville, 1843; Brachymetra albinervis Nieser, 1970, spelling error. Two subspecies have been described: Brachymetra albinerva incisa Shaw, 1933, known only from Bolivia, and B. a. albinerva, found at the other locations listed above. The Bolivian subspecies has a shorter first antenna segment, which just reaches the rear margin of the eye when directed posteriad, and the posterior margin of its first abdominal tergite is deeply rather than shallowly incised. Apterous adult B. a. incisa are reportedly 5.4 to 5.6 mm long, while macropterous adults are 6.0 to 6.4 mm. Macropterous adult B. a. albinerva are reported to be 6.0 to 6.1 mm. Subfamily Gerrinae Key to the species of adult Tachygerris in South America Information for the key was provided by Drake and Harris (1934), Matsuda (1960), Padilla and Nieser (2003), and Morales-Castaño and Molano-Rendón (2009). 1. The posterior ventral margin of the female abdomen is broadly rounded, obviously convex medially, and curving to form posterolateral corners slightly

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more acute than 90º and not extended to form posterolateral spines (Fig. 6.121). Length of male: 8.0 to 11.7 mm; female: 10.5 to 12.3 mm. ...................................................................Tachygerris spinulatus (Kuitert, 1942) (Pará). - There are distinct spines formed at the posterolateral corners of the apical ventral margin of the female abdomen, or, if the spines are indistinct, the posterior margin is sinuous (Fig. 6.122). .............................................................2

Fig. 6.121 Tachygerris spinulatus (left to right): apex of the female abdomen in ventral view, apical segment of the endosoma, and the middle (above) and hind tarsus (below). Based on Matsuda (1960).

Fig. 6.122 Tachygerris surinamensis (left to right): apex of the male abdomen in ventral and lateral view, and the apex of the female abdomen in ventral view. Based on Matsuda (1960) and Nieser (1970a). 2. (1) The posterior ventral margin of the female abdomen is sinuous without a small convex projection in the middle and has two broad convex lobes between the middle and the posterolateral corners; the spines at these corners are short and may be indistinct (Fig. 6.122). Length of male: 5.8 to 6.0 mm; female: 6.2 to 7.2 mm. ....................................Tachygerris surinamensis Hungerford and Matsuda, 1958 (Surinam, Amazonas, Pará). - The posterolateral corners of the apical ventral margin of the female abdomen are distinctly produced into large spines, and the margin itself is not sinuous (Fig. 6.123). ..........................................................................................................3

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Fig. 6.123 Tachygerris dentiferus: fore-femur of a male (lower left center), apex of the abdomen of a male in ventral (left) and lateral view (left center), and apex of the abdomen of a female in ventral (right center) and lateral view (right). Based on Padilla and Nieser (2003). 3. (2) There is a raised plate in the center of the metasternum, which is better developed in the male than in the female. The spines at the posterolateral angles of the connexiva do not extend beyond the apex of the female abdomen. The apical abdominal segment of the female is subquadrate in outline (Fig. 6.123). The seventh abdominal sternite of the male is longer than the combined length of the fifth and sixth. The coxae, trochanters, and femora of all legs are light green. The head is brown with darker markings between the eyes. The pronotum is black with a pale midline stripe on the anterior lobe. Length of male: 6 to 8 mm; female: 7 to 9 mm. ...................................................Tachygerris dentiferus Padilla and Nieser, 2001 (Colombia). - There is no raised plate in the center of the metasternum (Fig. 6.124). ...........4 4. (3) The head and anterior lobe of the pronotum have black markings. The first antennal segment is slightly longer than the width of the head through the eyes. The sixth sternite of the male is notched deeply in the middle of the posterior border. That of the female is broadly and convexly rounded (Fig. 6.124). Length: 6.2 to 11.0 mm. .......................................................Tachygerris quadrilineatus (Champion, 1898) (Mexico, Central America, Paraguay). Syn: Limnometra quadrilineata Champion, 1898; Tenagogonus quadrilineatus Kirkaldy and Torre-Bueno, 1908. - The head and anterior lobe of the pronotum are not or are only indistinctly marked with darker lines, which, if present, are never black. The first antennal segment is as long as the width of the head through the eyes. The sixth sternite of the male is not distinctly notched in the middle of its posterior border (Fig. 6.125). ...................................................................................................................5

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Fig. 6.124 Tachygerris quadrilineatus female: apical segment of the endosoma (left) and apical segments of the abdomen in ventral view (right). Based on Matsuda (1960).

Fig. 6.125 Tachygerris celocis (left to right): fore-femur of a male and apex of a male and a female abdomen in ventral view. Based on Matsuda (1960) and García (2002). 5. (4) There is a prominent tubercle near the base on the inner side of the male fore-femur. The apex of the female sixth sternite is subquadrately produced on the posterior edge (Fig. 6.125). Length: 6.3 to 8.1 mm. .......................................................Tachygerris celocis (Drake and Harris, 1930) (Colombia, Venezuela, Peru, Bolivia, Minas Gerais, Mato Grosso). Syn: Tenagogonus celocis Drake and Harris, 1930; Tachygonus celocis (Drake and Harris, 1930). - There is no tubercle near the base, on the inner side of the fore-femur of the male. The apex of the female abdomen has three processes, the middle of which is slightly longer than the other two (Fig. 6.126). .....................................6 6. (5) The fore-femur of the male has no structural modification. ..................................................................Tachygerris opacus (Champion, 1901) (Central America, Colombia, Ecuador, Venezuela, Guyana, Amazonas, Pará). Syn: Limnometra opaca Champion, 1901; Tenagogonus opacus Kirkaldy and Torre-Bueno, 1908. - The fore-femur of the male has a modification on the anterior third consisting of a dense group of tall setae. The apex of the female abdomen has two lateral

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processes but no process in the middle (Fig. 6.51). Length of male: 6.7 to 7.0 mm; female: 7.5 to 7.8 mm. ......................................................................Tachygerris adamsoni (Drake, 1942) (West Indies, Surinam, French Guiana, Venezuela, Colombia, Peru, Paraguay, Bolivia, Ecuador, Pará, Amazonas, Mato Grosso, Minas Gerais, Rio de Janeiro). Syn: Tenagogonus adamsoni Drake, 1942; Tenagogonus duolineatus Kuitert, 1942.

Fig. 6.126 The apex of the male abdomen of Tachygerris opacus in lateral view. Based on Nieser (1970a). Key to the species of adult Neogerris in the Neotropics Information for the key was provided by Nieser and Alkins-Koo (1991), Nieser (1994), and García (2002). 1. The apterous form is at least 7.5 mm long, and the macropterous form is longer than 8.2 mm. The pronotum of the apterous form covers almost the entire mesonotum (Fig. 6.127). ............................................................................Neogerris magnus (Kuitert, 1942) (Colombia, Venezuela, Guyana, Surinam, Bolivia, Paraguay, Pará, Amazonas). Syn: Limnogonus celeris magnus Kuitert, 1942. - The apterous form is shorter than 7 mm, and the macropterous form is shorter than 8 mm. ............................................................................................................2

Fig. 6.127 The apex of the male abdomen of Neogerris magnus in dorsal view (left) and the apex of a female abdomen in lateral view (right). Based on Nieser (1994).

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2. (1) Apterous: adults without wings (Fig. 6.55). ...............................................3 - Macropterous: adults with fully developed wings (Fig. 6.128). ......................9 3. (2) The pronotum lacks all but a trace of a posterior lobe (Fig. 6.55). ............4 - The pronotum is obviously divided into an anterior and posterior lobe, and it is wider than long (Fig. 6.128). ............................................................................6

Fig. 6.128 Neogerris lubricus macropterous male: dorsal view of habitus (left); fore (upper right) and hind wing (lower right). Based on Matsuda (1960). 4. (3) The width of the head and length of the first segment of the antenna are subequal. The male abdomen is short; the part of the abdomen posterior to the acetabula of the hind legs is 0.20 to 0.22 of the hind femur length (Fig. 6.55). ................................................................Neogerris hesione (Kirkaldy, 1902) pars (North and Central America to Panama, West Indies). - The width of one compound eye plus the interocular space is subequal to the length of the first segment of the antenna. The part of the male abdomen posterior to the acetabula of the hind legs is 0.35 to 0.40 of the hind femur length (Fig. 6.129). ...............................................................................................5 5. (4) The width:length ratio of the mesonotum is 0.65:1. The first genital segment of the male has a tuft of pale, hair-like setae on the dorsolateral edge. The apical segment of the endosoma is elongate and narrow (Fig. 6.129). Length: 5.0 to 5.8 mm. ...........................................................Neogerris visendus (Drake and Harris, 1934) (Peru, Colombia, Surinam, Roraima, Pará, Amazonas, Mato Grosso, Rondônia). Syn: Limnogonus visendus Drake and Harris, 1934. - The width:length ratio of the mesonotum is about 0.8:1. The first genital segment of the male lacks a tuft of pale hair-like setae (Fig. 6.130). Length: 3.9 to 5.2 mm. A macropterous form of this species is not known. ................................................................................Neogerris kontos Nieser, 1994 (Minas Gerais).

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Fig. 6.129 Apical segment of the endosoma of Neogerris visendus. Matsuda (1960).

Based on

Fig. 6.130 Neogerris kontos (left to right): apex of the male abdomen in dorsal and lateral view and apex of the female abdomen in dorsal and lateral view. Based on Nieser (1994). 6. (3) The mesonotum is almost completely covered by the pronotum, which reaches as far posteriad as the metanotum (Fig. 6.128). ......................................7 - The pronotum covers only about half of the mesonotum (Fig. 6.55). .............8 7. (6) The first genital tergite of the male is narrow, and its sides converge toward the apex; its posterior margin is barely notched in the middle, if at all. The fore-femur is about five times as long as wide and only slightly curved. The anterior margin of the pronotum of macropterous individuals is slightly concave (Fig. 6.128). Length: 3.5 to 5.1 mm. The pronotum is black with an ochraceous spot on the disc of the anterior lobe; the margin of the posterior lobe is of the same color. ....................................................................Neogerris lubricus (White, 1879) pars (West Indies, Guyana, Surinam, Colombia, Ecuador, Peru, Bolivia, Argentina, Minas Gerais, Pará, Amazonas, Rondônia, Mato Grosso). Syn: Limnogonus lubricus White, 1879; Neogerris celeris Nieser and Alkins Koo, 1991, partim, misidentified. See Couplet 12 for the macropterous form. - The first genital tergite of the male is wide and nearly parallel-sided; it has a deep notch along the mid-line (Fig. 6.131). The fore-femur is 4.5 times as long as wide and distinctly curved in dorsal view. A macropterous form of this species is not known. Length: 5.0 to 5.7 mm. .......................................................Neogerris celeris Drake and Harris, 1934 pars (Paraguay, Mato Grosso).

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Fig. 6.131 Apex of the male abdomen of Neogerris celeris in dorsal view. Based on Drake and Harris (1934). 8. (6) Length of apterous male: 5.5 to 5.9 mm; macropterous male: 6.2 to 6.6 mm; apterous female: 5.7 to 6.1 mm; macropterous female: 6.5 to 7.1 mm. The eyes of the apterous form, viewed laterally, are almost round, not longer than high. The eyes of the macropterous form are globose, not longer than high. The pronotum is strongly constricted laterally between the two lobes. The fore-femur has a black, oval spot on the proximal 2/3 of the posterior surface. The apical segment of the male abdomen appears triangular with a broadly rounded apex in dorsal view (Fig. 6.132). .........................................................................Neogerris lotus (White, 1879) pars (Peru, Colombia, Guyana, French Guiana, Surinam, Pará, Amazonas, Mato Grosso, Goiás). Syn: Limnogonus lotus White, 1879; Limnogonus celeris Nieser, 1970 misidentified; Neogerris celeris Nieser and Alkins Koo, 1991, partim misidentified. - Length: 4.1 to 4.2 mm. The eyes viewed laterally appear flattened and are obviously longer than high. A macropterous form of this species is not known. ..........................................................Neogerris genticus (Drake and Harris, 1934) (Pará). Syn: Limnogonus genticus Drake and Harris, 1934. 9. (2) The length is greater than 6.0 mm. ...........................................................10 - The length reaches no more than 5.8 mm. .....................................................11 10. (9) The maximum width is greater than 2.0 mm. The fore-femur has a black, oval spot on the proximal 2/3 of the posterior surface. The width of the fore-femur and one compound eye are subequal. The first segment of the antenna is about as long as the width of one compound eye plus the interocular space. The apical segment of the male abdomen appears triangular with a broadly rounded apex in dorsal view (Fig. 6.132). Length: 5.6 to 6.7 mm. .........................................................................Neogerris lotus (White, 1879) pars (West Indies, Colombia, Guyana, Surinam, Peru, Pará, Amazonas, Mato Grosso, Goiás, Distrito Federal). Syn: Limnogonus lotus White, 1879; Limnogonus

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celeris Nieser, 1970 misidentified; Neogerris celeris Nieser and Alkins Koo, 1991, partim misidentified. - The maximum width of the body does not exceed 2.0 mm. The fore-femur is usually mainly black on the apical half and is only about ¾ as wide as one compound eye (Fig. 6.55). The length of the first segment of the antenna is subequal to the entire width of the head across both compound eyes. ................................................................Neogerris hesione (Kirkaldy, 1902) pars

Fig. 6.132 Apex of the abdomen of a male Neogerris lotus in dorsal view. Based on Nieser (1970a). 11. (9) There is a tuft of hair-like setae on each side of the second genital segment of the male. The apices of the female connexiva are drawn out into distinct, obtuse spines (Fig. 6.129). Length: 5.0 to 5.8 mm. ...........................................................Neogerris visendus (Drake and Harris, 1934) (Peru, Colombia, Surinam, Pará, Amazonas, Mato Grosso, Rondônia). Syn: Limnogonus visendus Drake and Harris, 1934. - There are no tufts of hair-like setae on the second genital segment of the male (Fig. 6.131). The apices of the female connexiva are truncate and do not resemble spines. ..................................................................................................12 12. (11) The median length of the first genital tergite of the male is slightly less than half its basal width, and the lateral margins of the tergite are parallel with rounded posterolateral angles; it is distinctly notched at the midline (Fig. 6.131). Length: 5.0 to 5.7 mm. These features are presumed because a macropterous form of this species has not yet been described. .....................................................Neogerris celeris (Drake and Harris, 1934) pars (Paraguay, Mato Grosso). Syn: Limnogonus celeris Drake and Harris, 1934. - The median length of the first genital tergite of the male is about 2/3 of its basal width, and the lateral margins of the tergite converge toward the apex. The first genital tergite of the male has no more than a slight incision along the midline (Fig. 6.128). ....................................................................Neogerris lubricus (White, 1879) pars (Colombia, Guyana, French Guiana, Surinam, Peru, Ecuador, Bolivia, Argentina, Pará, Amazonas, Mato Grosso). Syn: Limnogonus lubricus White, 1879; Neogerris celeris Nieser and Alkins Koo, 1991, partim, misidentified. See Couplet 7 for the apterous form.

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Key to the species of adult Limnogonus in South America Information for the key was provided by Drake and Harris (1933a, 1934), Andersen (1975), Bachmann (1962e), and García (2002). Most specimens are macropterous. 1. The first antennal segment is shorter than the width of the head through the eyes. The mesopleurae are broadly marked with reddish or russet brown along the dorsal edges. In lateral view, the female abdomen appears to taper evenly to the final segment (Fig. 6.133). Length: 6.5 to 9.4 mm. An apterous form has not been described. ..........................................................Limnogonus ignotus Drake and Harris, 1934 (Guyana, French Guiana, Surinam, Colombia, Paraguay, Bolivia, Peru, Argentina, Uruguay, Pará, Minas Gerais, Mato Grosso, Mato Grosso do Sul, Distrito Federal). - The first antennal segment is as long as or longer than the width of the head through the eyes. The mesopleurae are brown or black dorsally along the pronotal margin; this color is sometimes interrupted by yellowish or silvery markings (Fig. 6.53). ............................................................................................2

Fig. 6.133 Limnogonus ignotus: apex of the female abdomen in lateral view. Based on Matsuda (1960).

Fig. 6.134 Limnogonus profugus (left to right): apex of the male abdomen in ventral and lateral view and the apex of the female abdomen in ventral view. Based on García (2002). 2. (1) Males (Fig. 6.134). .....................................................................................3 - Females (Fig. 6.134). .......................................................................................7 3. (2) The first genital segment is never produced ventrally at its apex (Fig. 6.134). ...................................................................................................................4 - The first genital segment is produced ventrally at apex (Fig. 6.135). ..............5

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Fig. 6.135 Limnogonus hyalinus: head in lateral view (left), apex of the male abdomen in ventral (upper center) and lateral view (right), and apex of the female abdomen in ventral view (lower center). Based on Nieser (1970a).

Fig. 6.136 Limnogonus franciscanus: habitus of an apterous male in dorsal view (below) and the apical segments of the abdomen of a male in right lateral view (upper left) and a female in left lateral view (upper right). Based on Nieser and Alkins-Koo (1991) and García (2002). 4. (3) The metasternum is flat and not swollen. The ventral part of the first genital segment of the male is distinctly constricted across the middle and

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broader than the distance between the apices of the connexiva (Fig. 6.134). Length: 7.5 to 10.6 mm. ................................................Limnogonus profugus Drake and Harris, 1930 pars (Colombia, Paraguay, Argentina, Pará, Amazonas, Mato Grosso, Goiás, Rio de Janeiro, São Paulo). - The metasternum is distinctly swollen. The ventral part of the first genital segment is only slightly impressed on each side (Fig. 6.136). Length: 5.0 to 8.75 mm. ............................................................Limnogonus franciscanus (Stål, 1859) pars (North and Central America, West Indies, Peru, Colombia, Venezuela). Syn. Gerris marginatus auctt., nec Say, 1832; Gerris franciscanus Stål, 1859; Gerris guerini (Lethierry and Severin, 1896); Limnogonus marginatus Champion, 1898; Limnogonus guerini Drake and Harris, 1934, nec Lethierry and Severin, 1896. 5. (3) The first genital segment has a tumid, almost globular swelling ventrally; its apex is produced medially into a prominent, recurved spine (Fig. 6.137). Length: 8.0 to 9.2 mm. ................................................Limnogonus recurvus Drake and Harris, 1930 pars (Bolivia, Pará, Amazonas, Mato Grosso, Goiás). - The first genital segment lacks such a swelling (Figs. 6.53, 6.135). ...............6 6. (5) First genital segment has an enormous ventral keel, which extends as far posteriad as the dorsal part of the segment (Figs. 6.53, 6.135). Length: 8.6 to 8.7 mm. A macropterous form has not been described. ...........................................................Limnogonus hyalinus (Fabricius, 1803) pars (Trinidad, Panama, Colombia, Guyana, Surinam, French Guiana, Ecuador, Bolivia, Pará, Amazonas). Syn: Hydrometra hyalina Fabricius, 1803; Gerris hyalinus Kirkaldy, 1899. - The first genital segment lacks an enormous keel but has a short, recurved tooth at the apex (Fig. 6.37). Length: 8.0 to 11.3 mm. The pronotum of the apterous form has raised humeri and two short fulvous stripes; it is black with an indistinctly fulvous margin and two short ochraceous lines on the anterior disc. The hemielytra of the macropterous form are brownish black. .................................................Limnogonus aduncus Drake and Harris, 1933 pars (Panama, Trinidad, Colombia, Venezuela, Guyana, French Guiana, Surinam, Ecuador, Peru, Bolivia, Paraguay, Argentina, Minas Gerais, Pará, Amazonas, Mato Grosso). 7. (2) The connexivum is convex at the apex and does not extend beyond the adjacent abdominal tergite (Fig. 6.53, 6.135). Length: 8.6 to 8.7 mm. A macropterous form has not been described. ...........................................................Limnogonus hyalinus (Fabricius, 1803) pars (Trinidad, Panama, Colombia, Guyana, Surinam, French Guiana, Ecuador, Pará, Amazonas). Syn: Hydrometra hyalina Fabricius, 1803. - The connexivum is somewhat produced at its apex and clearly extends beyond the lateral margin of the last abdominal tergite (Fig. 6.37). Length: 8.6 to 8.7 mm. .............................................................................................................8

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8. (7) The last abdominal sternite is sharply produced medially along its apical margin. The connexival spines at the posterolateral corners of the laterotergites are moderately long (Fig. 6.37). Length: 8.0 to 11.3 mm. .................................................Limnogonus aduncus Drake and Harris, 1933 pars (Panama, Trinidad, Colombia, Venezuela, Guyana, Surinam, Ecuador, Peru, Bolivia, Paraguay, Argentina, Pará, Amazonas, Mato Grosso, Minas Gerais, Espirito Santo, São Paulo, Rio de Janeiro). - The last abdominal sternite is broadly rounded along the posterior margin. The laterotergites of the connexiva are not produced into strong spines at their posterolateral corners (Fig. 6.134). ......................................................................9

Fig. 6.137 The apex of the male abdomen of Limnogonus recurvus in lateral view. Based on Nieser (1970a). 9. (8) The dorsal portions of the mesopleurae are largely reddish brown. and the areas of this color are not divided by paler longitudinal stripes. Length: 9.1 to 9.2 mm. The pronotum of the apterous form is fuscous or black with two short ochraceous or yellow stripes on the anterior lobe and a median stripe and margins of the posterior lobe of the same color. A macropterous form has not been described. ................................................Limnogonus recurvus Drake and Harris, 1930 pars (Bolivia, Pará, Amazonas, Mato Grosso, Rondônia, Goiás, Minas Gerais). - The dorsal portions of the mesopleura are black, divided by a yellowish brown stripe (Fig. 6.134). ...................................................................................10 10. (9) The last pre-genital abdominal sternite is strongly produced along its posterior margin so that it extends in the center slightly beyond the connexival processes. The yellowish brown stripe on the mesopleura is expanded, arched dorsally, and broader than the black stripe (Fig. 6.134). Length: 7.5 to 10.6 mm. ................................................Limnogonus profugus Drake and Harris, 1930 pars (Colombia, Venezuela, Guyana, Surinam, Peru, Bolivia, Paraguay, Argentina, Pará, Amazonas, Mato Grosso, Mato Grosso do Sul, Goiás, Minas Gerais, Rio de Janeiro, São Paulo, Rio Grande do Sul).

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- The last pre-genital abdominal sternite is only slightly produced along its posterior margin. The yellowish brown stripe dividing the black areas on the mesopleura is narrow, and its margins are nearly parallel. There is a broad, uninterrupted longitudinal yellowish brown stripe on the dorsal side of the connexivum (Fig. 6.136). Length: 5 to 8.75 mm. ............................................................Limnogonus franciscanus (Stål, 1859) pars (North and Central America, West Indies, Peru, Colombia, Venezuela). Syn. Gerris marginatus aucc, non Say, 1832; Gerris franciscanus Stål, 1859; Gerris guerini (Lethierry and Severin, 1896); Limnogonus guerini (Lethierry and Severin, 1896). Key to the species of adult Eurygerris in South America Information for the key was provided by Drake and Harris (1934), Kuitert (1942), Mazzucconi and Bachmann (1993b), and García (2002). Eurygerris has long been treated as a subgenus of Gerris, under which name the species appear in older literature. 1. There is no median carina on the metasternum or elsewhere on the venter. The pronotum of the apterous form extends only slightly posteriad and does not cover the mesonotum. The first genital segment is short. In ventral view, the seventh abdominal segment of the male is slightly longer than the sixth and the eighth segment. The anterior femur of the male is curved, but it lacks any obvious black enlargement on the ventral side (Fig. 6.54). Length: 5.2 to 10.0 mm. ............................................................Eurygerris flavolineatus (Champion, 1898) (Mexico, Central America, Ecuador, Colombia?). Syn: Gerris flavolineatus Champion, 1898. - There is a somewhat distinct median carina on the metasternum and elsewhere on the venter. The pronotum of the apterous form extends posteriad to cover at least part of the mesonotum. The first genital segment is longer than broad (Fig. 6.138). ........................................................................................2 2. (1) The fore-femur is flattened on the basal half and lacks a stout, oblong prominence on the ventral side. The connexivum of the female curves to form an angle of about 45°. The metanotum is elevated slightly above the plane of the mesonotum and the abdominal segments (Fig. 6.4, 6.138). Length: 6.3 to 9.5 mm. Micropterous forms occur. ........................................................................Eurygerris fuscinervis (Berg, 1898) (Argentina, Chile, Bolivia, Peru, Ecuador, Venezuela, Colombia). Syn: Brachymetra fuscinervis Berg, 1898; Gerris fuscinervis Drake and Harris, 1934; Gerris andromeda (Kirkaldy, 1899); Gerris perseus (Kirkaldy, 1898); Gerris kahli Drake and Harris, 1934; Eurygerris kahli (Drake and Harris, 1934).

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- The fore-femur is curved basally and expands greatly on its ventral side along its length to form a prominence resembling a mound or tubercle (Fig. 6.139). The omphalium is usually prominent. ..................................................................3

Fig. 6.138 Habitus of the dark form of Eurygerris fuscinervis. Based on Drake (1963). 3. (2) The ventral prominence on the fore-femur is at the distal third (Fig. 6.139). In the apterous form, the anterior lobe of the pronotum is greatly elevated. Length: 8.0 to 10.5 mm. ............................................................Eurygerris beieri (Drake and Harris, 1934) (Mexico, West Indies, Colombia). Syn: Gerris beieri Drake and Harris, 1934. - The ventral prominence on the fore-femur is approximately at its mid-length. In the apterous form, the anterior lobe of the pronotum is only moderately elevated (Fig. 6.140). ............................................................................................4 4. (3) The mesonotum is distinctly elevated above the plane of the metonotum and abdominal segments. The fore-femur of the male is nearly parallel-sided, only slightly curved, and bears a low, broad tubercle at about midlength (Fig. 6.141). The connexiva of the female is vertical. The seventh abdominal segment of the female is about twice as long as the sixth. Length: c. 9 mm. ...............................................................................Eurygerris atrekes Drake, 1963 (Colombia).

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- The mesonotum is only slightly elevated above the plane of the metanotum and the abdominal segments. The omphalium is well developed and forms an oblong prominence (Fig. 6.140). Length: 7.5 to 10.5 mm. .............................................................Eurygerris carniventris (Champion, 1898) (Mexico, Central America, Ecuador). Syn: Gerris carniventris Champion, 1898.

Fig. 6.139 Eurygerris beieri male: fore-femur (above) and sclerites of the vesicle (below). Based on Moreira and Ribeiro (2009).

Fig. 6.140 Eurygerris carniventris: fore-wing (left) and the fore-femur of a male (right). Based on Matsuda (1960).

Fig. 6.141 Eurygerris atrekes male: fore-femur (above) and sclerites of the vesicle (below). Based on Moreira and Ribeiro (2009).

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Veliidae Supplemental key to the subfamilies of adults Information for the key was provided by Andersen (1982). 1. The middle tarsus is deeply cleft and bears a plumose swimming fan and leaflike claws arising from the cleft (Fig. 6.142). ..............................................................................................Rhagoveliinae..p. 159 - The middle tarsus is not deeply cleft, or, if it is, two unmodified, hook-like claws or up to four leaf-like lamellae arise from the cleft (Fig. 6.143). ..............2

Fig. 6.142 Apex of a Rhagovelia sp. tarsus showing the swimming fan in dorsal view (upper left), unfolded in lateral view (lower center), and folded (upper right). Based on Andersen (1976).

Fig. 6.143 Oiovelia cunucunumana: hind tibia and tarsus (upper left) showing the enlargement of the apical tarsal segment (lower left), male genital capsule (center), and paramere (right). Based on Drake and Capriles (1952) and Spangler (1986).

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2. (1) All tarsi consist of three segments, although the first segment may be very small and visible only in lateral or ventral view (Fig. 6.143). There are four closed cells in each hemielytron. ........................................................................................................Veliinae..p. 209 - The fore-tarsus consists of one segment, while the middle and hind tarsi consist of two (Fig. 6.144). ...............................................................................................Microveliinae..p. 239

Fig. 6.144 Microvelia timida male: habitus in dorsal view (center) and the foretibia and tarsus showing the small projection at the apex of the tibia (upper left). Based on Drake and Roze (1958). Key to the genera in South America Information for the key was provided by Hungerford (1929a, b, c, d), Usinger (1963), Drake (1957c), Drake and Lauck (1959a), Drake and Menke (1962), Andersen (1982), Nieser and Alkins-Koo (1991), Bachmann and Mazzuconi (1995), Mazzucconi and Bachmann (1997a, b), and Pereira et al. (2007). Many South American veliid species were formerly placed in the Old World genus Velia Latreille, 1804, but all have since been transferred to Neotropical genera. 1. The middle tarsus is deeply cleft and bears a plumose swimming fan and leaflike claws arising from the cleft (Fig. 6.145). ...............................................................................Rhagovelia Mayr, 1865..p. 159 - The middle tarsus is not deeply cleft, or, if it is, only two unmodified, hooklike or leaf-like claws and sometimes two modified setae arise from the cleft (Fig. 6.146). ..........................................................................................................2

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2. (1) All tarsi consist of three segments, although the first segment may be very small and visible only in lateral or ventral view (Fig. 6.147). There are four closed cells in each hemielytron. ..........................................................................3 - The fore-tarsus consists of one segment, while the middle and hind tarsi consist of two (Fig. 6.148). ..................................................................................9

Fig. 6.145 Rhagovelia merga male: habitus of an apterous specimen in dorsal view (left), proctiger (upper right), and paramere (lower right). Based on Polhemus (1997).

Fig. 6.146 Stridulivelia tersa: habitus of an apterous male in dorsal (left) and lateral view without appendages (upper right center), and the femur of a male (lower right). Based on Drake and Menke (1962).

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Fig. 6.147 Steinovelia virgata: habitus of a macropterous female (left) and head in lateral view ( upper right), and head and thorax in ventral view (lower right), showing the metathoracic tubercles directly opposite the coxae. Based on Andersen (1982).

Fig. 6.148 Husseyella halophila (left to right): head and thorax of a cleared specimen in dorsal view, antenna, and the apical segment of the middle tarsus. Based on Drake (1958c). 3. (2) The apical segment of the middle tarsus bears four narrow leaf-like laminae, representing modified claws and setae. The shape may vary from elongate to elliptical (Fig. 6.149). ........................................................................4 - The apical segment of the middle tarsus bears a pair of falcate claws and sometimes unmodified setae (Fig. 6.147). The length is greater than 3 mm. ......5

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Fig. 6.149 Euvelia concava female: habitus of an apterous specimen in dorsal view (left) and lateral view of the head and thorax showing the location of the glabous area on the pleurae in black and the relative positions of the coxae. Based on Polhemus and Polhemus (1984a). 4. (3) Individuals are typically tiny and apterous, not exceeding 2.5 mm in length. The apical segment of the middle tarsus is longer than the basal segment and has a shallow impression in the basal third in which four narrow membranous laminae are inserted, which are 2/3 the length of the tarsal segment. The middle coxae are inserted closer to the hind coxae than to the fore-coxae (Fig. 6.149). ....................................................................................Euvelia Drake, 1957..p. 209 - The minimum length is 7 mm. The third tarsal segments of the middle and hind legs each bear four broad, thin, laminate or blade-like structures. The male parameres are conspicuous and held in a near vertical position (Fig. 6.150). ..................................................................................Veloidea Gould, 1934..p. 211

Fig. 6.150 Veloidea gigantea (left to right): apex of the male abdomen in lateral view, right male paramere, anal segment of a male, and the right femur of a female. Based on Drake and Lauck (1959a).

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5. (3) There are furrows between some of the abdominal segments. The surface of the body is covered by many round depressions, which do not bear setae. The pleurae of the second and third abdominal segments, and sometimes the fourth and fifth as well, have broad, shiny sulci. A stridulatory apparatus on the hind femur and on the ventral side near the upper border of the connexiva of the second, third, and fourth abdominal segments is sometimes evident (Fig. 6.146). ....................................................................Stridulivelia Hungerford, 1929..p. 211 - Intersegmental furrows are absent. The surface of the body is not covered by many round, depressed structures, which do not bear setae. The pleurae of the second and third abdominal segments do not have broad sulci. Rarely, there may be a stridulatory apparatus on a trochanter and abdominal segments, but never on the hind femur. The apical segments of the tarsi are either widened laterally or subcylindrical. The second segment of the middle tarsus is significantly longer than the first (Fig. 6.151). ....................................................6

Fig. 6.151 Paravelia paxilla: habitus of a male (lower left) and a female without appendages (middle right), head of a male in lateral view (upper left), fore-leg of a male in anterior view (upper right), and apex of a male abdomen in ventral view (lower right). Based on Mazzucconi and Polhemus (2003). 6. (5) The apical segments of all tarsi are widened laterally, making them suboval. The second segment of the middle tarsus is very short, not much longer than the tiny first segment. The apical segment of the middle tarsus is cleft, and the cleft bears unmodified, falcate claws (Fig. 6.152). .............................................................Oiovelia Drake and Capriles, 1952..p. 217 - The apical tarsal segments are subcylindrical (Fig. 6.151). ..............................7

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Fig. 6.152 Oiovelia rivicola (left to right): head and thorax in dorsal view, apex of the fore-tarsus, and the male genitalia in lateral view. Based on Spangler (1986).

Fig. 6.153 Platyvelia brachialis male: habitus of a macropterous North American specimen (left), head and thorax in oblique ventral view showing the opposing mesoacetabular and metathoracic tubercles (lower right), and the parameres of specimens from Santa Catarina and Texas (upper and middle right, respectively). Based on Polhemus and Polhemus (1993b). 7. (6) The anterior border of the metasternum does not bear a pair of tubercles (Fig. 6.151). .............................................................................Paravelia Breddin, 1898..p. 219 - The anterior border of the metasternum bears one pair of tubercles (Fig. 6.153). ...................................................................................................................8 8. (7) The tubercles on the anterior border of the metasternum are situated directly posterior to a pair of mesoacetabular tubercles on posterior border of the

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mesosternum. The femora, especially those on the hind legs, are robust (Fig. 6.153). ...............................................Platyvelia Pohlhemus and Pohlhemus, 1993..p. 236 - The tubercles on the anterior border of the metasternum are situated directly posterior to the middle coxae. The femora are narrow (Fig. 6.147). .............................................Steinovelia Pohlhemus and Pohlhemus, 1993..p. 237 9. (1) The middle tarsi bear only unmodified claws and lack laminae. The middle legs are inserted nearly equidistant from the other two pairs (Fig. 6.154). .......................................................................Microvelia Westwood, 1834..p. 239 - The tarsal claws and ariolae on the middle tarsi are modified into four broad, membranous plates (Fig. 6.148). ........................................................................10

Fig. 6.154 Habitus of Microvelia ashlocki. Based on Froeschner (1985).

Fig. 6.155 The male right paramere of Xiphovelia lacunana. Polhemus (1977).

Based on

10. (9) The second tarsal segment bears four membranous laminae. The middle coxae are inserted closer to the hind coxae than to the fore-coxae (Fig. 6.148). Marine species. ............................................................................Husseyella Herring, 1955..p. 254 - The second tarsal segment bears three membranous laminae. The male paramere has a swollen base bearing three long setae (Fig. 6.155). Length of only known species: c. 2.4 mm. Width across humeri: c. 1.0 mm. The habitat is inland freshwater bodies. .....................................................................................Xiphovelia Lundblad, 1933 The only South American species reported is Xiphovelia lacunana (Drake and Plaumann, 1955), from Mato Grosso. Syn: Microvelia lacunana Drake and Plaumann, 1955.

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Subfamily Rhagoveliinae Key to the species of adult Rhagovelia in South America Information for the key was provided by Drake and Harris (1935b), Drake (1953a, 1957d, 1958d), Drake and Carvalho (1955), Bacon (1956), Matsuda (1956), Drake and Doesburg (1966), Polhemus (1997), Nieser and Polhemus (1999), and Moreira and Ribeiro (2009). In some cases, adequate descriptions only of apterous or only of macropterous males or females are available, making identification difficult until all forms of each species have been described or until it is determined that one or more wing morphs of individual species do not occur. In most cases, apterous specimens are required to identify the species, but in a few cases only macropterous specimens can be identified. This key will therefore not permit identification of those wing morphs of known species which have not yet been described. The illustrations of the male paramere, also called clasper, are not standard in orientation or in depiction of the setae. The key includes the species formerly placed in the genus Trochopus Carpenter, 1898. Polhemus (1997) considered these species to be members of a subgroup within Rhagovelia, but he withheld judgement on the validity of proposed subgeneric status for this taxon. 1. The middle and hind tarsi, and sometimes the fore-tarsus, as well, consist of two segments. The adults are apterous (Fig. 6.156). All species are marine and had formerly been placed in a separate genus, Trochopus Carpenter, 1898, but are now considered by most to be species of Rhagovelia. ...................................2 - All tarsi consist of three segments. The adults are apterous or macropterous (Fig. 6.157). The species inhabit freshwater bodies, usually lotic ones. .............6

Fig. 3.1.156 Habitus of an apterous female Rhagovelia ephydros (left) and the right paramere of a male (right). Based on Nieser (1975).

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Fig. 6.157 Rhagovelia aiuruoca: habitus of a male (upper left) and female (lower left) with the legs removed in dorsal view; apical segments of the abdomen of a male (upper right center) and a female in ventral view (center); male paramere (left of center); fore, middle, and hind leg of a male (upper right, top to bottom); and fore (lower center), middle (lower right center), and hind leg of a female (lower right). Based on Moreira and Ribeiro (2009).

Fig. 6.158 Rhagovelia salina (left to right): outline of pronotum of a male in dorsal view, right fore-leg of a male, left hind femur of a male, and apex of the left ovipositor of a female specimen. Based on Drake and Doesburg (1966).

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2. (1) The posterior margin of the male pronotum appears almost truncate; that of the female may be truncate or convexly rounded (Fig. 6.158). .......................3 - The posterior margin of the pronotum does not appear truncate; it is excavated to form one or two concave indentations (Fig. 6.159). .......................................5 3. (2) The distal half of the fore-femur of the apterous male is bent more than 45º. The female lacks long dorsal setae. The apterous female has deep depressions on each side of the dorsal midline of the metanotum and first abdominal tergite (Fig. 6.160). Length of male: c. 3.0 to 3.1 mm; female: c. 4.2 to 4.3 mm. It typically inhabits estuaries and mangrove swamps. Macropterous forms are unknown. ..............................................Rhagovelia arcuatus Polhemus and Manzano, 1992 (Colombia). - The fore-femur of the male is nearly straight. The metanotum and first abdominal tergite are not depressed (Fig. 6.161). ................................................4 4. (3) The entire pronotum behind the vertex of the head is yellow or orange. Stripes are lacking on the basal segments of the antennae and all femora. There is a thick, blunt, tubercle-like spine on the ventral surface of the male foretrochanter (Fig. 6.158). The legs have a moderate coat of long, black, hair-like setae. The hind femur of the male has a long row of prominent spines, with two of the spines near the middle of the row longer than the rest. The long dorsal setae of the female are as long as the width of the hind tibia. The widest point of the female body is at the third abdominal tergite. Length: 3.50 to 3.73 mm. .......................................................................Rhagovelia salina (Champion, 1898) (Marine habitats off Panama). Syn: Trochopus salinus Champion, 1898. - The dorsal coloration is predominantly black. The fore-trochanter of the male lacks a blunt spine of the ventral surface. The female lacks long dorsal setae, and the widest point of the body is across the metanotum (Fig. 6.161). ........................................Rhagovelia colombianus Polhemus and Manzano, 1992 (Colombia). 5. (2) The posterior margin of the pronotum is deeply excavated in the middle. There is a black, longitudinal stripe on each femur and on the basal segments of the antenna (Fig. 6.156). The hind femur of the male has a row of spines on the ventral side for about ¾ of its length. The hind femur of the female has a row of about four to seven small spines on its apical half. ..................................................Rhagovelia ephydros (Drake and Doesburg, 1966) (Coastal waters of Surinam). Syn: Trochopus ephydros Drake and Doesburg, 1966. - The posterior margin of the pronotum is sinuate, forming two excavations (Fig. 6.159). The legs lack long, hair-like setae. The male bears three to five spines on the apical half. The femora of the female lack spines. ............................................................................Rhagovelia plumbea Uhler, 1894 (Marine habitats off Florida, Belize, West Indies). Syn: Trochopus plumbeus (Uhler, 1894); Trochopus marinus Carpenter, 1898; Taochopus (sic) plumbeus Kirkaldy, 1900; Trachopus (sic) marinus Van Duzee, 1917; Trochopus maritimus Drake and Harris, 1935; Trochopus plumbea Cobben, 1960.

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6. (1) The pronotum of apterous adults, measured along the midline, is obviously shorter than three times the length of the exposed part of the mesonotum (Fig. 6.162). For macropterous specimens, continue through both options in this couplet separately. Find the closest descriptions of the specimens, and refer to the original description and later descriptions in the literature cited. Until more information is available, undescribed winged or wingless forms of the known species may not be identifiable. ..........................................................7 - The pronotum of apterous adults, measured along the midline, is longer than three times the length of the exposed portion of the mesonotum. In many cases, the pronotum completely covers the mesonotum (Fig. 6.157). .........................54

Fig. 6.159 Rhagovelia plumbea (left to right): pronotum of a female, right hind femur of a male, and the apex of the left ovipositor. Based on Drake and Doesburg (1966).

Fig. 6.160 Rhagovelia arcuatus: habitus without appendages of a male (upper left) and a female (lower left), the fore (center) and hind leg of a male (right center), and the male paramere (right). Based on Polhemus and Manzano (1992). 7. (6) The pronotum of apterous adults, measured along the midline, is shorter than the dorsal length of a compound eye; its posterior margin is usually straight or slightly concave, making it somewhat shorter along the midline than lateral

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to it (Fig. 6.163). For macropterous specimens, check both options in this couplet separately. ................................................................................................8 - The pronotum of apterous adults, measured along the midline, is longer than the dorsal length of a compound eye; its posterior margin is convex, so that it extends farthest posteriad at its dorsal midline (Fig. 6.164). ............................39 8. (7) There are three closed cells in each hemelytron (Fig. 6.165), and the length ranges from 2.3 to 4.2 mm. ........................................................................9 - The hemelytron has four closed cells (Fig. 6.164), or only wingless specimens are available for examination. ...........................................................13

Fig. 6.161 Rhagovelia colombianus: habitus without appendages of a male (upper left) and a female (lower left), the fore (center) and hind leg of a male (right center), and the male paramere. Based on Polhemus and Manzano (1992).

Fig. 6.162 Rhagovelia agra: habitus of a female in dorsal view (left), male antenna (right), and the hind femur of a male (upper center). Based on Polhemus (1997).

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9. (8) The second segment of the antenna is distinctly shorter than the third. The middle coxae are at least partially dark brown or blackish ................................10 - The second antenna segment is subequal to or longer than the third. .............12

Fig. 6.163 Rhagovelia sbolos: female habitus without appendages in dorsal view (left), hind trochanter and femur of a male (upper center), male proctiger (right), and male paramere (lower center). Based on Nieser and Polhemus (1999).

Fig. 6.164 Rhagovelia cali: habitus without appendages in dorsal view (left), hemielytron (right center), male proctiger (right), and a male paramere (lower left center). Based on Polhemus (1997).

Fig. 6.165 Rhagovelia versuta (left to right): right paramere in lateral view, basal and apical segments of a macropterous female’s abdomen, and a hemielytron (above) and hind wing (below). Based on Bacon (1956) and Matsuda (1956).

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Fig. 6.166 Right paramere of a male Rhagovelia hambletoni in lateral view. Based on Bacon (1956). 10. (9) The right paramere is short and blunt with a shallow excavation between the apical end of its dorsal margin and its small apical protuberance (Fig. 6.166). The second tarsal segment of the middle leg is subequal to the third, or at least more than 4/5 as long. The hind femur length of the male is equal to the total length of the insect or slightly less. The pronotum and mesonotum are convex dorsally. The venter is bluish gray. The coxae on the middle legs are dark brown or black. The posterior femur of the female is armed. The central parts of the sixth and seventh abdominal tergites of the female are shiny. Length: 2.93 to 3.40 mm. .....................................................Rhagovelia hambletoni Drake and Harris, 1933 (Minas Gerais, Rio de Janeiro). - The male paramere does not have an apical excavation, although it may have a subapical excavation along its dorsal margin. The second tarsal segment of the middle leg is 4/5 or less as long as the third. The apex of the hind tibia is unarmed, but there are sometimes spines along the posterior margin (Fig. 6.167). The apterous female has shiny black spots on the dorsal surfaces of the last two unmodified abdominal segments. .........................................................11

Fig. 6.167 Rhagovelia vonprahli male: hind femur (left) and paramere (right). Based on Manzano et al. (1995). 11. (10) The length of the male paramere is at least three times as long as its width at mid-length. Ratios of antennal segments I through IV: 58:35:39:41. The hind femur of the male is armed with a strong spine on the posterior margin just beyond the mid-length followed by three to four small spines (Fig. 6.167). The hind femur of the female is also armed with a strong spine followed by two very small spines. The seventh and eighth abdominal segments are shiny in the

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center. Length of apterous male: 2.4 to 2.6 mm; apterous female: 2.6 to 2.8 mm. Length of macropterous male: 2.9 to 3.1 mm; macropterous female: 3.0 to 3.3 mm. Color: mainly black with yellow on the basal third of the first antennal segment, base of the fore-femur, bases of the coxae, and the trochanters. ..................................Rhagovelia vonprahli Manzano, Nieser, and Caicedo, 1995 (Colombia). - The male paramere has a swelling from just beyond its base to just beyond its midlength, so its length is about twice as long as its width measured at midlength (Fig. 6.168). Length: 2.86 to 3.66 mm. ............................................................Rhagovelia tantilla Drake and Harris, 1933 (Central America, Peru).

Fig. 6.168 Rhagovelia tantilla (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 12. (9) All legs, including the fore-femur, are entirely black. The apex of the male paramere is truncate (Fig. 6.165). The second antennal segment is usually subequal to the third but may be less than 10% shorter, while the third and fourth segments are subequal. Length: 3.4 to 4.1 mm. ............................................................Rhagovelia versuta Drake and Harris, 1935 (Peru, Bolivia, Argentina). Syn: Rhagovelia yacuivana Drake, 1958. - The fore-femur is yellow on the basal half, and the fore and hind coxae and trochanters are yellowish or stramineous. In lateral view, the ventral margin of the right paramere of the male appears strongly sinuous, and the apex is acute (Fig. 6.169). Length: 3.1 to 3.6. ............................................................Rhagovelia velocis Drake and Harris, 1935 (Peru).

Fig. 6.169 Right paramere of a male Rhagovelia velocis in lateral view. Based on Bacon (1956).

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13. (8) The hind femora of males and females are not armed with spines. ........14 - The hind femur of the male, and sometimes that of the female, is armed with spines (Fig. 6.170). .............................................................................................18

Fig. 6.170 Rhagovelia ochroischion (left to right): hind trochanter and femur of a male, male proctiger, paramere of a male, and the apex of the female abdomen in lateral view. Based on Nieser and Polhemus (1999). 14. (13) The hind tibia is less than 1/3 longer than the hind femur. Length of male: c. 2.3 to 2.8 mm; female: c. 3.0 mm. ........................................................15 - The hind tibia is more than 1/3 longer than the hind femur. The length of both males and females is at least 3 mm. ...................................................................17 15. (14) The color is mainly dark slate gray with yellow pubescence, and the legs are brownish on the coxae and otherwise bluish black with yellowish brown setae. The head has a pair of impressed points along the posterior margin and a deeply impressed arrow head pointing posteriad on the notocephalon. The ratios of antennal segments I to IV are approximately 15:10:8:12, respectively. The abdomen of the female lacks a ventral carina. Length: c. 2.6 mm. Only the apterous form has been described. ...........................................................................Rhagovelia festae Kirkaldy, 1899 (Ecuador). Although this name has been regarded by more recent authors as a nomen dubium, the description by Kirkaldy (1899) is thorough enough to place it tentatively in the key until its identity can be confirmed. - The color is blackish with a bluish tinge. There is a yellow marking on the pronotum, which is interrupted along the dorsal midline. The pronotum is very short and not at all produced over the mesonotum. The legs are black or brownish. ............................................................................................................16

Fig. 6.171 Apex of the abdomen of a female Rhagovelia novana in lateral view. Based on Moreira and Ribeiro (2009).

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16. (15) The length of the middle femur of the male is obviously more than 2 mm. The seventh abdominal tergite is shiny. The ventral side of the body is bluish gray. The coxae are dark brown or black. Length of male: c. 2.7 to 2.8; female c. 3.0 mm. ..............................................................................Rhagovelia rioana Drake, 1953 (Rio de Janeiro). - The middle femur is shorter than 2 mm. The abdominal tergites are usually completely matt without shiny spots. All tergites of the abdomen, except the gential segment, lack shiny patches or areas. The pronotum is black with a bluish tinge and a yellow marking that is interrupted along the dorsal midline. The entire ventral side of the body is dark bluish. The legs are brownish black, and the anterior coxae are testaceous. The abdomen tapers strongly; that of the male is reflexed obliquely dorsad near the apex, while that of the female is strongly constricted just proximal to the seventh abdominal segment, so that the segment forms a nearly vertical plate, which is directed caudad (Fig. 6.171). The connexival margins of the male are straight and taper evenly. The middle femur of the male is less than 2 mm long. Length of male: c. 2.3 mm; female: c. 2.5 mm. Winged specimens have not been found. .............................................................................Rhagovelia novana Drake, 1953 (Argentina, Uruguay, Rio Grande do Sul, Santa Catarina, Rio de Janeiro, São Paulo). 17. (14) The second antennal segment is distinctly shorter than the third, at least in the apterous female. The parameres of the male are narrowed toward the tip (Fig. 6.172). Length: 3.0 to 3.5 mm. ...........................................................................Rhagovelia longipes Gould, 1931 (Peru, Ecuador, Colombia). - The second and third antennal segments are subequal in length. The parameres of the male are broad and shovel-shaped (Fig. 6.173). Length of apterous form: 3.4 to 4.1 mm; macropterous form: 3.8 to 4.4 mm. ...........................................................................Rhagovelia imitatrix Bacon, 1948 (Peru, Argentina).

Fig. 6.172 Rhagovelia longipes (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

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Fig. 6.173 Rhagovelia imitatrix (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

Fig. 6.174 Rhagovelia spinosa (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 18. (13) There is a stout, slightly curved spine at the base of the apical abdominal sternite of the male (Fig. 6.174). The lengths of the second, third, and fourth segments of the antenna are subequal. Length: 2.6 to 3.2 mm. .............................................................................Rhagovelia spinosa Gould, 1931 (Honduras, Ecuador, Peru). - There is no spine at the base of the apical abdominal sternite of the male (Fig. 6.164). .................................................................................................................19 19. (18) There is an unusually large swollen area at the base of the abdomen on the ventral side of the male. The pronotum of the apterous adult is sutured off from the mesonotum. Length: 3.2 to 3.8 mm. Color: generally black with a bluish luster. ....................................................Rhagovelia culebrana Drake and Capriles, 1955 (Venezuela). - There is no large swollen area at the base of the abdomen (Fig. 6.175). ........20 20. (19) The hind femur of the male bears a single, small ventral spine. The length of the seventh abdominal segment is equal to its width at the base. The legs, including the middle coxae, are entirely black. The second segment of the antenna is clearly longer than the third. The connexiva of the female are

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oriented nearly vertically on their posterior halves and thickened toward their posterior apices; they are strongly and continuously convergent and nearly contact each other above the seventh abdominal segment. Length: 3.0 to 3.2 mm. ..........................................................................Rhagovelia occulcata Drake, 1959 (Paraná, São Paulo). - The hind femur of the male usually bears at least two ventral spines (Fig. 6.170), but if only one is present, then the length of the seventh abdominal segment is shorter than its width at the base. The connexiva of the female are not oriented nearly vertically, or if they are, they are not distinctly thickened toward their posterior apices and strongly and continuously convergent so that they are close but not in contact with each other above the seventh abdominal segment. ..............................................................................................................21

Fig. 6.175 Rhagovelia tenuipes (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

Fig. 6.176 Rhagovelia fontanalis (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 21. (20) All abdominal tergites of the apterous male, and often the female, as well, have shiny black central areas, although that on the first tergite may be very small. ..........................................................................................................22

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- No more than six of the tergites of the male have shiny black areas in the middle, and most species have fewer tergites so marked. ..................................24 22. (21) The middle coxae are shiny black. The hind femora of both sexes project beyond the apex of the genital segments; that of the female bears about seven ventral spines. The right male paramere is longer than 2½ times its maximum basal height, measured in lateral view (Fig. 6.175). Length: 3.1 to 4.5 mm. .............................................................Rhagovelia tenuipes Champion, 1898 pars (North and Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Argentina, Rio de Janeiro, Minas Gerais, Amazonas, Pará). Syn: Rhagovelia confusa Gould, 1931. - All coxae and acetabula, as well as the fore and hind trochanters are yellow or yellowish brown. The second segment of the antenna is considerably shorter than the third. ......................................................................................................23

Fig. 6.177 Rhagovelia calopa (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 23. (22) Length: 3.45 to 4.45 mm. The middle trochanter is dark. The hind femur does not project beyond the apex of the genital segments. The right male paramere is slightly more than twice as long as its maximum height, as measured in lateral view (Fig. 6.176). .........................................................................Rhagovelia fontanalis Bacon, 1948 (Peru). - Length of apterous male: 2.6 to 2.8 mm; apterous female: 3.1 to 3.2 mm. The basal part of the first antennal segment and all acetabula, coxae, trochanters, and bases of the femora are pale yellow. The hind femur of the male is five times as long as wide; it is slightly wider than the middle femur and bears seven to nine spines on its ventral surface (Fig. 6.170). The connexiva of the female is oriented horizontally or only slightly angled dorsad. The shiny black ground color is covered by a dark gray pruinosity. The anterior part of the pronotum is crossed by a transverse yellow-orange band, which is also covered by dark gray pruinosity. The ventral surface of the male abdomen is covered by yellowish pilosity. A macropterous form has not been described. ............................................Rhagovelia ochroischion Nieser and Polhemus, 1999 (Minas Gerais).

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24. (21) There is a conspicuous brownish spot along the anterior margin of each segment of the connexiva, except occasionally the first one or two. The hind femur of most males is greatly incrassate. The hind trochanter bears several teeth. The male paramere has an narrow, rounded apex (Fig. 6.177). Length: 3.4 to 3.9 mm. ............................................................Rhagovelia calopa Drake and Harris, 1927 (Mexico, Central America, Trinidad, Colombia, Venezuela). - The hind femur of the male is never more than moderately incrassate. The hind trochanter bears no teeth. ............................................................................25 25. (24) The male bears a prominent carina on the ventral side of the abdomen, and alongside the carina are two troughs formed by depressions in the abdomen on at least the three terminal segments. ..............................................................26 - If a carina is present, it is scarcely prominent, or it is limited to the terminal segment. ..............................................................................................................28 26. (25) The hind femur of the apterous male extends only to the tip of the genital segment. The last five segments of the female abdomen are concave on the dorsal surface. The right paramere of the male is arched and has a prominent ventral swelling near the base (Fig. 6.178). Length: 4.2 to 4.5 mm. ..............................................................................Rhagovelia viriosa Bacon, 1956 (Peru). - The hind femur of the apterous male extends well beyond the tip of the genital segment. The last five segments of the female abdomen are convex or flat on the dorsal surface. ...............................................................................................27

Fig. 6.178 Right paramere of a male Rhagovelia viriosa in lateral view. Based on Bacon (1956).

Fig. 6.179 Right paramere of a male Rhagovelia callida in lateral view. Based on Bacon (1956). 27. (26) The hind femur bears no spines longer than the diameter of the posterior tibia. The hind femur of the male is abruptly narrowed shortly before the apex.

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The right paramere of the male is arched and has a slight dorsal swelling near the base (Fig. 6.179). Length: 3.4 to 4.1 mm. A winged form has not been described. ............................................................Rhagovelia callida Drake and Harris, 1935 (Peru). - The hind femur bears at least one spine clearly longer than the diameter of the posterior tibia. The hind femur of the male is more than half as long as the body and tapers gradually to the apex (Fig. 6.175). The hind trochanter of the male is black. The connexival margin of the male curves broadly outward. The fifth through seventh abdominal tergites of the male are usually shiny black. The hind femur of the female bears about seven ventral spines. The posterior tibiae of both males and females are armed with subequal teeth along their entire lengths. The mesonotum of the female is matt and lacks a shiny black area. Length: 3.1 to 4.5 mm. .............................................................Rhagovelia tenuipes Champion, 1898 pars (Mexico, Central America, Trinidad, Colombia, Venezuela, Ecuador, Peru, Argentina, Rio de Janeiro, Minas Gerais, Amazonas, Pará). Syn: Rhagovelia confusa Gould, 1931.

Fig. 6.180 Right paramere of a male Rhagovelia deminuta in lateral view. Based on Bacon (1956). 28. (25) The connexiva is margined in brown orange. The posterior femur is black or brown in the middle and yellow or brownish yellow on both proximal and distal ends. The right paramere of the male is small and has evenly curved margins without processes visible in lateral view, except for a small ventral tooth about 2/3 to 3/4 of the way from the base to the apex (Fig. 6.180). Length: 2.42 to 2.80. A macropterous form has not been described. ..........................................................................Rhagovelia deminuta Bacon, 1948 (Guyana). - The margins of the connexiva are not orange. The femur is light in color only at the proximal end, if at all. ...............................................................................29 29. (28) The middle coxae of the male and all coxae of the female are yellow. The hind femur of the male is six times as long as wide and bears seven to nine spines; that of the female is unarmed. The second antennal segment is shorter than the third. Only the anterior half of the pronotum is marked with yellow or orange. The seventh abdominal and genital tergites of the male are the only ones that are sometimes shiny. The last segments of the connexiva are oriented vertically, reflexed, and contiguous, completely hiding the fifth through seventh

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abdominal segments in dorsal view (Fig. 6.181). The pronotum and mesonotum are convex dorsally. The venter is bluish gray. The second tarsal segment of the middle leg is subequal to the third, or at least more than 4/5 as long. Length of winged specimens: 3.9 to 4.2 mm; apterous male: c. 3.2 mm; apterous female: 3.6 to 3.7 mm. ..............................................................................Rhagovelia janeira Drake, 1953 (Argentina, Minas Gerais, Rio de Janeiro, Santa Catarina, Rio Grande do Sul). - The middle coxae of the male and one or more coxae of the female are blackish (Fig. 6.182). ..........................................................................................30

Fig. 6.181 Rhagovelia janeira (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 30. (29) The fore-trochanter of the male is black and bears a small tooth directed apicad (Fig. 6.182). ............................................................................................31 - The fore-trochanter of the male does not bear a spur or small tooth directed apicad (Fig. 6.163). The length is shorter than 4.0 mm, or, if the apterous female is longer than 4.0 mm, then the connexiva are horizontally oriented or angled only slightly dorsad when the abdomen is not swollen with eggs. .........32 31. (30) The second and third segments of the antenna are subequal in length. The fore-trochanter of the male is black and bears a small tooth directed apicad (Fig. 6.182). All abdominal tergites are shiny in the center. There is a row of three or four ventral spines on the distal third of the hind femur of the male; the proximal spine is about half the width of the femur, and those distal to it decrease progressively in length toward the apex. The female has one to three spines on the ventral margin of the hind femur. The connexiva of the female is nearly vertical on its posterior half but does not thicken or converge strongly near its apex and may rotate to become horizontal when the female swells with eggs. Length of apterous male: 3.6 to 3.9 mm; macropterous male: c. 4.3 mm; apterous female: 4.0 to 4.4 mm; macropterous female: c. 4.6 mm. Color: black with a blue-gray pruinosity of variable extent, a transverse yellow-orange band across the anterior part of the pronotum, and pale yellowish on the proximal third of the first antennal segment, the anterior and posterior parts of the middle and hind acetabula, and the fore and hind coxae. ....................................................Rhagovelia turmalis Nieser and Polhemus, 1999 (Minas Gerais).

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- The second segment of the antenna is longer than the third. The anterior trochanter of the male is armed with a stout spur near the apex that is directed apicad. The mesonotum of the female is depressed on the posterior third. There is a dorsal carina on the midline of the abdomen of the female (Fig. 6.183). The second antenna segment is subequal to or longer than the third. Length: 2.3 to 3.0 mm. ............................................................................Rhagovelia paulana Drake, 1953 (Argentina, São Paulo, Minas Gerais, Santa Catarina).

Fig. 6.182 Rhagovelia turmalis: fore-trochanter of a male (upper left), hind femur of a male (upper left center), male proctiger (right center), apex of the male (lower left) and female abdomen (lower left center), and the male paramere (right). Based on Nieser and Polhemus (1999).

Fig. 6.183 Rhagovelia paulana (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 32. (30) The middle trochanters are yellow or white. The apterous male has a shiny black marking on the last abdominal tergite and a small blue-gray triangle on the anterior margin of the last abdominal sternite (Fig. 6.184). The second

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antennal segment is shorter than the third. Only the anterior half of the pronotum is marked with yellow or orange. Length: 2.5 to 3.0 mm. ...............................................................................Rhagovelia evidis Bacon, 1948 (Peru, Amazonas). - The middle trochanters are black or brown. The last abdominal sternite of the male bears no blue-gray triangle, and there are often shiny black markings on more than just one tergite. ..................................................................................33

Fig. 6.184 Right paramere of a male Rhagovelia evidis in lateral view. Based on Bacon (1948). 33. (32) Length: c. 2.2 to 2.3 mm. The hind femur of the male has five or six ventral spines. The hind tibia is 1.2 times as long as the hind femur. The seventh, sometimes the sixth, and the genital segments are shiny black dorsally. The connexival margins of the male are slightly sinuous, and the abdomen becomes narrower in its posterior half. The apical portion of the male paramere is almost triangular (Fig. 6.185). The posterior parts of the connexiva of the female converge strongly and almost completely cover the sixth and seventh abdominal segments with their setiferous processes. .........................................................................Rhagovelia thaumana Drake, 1958 (Argentina, Santa Catarina). - The length is greater than 2.5 mm. ..................................................................34

Fig. 6.185 The male paramere of Rhagovelia thaumana. Based on Nieser and Polhemus (1999). 34. (33) There are no more than three spines along the ventral margin of the male hind femur. The hind femur of the female is not armed with ventral spines, and either it is long and narrow, with a length:width ratio of 1.70:0.19, or the female abdomen is especially short and wide (Fig. 6.163). ...............................35

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- There are at least five spines along the ventral margin of the male hind femur (Fig. 6.186). The female abdomen is not especially short and wide, and the hind femur of the female is shorter and broader. ........................................................36

Fig. 6.186 Rhagovelia sabrina (left to right): hind femur of a male, male proctiger, paramere of a male, and the apex of the female abdomen in lateral view. Based on Drake (1958) and Nieser and Polhemus (1999). 35. (34) The margins of the male acetabula are dark, and the fore and hind coxae and trochanters are light brown or yellow. The abdomen of the female is remarkably short and wide (Fig. 6.163). The color is dull black with shiny areas covering the seventh abdominal and genital segments and pale coloration on the basal third of the first antennal segment and a central marking on the pronotum, which is obscured by pruinosity. The hind femur of the male bears one to three ventral spines. Length of apterous male: 3.0 to 3.2 mm; apterous female: 2.95 to 3.15 mm. Macropterous specimens have not been described. .......................................................Rhagovelia sbolos Nieser and Polhemus, 1999 (Minas Gerais). - The edges of the acetabula are yellow. There are one or two ventral spines on the male hind femur, the diameter of which is less than that of the middle femur. It is armed well apicad from the middle with a small spine followed by only two or three very small ones aligned toward the apex. The right paramere of the male is obviously truncate at the apex (Fig. 6.187). The connexiva on the posterior half of the abdomen of the female and sometimes part of the anterior half, as well, are oriented almost vertically. The hind femur of the female is not armed with ventral spines, and either it is long and narrow, with a length:width ratio of 1.70:0.19, or the female abdomen is especially short and wide. Length: 3.15 to 3.45 mm. ...........................................................................Rhagovelia modesta Bacon, 1956 (Rio de Janeiro, São Paulo). There is a similar species known only from Mexico and Central America: R. bisignata Bacon, 1948, distinguished most easily by the spines on the posterior femur of the female, which are absent in R. modesta. 36. (34) Length: c. 3.5 mm. The mesonotum of the male has a shiny, heartshaped area covering most of it. The central area on the dorsal surface of the female mesonotum is shiny black. The connexiva of the female are oriented

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horizontally or only slightly inclined dorsad (Fig. 6.186). There are at least five spines along the ventral margin of the male hind femur but no spines on the hind femur of the female. .............................................................................Rhagovelia sabrina Drake, 1958 (West Indies, Minas Gerais, Paraná, Santa Catarina, São Paulo, Rio de Janeiro). - The length of the male is no more than about 3 mm, while the length of the female may reach c. 3.3 mm. ..............................................................................37

Fig. 6.187 Right paramere of a male Rhagovelia modesta in lateral view. Based on Bacon (1956). 37. (36) The apex of the hind tibia is armed with a spur, at least in the apterous male. The hind femur of the apterous male is slightly incrassate and armed with one long spine inserted 3/5 of its length from its joint with the trochanter followed by 8 to 10 much smaller spines, which decrease in size toward the apex. The second segment of the antenna is longer than the third. The apterous female has shiny black spots on the dorsum of the last four or five abdominal segments (Fig. 6.188). The second tarsal segment of the middle leg is 4/5 or less as long as the third. Length: 3.0 to 3.2 mm. .........................................................................Rhagovelia angustipes Uhler, 1894 (Mexico, Central America, West Indies, Venezuela, Ecuador). - There is no spur at the apex of the male hind tibia. There are shiny black markings on no more than the last three abdominal tergites. The hind femur of the female has at least six ventral spines. The female connixiva are horizontally oriented or angled only slightly dorsad. ............................................................38

Fig. 6.188 Rhagovelia angustipes (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

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38. (37) The central area on the dorsum of the seventh abdominal segment is usually shiny, and a smaller shiny spot is sometimes present on the sixth segment, as well. The second segment of the male antenna is longer than the third, but that of the female is shorter than the third. At about mid-length of the hind femur of the male, there is a large spine followed by a row of much smaller spines, which decrease continually in length toward the apex. The coxae and trochanters of the fore and hind legs are testaceous. Length: 2.6 to 3.2 mm. ...................................................................................Rhagovelia zela Drake, 1959 (West Indies, Argentina, Rio de Janeiro, Santa Catarina). - There are shiny central areas on the dorsal surfaces of the fifth through seventh abdominal segments. The hind tibia of the male is 1.1 times as long as the hind femur. There are nine ventral spines on the hind femur of the male and about six on the hind femur of the female. The width of the abdomen does not decrease on its posterior half. The connexival margin of the male curves outward and is not sinuate. The second antennal segment is subequal or slightly longer than the third. The color is mainly brownish black with fine golden setae. The legs are black with dull yellow coxae and fore and hind trochanters and fine golden setae. The ventral margin of the male paramere is only slightly concave (Fig. 6.189). Length of apterous male: 2.7 to 2.9 mm; macropterous male: c. 3.3 mm; apterous female: c. 3.1 to 3.2 mm; macropterous female: 3.8 to 3.9 mm. .............................................Rhagovelia rivulosa J. T. and D. A. Polhemus, 1985 (Minas Gerais, Santa Catarina). 39. (7) The ventral sutures on the abdomen are modified to form broad, glabrous channels (Fig. 6.190). The hemielytra of winged forms each have two small cells that do not extend onto the distal half. .......................................................40 - The ventral sutures on the abdomen are simple and not notably modified (Fig. 6.191). The two distal cells on the hemielytron extend onto its distal half. ......44

Fig. 6.189 The left paramere of a male Rhagovelia rivulosa. Polhemus and Polhemus (1985).

Based on

40. (39) The connexiva of apterous females converge strongly and become adpressed to each other posterior to the third abdominal segment. They are folded inward above the distal segments of the abdomen, concealing the segments from dorsal view. Length: 2.75 to 3.0 mm. The color is mainly blackish. The entire anterior portion of the pronotum, including a short, transverse orange band, is coated by a frosty white pubescence (Fig. 6.190). ...................................................Rhagovelia pacayana Drake and Carvalho, 1955 (Peru).

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- The connexiva of apterous females converge gradually and do not become adpressed on each other posterior to the third abdominal segment. All abdominal tergites are visible in dorsal view (Fig. 6.192). ................................41

Fig. 6.190 Rhagovelia pacayana: habitus of a female in dorsal view without appendages (left) and the male paramere (right). Based on Polhemus (1997). 41. (40) Measured along the midline, the exposed portion of the mesonotum is clearly longer than the pronotum (Fig. 6.192). ..................................................42 - Measured along the midline, the exposed portion of the mesonotum is equal to or shorter than the pronotum (Fig. 6.193). .........................................................43 42. (41) On the fourth through sixth abdominal sternites of the male, there is a low median carina, and the seventh sternite has a median longitudinal sulcus. The length and width of the eighth abdominal tergite of the female are subequal (Fig. 6.192). The posterior trochanter of the male is armed with several dark teeth. Length: 3.67 to 4.45 mm. ...............................................................................Rhagovelia vivata Bacon, 1948 (Peru, Bolivia). - There is no median carina on the fourth through sixth abdominal sternites of the male, and the seventh sternite has no median longitudinal sulcus. The eighth abdominal tergite of the female is shorter than wide. The posterior trochanter of the male is unarmed. The male paramere is truncate at the apex (Fig. 6.194). Length: 3.40 to 4.84 mm. A winged form has not been described. ...........................................................................Rhagovelia torquata Bacon, 1948 (Peru). 43. (41) In dorsal view, the male parameres extend posteriad beyond the apex of the proctiger. The hind trochanter of the male bears many small dark teeth. The eighth abdominal segment of the female lies in a horizontal plane and does not appear to be retracted into the seventh sternite (Fig. 6.193). Length: 4.0 to 4.8 mm. ............................................................................Rhagovelia abrupta Gould, 1934 (Peru). Syn: Rhagovelia hungerfordi Gould, 1933 (preoccupied).

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- In dorsal view, the male parameres do not extend posteriad beyond the apex of the proctiger and are not visible in dorsal view. The hind trochanter of the male does not bear small dark teeth. The eighth abdominal segment of the female lies at a 45º angle from the horizontal plane and appears retracted into the seventh sternite, making the abdomen appear truncate in lateral view. Length: 4.0 to 4.8 mm. The third antennal segment of the male is not wider than the second. The male paramere is thick and bluntly rounded at the apex (Fig. 6.195). Length: 3.30 to 4.25 mm. ................................................................................Rhagovelia trista Gould, 1931 (Ecuador, Peru, Argentina,).

Fig. 6.191 Rhagovelia trianguloides: apex of the female abdomen in dorsal view (upper center), hind femur of a male (left), male proctiger (right), and the male paramere (lower center). Based on Nieser and Polhemus (1999).

Fig. 6.192 Rhagovelia vivata (left to right): right paramere in lateral (above) and dorsal view (below); and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

Fig. 6.193 Rhagovelia abrupta (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

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Fig. 6.194 Right paramere of a male Rhagovelia torquata in lateral view. Based on Bacon (1948).

Fig. 6.195 Rhagovelia trista (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 44. (39) The dorsal abdominal carinae of winged females appear long and extend at least to the posterior margin of the second abdominal tergite (Fig. 6.164). .................................................................................................................45 - A dorsal abdominal carina is not evident on the second and third abdominal tergites (Fig. 6.196). ...........................................................................................47 45. (44) The middle femur of the female lacks a medial transverse constriction, but it has a thin section and a section of lighter colored integument on the ventral side near the mid-length. The male paramere is acutely pointed at the apex (Fig. 6.197). Length of wingless male: c. 3.8 mm; wingless female: c. 4.3 to 4.4 mm. ..................................................................Rhagovelia antioquiae Polhemus, 1997 (Colombia). - There is a medial transverse constriction on the middle femur of the female. The male paramere is blunt at the apex (Fig. 6.164). .........................................46 46. (45) The color is mainly orange brown. The male paramere forms a short, adpressed process at the apex (Fig. 6.198). Length of the wingless male: c. 3.7 to 3.8 mm; wingless female: 3.8 to 3.9 mm; winged male: 4.7 to 4.8 mm; winged female: 5.7 to 5.8 mm. ....................................................................Rhagovelia manzanoi Polhemus, 1997 (Colombia). - The color is mainly blackish grey (Fig. 6.164). Length of the wingless male: c. 3.8 to 3.9 mm; wingless female: 4.0 to 4.1 mm; winged male: 4.2 to 4.3 mm; winged female: 4.6 to 4.7 mm. .............................................................................Rhagovelia cali Polhemus, 1997 (Colombia).

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Fig. 6.196 Abdomen of a female Rhagovelia triangula in dorsal view (left), hind femur of a male (upper center), male proctiger (lower center), and male paramere (right). Based on Nieser and Polhemus (1999) and Moreira and Ribeiro (2009).

Fig. 6.197 The paramere of a male Rhagovelia antioquiae. Based on Polhemus (1997).

Fig. 6.198 The paramere of a male Rhagovelia manzanoi. Based on Polhemus (1997). 47. (44) There is a small, orange spot near the middle of the pronotum, and the rest of the dorsum is uniformly blackish. Sometimes, the third antennal segment has a greater diameter than the second and fourth segments. Some females possess a dorsal carina on the abdomen (Fig. 6.196). ........................................48 - There are distinctly contrasting markings on both the pronotum and the connexiva. The third antennal segment is always the same diameter as the second and fourth segments. The female always lacks a dorsal carina on the abdomen (Fig. 6.199). ........................................................................................51 48. (47) The connexiva have orange bands. There is no dorsal carina on the female abdomen, and the first abdominal segment is not notably inflated. The hind femur of the male has an irregular row of small black warts on its proximal half, a long spine near the mid-length, and a distal row of small spines that become progressively smaller toward the apex (Fig. 6.191). The third antennal segment shows no tendency to be slightly more robust than the other segments.

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Length of apterous male and female: 3.3 to 3.6 mm; macropterous specimens: 4.1 to 4.5 mm. Color: blackish with orange yellow and yellowish brown markings. ............................................Rhagovelia trianguloides Nieser and Polhemus, 1999 (Minas Gerais). - The connexiva are uniformly dull gray. There is a dorsal carina on some segments of the female abdomen, or, if none is evident, the first abdominal segment is obviously inflated (Fig. 6.196). The third antennal segment of some specimens shows a tendency to be more robust than the other segments. .........49

Fig. 6.199 Rhagovelia itatiaiana male: habitus without appendages in dorsal view (upper left), antenna (left center), apex of the abdomen in lateral view (lower left), hind leg (right center), and male paramere (right). Based on Polhemus (1997). 49. (48) The hind femur of the male lacks spines on its basal half. The third antennal segment of the male is relatively slender. There is a sigmoid indentation near the lateral edges of the seventh abdominal segment of the female (Fig. 6.196). Length of apterous male: 2.8 to 3.0 mm. ..........................................................................Rhagovelia triangula Drake, 1953 (São Paulo, Rio de Janeiro, Minas Gerais). - The hind femur of the male has a row of scales on the ventral side of its basal half. The third antennal segment of the male is obviously widened. There is no sigmoid indentation near the lateral edges of the seventh abdominal segment of the female (Fig. 6.162). ......................................................................................50 50. (49) The first abdominal segment of the female is considerably inflated; in lateral view, it projects over the metanotum. Median longitudinal carinae are absent on the second through fourth abdominal tergites of the female (Fig. 6.162). ..................................................................................Rhagovelia agra Drake, 1957 (Rio de Janeiro). - The first abdominal segment of the female is not inflated enough to project above the metanotum in lateral view. There is a prominent median carina on the

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second through fourth tergites of the female. The third antennal segment of the male is flattened and dilated, making it wider than the second. The male paramere is hooked at the apex (Fig. 6.200). Length: c. 3.6 mm. ...............................................................................Rhagovelia lucida Gould, 1931 (Argentina, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul).

Fig. 6.200 Rhagovelia lucida male: paramere (left) and proctiger (right). Based on Polhemus (1997). 51. (47) The seventh abdominal sternite lacks a tuft of setae near the middle. The outline of the male paramere is strongly sigmoid (Fig. 6.201). Length: c. 4.5 mm. ..........................................................................Rhagovelia accedens Drake, 1957 (São Paulo, Rio de Janeiro, Minas Gerais). - The seventh abdominal sternite has a tuft of setae near the middle (Fig. 6.199). The total length is less than 4 mm. .....................................................................52

Fig. 6.201 Rhagovelia accedens male: paramere (left) and proctiger (right). Based on Polhemus (1997).

Fig. 6.202 Right paramere of a male Rhagovelia trepida in lateral view. Based on Bacon (1948).

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52. (51) The color of the mesonotum is uniformly dark. The posterior margin of the pronotum is yellow. The posterior tibia of the apterous male usually bears an enlarged tooth 1/4 of the distance from the apex. The male paramere curves slightly near the apex (Fig. 6.202). Length: 3.72 to 3.78 mm. .............................................................................Rhagovelia trepida Bacon, 1948 (Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul). - The mesonotum is dark with dark yellow or orange markings in the center, which may be extensive enough to extend for its whole length (Fig. 6.199). ....53 53. (52) The seventh abdominal tergite of the male and the seventh and eighth of the female have orange markings. The seventh abdominal tergite of the female is elevated so that it partially covers the eighth. The male paramere is elongate and sinuous (Fig. 6.203). .........................................................Rhagovelia macta Drake and Carvalho, 1955 (Minas Gerais, Rio de Janeiro). - The color of all abdominal tergites of both males and females is uniform. The seventh abdominal tergite of the female is not elevated and does not partially cover the eighth. There is a median ventral carina on the anterior half of the last abdominal segment of the male. The male paramere is bluntly triangular and not at all recurved at the apex (Fig. 6.199). Length: 3.15 to 3.40 mm. ..........................................................................Rhagovelia itatiaiana Drake, 1953 (Rio de Janeiro).

Fig. 6.203 Male paramere of Rhagovelia macta. Based on Polhemus (1997).

Fig. 6.204 Rhagovelia bocaina: habitus of a male (middle left) and a female (lower left) in dorsal view without legs; fore (upper center), middle (upper right), and hind leg of a male (middle right); hind leg of a female (lower right), and male paramere (upper left). Based on Moreira and Ribeiro, 2009.

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54. (6) A small part of the mesonotum of apterous specimens is left exposed beyond the pronotum. There are elevated projections on the pronotum of the macropterous female. At midlength, the middle femur of the female is flattened and blade-like. The eighth abdominal sternite of the female bears dense tufts of setae on its lateral margins (Fig. 6.157). ....................................................Rhagovelia aiuruoca Moreira and Ribeiro, 2009 (Minas Gerais, São Paulo, Rio de Janeiro). - The mesonotum of the apterous form is completely covered by the pronotum, and there are no elevated projections on the pronotum of the female. At midlength, the middle femur of the female is not flattened or blade-like. The eighth abdominal sternite of the female lacks dense tufts of setae on its lateral margins (Fig. 6.204). ..........................................................................................55

Fig. 6.205 Rhagovelia impensa (left to right): apex of the pronotum in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Matsuda (1956). 55. (54) There are usually many but occasionally only a few large black denticles on the posterolateral borders of the seventh abdominal segment where it surrounds the genital cavity. In some species, the middle femur of the female is ventrally flattened on the basal third. The apex of the pronotum of the macropterous female is broadly angulate and bears no posterior elevated projection (Fig. 6.205). .......................................................................................56 - There are no large black denticles on the posterolateral borders of the seventh abdominal segment where it surrounds the genital cavity (Fig. 6.204). ............68

Fig. 6.206 Rhagovelia humboldti male: paramere (left) and proctiger (right). Based on Polhemus (1997).

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Fig. 6.207 Rhagovelia cauca: habitus of a female without appendages in dorsal view (upper left), apex of the female abdomen in lateral view (lower left), pronotum of a winged female in dorsal (upper center) and lateral view (lower right center), male paramere (lower left center), and hind leg of a male (right). Based on Polhemus (1997). 56. (55) There is no elevated process at the posterior apex of the female pronotum. The connexiva of wingless females do not strongly converge posterior to the third abdominal tergite (Fig. 6.206). Length of wingless male: c. 4.0 to 4.1 mm; wingless female: c. 4.0 to 4.1 mm. Color: black on the dorsum with brown or yellow brown on the bases of the antennae, connexival margins, and anterior and posterior parts of the pronotum. ...................................................................Rhagovelia humboldti Polhemus, 1997 (Venezuela). - There is an elevated process at the posterior apex of the apterous female pronotum. The connexiva of wingless females sometimes strongly converge posterior to the third abdominal tergite. The fore-tibia of the male is expanded, resulting in the distal half of its ventral margin being concave. The hind tibia is considerably longer than the hind femur (Fig. 6.207). The distal lobe of the male paramere is considerably elongated. Macropterous individuals have dorsal abdominal carinae, the posterior sections of which are parallel. . .....................57 57. (56) The connexiva are folded over the fourth through seventh abdominal segments with their margins in contact or almost touching above the fourth tergite and then slightly diverging posteriad, concealing the sixth tergite from dorsal view. The middle portions of the seventh abdominal segment can be seen only as a narrow strip between the adpressed margins of the connexiva (Fig. 6.205). .................................................................................................................58 - The connexiva do not extend over the abdomen, or if they are folded, they are not closely adpressed, leaving ample space between their margins to reveal the entire length of the fourth tergite to dorsal view. At most, only the lateral

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margins of the fifth through seventh abdominal segments are covered by folded portions of the connexiva (Fig. 6.207). The posterior part of the central portion of the seventh abdominal segment is not tumescent, so the whole tergite is flat and below the level of the connexival margins. .................................................59 58. (57) The ground color is orange brown, sometimes with dark fuscous tergites (Fig. 6.208). There is a moderately dense coat of erect golden pubescence on the thorax and abdomen, which is denser on the male. Length: c. 5.2 mm. The pronotum of the winged female bears a strong, cylindrical horn at the apex, which is hairy and bends sharply downward in the apical third. The connexiva of the apterous female are broad and strongly reflexed so that their outer edges meet above the three apical abdominal tergites. Their apices flare from above the tergites and bear dense tufts of long, brown, spine-like hairs. ................................................................................Rhagovelia citata Drake, 1953 (Panama, Venezuela). - The color is blackish gray with a yellowish anterior band on the pronotum and strongly contrasting yellow markings on the margins of the connexiva. There are no minute, black, conical setae on the proepisternum and venter of the abdomen, or such setae are very sparce (Fig. 6.205). Length: 4.36 to 5.30 mm. ...........................................................................Rhagovelia impensa Bacon, 1956 (Peru).

Fig. 6.208 Rhagovelia citata (left to right): habitus of a female without appendages in dorsal view, apex of the female abdomen in lateral view, and male paramere. Based on Polhemus (1997). 59. (57) The length is always less than 6.5 mm. ................................................60 - The length is always greater than 6.5 mm. ......................................................66 60. (59) The entire eighth abdominal tergite is clothed in long, erect black setae (Fig. 6.209). Length of wingless male: c. 4.9 to 5.0 mm; wingless female: c. 5.3 to 5.4 mm; winged male: c. 5.1 mm; winged female: c. 5.2 to 5.3 mm. ..................................................................Rhagovelia yanomamo Polhemus, 1997 (Venezuela, Guyana). - If long, black setae are present on the eighth abdominal segment at all, they are confined to the anterior margin or to tufts along the lateral margins (Fig. 6.207). .................................................................................................................61

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Fig. 6.209 Rhagovelia yanomamo: habitus of a female without appendages in dorsal view (upper left), apex of the female abdomen in lateral view (lower left), pronotum of a winged female in dorsal (upper center) and lateral view (lower center), and the male paramere (right). Based on Polhemus (1997). 61. (60) The insccts are uniformly orange brown, but there is a band across the anterior part of the pronotum slightly lighter than the rest of the dorsum (Fig. 6.207). .................................................................................................................62 - The insccts are mainly dark brown, blackish, or a mixture of black and brown, usually with strongly contrasting yellow or orange brown margins on the connexiva and a band across the anterior part of the pronotum (Fig. 6.210). ....63

Fig. 6.210 Rhagovelia macarena: habitus of a female without appendages in dorsal view (upper left), male antenna (lower left), apex of a female abdomen in lateral view (lower right center), and male paramere (right). Based on Polhemus (1997). 62. (61) The posterior margin of the seventh abdominal tergite bears a thick tuft of long, black setae. The ventral border of the middle femur is concave, and the basal part of the femur is very flat (Fig. 6.207). Length of wingless male: c. 4.7 to 4.8 mm; wingless female: c. 5.5 mm; winged male: c. 5.4 mm; winged female: c. 5.5 to 5.6 mm. ..................................................................Rhagovelia cauca Polhemus, 1997 pars (Colombia).

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- The posterior margin of the seventh abdominal tergite does not bear a thick tuft of black setae, although there may be a thin fringe of fine, black setae. The ventral border of the middle femur is not concave, and the basal part of the femur tends, at most, to be weakly flattened (Fig. 6.211). The anterolateral surfaces of the first seven abdominal tergites are not at all or only slightly pruinose. Length of wingless male: c. 5.4 mm; wingless female: c. 6.2 to 6.3 mm; winged male: c. 5.7 to 5.8 mm; winged female: c. 6.2 to 6.3 mm. .......................................................................Rhagovelia malkini Polhemus, 1997 (Colombia).

Fig. 6.211 Rhagovelia malkini: habitus of a female without appendages in dorsal view (upper left), apex of the female abdomen in lateral view (lower left center), pronotum of a winged female in dorsal (upper center) and lateral view (lower center), male proctiger (upper right), and male paramere (lower right). Based on Polhemus (1997). 63. (61) A fringe of long, dark setae is absent from the posterior margin of the seventh abdominal segment, but the margin may be concealed by groups of black setae on the infolded connexival segments. At most, the setae on the margin will be short and adpressed. The posterolateral angles of the terminal segments of the connexiva fold inward to cover the posterior margin of the seventh abdominal tergite and the base of the eighth (Fig. 6.210). Length of wingless male: c. 5.5 mm; wingless female: c. 5.3 to 5.4 mm; winged male: c. 5.3 to 5.4 mm; winged female: c. 6.1 to 6.2 mm. ...................................................................Rhagovelia macarena Polhemus, 1997 (Colombia). - There is always a fringe of long, dark setae on the posterior margin of the seventh abdominal segment (Fig. 6.212). ..........................................................64 64. (63) There are small, black denticles on the anteromedial surface of the proepisternum, where it contacts the rostrum (Fig. 6.212). The fourth abdominal tergite is not greatly narrowed; its length is 1.4 times its width. Length of wingless male: c. 5.2 mm; wingless female: c. 5.7 to 5.8 mm. Specimens with wings have not been reported. ..........................................................................Rhagovelia perija Polhemus, 1997 (Colombia).

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- There are no small, black denticles on the anteromedial surface of the proepisternum, where it contacts the rostrum (Fig. 6.205). ...............................65

Fig. 6.212 Rhagovelia perija: habitus of a female without appendages in dorsal view (upper left), apex of a female abdomen in lateral view (lower left), male proctiger (center), male hind femur in dorsal view (right), and male paramere (lower left center). Based on Polhemus (1997). 65. (64) The posterior margin of the seventh abdominal tergite bears a thick tuft of long, black setae. The tuft conceals the basal portion of the eighth tergite. There are no tufts of black setae at the apices of the adjacent connexiva. The pronotum lacks a median carina on the posterior lobe. The dorsal coloration is usually somewhat orange brown with the margins of the connexiva not obviously lighter (Fig. 6.207). Length of wingless male: c. 4.7 to 4.8 mm; wingless female: c. 5.5 mm; winged male: c. 5.4 mm; winged female: c. 5.5 to 5.6 mm. ..................................................................Rhagovelia cauca Polhemus, 1997 pars (Colombia). - The posterior margin of the seventh abdominal tergite bears only a thin fringe of black setae, which does not conceal the basal portion of the eighth tergite. There are tufts of black setae at the apices of the adjacent connexiva, which may cover the fringe in dorsal view. The dorsal coloration is blackish or greyish with the margins of the connexiva a much lighter yellow (Fig. 6.205). Length: 4.36 to 5.30 mm. ...........................................................................Rhagovelia impensa Bacon, 1956 (Peru). 66. (59) There are two rows of 15 black teeth on the posterior margin of the hind tibia of the male, which increase in size toward the apex. There are about 13 short, black teeth on the jugum and 10 on the adjacent proepisternum. The third and fourth abdominal segments are subequal in length. The apex of the distal cone of the male proctiger appears polygonal, and the lateral lobes on the basal section are moderately developed (Fig. 6.213). The dorsal margin of the male

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paramere is bent at an angle of about 130º. The eighth tergite of the female abdomen has tufts of black setae at the posterolateral corners and a median longitudinal carina bordered by flattened and depressed areas. The first abdominal tergite of the female is longer than the second, and the second and third are subequal in length. The hemielytra of macropterous specimens each have four closed cells. Length of apterous male: c. 6.9 mm; winged and wingless female: c. 8.2 to 8.3 mm. The dorsum is mainly brown with yellowish brown on a band across the pronotum, the margins of the connexiva, and legs. The dorsum is covered by short gold setae intermixed with a few long black setae. The winged male has not been described. ....................................................................Rhagovelia oporapa Padilla-Gil, 2009 (Colombia). - One or two rows of 21 teeth are present on the posterior margin of the hind tibia of the male (Fig. 6.214). .............................................................................67

Fig. 6.213 Rhagovelia oporapa: body of a female without appendages in dorsal view (upper left), pronotum of a macropterous female in dorsal (upper right) and lateral view (middle right), male hind femur (middle left), male paramere (lower left), and female abdomen in lateral view (lower right). Based on Padilla-Gil (2009). 67. (66) The proepisternum lacks denticles, but the jugum bears about 15 black ones. The apex of the distal cone of the male proctiger is triangular in shape, and its base has greatly developed lobes (Fig. 6.214). The dorsal margin of the male paramere is bent at an angle of about 100º. The eighth tergite of the female abdomen has tufts of yellow setae along the posterolateral margins. The first, second, and third abdominal tergites of the female are subequal in length.

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Length of apterous male: c. 6.9 mm; winged male: c. 7.4 to 7.5 mm; winged and wingless female: c. 8.0 to 8.2 mm. The dorsum and legs are mainly brown with yellowish brown on a band across the pronotum and the margins of the connexiva. The ventral surface is orange yellow. The dorsum is covered by short gold setae but lacks long black setae. .........................................................................Rhagovelia huila Padilla-Gil, 2009 (Colombia). - The proepisternum bears 4 or 5 black denticles, and the jugum bears more than 15. The apex of the distal cone of the male proctiger is rounded, and its base has relatively small lobes (Fig. 6.215). The dorsal margin of the male paramere is bent at an angle of about 115º. The eighth tergite of the female abdomen lacks tufts of setae near the posterolateral corners, but it has a distinct median longitudinal carina. The first abdominal segment of the female is shorter than the second, and the second and third are subequal in length. Length of apterous male: c. 7.0 to 7.1 mm; winged male: c. 7.3 to 7.4 mm; wingless female: c. 8.4 mm. The dorsum is mainly brown with yellowish brown on a band across the pronotum and the margins of the connexiva. The ventral surface is orange brown. The dorsum is covered by short gold setae intermixed with a few long black setae. ......................................................................Rhagovelia azulita Padilla-Gil, 2009 (Colombia).

Fig. 6.214 Rhagovelia huila (above, left to right): body of a female without appendages in dorsal view, pronotum of a macropterous female in dorsal view, male paramere, and (below, left to right): male femur, pronotum of a macropterous female in lateral view, and abdomen of a female in lateral view. Based on Padilla-Gil (2009). 68. (55) There is a carina along the dorsal midline of at least one abdominal sternite of the wingless female, but it may continue onto the other sternites (Fig. 6.216). The lengths of the species range from 3.9 mm to 5.5 mm. ...................69

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- There is no carina along the dorsal midline of the abdomen of the wingless female. The middle femur of the female is cylindrical and never flattened ventrally on the basal third (Fig. 6.217). The seventh abdominal tergite of the female bears lateral tufts of stiff black setae but no tufts are present on the proctiger, although it may have a fringe of black setae. .....................................74

Fig. 6.215 Rhagovelia azulita: pronotum of a macropterous female in dorsal (upper left) and lateral view (lower left), male femur (upper center), male paremere (lower center), and abdomen of a female in lateral view (right). Based on Padilla-Gil (2009).

Fig. 6.216 Rhagovelia gaigei: habitus without appendages (left) and male paramere (right). Based on Polhemus (1997).

Fig. 6.217 Rhagovelia elegans: right paramere in lateral view (left) and the apex of the hind tibia, sickle-shaped spur, and the tarsus of a female, shown diagrammatically (right). Based on Bacon (1956) and Matsuda (1956), who referred to the species by its synonym, Rhagovelia costalimai.

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69. (68) The color is fairly uniform reddish brown, orange-brown, or orange-red. The hind femur of the female is usually armed with five to seven spines along its posterior margin, and the male is armed with more (Fig. 6.218). ......................70 - The color is mainly black, sometimes with orange-brown markings. ............72 70. (69) The hind femur of the male is moderately swollen, so that it is about 3.5 times as long as wide. It bears a row of minute teeth on the basal 2/5 and two rows of spines on the apical 3/5, with eight or nine small spines in the anterior row and a posterior row with the largest spine at the basal end and eight or nine teeth decreasing progressively in length toward the apex of the femur. There are several small denticles on the hind trochanter. The hind femur of the female is only slightly swollen and about five times as long as wide. It bears a posterior row of one moderately large spine 3/5 of the way from the base to the apex followed by four to six smaller spinules toward the apex and sometimes three or four small spinules representing the anterior row. The carina along the dorsal midline of the apterous female extends from the third to the sixth segment (Fig. 6.216). The hemielytra of the macropterous form are blackish fuscous with darker veins. Length of male: c. 4.2 mm; female: c. 4.7 to 4.8 mm. Maximum width: c. 1.45 mm. Color: uniform reddish brown. ............................................................Rhagovelia gaigei Drake and Hussey, 1957 (Colombia). - The hind femur of the male bears only five or six spines on the posterior margin. The spines of the hind femur of both the male and the female show only a gradual decrease in size, and one of them is not considerably larger than the others (Fig. 6.218). .......................................................................................71

Fig. 6.218 Rhagovelia gorgona (left to right): head and thorax in lateral view showing the locations of circular pits, hind femur of a male, and male paramere. Based on Manzano et al. (1997). 71. (70) The ground color is orange brown without any markings on the pronotum or the margins of the female connexiva. The middle femur of the female is cylindrical. On the apical fourth of the female hind femur, there is a row of five short spines along the ventral margin, which decrease progressively in length. The male paramere is elongate and curves more than 45° (Fig. 6.219).

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The seventh abdominal tergite of the female bears lateral tufts of stiff black setae, but there are no tufts on the proctiger. Only apterous specimens have been described. Length: 4.4 to 5.0 mm. .........................................................................Rhagovelia henryi Polhemus, 1997 (Central America?, São Paulo, Rio de Janeiro). - The ground color is orange-red. The pronotum is reddish orange with a pale mid-dorsal line. The hind femur of both the male and the female is armed with a row of six spines along the posterior margin, beginning proximal to mid-length. The second spine is longest, and the apical four decrease progessively in length (Fig. 6.218). There is a mid-ventral carina on the first abdominal sternite of both males and females; this carina continues weakly onto the other abdominal segments of the male but is absent from the other segments of the female. Length of apterous male: 3.9 to 4.4 mm; apterous female: 3.9 to 4.3 mm. Length of macropterous male: 4.6 to 4.9 mm; macropterous female: 4.4 to 4.7 mm. ....................................Rhagovelia gorgona Manzano, Nieser, and Caicedo, 1997 (Colombia).

Fig. 6.219 Male paramere of Rhagovelia henryi. Based on Polhemus (1997).

Fig. 6.220 Rhagovelia tijuca: female habitus without appendages in dorsal view (left) and male paramere (right). Based on Polhemus (1997).

Fig. 6.221 Right paramere of a male Rhagovelia femoralis in lateral view. Based on Bacon (1956).

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72. (69) The color is mainly black with large, orange brown markings on the anterior part of the pronotum and on the margins of the connexiva of the female. There are only two or three tiny black spines along the distal part of the ventral margin of the female hind femur. The male paramere curves sharply beyond its mid-length. The female has tufts of setae at the posterolateral corners of the mesonotum (Fig. 6.220). Length of wingless male: c. 4.3 to 4.4 mm; apterous female: c. 4.8 to 4.9 mm. Macropterous forms have not been described. ..........................................................................Rhagovelia tijuca Polhemus, 1997 (Rio de Janeiro, São Paulo). - The insects are dark gray or blackish and heavily pruinose on the pronotum and underside. Extensive orange markings are lacking. The male paramere is not sharply hooked beyond its mid-length (Fig. 6.221). ....................................73 73. (72) The first through eighth abdominal sternites of the male usually have a median longitudinal carina. The minute, black, conical setae on the proepisternum are elongate and appear almost like bristles. The basal 3/5 of the male paramere is inflated on the dorsal surface, and the apical 2/5 is straight and finger-like (Fig. 6.221). The margins of the female connexiva do not have an inwardly curved, glabrous area adjacent to the third through fifth abdominal tergites. Length: c. 5.05 mm. ...................................................................Rhagovelia femoralis Champion, 1898 (Panama, Colombia). - The first through sixth abdominal sternites of the male lack a median longitudinal carina. The paramere of the male is evenly curved to form a small excavation on the inner side of the curve (Fig. 6.222). The margins of the female connexiva are considerably thickened adjacent to the third through fifth abdominal tergites. Length of apterous male: 4.4 to 4.5 mm; apterous female: 4.8 to 4.9 mm. Winged forms have not been reported. ........................................................................Rhagovelia baconi Polhemus, 1997 (São Paulo). 74. (68) There is a strongly curved spur at the apical end of the hind tibia (Fig. 6.217). .................................................................................................................75 - If any spur at all is present at the apex of the hind tibia, it is straight (Fig. 6.204). .................................................................................................................77

Fig. 6.222 Male paramere of Rhagovelia baconi. Based on Polhemus (1997). 75. (74) The apical genital segment terminates in a spine-like process (Fig. 6.223). Length: 4.86 to 5.05 mm. ....................................................................Rhagovelia uncinata Champion, 1898 (Central America, French Guiana).

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- The apical genital segment is rounded or triangular but not spine-like at the apex (Fig. 6.145). ...............................................................................................76

Fig. 6.223 Rhagovelia uncinata (left to right): right male paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956). 76. (75) The hind femur bears three rows of irregular spines and has one large spine near the center that is located dorsal to the rows. The hind tibia is armed for its entire length. The ventral margin of the male right paramere has a semicircular emargination near the middle, seen best in lateral view (Fig. 6.145). Length: c. 5.05 mm. ..............................................................................Rhagovelia merga Bacon, 1956 (Panama; not yet reported from South America). - The hind femur is armed with a row of about six spines. The hind tibia is armed with knob-like teeth on its proximal 2/3 and with its sickle-shaped spur at the apex (Fig. 6.217). Length: 5.2 to 5.5 mm. ..............................................................................Rhagovelia elegans Uhler, 1894 (Central America, West Indies, Venezuela, Colombia, French Guiana, Ecuador, Amazonas, Pará, Rio de Janeiro). Syn: Rhagovelia insularis Champion, 1898; Rhagovelia costalimai Drake, 1948; Rhagovelia trinidalis Drake, 1948. 77. (74) The hind tibia of the male is cylindrical and sinuate; it bears two rows of about 20 small black teeth parallel to its posterior margin. Those in the ventral row are small and similar in size. Those in the dorsal row progressively enlarge in the apical ¼ of the tibia. There is a short conical spur at the apex. The hind tibia of the female is straight and bears a single row of about 20 small, black, subequal teeth on the inner face. The hind femur of the male is incrassate and bears a row of six to eight small teeth on its basal quarter and two parallel rows of teeth along its apical 2/3 (Fig. 6.224). The wingless female bears a dense tuft of long black setae, and the lateral portions of the proctiger bear fringes of such setae. Length of apterous male: c, 6.5 mm; winged male: c. 6.6 to 6.7 mm; apterous female: c. 7.25 mm. The winged male has been described, but winged females are unknown. Polhemus (1997) speculated that the winged female, when found, might have a posterior process on the pronotum. .......................................................................Rhagovelia roldani Polhemus, 1997 (Colombia).

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- The teeth on the hind tibia of the male are either all small and subequal, or some are considerably larger than the others without enlarging progressively along the row (Fig. 6.204). .................................................................................78

Fig. 6.224 Rhagovelia roldani: habitus of a wingless female without appendages (upper left), hind femur of a male (lower left center), proctiger of a male (right center), and the male paramere (right). Based on Polhemus (1997).

Fig. 6.225 Rhagovelia traili (left to right): habitus, right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Polhemus (1997) and on illustrations labelled Rhagovelia perfidiosa by Bacon (1956) and Matsuda (1956). 78. (77) There are small teeth of subequal size, usually peg-like, covering parts of the hind tibia, together with a single long tooth, directed posteriad, at the apex (Fig. 6.225). ........................................................................................................79

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- The denticulation of the posterior tibia of the male includes some teeth notably larger than the others, usually located on the distal part of the tibia but proximal to the apex, and a long apical spur (Fig. 6.204). Some of the larger teeth may resemble conical spikes. .....................................................................................85 79. (78) There are no teeth on the hind trochanter, but occasionally, there might be one small black peg present. The posterior femur bears two parallel rows of teeth near the apex, along the posterior margin. The size of the teeth decreases regularly apicad. The margins of the male paramere are sinuous (Fig. 6.226). The length is shorter than 3.5 mm. The color is mainly uniform, from reddish brown to black, but with a small dark yellow spot on the anterior surface of the pronotum. ................................................................................Rhagovelia zeteki Drake, 1953 (Panama, Colombia). - At least two or three small teeth are on the hind trochanter. ...........................80

Fig. 6.226 Male paramere of Rhagovelia zeteki. Based on Polhemus (1997). 80. (79) In addition to several small subequal teeth, the hind trochanter bears one tooth that is much larger than the others. .....................................................81 - There are two to four small teeth on the posterior trochanter, which are nearly equal in size. .......................................................................................................82 81. (80) There are no black denticles on the lateral parts of the mesosternum. A dorsal row of spines on the hind femur encompasses two especially large teeth separated by four or five smaller tooth-like spines. One of the large teeth is located 1/3 of the way from the base, and the longer one is beyond midlength; a series of small spines of decreasing size runs toward the apex. The anterior band on the pronotum of the female is not distinctly separated from the disc of the pronotum. The male paramere is sharply angled near mid-length and truncate at the apex (Fig. 6.227). The dorsum of the first genital segment of the female is longer than wide. Length: reported by different authors to be 3.56 to 3.69 mm and 4.35 to 5.40 mm. .............................................................................Rhagovelia sinuata Gould, 1931 (Ecuador, Peru). - Black denticles are present on the lateral parts of the mesosternum. A dorsal row of spines on the hind femur encompasses two especially large teeth separated by two smaller teeth. The male paramere is notched near mid-length and broadly rounded at the apex (Fig. 6.228). ..........................................................................Rhagovelia rubra Polhemus, 1997 (Colombia).

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Fig. 6.227 Rhagovelia sinuata (left to right): right paramere in lateral view, the basal and apical segments of the abdomen of a macropterous female, and the hind femur. Based on Bacon (1956) and Matsuda (1956).

Fig. 6.228 Male paramere of Rhagovelia rubra. Based on Polhemus (1997). 82. (80) There are minute black denticles along the inner margins of the proepisternum. ....................................................................................................83 - There are no minute black denticles on the proepisternum. ............................84 83. (82) The lateral parts of the fifth and sixth abdominal segments are pruinose, and the central parts have ovate brown spots. There is a dark yellowish spot in the center of the seventh abdominal tergite. The hind femur has a long, pointed tooth on the dorsal surface and two rows of teeth along the posterior edge. The apex of the male paramere is truncate (Fig. 6.225). Length: 2.98 to 3.52 mm. .................................................................................Rhagovelia traili White, 1879 (Peru, Venezuela, French Guiana, Surinam, Amazonas, Pará, Roraima). Syn: Rhagovelia perfidiosa Bacon, 1948. - The fifth and sixth abdominal tergites are dull black without ovate brown spots. The seventh abdominal tergite is uniformly dull black. The hind femur does not have a long, pointed tooth on the dorsal surface. The apex of the male paramere is slightly hooked (Fig. 6.229). ..................................................................Rhagovelia plaumanni Polhemus, 1997 (Paraguay, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul).

Fig. 6.229 Male paramere of Rhagovelia plaumanni. (1997).

Based on Polhemus

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Fig. 6.230 Male paramere of Rhagovelia williamsi. Based on Polhemus (1997).

Fig. 6.231 Right paramere of a male Rhagovelia amazonensis in lateral view. Based on Bacon (1956). 84. (82) The hind femur has one moderately long spine on the basal third, a larger spine near its mid-length, and a row of spines decreasing in size on its apical half. The male paramere is distinctly hooked at its apex (Fig. 6.230). ..........................................................................Rhagovelia williamsi Gould, 1931 (Ecuador). - The hind femur does not have a moderately long spine on the basal third but has only a larger spine near its mid-length, and a row of spines decreasing in size on its apical half. The dorsum of the first genital segment of the female is wider than long. The apex of the male right paramere forms a thickened hook (Fig. 6.231). Length: 3.20 to 3.65 mm. ....................................................................Rhagovelia amazonensis Gould, 1931 (Amazonas, Mato Grosso). 85. (78) The hind trochanter is not armed with any teeth (Fig. 6.232). ............................................................Rhagovelia equatoria Polhemus, 1997 pars (Venezuela). Also see Couplet 97. - The hind trochanter bears at least one tooth (Fig. 6.204). ..............................86

Fig. 6.232 Male paramere of Rhagovelia equatoria. Based on Polhemus (1997). 86. (85) The hind trochanter bears only one tooth, which is larger than any tooth on the hind tibia, or it bears several teeth, among which one is much larger than the others (Fig. 6.233). .......................................................................................87

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- The hind trochanter bears one tooth that is subequal to or smaller in size than one or more on the hind tibia, or it bears several subequal teeth (Fig. 6.204). .........91

Fig. 6.233 Rhagovelia whitei male: right paramere in lateral view (upper left), a paramere of another specimen (lower left), and the hind leg (right). Paramere at upper left based on Bacon (1956), and the rest, on Polhemus (1997). 87. (86) There are no minute denticles on the proepisternum. The male paramere forms a small subapical barb (Fig. 6.234). ...........................................................................Rhagovelia castanea Gould, 1931 (Ecuador). - There are minute, black, conical setae on the proepisternum and sometimes other parts of the underside. ...............................................................................88 88. (87) There are minute, black, conical setae in a ventrolateral cluster on the posterior third of the apical abdominal segment (Fig. 6.235). ...........................89 - The ventrolateral setae on the apical abdominal segment are all hair-like. ....90

Fig. 6.234 Right paramere of a male Rhagovelia castanea in lateral view. Based on Bacon (1956).

Fig. 6.235 Rhagovelia palea (left to right): right paramere in lateral view and the basal and apical segments of the abdomen of a macropterous female. Based on Bacon (1956) and Matsuda (1956).

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Fig. 6.236 Rhagovelia jubata male: hind femur (left) and paramere (right). Based on Polhemus (1997).

Fig. 6.237 Right paramere of a male Rhagovelia scitula in lateral view. Based on Bacon (1956). 89. (88) The middle trochanter is dark brown or black. The hind femur of the male bears teeth of varying sizes (Fig. 6.236). Length: 4.55 to 4.72 mm. ..............................................................................Rhagovelia jubata Bacon, 1948 (Peru). - The middle trochanter is yellow (Fig. 6.235). Length: 4.05 to 4.66. ................................................................................Rhagovelia palea Bacon, 1956 (Peru, Bolivia). 90. (88) There are small black denticles on the jugum, proepisternum, mesosternum, and on the middle and hind acetabula. The hind trochanter of the male bears one large spine, which is longer than any spine on the hind tibia. The distal spines on the hind tibia of the male are no more than slightly longer than basal spines (Fig. 6.237). There are yellow-brown spots in the middle of the apical three or four abdominal tergites. The hind tibia of the male is cylindrical. Length: 3.83 to 4.16 mm. ..............................................................................Rhagovelia scitula Bacon, 1956 (Peru, São Paulo, Rio de Janeiro, Minas Gerais). - There are dark brown spots in the middle of the apical three abdominal tergites. The hind tibia of the male is flattened (Fig. 6.233). Length: 4.00 to 4.11 mm. ..........................................................................Rhagovelia whitei (Breddin, 1898) (Minas Gerais, Mato Grosso). 91. (86) At the base of the jugum and the adjacent proepisternum, there are small black denticles, which may only be visible under high magnification. ....92 - There are no small black denticles at the base of the jugum or on the proepisternum. The male hind tibia bears only the distal spine and a few small black pegs (Fig. 6.238). ......................................................................................96

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Fig. 6.238 Rhagovelia guianana male: hind leg (left) and paramere (right). Based on Polhemus (1997). 92. (91) The color is mainly orange brown with a silvery pruinosity forming an apical band on the pronotum (Fig. 6.204). The middle trochanter is yellowish brown, yellow, or pale. .......................................................................................93 - The color is mainly black with a dark yellow apical band on the pronotum. The middle trochanter is dark brown or black. ...................................................95

Fig. 6.239 Rhagovelia venezuelana: Habitus without appendages of a male (upper left) and female in dorsal view (lower left), male proctiger (center), and right paramere (right). Based on Polhemus (1997). 93. (92) There is no large spine on the inner margin of the hind tibia, but only two or three teeth slightly larger than the others. The hind femur bears two parallel rows of teeth but lacks irregular rows. There are numerous small black denticles on the central parts of the mesosternum, metasternum, and first seven abdominal tergites. The male paramere is truncate at the apex (Fig. 6.239). ...............................................................Rhagovelia venezuelana Polhemus, 1997 (Venezuela).

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- There is a large conical spine on the inner margin near the apex of the hind tibia, at least of the male. If the female lacks such a spine, then a smaller apical spine is present at the apex of the tibia (Fig. 6.204). ..........................................94 94. (93) The male paramere has a sinuate dorsal and ventral margin (Fig. 6.240). The hind femur bears two parallel rows of teeth along its hind margin and two irregular rows of teeth. There are no small black denticles on the central parts of the mesosternum, metasternum, and first seven abdominal tergites. The hind femur of the male is sometimes greatly incrassate. ..........................................................Rhagovelia calcaris Drake and Harris, 1935 (Peru). - The male paramere appears strongly folded and truncate (Fig. 6.204). Length: 3.7 to 4.1 mm. Winged forms have not yet been found. Color: mainly orange brown with yellowish markings. ....................................................Rhagovelia bocaina Moreira and Ribeiro, 2009 (São Paulo).

Fig. 6.240 The male paramere of Rhagovelia calcaris. Based on Polhemus (1997).

Fig. 6.241 Right paramere of a male Rhagovelia robusta in lateral view. Based on Bacon (1956).

Fig. 6.242 Right paramere of a male Rhagovelia relicta in lateral view. Based on Bacon (1956). 95. (92) The middle trochanter is yellow or yellowish brown. The hind femur of the male is armed with four or five irregular rows of spines. The male paramere has a prominent apical hook (Fig. 6.241). Length: 4.45 to 4.75 mm. .............................................................................Rhagovelia robusta Gould, 1931 (Peru, Paraguay, Argentina, Santa Catarina, Rio de Janeiro, Minas Gerais, Goiás).

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- The middle trochanter is dark brown or black. The male hind femur is armed with two or three rows of spines. The right male paramere is broadly truncate at the apex and lacks a prominent hook (Fig. 6.242). Length: 3.80 to 3.85 mm. ..............................................................................Rhagovelia relicta Gould, 1931 (Argentina, São Paulo). 96. (91) There are many long, erect, pilose setae and stout, erect, black setae on the anterior margin of the hind femur. There is a long spine on the inner margin of the hind tibia quite close to its apical end (Fig. 6.238). ....................................................................Rhagovelia guianana Polhemus, 1997 (Venezuela, Surinam, Amazonas). - Long, erect, pilose setae are not numerous on the anterior margin of the hind femur, but stout, erect, black setae may be scattered there. There is a long spine on the inner margin of the hind tibia separated from the apical end by a distance equal to or greater than 1/5 of the tibia length (Fig. 6.243). ..............................97 97. (96) The dorsal ground color is black with pale markings on the anterior part of the pronotum, connexiva margins, and legs, creating a strongly contrasting pattern. There is sometimes one small tooth on the hind trochanter (Fig. 6.232). ............................................................Rhagovelia equatoria Polhemus, 1997 pars (Venezuela). Also see Couplet 85. - The dorsal color is uniform orange brown without strongly contrasting markings. The hind trochanter bears several small teeth (Fig. 6.243). .............98 98. (97) There are fairly large, peg-like teeth on the basal 1/3 of the hind femur, irregularly arranged or may appear to form several rows. The male paramere forms a slight hook at the apex (Fig. 6.243). ....................................................................Rhagovelia boliviana Polhemus, 1997 (Bolivia, Argentina). - On the hind femur, there is a single ventral row of small, black, peg-like teeth along the basal 1/3. A large, spike-like tooth is inserted on the hind tibia 2/3 of the way from the base to the apex. There are no minute, black, conical setae on the jugum of the head or on the proepisternum. The first seven abdominal segments have a combined length 1.8 times the greatest abdominal width. The paramere is broad at or near its mid-length (Fig. 6.244). Length: 5.5 to 6.5 mm. ..............................................................................Rhagovelia ornata Bacon, 1948 (Bolivia, Argentina).

Fig. 6.243 Rhagovelia boliviana male: hind leg (left) and paramere (right). Based on Polhemus (1997).

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Fig. 6.244 Right paramere of a male Rhagovelia ornata in lateral view. Based on Bacon (1956). Subfamily Veliinae Key to the species of adult Euvelia in South America Information for the key was provided by Polhemus and Polhemus (1984a). 1. There are silvery patches of hair-like setae on the dorsal surface of the first and fourth connexival segments, and there are no pruinose areas on the connexiva. The combined length of the thorax and abdomen along the mid-line is only slightly greater than the maximum width (Fig. 6.245). Length of apterous male: 1.4 to 1.5 mm. Maximum width: 1.0 to 1.5 mm. .....................................................Euvelia discala J. T. and D. A. Polhemus, 1984 (Peru, Amazonas). - There are no patches of silvery setae on the connexiva, which have pruinose areas at least on the second and the fourth through seventh segments. The combined length of the thorax and abdomen is considerably greater than the maximum width (Fig. 6.149). ...............................................................................2 2. (1) The trochanter on the fore-leg of the male has a long spur (Fig. 6.246). The width of the female at the humeri is about 1.5 times the width of the head at the compound eyes. There are no glabrous areas on the pleurae of the females. Length of the apterous male: 1.3 to 2.0 mm ....................................................................................Euvelia advena Drake, 1957 (Peru, Bolivia, Rondônia, Mato Grosso, Amazonas, Goiás). - The trochanter on the fore-leg of the male does not bear a long spur (Fig. 6.247). Either the width of the female at the humeri is almost twice the width of the head at the compound eyes, or there is a glabrous areas on the pleurae of the females. .................................................................................................................3 3. (2) The male has neither a spur nor tuft of stiff setae at the apical end of the fore-trochanter. The female has a depressed glabrous area on the pleurae, which do not bear stout setae (Fig. 6.149). Length of apterous male: 1.2 to 1.3 mm. Maximum width: 0.7 to 0.8 mm. ...................................................Euvelia concava J. T. and D. A. Polhemus, 1984 (Mato Grosso). - The male has a spur or tuft of stiff setae at the apical end of the fore-trochanter (Fig. 6.247). The females have no glabrous areas on the pleurae, which bear

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stout setae or various lengths. Length of apterous male: 1.3 to 1.4 mm. Maximum width: 0.9 to 1.0 mm. ..........................................................Euvelia lata J. T. and D. A. Polhemus, 1984 (Amazonas).

Fig. 6.245 Habitus of an apterous female Euvelia discala. Based on Polhemus and Polhemus (1984a).

Fig. 6.246 Fore-trochanter of Euvelia advena. Polhemus (1984a).

Based on Polhemus and

Fig. 6.247 Fore-trochanter of Euvelia lata. Based on Polhemus and Polhemus (1984a).

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Key to the species of adult Veloidea in South America Information for the key was provided by Drake and Lauck (1959a). 1. The median length of the pronotum is about equal to the humeral width. The pronotum is slightly raised at the humeral angles, but spines are not present (Fig. 6.150). Length: 8.4 mm. ............................................................................Veloidea gigantea (Gould, 1928) (Colombia). - The median length of the pronotum is greater than the humeral width. The pronotum is covered with coarse punctures and not at all produced at the humeral corners (Fig. 6.248). Length: 7.25 mm. ...........................................................Veloidea venezolana Drake and Roze, 1955 (Venezuela). There are two additional species known only from Central America, both with spines at the humeral angle.

Fig. 6.248 Veloidea venezolana male (left to right): apex of the abdomen in lateral view, right paramere, and the anal segment. Based on Drake and Lauck (1959a). Key to the species of Stridulivelia in South America Information for the key was provided by Drake and Harris (1938, 1941), Drake (1957e), Polhemus and Polhemus (1984b, 1984c), Nieser and Alkins-Koo (1991), Polhemus and Spangler (1995), and Melo and Nieser (2004). 1. No stridulatory structures are evident. There is a spine, which is usually long and stout but can vary considerably in size, on the hind trochanter of the male. It is rarely absent. The parameres have a complex structure (Figs. 6.30, 6.249). ........................................Stridulivelia (Aenictovelia) cinctipes (Champion, 1898) (Mexico, Central America, Colombia, Venezuela, Guyana, Pará). Syn: Velia cinctipes Champion, 1898; Velia (Stridulivelia) cinctipes Hungerford, 1929. - Stridulatory structures are present on the hind femora and abdomen (Fig. 6.250). ...................................................................................................................2

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Fig. 6.249 The right paramere of a male Stridulivelia cinctipes. Based on Drake and Menke (1962), who referred to the species as Velia cinctipes. 2. (1) There are glabrous transverse grooves on each side of the second through fifth abdominal sternites, which are distinct from the sutures separating the segments (Fig. 6.146). The first segment may be concealed, making it appear that the grooves are on the first four segments. The fourth groove of the four on the female may be reduced to short spots. The third and fourth grooves on the male may be short. There are prominent spurs on the hind trochanters. .............3 - There are glabrous transverse grooves on no more than three abdominal sternites (Fig. 6.250). If a spur is present on the hind trochanter, it is inconspicuous. ......................................................................................................4

Fig. 6.250 Stridulivelia stridulata: habitus of a micropterous female in dorsal view (lower center to right), left hind leg of a male (upper left) and the stridulatory area on the femur enlarged (lower left), male abdomen in lateral view (upper center), and left male paramere (upper right). Based on Hungerford (1929a) and Polhemus and Spangler (1995).

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Fig. 6.251 Stridulivelia transversa male: hind trochanter, femur, and tarsus (left); left paramere (right). Based on Hungerford (1929a).

Fig. 6.252 Stridulivelia alia: habitus of a female in dorsal view (left), hind femur with stridular denticles (upper right), and connexiva in lateral view (lower right). Based on Polhemus and Spangler (1995). 3. (2) There are glabrous grooves only on the second through fifth abdominal segments of the female, but the male also has a small groove on the sixth segment. Length of macropterous male: 4.7 to 5.0 mm. Both males and females possess stridulatory structures on the hind femora and connexiva. The male paramere is broadly and evenly rounded at the apex (Fig. 6.146). Color: apterous form dark reddish brown to chocolate with yellowish brown connexiva and appendages; the hemielytra of the macropterous form are dark fuscous with small silvery setae near the bases. ....................................Stridulivelia (Stridulivelia) tersa (Drake and Harris, 1941) (Trinidad, Surinam, Guyana, Venezuela, Bolivia, Peru, Amazonas, Toncantins, Minas Gerais). Syn: Velia tersa Drake and Harris, 1941; Velia nama Drake, 1957.

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- There are glabrous transverse grooves on the second through sixth abdominal segments, and a small grove on the seventh segment of the male, as well. Length: c. 3.2 mm. There is a spur on the hind trochanter (Fig. 6.251). The female abdomen is strongly narrowed. Color: light brown with a pair of silvery, roughly triangular patches on the anterior part of the pronotum. The ventral surface is tan frosted with gray. ....................................Stridulivelia (Stridulivelia) transversa (Hungerford, 1929) (Guyana, French Guiana, Surinam, Amazonas). Syn: Velia transversa Hungerford, 1929. 4. (2) The humeral angles of the pronotum are produced to form large conspicuous spines (Fig. 6.252). Length of antenna segments I to IV: 0.19:0.65:0.63:0.53 mm, respectively. Length of micropterous male: 3.7 to 4.5 mm; micropterous female: 4.0 to 4.3 mm; macropterous male: 4.3 to 4.5 mm; macropterous female: 4.2 to 4.7 mm. Maximum width of micropterous male: 1.0 to 1.2 mm; micropterous female: c. 1.1 to 1.2 mm; macropterous male: 1.4 to 1.5 mm; macropterous female: 1.3 to 1.5 mm. Color: mainly orange brown dorsally and lighter ventrally, with yellow to light brown antennae and legs, black apical segments of the beak, and triangular silvery spots on the anterior parts of the pronotum. .........................................................Stridulivelia (Stridulivelia) alia (Drake, 1957) (Venezuela, Guyana, Surinam, Amazonas). Syn: Velia alia Drake, 1957. - The humeral angles of the pronotum are not produced to form spines (Fig. 6.250). ...................................................................................................................5 5. (4) Only the two anteriormost abdominal sternites have glabrous transverse grooves (Fig. 6.250). ............................................................................................6 - The three anteriormost abdominal sternites have glabrous transverse grooves. There is no ventral protuberance on the first genital segment of the male (Fig. 6.253). ...................................................................................................................7

Fig. 6.253 Stridulivelia quadrispinosa: hind leg (left), male left paramere (center), and abdomens of a male in lateral view (lower right) and a female in dorsal view (upper right). Based on Hungerford (1929a).

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6. (5) There is a ventral protuberance on the first genital segment of the male (Fig. 6.250). There are two spine-like processes along the posterior margin of the seventh abdominal segment of the female. Length: c. 5.4 mm. .....................................Stridulivelia (Stridulivelia) stridulata (Hungerford, 1929) (Peru, Colombia, Venezuela, Guyana, French Guiana, Surinam, Amazonas, Mato Grosso). Syn: Velia stridulata (Hungerford, 1929). - There is no ventral protuberance on the first genital segment of the male (Fig. 6.254). The female lacks a spine-like process along the posterior margin of the seventh abdominal segment. Length of brachypterous male: 3.8 to 5.0 mm; brachypterous female: 4.7 to 5.2 mm; macropterous female: 5.2 to 5.4 mm. No macropterous males have been described. .....................Stridulivelia (Stridulivelia) ayacucho Polhemus and Spangler, 1995 (Guyana, Venezuela, Paraguay, Peru, Pará, Minas Gerais).

Fig. 6.254 Stridulivelia ayacucho male: genital capsule in lateral view (left) and the male paramere in anterior (upper right), posterolateral (middle right), and lateral view (lower right). Based on Polhemus and Spangler (1995).

Fig. 6.255 Stridulivelia strigosa male: abdomen in lateral view (left) and left paramere (right). Based on Hungerford (1929a). 7. (5) The first segment of the antenna is about 1.5 times as long as the width of the head across the eyes. Males lack projections on the posterior margins of the

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first genital segment. The hind femora are not incrassate (Fig. 6.255). Length: c. 4.5 mm. .........................................Stridulivelia (Stridulivelia) strigosa (Hungerford, 1929) (French Guiana). - The first segment of the antenna is not longer than 1.2 times the width of the head across the eyes. The hind femur is incrassate (Fig. 6.253). The scutellum is usually exposed. ................................................................................................8 8. (7) The first segment of the antenna is about 1.2 times the width of the head across the eyes. The males bear a pair of prominent projections on the posteriormost abdominal sternite (Fig. 6.256). The hind femur has a stridulatory patch consisting of 26 to 32 sharp pegs, scattered and not arranged in rows. Length of brachypterous male: 3.7 to 4.5 mm; macropterous male: 4.3. to 4.5 mm; brachypterous female: 4.0 to 4.3 mm; macropterous female: 4.2 to 4.7 mm. .............................Stridulivelia (Stridulivelia) anta Polhemus and Spangler, 1995 (Venezuela, Amazonas). - The first segment of the antenna is subequal to or shorter than the width of the head across the eyes. .............................................................................................9

Fig. 6.256 Stridulivelia anta male: genital capsule in lateral view (left) and the male paramere in anterior (upper right), posterolateral (middle right), and lateral view (lower right). Based on Polhemus and Spangler (1995). 9. (8) The length of the first antenna segment equals about 0.8 times the width of the head across the eyes. Males lack prominent projections along the posterior margin of the posteriormost abdominal sternite. In dorsal view, the female genital segment appears triangular with its length subequal to its width at the base. About 17 rows of tiny spinules form the stridulatory patch on the hind femur. .................................Stridulivelia (Stridulivelia) astralis (Drake and Harris, 1938) (Argentina, Mato Grosso do Sul). Syn: Velia astralis Drake and Harris, 1938.

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- The length of the first antenna segment is about equal to the width of the head across the eyes. Males have a pair of prominent projections along the posterior margin of the posteriormost abdominal sternite. In dorsal view, the female genital segment appears rounded or triangular with the width at the base obviously greater than its length (Fig. 6.253). ...................................................10 10. (9) The stridulatory patch on the hind femur consists of about 40 sharp pegs scattered rather than in rows. Opposite the patch is a row of fine vertical ridges on the margin of the connexivum. The seventh abdominal segment has prominent projections on the posterior margin of the seventh abdominal segment, which are on the dorsal side of the female and ventral side of the male (Fig. 6.253). Length: c. 4.8 mm. ..............................Stridulivelia (Stridulivelia) quadrispinosa (Hungerford, 1929) (Guyana, Venezuela, Peru, Bolivia, Amazonas, Pará). Syn: Velia quadrispinosa Hungerford, 1929. - The stridulatory patch on the hind femur consists of about 17 rows of minute spinules. This is opposed by a thin row of tiny pegs along the margin of the connexivum. The male has prominent projections along the posterior margins on the ventral side of the seventh abdominal segment, but the female lacks any such projections anywhere on this segment (Fig. 6.257). Length: c. 5.0 mm. ...........................................Stridulivelia (Stridulivelia) raspa (Hungerford, 1929) (Amazonas). Syn: Velia raspa Hungerford, 1929.

Fig. 6.257 Stridulivelia raspa: hind femur of a male (upper left) and a female (upper right) and the female abdomen in lateral view (below). Based on Hungerford (1929a). Key to the species of adult Oiovelia in South America Information for the key was provided by Spangler (1986) and Mazzucconi and Bachmann (1997a). 1. The pronotum of the apterous form is entirely creamy yellow, and that of the winged form is creamy yellow only on the posterior half. Only the anterior fourth

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of the pronotum bears a few long, brownish setae. The antennae are intensely tomatose and brownish yellow with a creamy yellow last segment in apterous forms. The femora are creamy yellow with a narrow brown ring at the apex. The male paramere tapers distally and ends in an acute apical hook (Fig. 6.258). Length of apterous males: 3 mm; winged females: 3.2 to 3.5 mm. .........................................................................Oiovelia spumicola Spangler, 1986 (Venezuela). - The pronotum is reddish brown with numerous long, dark brown setae, especially numerous on the anterior third and along the margins and sparcer on the discal areas. The antennae are not densely tomentose. The femora are reddish brown. The paramere of the male is not hooked (Fig. 6.152). ...............2

Fig. 6.258 Oiovelia spumicola: habitus of a male (upper left) and a female (lower left) in dorsal view, head and pronotum in dorsal view (lower right), and the genital capsule of a male (upper right). Based on Spangler (1986). 2. (1) The ratio of the antennal segments is 30:27:19:22. Total length: 3.16 to 3.89 mm. There are narrow black band-like markings on the dorsolateral margin of the metasternum. The male paramere is broadened preapically (Fig. 6.152). .............................................................................Oiovelia rivicola Spangler, 1986 (Venezuela). - The ratio of the antennal segments is 30:22:15:19. Total length: 3.3 to 4.4 mm. The male paramere is not broadened preapically when viewed as part of the genitalia, but it is enlarged when viewed intact. Larval stages have been described (Fig. 6.143, 6.259). ................................................Oiovelia cunucunumana Drake and Capriles, 1952 (Venezuela, Peru, Paraguay, Santa Catarina).

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Fig. 6.259 Oiovelia cunucunumana larva: instars showing the dorsal habitus without appendages (above), thorax and abdomen in lateral view (below), and antenna between the two, of the second (upper left), fourth (lower left), and fifth instar (right). Based on Mazzucconi and Bachmann (1997b). Key to the South American species of adult Paravelia Information for the key was provided by Osborn (1904), Hungerford (1930a, b), Drake (1951a), Polhemus (1976a), Polhemus and Polhemus (1984b, c; 1991), Spangler (1989), and Iglesias and Crespo (1999). The males of several species have not yet been described. 1. Only the third and fourth sternites bear a transverse row of 17 tiny black pegs on each side, which rub against the plectrum, consisting of a field of pegs on the anterior surface of the trochanter, forming a stridulatory organ (Fig. 6.260). Length: 3.5 to 3.6 mm. Color: mainly orange brown. ............................................Paravelia stenoptera J.T. and D.A. Polhemus, 1984 (Surinam). - There is no stridulatory organ present on the trochanter or any other part of the leg (Fig. 6.261). ....................................................................................................2

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Fig. 6.260 Paravelia stenoptera (left to right): the hind trochanter in anterior view showing the stridulatory mechanism, the abdomen in lateral view, and the male paramere. Based on Polhemus and Polhemus (1984b).

Fig. 6.261 Paravelia biae male: habitus of a macropterous specimen (upper left), genital capsule in lateral view (upper right), left paramere (lower left), and apex of the abdomen in ventral view (lower right). Based on Spangler (1989). 2. (1) Each side of the pronotum bears a large, prominent spine on the humeral angle (Fig. 6.262). Length: 3.3 to 3.4 mm. Color: mainly orange brown. .................................................Paravelia spinifera J.T. and D.A. Polhemus, 1984 (Surinam). - There are no spines at the humeral angles of the pronotum (Fig. 6.263). ........3 3. (2) The posteriormost pregenital segment of the male has two ventral processes, pointed at the apices, with the points directed generally posteriad but slightly divergent. The left paramere is nearly straight with a bluntly rounded apex; it is constricted at the base but bears a lateral tumescence at the

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constriction (Fig. 6.264). The general color is black with only two yellow spots on the hemielytra. Better descriptions of this species are urgently needed. At present, the species can only be identified by examination of the male abdomen. ...........................................................................Paravelia basalis (Spinola, 1837) (Brazil). Syn: Velia basalis Spinola, 1937; Paravelia bicolor (Blanchard, 1843); Paravelia brasiliensis (Herrich-Schäffer, 1850). - If there are processes on the posteriormost pregenital segment of the male, their shape or orientation is different (Fig. 6.265). The left paramere of the male is not nearly straight, or a basal process, if persent, does not appear tumescent. The ground color is not black, or, if it is, there are pale markings other than two yellow spots on the hemielytra. ............................................................................4

Fig. 6.262 Paravelia spinifera: habitus in dorsal view without legs (left) and the paramere of a male (right). Based on Polhemus and Polhemus (1984b).

Fig. 6.263 Paravelia inveruglas male: habitus without appendages in dorsal view (left), genital segments without the supra-anal hooks in lateral view (center), hooks on the supra-anal plate (upper right), the flat side of the left paramere (lower right), and the apex of the paramere in posterior view (upper right center). Based on Hungerford (1930a).

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Fig. 6.264 Paravelia basalis male: apex of the abdomen in ventral view (left) and left paramere (right). Based on Hungerford (1930b).

Fig. 6.265 Paravelia confusa male: apex of the abdomen in ventral view (left) and left paramere (right). Based on Hungerford (1930b). 4. (3) The length is greater than 7.0 mm. If slightly shorter, then the male genital segments appear very hairy, and there are no hooks on the supra-anal plate (Fig. 6.266). .................................................................................................5 - The length is less than 6.5 mm. .......................................................................13

Fig. 6.266 Paravelia correntina (left to right): pronotum of a female in lateral view showing the depressions on the surface, left male paramere, apex of a male abdomen in dorsal and ventral view. Based on Iglesias and Crespo (1999). 5. (4) The length of the male is c. 7.6 mm or shorter, and the width across the humeri is c. 2.5 mm or less. .................................................................................6 - The length is much greater than 7.6 mm, or, if it is only slightly greater, the maximum width is greater than 1/3 of the length. ................................................8

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6. (5) The maximum width of the body is almost 1/3 of the total length. Length of male: c. 7.6 mm; width at humeri: c. 2.5 mm. The lengths of antenna segments I to IV in mm are 1.30:1.20:0.86:1.00, respectively. Color: dark brown with a testaceous posterior margin of the pronotum and testaceous humeral angles. The appendages are uniformly brown, or the dorsal surfaces of the femora are darkened. The ventral surface is uniformly brown. ............................................................................Paravelia amoena (Drake, 1957) (Venezuela). Syn: Velia amoena Drake, 1957. - The maximum width of the body is 1/4 or less of the total length. The pronotum is coarsely punctured (Fig. 6.263). ......................................................7 7. (6) The length ratios of antenna segments I to IV are 50:40:37:40, respectively. The male lacks supra-anal hooks, although its genital segments are so densely hairy that hooks may appear to be there. Length of male: c. 7.2 mm; maximum width of male: c. 1.6 mm. The color is mainly brown with the pronotum yellowish and the apices of all femora and bases of all tibiae blackish. The species was described from one male specimen. ................................................................Paravelia willei (Drake and Harris, 1940) (Peru). Syn: Velia willei Drake and Harris, 1940. - The ratios of antenna segments I to IV are 16:14:12:15. The pronotum is slightly constricted near the middle and has the trace of a longitudinal carina. The supra-anal hooks of the male are directed anterolaterad and extend beyond the lateral margins of the abdomen (Fig. 6.263). The genital segments of the male do not appear densely hairy. Color: black with reddish black antennae, rostrum, and legs; the pronotum is pale violet brown with reddish violet margins. Length of male: c. 7.2 to 7.3 mm; maximum width of male: c. 1.8 mm. ....................................................................Paravelia inveruglas (Kirkaldy, 1899) (Ecuador). Syn: Velia inveruglas Kirkaldy, 1899. 8. (5) Black is a predominant color, although the posterior part of the pronotum may be yellowish. .................................................................................................9 - Black is not a predominant color, or, if it is, the antennae and legs are brown, and there may also be prominent pale markings on the hemielytra, or the entire margin of the pronotum is yellowish. In case of doubt, the length is about 9 mm and the connexivum is produced slightly caudad (Fig. 6.267). ..........................10

Fig. 6.267 Paravelia albotrimaculata female: habitus in dorsal view without appendages (left) and the hind leg (right). Based on Hungerford (1930a).

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9. (8) The head, first antenna segment, entire anterior part of the pronotum, femora, abdominal tergites, and connexivum are chocolate brown, while the three apical antenna segments, beak, tibia, tarsi, and major part of the ventral surface are black. The elytra are velvety black. Only the posterior margin of the pronotum is yellow and not covered by the fine, velvety pruinosity that covers the rest of the body. The second and fourth antenna segments are subequal and longer then the third. The posterior margin of the pronotum is broadly bisinuate. The hind femora lack spines (Fig. 6.268). Length: c. 8 mm. ..................................................................Paravelia osborniana (Kirkaldy, 1909) (Bolivia). Syn: Velia osborniana Kirkaldy, 1899; Velia brunnea Osborn, 1904 (preoccupied). - The head, including the antennae and base of the rostrum, and the legs are blackish with a dense covering of short, recumbent, gray setae. Much of the posterior part of the pronotum is yellowish. The connexivum is elevated perpendicular from the horizontal plane of the body. Length: c. 9 mm. The original description was based on an apterous female. .........................................................................Paravelia boliviana Breddin, 1898 (Bolivia).

Fig. 6.268 Paravelia osborniana male: hooks on the supra-anal plate (left), genitalia in lateral view (center), and apex of a paramere in posterior view (right). Based on Hungerford (1930a). 10. (8) The total length is greater than three times the maximum width (Fig. 6.267). .................................................................................................................11 - The length is equal to or shorter than three times the width across the humeri (Fig. 6.261). ........................................................................................................12 11. (10) Length: c. 9 mm; width at widest point: c. 2.8 mm. The connexivum is produced caudad only very slightly. The supraanal hooks of the male are directed anterolaterad and do not reach to the lateral margins of the abdomen (Fig. 6.269). The color is black with an orange yellow posterior margin of the pronotum. A silvery pubescence covers the head and lateral margins of the pronotum. Three is a golden pubescence on the abdomen, most evident on the connexiva. The antennae and legs are brown. ...........................................................Paravelia columbiensis (Hungerford, 1930) (Colombia). Syn: Velia columbiensis Hungerford, 1930.

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- Length of the female: c. 8.2 mm; width across the humeri: 2.6 mm. According to the original description by Kirkaldy (1899), the length is 7 mm, and the width across the humeri is 3 mm, but Hungerford (1930a) reexamined the type specimen and found the dimensions to be those reported above. The anterior half of the pronotum is carinate, and its surface is pitted. The hind femur of the female bears six strong spines of about equal length along the distal 3/5 of the posterior margin (Fig. 6.267). The ground color on the dorsum and venter of the thorax is dark with three nearly round, white spots evident on the folded hemielytra. The abdomen has an orange brown tinge, and the coxae, trochanters, and a ring at the base of the legs varies from lemon yellow to hornyellow. ...........................................................Paravelia albotrimaculata (Kirkaldy, 1899) (Venezuela). Syn: Velia albotrimaculata Kirkaldy, 1899.

Fig. 6.269 Paravelia columbiensis male: hooks on the supra-anal plate in dorsal view (left) and the left paramere (right). Based on Hungerford (1930a). 12. (10) Length: c. 7.9 mm; width at widest point: c. 5.6 to 5.7 mm. The head and thorax are uniformly brown or light brown. The venter and legs are all dark brown, or the legs are fuscous. The connexivum is not bordered by orange yellow. The hind trochanter of the male bears small spines, and the femur bears small, well-spaced spines for almost its whole length (Fig. 6.270). Each hemielytron is chocolate brown with a small white spot near the base and a larger one on the membrane. The posterior part of the pronotum is deeply pitted. ..............................................................................Paravelia atra (Polhemus, 1969) (Peru). Syn: Velia atra Polhemus, 1969. - Length: c. 8.4 mm; width across the humeri: c. 2.8 mm. The margin of the pronotum is orange yellow, forming a band that completely surrounds the blackish disc. The connexivum is also margined by the same color. The anterior margin of the pronotum is narrower than the head. The lateral margins become wider toward mid-length, where there is a constriction. The connexivum forms stout, sharp projections at its posterolateral corners. The male supra-anal plate hooks are flat, short, and inconspicuous (Fig. 6.271). .......................................................Paravelia flavomarginata (Hungerford, 1930) (Colombia). Syn: Velia flavomarginata Hungerford, 1930.

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Fig. 6.270 Paravelia atra male (left to right): hind femur and apical part of the trochanter with the hair-like vestiture removed, apical segments of the abdomen in ventral view, and the left paramere. Based on Polhemus (1969).

Fig. 6.271 Paravelia flavomarginata male: supra-anal plate hooks in dorsal view (left) and the left paramere (right). Based on Hungerford (1930a). 13. (4) The pronotum has a tall, almost conical process along the midline extending posteriad to a point between the mid-points of the two humeri, which are only slightly elevated. The pronotum is pitted. The third antenna segment is the longest. Length: c. 5 mm. Color: mainly chocolate brown with silvery patches at the following locations: a broad, slightly sigmoid spot divided by the mid-line of the pronotum; a long spot at the base of the hemielytra; three spots on the membrane; five transverse spots on the sutural lines of each connexivum. The legs are pale with one broad band on each femur and two narrow bands on each tibia. .....................................................................Paravelia conata (Hungerford, 1929) (French Guiana). Syn: Velia conata Hungerford, 1929. - The pronotum does not bear a tall, conical process (Fig. 6.261). ...................14 14. (13) The head and pronotum of the apterous female are covered by numerous tiny, conical spines (Fig. 6.151). The color is brown. ......................15 - Apterous specimens have not been described, or, it they have, there are no tiny conical spines on the head or pronotum, or, if such spines are present, they are few (Fig. 6.272). .................................................................................................20

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Fig. 6.272 Paravelia hungerfordi female: habitus without appendages in dorsal view (left) and the head in lateral view with the apical part of the beak omitted (right). Based on Mazzucconi (2000a). 15. (14) The first segment of the antenna is longest (Fig. 6.151). Forms with wings have not been found. The general color is yellowish brown, and the color of the short spines on the body is black. The body is also covered by a dark pilosity that appears light when illuminated at some angles. Length: 4.3 to 4.6 mm. Maximum width (at the fourth abdominal segment): 1.4 to 1.5 mm. ..............................................................................Paravelia paxilla (Drake, 1957) (Argentina, Mato Grosso). Syn: Velia paxilla Drake, 1957. - The fourth segment of the antenna is the longest. ...........................................16 16. (15) Length of macropterous male: c. 5.6 to 5.7 mm; width: c. 2.0 to 2.1 mm. The lengths of antenna segments I to IV in mm are 0.92:0.80:0.90:1.10, respectively (Drake, 1957e). These proportions are different from those in the original description by Drake and Harris (1933b), who reported ratios of 50:38:45:55. The legs are ringed. General color: brown to dark brown, with a darker pronotum. ...........................................................Paravelia cognata (Drake and Harris, 1933) (Maranhão). Syn: Velia capillata var. cognata Drake and Harris, 1933; Velia cognata Drake and Harris, 1933. - The length is less than 5.3 mm. .......................................................................17 17. (16) Length: c. 5 mm. Width across humeri: c. 1.9 mm. Only winged specimens have been described. There is a prominent forked process on the ventral side of what appears to be the sixth abdominal segment of the male (Fig. 6.265). The general color is light brownish yellow. ....................................................................Paravelia confusa (Hungerford, 1930) (Pará). Syn: Velia confusa Hungerford, 1930. - The length is less than about 4.5 mm. There is no prominent, forked process on the ventral side of the apparent sixth abdominal segment of the male (Fig. 6.273). The general color is brown or darker. ...................................................18

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Fig. 6.273 Paravelia rotundanotata male: apical abdominal segments in ventral view (left) and the left paramere (right). Based on Hungerford (1930b). 18. (17) Length: 2.3 to 3.4 mm; maximum width: 1.0 to 1.2 mm. The length ratios of antenna segments I to IV are 25:18:22:28, respectively. The pronotum of the apterous form is covered with coarse pits, and its posterior margin is broadly rounded. The color of both winged and wingless forms is mainly brown. Each hemielytron is smoky brown with an elongate white marking near the base and a round white spot near the middle of the membrane. The pronotum has a median ridge, which is prominent on the apterous form but indistinct on the winged form. ........................................................Paravelia splendoris (Drake and Harris, 1933) (Mato Grosso). Syn: Velia splendoris Drake and Harris, 1933. - The length is greater than 3.4 mm. ..................................................................19 19. (18) Winged and wingless specimens have been described. There are minute, peg-like points and darker, bristle-like hairs on the head, thorax, and connexiva. The points are most noticeable on the humeral elevations of the pronotum. There are no processes of any kind on the ventral side of what appears to be the sixth abdominal segment. There is a group of tiny spines on each of the elongated posterolateral corners of the seventh abdominal segment (Fig. 6.273). The general color is light brown. Length: c. 4 mm. Width across the humeri: c. 1.15 mm. .........................................................Paravelia rotundanotata (Hungerford, 1930) (Argentina, Minas Gerais). Syn: Velia rotundanotata Hungerford, 1930. - Only winged specimens have been described. The pronotum lacks peg-like points but is covered by fine, brownish hairs and coarse, deep pits. There is sometimes a trace of a median carina. The posterior margin of the pronotum is broadly rounded in the female but more narrowly angled in the male. The lengths of antenna segments I to IV in mm are 0.75:0.40:0.82:0.92, respectively. Length: c. 4.2 mm. Greatest width: c. 1.73 mm. General color: brown to dark brown, with a darker pronotum. ..........................................................Paravelia capillata (Drake and Harris, 1933) (Mato Grosso). Syn: Velia capillata Drake and Harris, 1933. 20. (14) The length is greater than 5.8 mm, and the maximum width is about 2.3 mm. Apterous specimens have not been described. ..........................................21 - The length is shorter than about 5.8 mm. ........................................................22

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21. (20) The length of the macropterous form is reported to be 6.45 mm, and its maximum width is c. 2.5 mm. The color is black with reddish brown on the base of the rostrum and coxae and conspicuous yellow markings on the hemielytra, including one at the base and one near the apex of each, so that the apical markings overlap when the wings are folded back to form one large yellow subapical spot. There are coarse pits on the pronotum, which are separated from each other by a distance equal to about five times their diameter. The paramere is sinuate and has a cluster of setae at the base and a row of erect, stiff setae on the dorsal margin (Fig. 6.261). ................................................................................Paravelia biae Spangler, 1989 (Pará) - The length of the macropterous form is reported to be 6 mm, and the maximum width is c. 2.3 mm. The width of the head across the eyes is c. 1.2 mm. The color is light chocolate brown with a pale tan transverse triangular marking at the base of each hemielytron, which extends to the apex of the scutellum. There is a round, pale spot on the membrane. The pronotum has a low median carina and is sparsely punctured. The paramere is sinuate only at the base (Fig. 6.274). .................................................................Paravelia williamsi (Hungerford, 1930) (Guyana). Syn: Velia williamsi Hungerford, 1930.

Fig. 6.274 Paravelia williamsi male: apex of the abdomen in ventral view (left) and left paramere (right). Based on Hungerford (1930b).

Fig. 6.275 Paravelia manausana: habitus without legs (upper left), head in lateral view (lower left), posterior view of the hind femur (lower center), and male genitalia in lateral view (right). Based on Polhemus and Polhemus (1991).

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22. (20) The pronotum is deeply and densely pitted or has a coarsely rugose surface, which, like the ventral part of the abdomen, bears numerous hairless foveae and pits (Fig. 6.272). ...............................................................................23 - The pronotum and the ventral part of the abdomen are not densely and deeply pitted or foveate (Fig. 6.275). .............................................................................30 23. (22) The ground color is dark brown or brownish black. The pronotum is deeply and densely pitted and has a median carina. The second antenna segment is usually shorter than or only barely longer than half the length of the third. All three species fitting this description are very poorly described and were never illustrated. Only macropterous specimens were described. The females of two species are apparently indistinguishable, and the males can only be tentatively identified until the species in this group are redescribed. ..................................24 - The color is orange to orange brown, chestnut brown, or reddish brown with yellow markings. The second segment of the antenna is considerably longer than half the length of the third. .........................................................................26 24. (23) The lateral edges of the connexiva are armed with many short black spinules. The head is black with impressed lines. The legs and antennae are brown. The proportions of the four antenna segments are 28:16:37:37. At the basal angles of the segments of the connexiva, there are pale markings. The male parameres are constricted at the base and beyond the midlength; they are curved and relatively broad, even at the apex. Length: c. 4 mm; maximum width: 1.2 to 1.4 mm. ................................................................Paravelia kahli (Drake and Harris, 1933) (Mato Grosso). Syn: Velia kahli Drake and Harris, 1933. - The lateral edges of the connexiva are not armed with short black spinules. The male parameres are pointed at the apex or darkened apically. ....................25 25. (24) The male parameres are long, curved, sharply bent preacipally, and pointed at the apex. The second segment of the antenna is less than half as long as the third. Length: c. 4 mm. Maximum width: 1.2 to 1.4 mm. ..............................................................Paravelia parilis (Drake and Harris, 1933) (Mato Grosso). Syn: Velia parilis Drake and Harris, 1933. - The male parameres are long, broad, stout, and hairy; they are curved inward, narrowed, and darkened toward the apex. The ratios of the lengths of the four antenna segments are 30:18:35:35, respectively. Length: c. 3.5 to 3.6 mm; maximum width: c. 1.35 mm. .................................................................Paravelia nexa (Drake and Harris, 1933) (Maranhão). Syn: Velia nexa Drake and Harris, 1933. 26. (23) The total length is less than 4 mm. .......................................................27 - The total length is greater than 4 mm. The general coloration is chestnut or reddish brown with yellow and white markings (Fig. 6.272). ...........................28 27. (26) The dorsum is nearly free of setae, except for a few scattered, upright ones. The male paramere is widest in the apical 1/8 (Fig. 6.276). Length: 3.3 to 3.4 mm. Color: Mainly orange brown with red eyes and dark brown antennae. .....................................................Paravelia foveata Polhemus & Polhemus, 1984 (Amazonas).

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- The body is covered by dense pubescence and a few longer setae. The male paramere is widest in the basal half (Fig. 6.266). The ratios of antennal segments I to IV are 0.57:0.48:0.35:0.39, respectively. Length: c. 3.6 to 3.7 mm. The color is mainly testaceous orange, becoming darker toward the posterior part of the body. The abdominal segments are testaceous ventrally. .....................................................Paravelia correntina Iglesias and Crespo, 1999 (Argentina).

Fig. 6.276 The outline of the male paramere of Paravelia foveata. Based on Polhemus and Polhemus (1984c). 28. (26) The general coloration is chestnut brown with whitish pilosity on the ventral surface of the abdomen. There is a broad mid-ventral keel on the seventh abdominal segment (Fig. 6.277). Length: 4.6 to 5.0 mm. .............................................................................Paravelia platensis (Berg, 1883) (Uruguay, Argentina). Syn: Velia platensis Berg, 1883. - The general coloration is reddish brown with yellowish brown markings, two whitish markings on the hemielytra, and bright red eyes. The whitish marking at the apex of the hemielytra is elongated (Fig. 6.272). .........................................29

Fig. 6.277 Paravelia platensis male: left paramere (lower left), right paramere (upper left), and apex of the abdomen in ventral view (right). Based on Hungerford (1930b).

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29. (28) The anterolateral margins of the pronotum are constricted anterior to the humeri. The ratios of antennal segments I to IV are 32:26:28:33. The entire dorsum of the macropterous female is covered with setae longer than the width of the first antennal segment. The apical marking on the hemielytra is only weakly constricted near its mid-length (Fig. 6.272). Length of female: c. 4.8 mm. The male has not been described. .......................................................Paravelia hungerfordi (Drake & Harris, 1933) (Mato Grosso?). Syn: Velia hungerfordi Drake & Harris, 1933. - The anterolateral margins of the pronotum are not notably constricted anterior to the humeri. The apical marking on the hemielytra is strongly constricted near its mid-length (Fig. 6.278). Length: 5.4 to 5.7 mm. The male and female appear alike, except for the genitalia. ........................................................................Paravelia anta (Mazzucconi, 2000) (Argentina, Paraguay).

Fig. 6.278 Paravelia anta (above, left to right): habitus without appendages in dorsal view; apex of the male abdomen in ventral view; fore, middle, and hind legs of a male, and (below, left to right): antenna, hemielytron of a male and a female. Based on Mazzucconi (2000a). 30. (22) The hemielytra are as long as the abdomen and leave part of the connexiva uncovered; each has a basal orange-yellow stripe that extends beyond the apex of the pronotum by about half its length. The pronotum and the venter are uniform chocolate brown. The apical connexival segments of neither sex are drawn out into spines; the apices are triangular. The ratios of antennal segments I to IV are 28:19:20:21. Length of macropterous form: c. 3.8 to 3.9 mm; maximum width: c. 1.6 to 1.7 mm. An apterous form has not been described. ...........................................................................Paravelia itatiayana Drake, 1951 (Rio de Janeiro). - The hemielytra lack basal orange-yellow stripes extending beyond the apex of the pronotum by half their own lengths. .............................................................31

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31. (30) The pronotum is not densely pitted or rugose, but it is indented with at least one pair of coarse pits (Fig. 6.279). ...........................................................32 - All pits on the pronotum are very fine (Fig. 6.280). .......................................33

Fig. 6.279 Paravelia dilatata: habitus (left) and the left paramere of the male (right). Based on Polhemus and Polhemus (1984c).

Fig. 6.280 Paravelia helenae: Habitus without appendages showing white as unmarked areas, black as solid areas, and bright orange red as stippled areas (upper left); middle (upper center) and hind femur (lower center) with the dense coating of hairy setae removed; apex of the abdomen (right); male paramere with its apex enlarged to its right (lower left center). Based in part on Polhemus (1969).

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32. (31) The pits on the pronotum are shallow and poorly defined (Fig. 6.281). The head is bright orange-red with a few small, yellowish spots. The hemelytra are black with yellow spots. Length: 5.2 to 5.3 mm. It inhabits bromeliads. ................................................Paravelia paolettii J. T. and D. A. Polhemus, 1991 (Venezuela). - The pits on the pronotum are well defined (Fig. 6.279). The color is mainly orange brown, and there are no strongly contrasting black and yellow markings. Length: 5.6 to 5.7 mm. ...............................................Paravelia dilatata J. T. and D. A. Polhemus, 1984 (Surinam, Amazonas).

Fig. 6.281 Paravelia paolettii male: habitus without appendages (upper left), head in lateral view (lower left), posterior view of the hind femur (lower left center), apex of the abdomen in ventral view (right center), paramere (upper right), and proctiger (lower right). Based on Polhemus and Polhemus (1991). 33. (31) There is a bubble-like structure at the base of each hemelytron, which is located between the costal and radial veins and is subdivided into about eight sections by fine, transverse brown lines. The posterior tibia of the male is not clavate (Fig. 6.282). Length: c. 4.0 mm. Color: Mainly orange brown. ............................................Paravelia bullialata J. T. and D. A. Polhemus, 1984 (Surinam, Guyana, Amazonas). - There is no bubble-like structure at the base of each elytron. The posterior tibia of the male is clavate (Fig. 6.280). The species is usually encountered in bromeliads. .........................................................................................................34 34. (33) The length is usually at least 5.6 mm, but if it is shorter, the posterior end of the abdomen forms a sharp, acute angle, or, if it is rounded, the color pattern is a strikingly bright combination of black, white, orange-red, and yellow (Fig. 6.280). ........................................................................................................35 - The length is shorter than 5.6 mm. The posterior end of the abdomen is always broadly rounded (Fig. 6.283). The color is mainly chocolate or orange brown. .................................................................................................................36

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Fig. 6.282 Paravelia bullialata: habitus in dorsal view (left) and a male paramere (right). Based on Polhemus and Polhemus (1984b).

Fig. 6.283 Paravelia recens (above, left to right): the hemielytra and outline of the thorax in dorsal view, male parameres of two specimens (above and below), first genital segment of a male in dorsal view, and (below, left to right): head in lateral view, male genitalia in lateral view, and the first genital segments of two specimens in lateral view. Based on Polhemus and Polhemus (1991). 35. (34) The posterior end of the pronotum forms a sharp, acute angle. The ventral process of the male is lacking. The male paramere is small and ovate (Fig. 6.284). Length: 5.6 to 5.7 mm. Color: mainly dark brown with the abdomen black. ...............................................Paravelia juruana J. T. and D. A. Polhemus, 1984 (Amazonas). - The posterior end of the pronotum is broadly rounded. The head, pronotum, and most of the underside are bright orange red with yellowish markings, contrasting strongly with the dorsal surface of the abdomen, which is blackish with three large, white spots when the wings are at rest (Fig. 6.280). The medial

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spot at the posterior end of the abdomen is formed from one white spot on each of the hemielytra, which become superimposed on each other when the wings are folded. The ratios of antenna segments I to IV are 48:40:45:44, respectively. Length: 5.6 to 5.9 mm. Width across humeri: c. 2.1 mm. The species is an inhabitant of bromelias. ....................................................................Paravelia helenae (Hungerford, 1929) (Peru). Syn: Velia helenae Hungerford, 1929.

Fig. 6.284 The paramere of a male Paravelia juruana. Based on Polhemus and Polhemus (1984c). 36. (34) Length: 3.5 to 3.6 mm. Color: mainly orange brown. The fourth antenna segment is shorter than the width of the head. The male paramere is blunt and ovate (Fig. 6.285). ...........................................Paravelia cupariana J. T. and D. A. Polhemus, 1984 (Pará). - The length is greater than about 4 mm. The color is chocolate brown (Fig. 6.283). .................................................................................................................37

Fig. 6.285 The paramere of a male Paravelia cupariana. Based on Polhemus and Polhemus (1984c). 37. (36) The apical white spot on the hemelytra is elongate and reaches the apex (Fig. 6.275). Length: 4.1 to 4.2 mm. Color: white markings on a dark brownish black ground color. It inhabits bromeliads. .........................................Paravelia manausana J. T. and D. A. Polhemus, 1984 (Amazonas). - The apical white spot showing on the folded hemelytra is round and does not reach the apex (Fig. 6.283). Length: c. 4 mm. The species inhabits bromeliads and was found in Guzmania brasiliensis Ule by Pereira et al. (2007). ...............................................................Paravelia recens Drake and Harris, 1935 (Central America, Colombia, Venezuela, Guyana, Peru, Bolivia, Amazonas, Pará).

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Key to the species of adult Platyvelia in South America Information for the key was provided by Drake (1951b), Polhemus and Manzano (1992), and Polhemus and Polhemus (1993b). 1. The posterior margin of the pronotum of the apterous form is nearly truncate and exposes a small, angulate corner of the mesonotum on each side (Fig. 6.286). When the pronotum is lifted at the posterior end, no traces of wing buds are detected. Length of apterous male: c. 4.1 mm. Color: blackish brown with a brown band across the anterior margin of the pronotum between the inner margins of the eyes and a pronotal disc somewhat lighter than the ground color. The habitat is marine. Only the apterous male has been described, and it was suggested that a winged form would not appear among such marine species. ..............................................Platyvelia maritima (Polhemus and Manzano, 1992) (Colombia). Syn: Paravelia maritima Polhemus and Manzano, 1992. - The posterior margin of the pronotum is not truncate in the apterous form and does not expose a only small section of the mesonotum at each posterolateral corner (Fig. 6.153), or forms at least bearing minute wing buds occur. ..............2

Fig. 6.286 Platyvelia maritima male: habitus without appendages in dorsal view (left) and paramere (right). Based on Polhemus and Manzano (1992). 2. (1) There is only one pair of large pits marked by silvery hairs on the anterior half of the pronotum. The width across the eyes is 0.8 mm. The pronotum also has some tufts of stout, slightly reddish brown bristles. The abdomen has a broad, U-shaped patch of silvery hairs on the three posterior tergites and small silvery intersegmental markings on the posterior half of the connexiva. The wings are vestigial, visible as very small white protrusions at the hind corner of the pronotum (Fig. 6.153). Length: c. 4.5 mm. Color: medium brown with lighter brown bands on the legs. .............................................................................Platyvelia brachialis (Stål, 1860) (North America, West Indies, southward to Argentina, Minas Gerais). Syn: Velia brachialis Stål, 1860; Velia stagnalis Uhler, 1894 (nec Burmeister); Velia australis Torre Bueno, 1916; Paravelia brachialis (Stål, 1860). - The pronotum of the macropterous form is coarsly pitted, or its posterior border is short and rounded. .................................................................................3

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3. (2) The pronotum is coarsely pitted and densely pubescent. The ratios of antenna segments I to IV are 42:27:25:30, respectively. The general color is brown with white basal and apical markings on the hemielytra. The legs are brown with testaceous bands. Length: c. 4.7 to 4.8 mm; maximum width: c. 1.7 to 1.8 mm. ..............................................................................Platyvelia verdica (Drake, 1951) (São Paulo, Amazonas). Syn: Velia verdica Drake, 1951. - The pronotum of the apterous form is coarsely pitted, while that of the macropterous form is much shorter with a rounded posterior border. The ratios of antenna segments I to IV are 56:38:40:37, respectively. The general color is brown to dark brown with a large elongate white spot at the base of the hemielytra and three white spots near the apex. The connexivum of the apterous form is broad and brown with silver spots between the segments. The legs are brown, densely covered with long, brown, hair-like setae, and banded with one indistinct ring near the apex of each femur and another at the base of each tibia. There is also a broad band across the middle of each tibia, and the ventral sides of the femora are yellowish fuscous. Length: c. 5.3 to 5.3 mm. ............................................................Platyvelia egregia (Drake and Harris, 1935) (Central America, Colombia). Syn: Velia egregia Drake and Harris, 1935. Key to the species of adult Steinovelia in South America Information for the key was provided by Drake (1951b) and Polhemus and Polhemus (1993b). The original descriptions of both species are sketchy, and there is little to distinguish them from one another because the original description of one was based on an apterous specimen of unspecified sex and the other was based on a macropterous female. 1. The hind tibia of the male has three dark bands. Length: c. 5 mm; width: c. 1 mm. (Fig. 6.147). The original description was based only on an apterous specimen. .............................................................................Steinovelia virgata (White, 1879) (Argentina, Paraguay, Amazonas). Syn: Velia virgata White, 1879; Paravelia virgata (White, 1879). - The hind tibia of the female has one dark band. The ratios between antenna segments I to IV are 35:32:33:32, respectively. The pronotum is deeply pitted, velvet chestnut brown becoming deeply fuscous on its posterior half, and displaying a length:width ratio is 62:55. Length: c. 4.8 mm; width: c. 1.38 mm. Color: mainly fuscous brown; the hemielytra are yellowish brown with pale brown veins, yellowish white cells, and a white, crescent-shaped marking at the base on each side. The antennae are mainly yellowish with a brown basal segment and pilose. Only a winged female was described. ............................................................................Steinovelia vinnula (Drake, 1951) (São Paulo). Syn: Velia vinnula Drake, 1951; Paravelia vinnula (Drake, 1951).

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Subfamily Microveliinae Key to the species of adult Microvelia in South America Information for the key was provided by Kirkaldy (1902), McKinstry (1937), Drake and Capriles (1952), Drake and Hussey (1955), Drake (1951c, d, 1952b, 1958), Drake and Hottes (1952), Drake and Roze (1958), Polhemus (1968a, 1972), Nieser and Alkins-Koo (1991), and Polhemus and Manzano (1992). Only the males of some species can be identified. 1. The metanotum contains deep, lunate furrows on each side, which are testaceous or black and very shiny (Fig. 6.287). Length: 2.4 to 2.65 mm. The color is very variable from testaceous to nearly completely black. ..................................................................................Microvelia arca Drake, 1958 (Santa Catarina). - There are no deep furrows in the metanotum (Fig. 6.288). ..............................2

Fig. 6.287 Thorax of Microvelia arca in dorsal view, showing the pair of sulcate furrows on the anterior part of the metanotum. Based on Drake (1958).

Fig. 6.288 Microvelia leucothea: habitus of a male (upper left) and a female (lower left) without appendages and two views of the male paramere (right). Based on Polhemus and Manzano (1992).

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2. (1) The fourth antenna segment of the winged female is slightly longer than the third, which is 1.25 times the length of the second. The first segment of the antenna is also distinctly longer than the second. The coloration is black with yellowish markings on the base of the first antenna segment, the basal ¾ of all femora, the anterior margin of the pronotum, and the ventral surface of the connexiva. The color of the rest of the legs, the apical parts of the antennae, and much of the hemielytra is dark sienna brown. There are extensive silvery white markings at the bases of the hemielytra, which also have seven or eight much smaller silvery spots distributed between the basal markings and the apices. The pronotum lacks a median carina, but on the head, there is a longitudinal sulcus along the midline of the vertex. Length of winged female: c. 2.2 mm. Maximum width: c. 0.8 mm. The description of this species is extremely sketchy, and a better description is urgently needed. Only the winged female has been described. ..........................................................................Microvelia stellata Kirkaldy, 1902 (West Indies, Ecuador, Peru, Bolivia, Paraguay, Brazil). - The proportions of the antenna, color pattern, size, or other features differ. For apterous specimens, continue to the next couplet. .........................................3 3. (2) In apterous and micropterous males, the pronotum is short and does not cover any part of the mesonotum (Fig. 6.288). The fourth segment of the antenna is longer than the third. The length varies from 1.7 to 3.0 mm. .............4 - Either the pronotum covers all or part of the mesonotum, or, if it does not, then the fourth segment of the antenna is less than ¾ as long as the third. If only macropterous specimens without orange markings on the pronotum are available, continue here. .......................................................................................6 4. (3) The color is black, somewhat obscured by dense, recumbent, silvery pubescence. The appendages are mostly testaceous. The pronotum is uniformly blackish. The eyes are only about ¼ as wide as the interocular space (Fig. 6.288). The length ratios of antenna segments I:II:III:IV are 19:19:27:34, respectively. Length of male: c. 1.31 mm; female: c. 1.47. Only wingless forms have been found. The habitat is the marine intertidal zone. ..............................................Microvelia leucothea Polhemus and Manzano, 1992 (Colombia). - The pronotum has markings paler than the ground color. The eyes are much wider than ¼ of the interocular width (Fig. 6.154). The habitat is primarily fresh water. ....................................................................................................................5 5. (4) The markings on the head and thorax are reddish brown. In apterous and micropterous males, the pronotum is about four times as wide as long. There are glabrous areas on some of the abdominal segments. The ratios of antenna segment lengths in order I:II:III:IV is 14:11:19:20 (Fig. 6.154). Length: 1.85 to 2.95 mm. The color is mainly blackish brown dorsally with velvety reddish brown areas on the head between the eyes and on a stripe along the vertex, as well as along a longitudinal stripe as wide as the head running the length of the pronotum, on broad median areas on both lobes of the mesonotum, and on the dorsum of the first three abdominal segments. The fourth through seventh

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abdominal segments are glabrous yellow brown dorsally, and the first laterotergites of the connexivum are yellowish or reddish brown. ......................................................................Microvelia ashlocki Polhemus, 1968 (Galapagos Islands). - The head of the male is black, and the marking on the pronotum is orange. In the macropterous form, the anterior part of the pronotum has an orange transverse band, usually interrupted in the middle and often obscured by silvery hairs. The pronotum of the male is not otherwise marked. The hind femur of the male is curved. The ratios of antenna segment lengths in order I:II:III:IV are 14:9:16:22 (Fig. 6.289). Length: 1.7 to 2.4 mm. Color: variable. ....................................................................Microvelia pulchella Westwood, 1834 (North America, West Indies, all South American countries except Chile; reported from Minas Gerais). Syn: Hydroessa pulchella Herrich-Schäffer, 1842; Velia (Microvelia) Pulchella Westwood, 1834. Two subspecies were recognized by Drake and Hussey (1955): Microvelia pulchella pulchella Westwood, 1834; and Microvelia pulchella incerta (W. F. Kirby, 1890). Synonyms for the subspecies include Microvelia borealis Torre-Bueno, 1916; Microvelia robusta Uhler, 1894; Microvelia marginata Uhler, 1894, partim.

Fig. 6.289 Microvelia pulchella (left to right): apterous male, apterous female, and macropterous female. Based on Cobben (1960a). 6. (3) The color pattern of the macropterous male is mainly black and fulvous, with blackish brown and grayish streaks on the hemielytra and flavous yellow on the following parts of the body: the posterior part of the head; posterolateral parts of the pronotum and a narrow transverse stripe across its anterior part; the venter along the midline, lateral margins, and posterior part; the connexivum except for its sutures; the coxae and trochanters; and the rostrum (Fig. 6.290). The ventral surfaces of the pleurae have a bluish gray pruinosity. The legs are brown. The apterous female appears short and fusiform with a ferrugineous color on the thorax and posterior parts of the abdomen. The two basal segments of the antenna are subequal in length; the third segment is much longer and

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subequal to the fourth. Length: c. 2.5 mm; width: c. 1 mm. The apterous male and macropterous female have not been described. ................................................................Microvelia panamensis Champion, 1898 (Panama, Colombia). - Color patterns are other than blackish with fulvous markings and brown legs for the macropterous male and dark with extensive ferrugineous markings for the apterous female. The length is usually significantly greater or less than 2.5 mm. .......................................................................................................................7

Fig. 6.290 Habitus of a macropterous male Microvelia panamensis. Based on Champion (1898). 7. (6) The length is at least 2.5 mm. .....................................................................8 - The length is no greater than 2.4 mm and usually much shorter. ...................13

Fig. 6.291 Microvelia isabelae: habitus of a male (left) and a female (right); right paramere (below center). Based on Bachmann (1979b).

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8. (7) The male is very dark brown with a reddish tinge and large yellow markings, sometimes darkened, on the dorsal surface of all segments of the thorax and abdomen, except the genital segments. The female has such yellow markings only on the thoracic segments (Fig. 6.291). Length of apterous male: c. 2.9 mm; apterous female: c. 2.6 mm. Maximum width of apterous male: c. 0.72 mm; apterous female: 0.88 mm. A macropterous form has not been described. .....................................................................Microvelia isabelae Bachmann, 1979 (South Isabela Island, Galapagos Islands). - Yellow markings are small or restricted to the thorax of both sexes (Fig. 6.292). ...................................................................................................................9 9. (8) The general color is dark grayish brown. A transverse band across the pronotum, interrupted in the middle, is located near the anterior margin. The hind femur of the male is nearly as long as or longer than the body. That of the female is about half of the body length, and it reaches the hind margin of the abdomen. The hind margin of the pronotum of apterous forms is deeply incised in the middle (Fig. 6.292). .............................................................................Microvelia longipes Uhler, 1894 (West Indies, Ecuador, Peru, Venezuela, Guyana, Paraguay, Argentina, Minas Gerais). Syn: Microvelia modesta Uhler, 1894 (females). - The general coloration is blackish, testaceous, or brown with yellow markings, and the transverse band across the pronotum is not interrupted in the middle (Fig. 6.293). ........................................................................................................10

Fig. 6.292 Habitus of an apterous male (left) and a macropterous female Microvelia longipes (right). Based on Cobben (1960a).

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Fig. 6.293 Microvelia ayacuchana: habitus of a male in dorsal view (upper left), antenna (lower center), male genitalia in dorsal (upper right) and lateral view (middle right), and the female genitalia in dorsal (lower left) and lateral view (lower right). Based on Spangler and Froeschner (1987). 10. (9) Length: 2.5 to 2.75 mm. ..........................................................................11 - The length is at least c. 3 mm. There is a keel on the last abdominal sternite of the male. .............................................................................................................12 11. (10) The abdomen is brown or testaceous ventrally, and there is a transverse yellowish or brownish band on a black or fuscous black background across the middle of the pronotum. The posteriormost sternite of the male is deeply notched. The connexiva of the female is strongly produced apically (Fig. 6.293). .................................................Microvelia ayacuchana Drake and Capriles, 1952 (Venezuela, Guyana). - The general coloration of the macropterous male is blackish with a short, transverse fulvous band near the anterior margin of the pronotum, which is almost pentagonal in shape. The hemielytra are dark fuscous with two silver white basal stripes and three to five silver spots on the posterior half. Length of macropterous male: c. 2.55 mm; width; c. 0.8 mm. The macropterous female has not been described. ...............................................Microvelia novana Drake and Plaumann, 1955 pars (Uruguay, Santa Catarina). See Couplet 38 for the apterous form. 12. (10) Ratios of antenna segment lengths in order I:II:III:IV are 20:18:31:33. Length of apterous form: 3.0 to 3.1 mm. Length of winged female: c. 3.55 mm.

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The general coloration is fuscous with yellow markings. The legs are dark with yellow bases. The apex of the pronotum is truncate (Fig. 6.294). The head is flavo-testaceous with a narrow black median line. ................................................................Microvelia braziliensis McKinstry, 1937 (Ecuador, Peru, Bolivia, Paraguay, Argentina, Minas Gerais, Rio de Janeiro). - Ratio of antenna segment length in order I:II:III:IV is 23:32:27:33. Length of winged and apterous forms: 3.8 to 4.1 mm. The general coloration is brown with rusty yellow markings. ...............................................................Microvelia hungerfordi McKinstry, 1937 (Paraguay, Argentina).

Fig. 6.294 Microvelia braziliensis: thorax and anterior abdominal segments in dorsal (left) and lateral view (right). Based on Drake and Hussey (1955). 13. (7) The basal parts of the hemielytra and a large, apical spot are pearly white, contrasting strongly with the blackish, fuscous, or testaceous ground color. The ventral surface is usually dark fuscous. The species inhabit bromeliads. .........................................................................................................14 - The color pattern is not mainly pearly white contrasting against a dark ground color; pale markings usually include yellow or dull white. ................................15 14. (13) The connexiva of the female are entirely dark fuscous. The ratios of antennal segments I to IV are 17:12:19:30, respectively. Length of the apterous female: c. 2.1 mm; maximum width: c. 0.75 mm. Males and apterous forms have not been described. .......................................................Microvelia ancona Drake and Chapman, 1954 (Panama, Colombia, Ecuador, Venezuela). - The parts of the connexiva beneath the pearly white parts of the hemielytra, that is, the first three segments, are testaceous on both the dorsal and ventral surfaces. Length: 1.9 to 2.3 mm. The body is mainly black, and the legs are straw colored. It commonly inhabits bromeliads. .........................................................................Microvelia distanti Lundblad, 1933 (West Indies, Venezuela). Syn: Microvelia insignis Distant, 1912 (preoccupied); probably Microvelia stellata Hynes, 1948 (nec Kirkaldy).

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15. (13) The large pronotum covers the entire dorsal part of the thorax. ...........16 - The pronotum does not cover the entire dorsal surface of the thorax, or, if it does, it is not truncate on its posterior margin and subrectangular in shape (Fig. 6.295). .................................................................................................................19

Fig. 6.295 An apterous male Microvelia argentata: habitus in dorsal view (left), outline of the pronotum in dorsal view (right), outline of first genital segment (upper left center), and enlarged second genital segment in dorsal view (right center). Based on Nieser and Alkins-Koo (1991). 16. (15) Coarse and deep pits are not prominent on the pronotum. The color is rufous brown with a whitish pubescence on the lateral parts of the mesonotum and bluish patches on the first two and apical three abdominal tergites. The connexiva are strongly reflexed, and the apical part of the abdomen is upturned. The bases of the hemielytra of both sexes are hairy. Length: 1.8 to 2.0 mm. ............................................................................Microvelia limaiana Drake, 1951 (Panama, Argentina, Santa Catarina). - Coarse and deep pits are prominent on the pronotum, at least on the posterior part. .....................................................................................................................17 17. (16) The male is black with transverse brownish to testaceous markings on the pronotum. The coloration of the female is entirely black without markings. Length of male: 1.65 to 1.75; female: c. 2.15 mm. ........................................................Microvelia duidana Drake and Capriles, 1952 (Venezuela, Guyana). - The known males lack transverse markings on the pronotum, and the known females are not uniformly black. The species in the following couplet are poorly known and in need of revision before identifications can be made with confidence. ..........................................................................................................18 18. (17) The ratios of antennal segments I to IV are 17:15:24:26. The pronotum of the female completely covers the rest of the thorax and has a subtruncate posterior margin, making it appear nearly rectangular with only slightly rounded corners. Posterior to the wavy line on the pronotum, the surface is distinctly

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pitted, contrasting with the smaller anterior area, which lacks punctures. Length of apterous female: c. 2.15 mm. The color is mainly brownish with testaceous tinges on the pronotum and testaceous markings on each side at the base of the abdomen and covering the three posteriormost abdominal segments. The ventral surface is mainly blackish. Only the apterous female has been described. ..............................................................Microvelia ioana Drake and Hottes, 1952 (Rio de Janeiro). - The ratios of antennal segments I to IV are 16:12:17:20, respectively. The color of the apterous male is brown, and there is no transverse marking on the pronotum. There are pale quadratic markings on each side of the first two abdominal tergites and pale central markings on the two posteriormost tergites. The abdomen appears especially hairy. There are many large, dark pits on the pronotum, including a pair near the base of the head. Length of apterous male: c. 1.85 mm; maximum width at base of abdomen: c. 0.75 mm.. Only the apterous male has been described. ............................................................................Microvelia recifana Drake, 1951 (Pernambuco). 19. (15) The ventral surface of the second genital segment of the male has prominent rounded ridges. The third genital segment does not have lateral spines. .................................................................................................................20 - The ventral surface of the second genital segment of the male lacks prominent rounded ridges (Fig. 6.295). ...............................................................................21 20. (19) The first antennal segment is pale at the base, slightly curved, and much thicker and almost 1.5 times as long as the second. The third and fourth antennal segments are slender; the third is almost as long as the combined length of the first and second but only about ¾ as long as the fourth. The middle femora are greatly enlarged. Length: c. 2.2 mm. Maximum width: 0.6 to 0.7 mm. Only the apterous male has been described. ..............................................................Microvelia sarpta Drake and Harris, 1936 (São Paulo). - The ratios of antennal segments I to IV are 16:10:22:33. Length of macropterous form: 1.9 to 2.2 mm; maximum width of macropterous form: 0.70 to 0.82 mm. An apterous form has not been encountered. ........................................................Microvelia parana Drake and Carvalho, 1954 (Pará, Mato Grosso). 21. (19) The pronotum only covers a very small part of the mesonotum. There is a transverse orange yellow marking on the pronotum, which is not connected to any stripe or spot on an adjacent part of the thorax of the macropterous form. There is a prominent first genital segment; the hypandrium and anal lid lack lateral projections or points. The left paramere is bigger than the right. The proportions of the first through fourth antennal segments are 14:9:19:21, respectively. Dorsally, the color pattern is dark brown with yellow markings; ventrally, it is lead-blue. ................................................................Microvelia peruviensis McKinstry, 1937 (Peru).

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- The pronotum covers at least almost half of the mesonotum (Fig. 6.295), or the hypandrium and anal lid bear obvious lateral projections or points, or the yellow or orange band on the pronotum adjoins similarly colored markings on the mesonotum or is dark or indistinct, or the second antennal segment is more than ¾ as long as the third. .................................................................................22 22. (21) The general coloration of both males and females is pale, typical of cave-dwelling organisms. The ground color is yellowish brown with brown areas dorsally on the head, posterior 2/3 of the pronotum, lateral parts of the tergites, genital segments, antennae, and legs except for the pale basal parts of the femora. The pronotum has evenly rounded lateral margins and completely covers the mesonotum. The female has a deep excavation between the seventh abdominal segment and the genital segments (Fig. 6.296). Length: 1.7 to 1.9 mm. ....................................................................Microvelia cavernula Polhemus, 1972 (Venezuela). - The markings are darker than those typical of pale, cave-dwelling species. The female does not have a deep excavation between the seventh abdominal segment and the genital segments (Fig. 6.295). The pronotum of the male is sometimes a different color on the posterior part than the anterior part. Some females may be entirely black. The abdomen is black ventrally, or, if not, the head is not black. ................................................................................................23

Fig. 6.296 Seventh abdominal segment and the genital segments of an apterous male Microvelia cavernula in ventral view. Based on Polhemus (1972). 23. (22) The male apterous form bears a large, pointed ventral tubercle on the second abdominal segment. The lateral horns of the apical genital segment project caudally. The orange transverse marking on the anterior part of the pronotum has a median longitudinal projection giving it a T-form. The dorsoposterior part of the head is light brownish. Length: slightly more than 2 mm. ..................................................................Microvelia albonotata Champion, 1901 (North and Central America, West Indies, Venezuela, Peru). Syn: Microvelia capitata Barber, 1939, nec Guérin.

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- The male apterous form does not have a large pointed ventral tubercle on the second abdominal segment, and the orange transverse marking on the pronotum, if present, does not have the shape of a T (Fig. 6.295). The length is usually less than 2 mm. ..........................................................................................................24 24. (23) In addition to the anterior orange band, the pronotum has broad orange or yellow markings at the posterior border. Length: 1.4 to 1.8 mm. Only macropterous forms are known. .........................................................................Microvelia marginata Uhler, 1893 (North and Central America, West Indies, Venezuela, Peru). Syn: Microvelia pudoris Drake and Harris, 1936. Two very similar West Indian species have been described, which are scarcely distinguishable from this species, so a revision of the group will be necessary to determine the actual ranges of each (Nieser and Alkins-Koo, 1991). - Markings on the posterior part of the pronotum, if present, are less extensive than a broad orange or yellow band (Fig. 6.295). ..............................................25 25. (24) The pronotum and connexiva lack prominent silvery pubescence and are uniformly dark fuscous brown or brown with obscure darker markings. The ventral surface is dark fuscous with a bluish luster or testaceous. .....................26 - The pronotum or connexiva of the male and usually the female are not uniform in color but rather have obvious pale or black markings or silvery pubescence (Fig. 6.295). In case of doubt, the first and third segments of the antenna are not subequal in length, or the first genital segment of the male is not excavated or truncate apically, or the marking on the pronotum is brick-red or orange with small silvery setae and interrupted along the midline. If the female is uniform in color, it is black and not dark fuscous. .........................................27 26. (25) The pronotum and connexiva are uniformly dark fuscous brown without prominent silvery pubescence. The ventral surface is dark fuscous with a bluish luster. The head is dark fuscous brown with a pair of broad longitudinal stripes between the eyes separated by a median line. The hemielytra are uniformly dark brown. The first genital segment of the male is deeply, broadly, and roundly excavated but convex apically. The second genital segment is mainly concealed by the first. The ratios of antenna segments I to IV are 11:7:11:17, respectively. The first antennal segment is stout; the second is enlarged distally, and the third is the most slender. Length: 1.5 to 1.7 mm; maximum width: 0.62 to 0.70 mm. Only the macropterous form has been described. The original description is cursory, and several important features are left undescribed, making identification difficult. ............................................................................Microvelia lujanana Drake, 1951 (Argentina). - The pronotum and connexiva of apterous specimens are fuscous brown or brown. The entire ventral surface is testaceous. The head is convex and either fuscous or brown, sometimes with wide bands separated by a median line. The ratios of antenna segments I to IV are 9:7:10:15, respectively. The male is oblong and narrows posteriorly, but the female is ovate. The pronotum covers about half of the mesonotum. Length of apterous male: c. 1.5 mm; apterous

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female: 1.25 to 1.50 mm. Maximum width of the male: c. 0.54 mm: female: 0.50 to 0.62 mm. Winged forms have not been described. ........................................................Microvelia inannana Drake and Hottes, 1952 (Argentina, Brazil). 27. (25) The male is usually shorter than about 1.6 mm. ...................................28 - The size of the apterous form is usually equal to or greater than 1.6 mm. The pronotum of macropterous males is generally marked with a different color on the posterior part than the ground color on the anterior part, or it has a broad fulvous, orange, or reddish stripe reaching laterad at least as far as the middle of the compound eyes (Fig. 6.295). ........................................................................32 28. (27) The general coloration is brown covered by velvety pruinosity, but there is a transverse yellow marking on the anterior part of the pronotum. The hemielytra are pale brown and lack prominent veins. The head, which has a median, indented, glabrous line, is withdrawn beneath the pronotum to the posterior part of the eyes, which are large. The ratios of antenna segments I to IV are 9:6:7:13, respectively. The pronotum is coarsely pitted, subtruncate at the apex, and has a length, measured along the mid-line, which equals about 3/5 of the width across the humeri. The ventral surface is dark. The legs are pale brown with darker apices of the tibiae and tarsi. Length: c. 1.40 to 1.45 mm; maximum width: c. 0.61 mm. The original description is cursory and inadequate for positive identification of this species. Only macropterous specimens have been described. .........................................................Microvelia summersi Drake and Harris, 1928 (West Indies, Panama, Guyana, Brazil). The reported range may not be fully accurate because of the difficulties distinguishing this species from several similar ones. - The ratios of the antenna segments to each other are different, or the color is not predominantly brown, or the pronotum lacks a yellow transverse marking on its anterior half, or the total length is clearly shorter or longer than 1.40 to 1.45 mm. For apterous and micropterous specimens, continue here to see whether it can be identified. ................................................................................................29 29. (28) The genital segments of the male are largely withdrawn beneath the sternites anterior to it. .........................................................................................30 - The genital segments of the male are largely visible and not withdrawn beneath the sternites anterior to it (Fig. 6.297). .................................................31 30. (29) The fourth segment of the antenna is obviously shorter than the combined length of the second and third. The first genital segment appears as a yellow semi-circle, and the second segment does not extend beyond its edges. The visible edges of the U-shaped emargination of the first genital segment bear about eight to ten long, yellow hairs, and its external portions are also covered with yellow hairs directed dorsad. The general coloration is dark plumbeous to black with a flavo-testaceous band on the pronotum. Fine silvery hairs are present. Length: 1.3 to 1.4 mm. ........................................................Microvelia venustatis Drake and Harris, 1933 (Peru, Paraguay, Argentina, Amazonas, Minas Gerais, Rio de Janeiro).

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- The fourth segment of the antenna is at least as long as the combined length of the second and third. The ratios of antenna segments I to IV are 12:9:11:22, respectively. The first genital segment is concavely excavated at its posterior end, and it bears short longitudinal rows of long, brown, bristle-like setae at its lateral margins. The general coloration is blackish with an orange-yellow band across the anterior part of the pronotum. Fine dark brown hairs are present. Length: 1.3 to 1.4 mm. Maximum width: c. 0.75 mm. The ventral surface is bluish black. Only the apterous male has been described. ..............................................................................Microvelia minima Drake, 1952 (Rio de Janeiro, Santa Catarina). 31. (29) In the micropterous and apterous forms, the male has small tubercles on the second and sixth abdominal sternites. The proximal genital sternite is widened to accomodate the laterally projecting horns of the apical genital segment (Fig. 6.297). In macropterous forms, the dorsoposterior part of the head is usually light brownish, and the anterior orange band is not interrupted in the middle. Posterior markings on the pronotum are lacking. Ratio of antenna segment lengths in order I:II:III:IV is about 7:5:8:14. Length of apterous male: 1.20 to 1.56 mm; apterous female: c. 1.95 mm; macropterous form: c. 1.75 mm. ..............................................................................Microvelia mimula White, 1879 (West Indies, Central America, Ecuador, Paraguay, Uruguay, Argentina, Amazonas, Minas Gerais, Mato Grosso, Mato Grosso do Sul). Syn: Microvelia capitata Uhler, 1894, nec Guérin; Microvelia mendozana Jensen-Haarup, 1920; Microvelia myersi McKinstry, 1937. - In the micropterous and apterous forms, the male lacks small tubercles on the second and sixth abdominal sternites. The proximal genital sternite is not greatly widened, and the horns of the apical genital segment project outward. The ratios of antenna segment length in order I:II:III:IV are about 7:4:8:12. Length: 1.3 to 1.5 mm. A macropterous form has not been described. ..................................................................................Microvelia hinei Drake, 1920 (North America, West Indies, southward to Argentina).

Fig. 6.297 The eighth through tenth abdominal segments of a male Microvelia mimula in dorsal view. The ninth and tenth are also referred to as the proximal and distal genital segments. Based on Drake and Carvalho (1954).

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32. (27) The pronotum has a broad fulvous, orange, or reddish stripe contrasting with the blackish ground color and reaching laterad at least as far as the middle of each compound eye (Fig. 6.295). ...................................................................33 - The pronotum lacks a broad fulvous stripe extending laterad as far as the middle of each compound eye. ...........................................................................35 33. (32) The band on the anterior part of the pronotum is brick-red or orange. The pronotum covers the mesonotum and leaves less than half of the metanotum uncovered (Fig. 6.295). The pronotum is dark gray with a transverse brick-red or orange transverse band across the anterior part. The first genital segment of the male is notched. There is a broad median depression in the middle of the ventral surface of the abdomen. The length ratios of antenna segments I:II:III:IV are 9:6:10:12. The ground color is dark gray or black with silvery setae covering much of the dorsal surface, uncluding the anterior transverse stripe on the pronotum. It inhabits bromeliads. Only the apterous male has been described. ...............................................Microvelia argentata Nieser and Alkins-Koo, 1991 (Trinidad). - The band on the anterior part of the pronotum is fulvous. ..............................34

Fig. 6.298 The ninth and tenth abdominal segments, also referred to as the proximal and distal or first and second genital segments, of a male Microvelia quieta in dorsal view. Based on Drake and Carvalho (1954). 34. (33) The ratios of the antenna segment lengths in order I:II:III:IV are 19:11:22:26. The pronotum has patches of silvery pubescence. Length of apterous male: c. 1.6 to 1.7 mm; macropterous form: c. 2.0 mm. A macropterous female has not been reported. ........................................................................Microvelia hambletoni Drake, 1951 (Peru, Argentina, Santa Catarina, Minas Gerais). - The length ratios antenna segments I to IV are 16:11:21:33, respectively. The anterior 1/3 to 2/3 of the pronotum is blackish fuscous with a yellowish transverse marking starting behind the anterior margin; the posterior part of the pronotum is brownish testaceous. The pronotum covers about half of the mesonotum and is somewhat concave at the midline along the posterior margin.

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Both genital segments of the male have distinctive lateral processes (Fig. 6.298). Length of apterous form: 1.8 to 2.0 mm; maximum width: 0.65 to 0.80 mm. A macropterous form is unknown. .........................................................Microvelia quieta Drake and Carvalho, 1954 (Minas Gerais). 35. (32) The pronotum does not reach posteriad as far as the metanotum; it covers only about ½ to ¾ of the mesonotum. The color is mainly testaceous with black or paler markings. .............................................................................36 - The pronotum reaches the metanotum and covers at least part of it. The second genital segment lacks spines or tubercles. ..............................................39 36. (35) The color of the male is reddish brown or orange with the second, third, fourth, and most of the fifth and sixth tergites blackish. The venter is testaceous. The pronotum conceals about half of the mesonotum, and only the hind corners of the metanotum are visible on both sides of the mesonotum. The second genital segment lacks spines or tubercles. The female is almost entirely testaceous. Length of apterous male: c. 1.9 mm. Winged forms are unknown. ..........................................................Microvelia chilena Drake and Hussey, 1955 (Chile). - The coloration is mainly testaceous with blackish markings. The ventral surface of the abdomen lacks spines and tubercles. ...........................................37 37. (36) The first genital segment of the male widens posteriad and is truncate along the posterior margin, where it is widely, roundly, and deeply excavated. The second genital segment of the male bears a long, slender, straight spine, which is directed laterad. The ratios for antennal segments I to IV are 16:11:20:32, respectively, for both sexes. The pronotum conceals about 3/5 of the mesonotum. The connexiva of the female are wide, erect, but not reflexed. Length of apterous male: c. 1.65 mm; maximum width: c. 0.58 mm. Length of apterous female: c. 1.85 mm; maximum width: 0.75 mm. Macropterous specimens have not been described. Color: mainly brownish testaceous to dark fuscous. ..................................................Microvelia aemulana Drake and Plaumann, 1955 (Argentina, São Paulo, Santa Catarina). - The first genital segment of the male is not deeply excavated along the posterior margin, and the second genital segment does not bear laterally directed spines. .................................................................................................................38 38. (37) The hind femur is not thicker than the other femora; it is convex ventrally and unarmed. Apterous males are mainly testaceous with black on the lateral parts of the pronotum, the anterior part of the head, and along the sutures of the thorax and connexivum. Spines and tubercles are lacking, and the second abdominal segment is completely flat on the ventral surface. Length of male: c. 2.0 mm. Females or forms with wings have not been described. .......................................................Microvelia costaiana Drake and Hussey, 1951 (Rio de Janeiro). - The hind femur of the male is somewhat bowed and strongly flattened ventrally, and it bears a fringe of pale hair-like setae on its external edge. The

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length ratios of antenna segments I to IV are 18:13:30:42 for the male and 16:12:22:36 for the female. Apterous males and females are mainly brownish testaceous with blackish fuscous markings. The abdomen has broad blackish transverse bands across the segments, or the abdominal segments may appear black. Length of apterous form: 2.0 to 2.25 mm. ...............................................Microvelia novana Drake and Plaumann, 1955 pars (Uruguay, Santa Catarina). See Couplet 11 for the macropterous form. 39. (35) The fore-tibia ends in a short, cylindrical, apical projection. Only macropterous specimens have been described. The general coloration is blackish brown with lighter brown on the anterior part of the pronotum. In dorsal view, the pronotum but no parts of the mesonotum or metanotum are visible (Fig. 6.144). The posterior margin of the apical sternite of the male is deeply and broadly excavated and lacks an apical notch. The first through fourth antenna segments measure 0.17 mm, 0.15 mm, 0.20 mm, and 0.30 mm, respectively. Length of males and females: c. 1.9 mm. ..............................................................Microvelia timida Drake and Roze, 1958 (Venezuela). - The fore-tibia lacks a short, cylindrical, apical projection. The metanotum is not covered completely by the pronotum, which lacks a median carina. The punctures on the pronotum are nearly concealed by a velvety pubescence. The anterior part of the pronotum is rufous brown. Length of male: 1.8 mm; female: 2.3 mm. Color: black fuscous, sometimes with a velvety sheen. The abdomen is black or bluish black dorsally and ventrally. ..............................................................................Microvelia potama Drake, 1958 (Bolivia). Key to the species of adult Husseyella in South America Information for the key was provided by Drake and Harris (1933b) and Drake (1958). 1. The body is almost entirely bluish black and thickly coated by pubescence and longer dark setae. The legs and antennae are dark brown with testaceous areas on the coxae, trochanters, and basal parts of the femora (Fig. 6.148). Length of male: c. 2.25 mm; female: c. 2.5 mm. ..........................................................................Husseyella halophila Drake, 1958 (Santa Catarina). - The body is mainly brownish black; the connexiva are reddish brown and covered by a moderate coat of long, dark setae. Length of apterous female: c. 2.5 to 2.6 mm. Maximum width: c. 1.2 mm. The original description was based on female specimens, and the male closely resembles the female. ........................................................Husseyella diffidentis Drake and Harris, 1933 (Goiás).

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Hydrometridae Key to the genera in South America The family Hydrometridae is represented in South America by two subfamilies. Heterocleptinae Villiers includes only the genus Veliometra, while Hydrometrinae Billberg encompasses Bacillometroides, Bacillometra, and Hydrometra. Information for the key was provided by Nieser (1970b), Andersen (1977), Bachmann (1977a), Froeschner (1981), and Polhemus and Polhemus (2010). 1. The first segment of the antenna is much longer than the second and extends beyond the apex of the head by more than half its own length. Its second segment arises subapically from the first. The pronotum is only slightly longer than the head, which is subcylindrical posterior to the eyes (Fig. 6.299). There is one scent gland opening in the metasternum. The length is 3 to 5 mm. .....................................................................................Veliometra Andersen, 1977 The only species in this genus is Veliometra schuhi Andersen, 1977, from Amazonas. - The first segment of the antenna is subequal to or shorter than the second and usually only barely extends beyond the apex of the head. The second segment arises apically from the first (Fig. 6.300). There are no scent gland openings in the metasternum. The length is at least 6 mm. ....................................................2

Fig. 6.299 Veliometra schuhi: habitus without appendages in dorsal view (below), left middle tarsus in lateral view with its ventral arolium enlarged to its right and its campaniform sensillum enlarged to its upper left (upper left), and the pygophore and parameres in dorsal view (right). Based on Schuh and Slater (1995). 2. (1) The middle coxae are inserted approximately midway between the fore and hind coxae. The body is covered with small black denticles. The length of

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the thorax measured along the dorsal midline is less than twice its maximum width. The lengths of antenna segments I through IV are 0.50:1.20:2.90:1.90, respectively. The fourth segment of the antenna is bulbous apically. The width of each mediotergite is greater than its length. The mesosternum has a median longitudinal sulcus, and the metasternum has two distinct longitudinal sutures that are prolonged onto the abdomen (Fig. 6.300). There is an elevated triangular area on the metanotum, just posterior to the pronotum. The trichobothria of the posterior pair on the head arise from small tumescences. The length of the macropterous female of the only species in this genus is approximately 9.2, and its maximum width is 1.2 mm. ........................................................................................Bacillometra Esaki, 1927 The only South American member of this genus is Bacillometra ventralis Esaki, 1927, found in French Guiana and Venezuela. - The middle coxae are closer to the fore-coxae than to the hind ones (Fig. 6.301). ...................................................................................................................3

Fig, 6.300 Habitus of Bacillometra ventralis in dorsal view with the fourth and apical part of the third antenna segment removed. Based on Hungerford and Evans (1934). 3. (2) The mesosternum and metasternum lack a sulcus or sutures (Figs. 6.29, 6.301). The scutellum is concealed. The second tarsal segment is longer than the third. ........................................................................Hydrometra Lamarck, 1801..p. 258 - The mesosternum has a median longitudinal sulcus, and the metasternum has two distinct longitudinal sutures that are prolonged onto the abdomen (Fig. 6.302). The scutellum is exposed. The setae covering the body are fine and hair-like. The length of the thorax measured along the dorsal midline is more than twice its maximum width. The fourth segment of the antenna is not

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bulbous apically. The width of each mediotergite is less than its length. The trichobothria of the posterior pair on the head arise from large tubercles. The length of the macropterous female of the only species in this genus is approximately 9.2 mm, and its maximum width is 1.2 mm. ......................................Bacillometroides J. T. and D. A. Polhemus, 2010..p. 265

Fig. 6.301 Hydrometra quadrispina: habitus of a male without appendages in dorsal view (upper left); head without the antenna in dorsal (lower left) and lateral view (lower left center); apex of the male abdomen in ventral (lower center) and lateral view (lower right center); apex of the female abdomen in dorsal (upper right), ventral (middle right), and lateral view (lower right). Based on Pérez Goodwyn (2001b).

Fig. 6.302 Bacillometroides woytkowskii: habitus of a macropterous male without appendages on the right side in dorsal view (lower left), apex of an apterous female without appendages (middle left) based on Hungerford (1935) and another without appendages on the left side (upper left) based on Aguilar et al. (1987), thorax and abdomen of a female in ventral view (upper center), ventral view of the apex of the abdomen of a male (lower right center) and a female (middle right), and a lateral view of the apex of a male abdomen (upper right). Based on Hungerford (1935) and Aguilar et al. (1987).

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Key to the species of adult Hydrometra in South America Information for the key was provided by Torre-Bueno (1926), Jaczewski (1928a), Hungerford and Evans (1934), Drake (1953b), Drake and Lauck (1959b), Mychajliw (1961), Nieser and Alkins-Koo (1991), Pérez Goodwyn (2001b), and Moreira et al. (2009). Both macropterous and brachypterous forms commonly occur. 1. The anterior part of the clypeus is excavated (Fig. 6.303). Thoracic pits are absent. The general color is reddish brown, but the head and venter are darker. A thin silvery line bordered with black runs along the posterior 66% of the pronotum, and silvery lateral stripes are present. The abdominal tergites are blackish with silvery lateral stripes. Length of male: c. 11.4 mm. Female: 11 to 13 mm. .................................................................Hydrometra comata Torre-Bueno, 1926 (West Indies, Colombia, Surinam). - The clypeus is not excavated (Fig. 6.301). .......................................................2

Fig. 6.303 Hydrometra comata male: clypeus (left) and right side of the apex of the abdomen in lateral view (right). Based on Nieser (1970). 2. (1) The clypeus appears broad and quadrate with a truncate, slightly rounded, or medially angulate anterior apex. The surface of the pleurae is dull and pitted (Fig. 6.29). ............................................................................................................3 - The clypeus appears narrow and elongate or conical (Fig. 6.301). ................12 3. (2) The antenna is slender and as long as the body. The fourth segment of the antenna is twice as long as the first. The fore and middle acetabula lack pits. The seventh segment of the male abdomen has a pair of broad ventrolateral depressions and two oval patches of dense setae (Fig. 6.304). Length: c. 18 mm. Descriptions of this species have been sketchy, but the length of the antenna is distinctive. ...........................................................................Hydrometra metator White, 1879 (Amazonas). - The antenna is distinctly shorter than the body, or the fourth segment is much more than twice as long as the first (Fig. 6.305). .................................................4

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Fig. 6.304 Hydrometra metator male (left to right): the seventh abdominal segment in ventral view with parts of the adjacent segments, and the arrangements of the pits on the fore, middle, and hind acetabula. Based on Drake (1954).

Fig. 6.305 Hydrometra olallai (above, left to right): apical segments of a male abdomen in dorsal, ventral, and lateral view, and (below, left to right): apical segments of a female abdomen in dorsal, ventral, and lateral view. Based on Mychajliw (1961). 4. (3) The head is mainly blackish; its anteocular length is between 2.5 and 2.75 times the length of the postocular part. The length ratios of antenna segments I to IV are 21:50:215:80, respectively. The pronotum is brown on the anterior part and flavous on the posterior lobe. Both lobes have a deep, median, longitudinal line, which is wider on the posterior lobe. The acetabula are brownish. Each acetabulum of a fore or middle leg has 8 or 9 deep pits anterior to the cleft and 10 or 12 pits posterior to it. Each acetabulum of a hind leg has 8 to 12 pits. The legs are brown with a reddish tinge and fuscous apices of all femora. The ventral side of the abdomen has several long hairs and a frosty stripe on each side. The pronotal and acetabular pits are large. The genital

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segment of the female ends dorsally in a short, straight, dark spine. Length: 12 to 14 mm. ...............................................................................Hydrometra zeteki Drake, 1952 (Mexico, Central America, Colombia). - If the length is between 12 and 14 mm, the anteocular length is not equal to between 2.5 and 2.75 times the postocular length, the color pattern varies, or the ratios of the antenna segments differ (Fig. 6.305). ...............................................5 5. (4) The anteocular length of the head equals between 2.8 and 3.2 times the postocular length. All acetabula have pits, which usually number at least 6 both before and after the cleft (Fig. 6.305). ..................................................................6 - The anteocular length of the head is not equal to between 2.8 and 3.2 times the postocular length (Fig. 6.306). .............................................................................8 6. (5) The sixth abdominal segment and first genital segment of the male each have a pair of small elevations covered with black setae. The first genital segment of the female has a supraanal plate ending is a sharp spine (Fig. 6.305). The length ratios of antenna segments I to IV of the female are 55:118:400:130, respectively. Each acetabulum of a fore-leg has 7 pits anterior to the cleft and 6 pits posterior to it. Those of a middle leg each have 7 pits anterior to the cleft and 8 pits posterior to it. Each acetabulum of a hind leg has 8 pits. Length of male: 14.5 to 15.0 mm. Female: 16 to 19 mm. The anterior part of the pronotum is brown, and the posterior lobe is usually yellowish with a purple band and sometimes a central silvery line. Brachypterous specimens with padlike wings occur. ......................................................................Hydrometra olallai Mychajliw, 1961 (Amazonas). - The sixth apical abdominal segment and the first genital segment of the male have sparse coverings of long setae, and large ventrolateral depressions may also be present. The very long legs are fuscous with predominantly brownish tibiae. The hind femur is much longer than the abdomen. ..............................................7

Fig. 6.306 Hydrometra fruhstorferi: apical segments of the male (upper left) and female abdomen in dorsal view (lower left) and the same segments of a male (upper right) and a female (lower right) in lateral view. Based on Hungerford and Evans (1934).

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7. (6) The length ratios of antenna segments I to IV are 13:19:95:41, respectively. Each acetabulum of the fore-leg has 9 or 10 pits anterior to the cleft and 12 or 13 pits posterior to it. Those of the middle leg each have 9 to 13 pits anterior to the cleft and 12 or 13 pits posterior to it. Each acetabulum of a hind leg has 15 to 17 pits. Length of macropterous specimens: 13.7 to 15.0 mm. The head is blackish fuscous with a brown base. The predominant color is brown, and the hemielytra are dark fuscous. .......................................................................Hydrometra fuanucana Drake, 1954 (Peru). - The length ratios of antenna segments I to IV of the female are 28:64:210:85, respectively. Each acetabulum of a fore-leg has 7 or 8 pits anterior to the cleft and 9 to 11 pits posterior to it. Those of a middle leg each have 7 to 11 pits anterior to the cleft and 10 or 11 pits posterior to it. Each acetabulum of a hind leg has 15 to 18 pits. Length: 14 to 16 mm. The head is fuscous black with a brown base. The predominant color is brownish fuscous. The clypeus is dark reddish brown, and the first segment of the antenna is black. .....................................................................Hydrometra huallagana Drake, 1954 (Peru). 8. (5) The anterior part of the pronotum is reddish brown. A median silvery line bordered with black runs for almost the whole length of the pronotum. The anteocular part of the head is 2.5 times as long as the postocular part. The lengths of antenna segments I to IV, not including the intersegmental parts, are 0.83:1.82:6.50:2.14, respectively. The clypeus has only a slight rounded projection (Fig. 6.29). Length of male: 12.2 to 14.0 mm. Female: 13 to 19 mm. ........................................................Hydrometra caraiba Guérin-Méneville, 1857 (West Indies, North and Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Amazonas). Syn: Limnobates caraiba Uhler, 1886; Gerris caraiba Lethierry and Servin, 1898; Hydrometra agenor Kirkaldy, 1902; Hydrometra championiana Torre-Bueno, 1926. - The anterior part of the pronotum is not reddish brown, or a median silvery line bordered by black is not evident on the pronotum (Fig. 6.306). ...................9 9. (8) The color is mainly dark brown to black, but there are lighter markings on the pronotum and abdomen. The male has a pair of lateral tufts of hair-like setae, each in the form of an incomplete semicircle, on the seventh abdominal sternite (Fig. 6.306). The anteocular part of the head is twice as long as the postocular part. The length ratios of antenna segments I to IV are 14:19:60:34, respectively. Each acetabulum of a fore-leg has 6 or 7 pits anterior to the cleft and 8 or 10 pits posterior to it. Those of a middle leg each have 5 to 8 pits anterior to the cleft and 8 or 10 pits posterior to it. Each acetabulum of a hind leg has 9 to 12 pits. The length is 12.0 to 13.3 mm. ...........................................Hydrometra fruhstorferi Hungerford and Evans, 1934 (Argentina, Santa Catarina). Syn: Hydrometra brasilana Drake, 1952. - The color is lighter, and there is often a silvery median stripe that does not reach the anterior part of the pronotum. (Fig. 6.307). The male is at least 14 mm long, and the female is somewhat longer. ..........................................................10

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10. (9) The male genital segments have long setae forming a distinctive pattern on the ventral surfaces. The pale median line runs the entire length of the pronotum. Where the median line crosses the posterior lobe of the pronotum, it contains and is bordered by many pits. The anterior lobe is free of pits. The acetabular pits are well defined. The anteocular part of the head is 2.5 to 2.6 times as long as the postocular part. The length ratios of antenna segments I, II, and IV are 70:155:235, respectively. The third segment of the antenna is very long, but its exact length was not reported. The acetabulum of the fore-leg has 7 well-defined pits anterior to the cleft and 10 pits posterior to it. The middle acetabulum has 8 pits anterior to the cleft and 9 pits posterior to it. The hind acetabulum has 15 pits (Fig. 6.307). Length: c. 15 mm. The predominant color is brownish, somewhat testaceous on the posterior lobe of the pronotum, and the hemielytra and sternum are dark fuscous. The base and apex of the head are black. ............................................................Hydrometra exalla Drake and Lauck, 1959 (Peru). Syn: Hydrometra excella Drake and Lauck, 1959 (different spelling used in the same publication that contained the description). - The male genital segments do not have long setae forming a distinctive pattern on the ventral surfaces, although long setae may be present on the dorsal surface of the male of one species (Fig. 6.308). There are curved lines of short setae on the ventrolateral surfaces of one or two of the abdominal segments. ................11

Fig. 6.307 Hydrometra exalla male: apical segments of the abdomen in lateral (upper left) and ventral view (upper right) and the fore, middle, and hind acetabular and propleural pits (below, left to right). Based on Drake and Lauck (1959b). 11. (10) The apical segment of the male abdomen visible in dorsal view is broadened laterally to form triangular processes. The sixth abdominal segment has a pair of U-shaped ridges bearing a short brush of setae. A thin silvery line bordered with black runs along the posterior 60% of the pronotum. The

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anteocular part of the head is about 3.1 to 3.2 times as long as the postocular part. The lengths of antenna segments I to IV of a brachypterous female, not including the intersegmental parts, are 0.54:1.75:6.08:2.60, respectively. The clypeus ends in a short, sharp point. The apical segment of the female abdomen ends in a narrow, acutely pointed process (Fig. 6.308). Length of male: 14.3 to 14.9 mm; female: 15.0 to 18.4 mm. The dominant color is reddish brown; the head is dark. ..............................................Hydrometra guianana Hungerford and Evans, 1934 (West Indies, Colombia, Venezuela, Guyana, Surinam, Peru, Pará, Amazonas, Mato Grosso). - The apical segment of the male abdomen that is visible in dorsal view has nearly straight sides and is not broadened laterally to form triangular processes. There is a pair of large oval depressions located on the ventrolateral surfaces and surrounded by short, inwardly-directed setae (Fig. 6.309). The anteocular part of the head is about 2.6 to 2.7 times as long as the postocular part. Length: c. 15.6 mm. ...............................................Hydrometra williamsi Hungerford and Evans, 1934 (Ecuador).

Fig. 6.308 Hydrometra guianana: apex of the male abdomen in dorsal (upper left) and lateral view (lower left); apex of the female abdomen in dorsal (upper right) and lateral view (lower right). Based on Hungerford and Evans (1934).

Fig. 6.309 Hydrometra williamsi male: apical segments of the abdomen in dorsal (left) and lateral view (right). Based on Hungerford and Evans (1934).

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Fig. 6.310 Hydrometra sztolemani: apical segments of the abdomen of a male (left) and a female (right). Based on Hungerford and Evans (1934). 12. (2) There are two large processes on each lateral edge of the sixth abdominal segment of the male, which overlap the posterior edge of the fifth segment. The terminal process of the female abdomen is blunt (Fig. 6.310). ................................................................Hydrometra sztolemani Jaczewski, 1928 (Colombia, Paraguay, Argentina, Minas Gerais, São Paulo, Paraná). - There are no large processes on the sixth abdominal segment of either sex. The terminal process of the female is acutely pointed (Fig. 6.311). ..................13 13. (12) The third segment of the antenna is more than 2 2/3 the length of the second. ................................................................................................................14 - The third segment of the antenna is only about twice as long as the second (Fig. 6.311). The length is at least about 9.5 mm. .............................................15 14. (13) Length: c. 8.5 mm. The first genital segment of the male is compressed laterally to form a ventral keel, which is expanded laterally at the apical end and armed with laterally-directed spines. The color of the head and thorax is mainly yellowish brown. The apterous form is lighter. Exposed tergites of the abdomen are reddish brown bordered by dark brown, while the connexivum is yellow with brownish margins. The hemielytra have yellow or brown veins. The female has not been described. ....................................................................Hydrometra thomasi Mychajliw, 1961 (Mexico, Central America, Colombia). - Length: c. 12.2 mm. The color is predominantly brown, slightly paler on the ventral surface of the abdomen. The ratio of the anteocular to the postocular head length is about 94:45. The ratios of antenna segments I through IV are 24:42:125:80, respectively. The propleurae and acetabula lack pits. The legs are long, slender, and brownish, with the hind femur of the female reaching the apex of the abdomen. The genital segment ends in a long, straight spine directed posteriad. Macropterous forms and the male have not been described. .............................................................................Hydrometra placita Drake, 1953 (Paraguay).

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Fig. 6.311 Hydrometra argentina (upper row, left to right): head of a male in dorsal view, anterior part of the thorax in lateral view, fore-wing, and (lower row, left to right): apex of the male abdomen in dorsal and ventral view, and the apex of the female abdomen in dorsal view. Based on Moreira et al. (2009). 15. (13) There is no pale mid-dorsal line in either sex. Thoracic pits are absent. There are only two spines on the seventh abdominal sternite of the male (Fig. 6.311). The dominant color is brownish with yellowish brown on the head and most of the thorax. Most of the body is covered by small black tubercles. The clypeus is conical. Length: 9.5 to 10.5 mm. ..........................................................................Hydrometra argentina Berg, 1879 (West Indies, Central America, Colombia, Venezuela, Surinam, Peru, Bolivia, Paraguay, Argentina, Uruguay, Chile, Amapá, Bahia, Pará, Amazonas, Minas Gerais, Mato Grosso, Mato Grosso do Sul, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul). Syn: Hydrometra mensor White, 1979 (nec Champion, 1898); Limnometra chilensis Reed, 1901; Hydrometra chilensis (Reed, 1901); Hydrometra kirkaldyana Torre-Bueno, 1926; Hydrometra husseyi Torre-Bueno, 1926. - Both sexes display a pale mid-dorsal line. There are four spines on the seventh abdominal sternite of the male (Fig. 6.301). Most of the body is covered by tiny brown tubercles. Length of male: 9.5 to 10.5 mm; female: 10.5 to 11.5 mm. .....................................................Hydrometra quadrispina Pérez Goodwyn, 2001 (Argentina).

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Key to the species of adult Bacillometroides in South America Information for the key was provided by Hungerford (1927a), Drake (1956), Nieser (1970b), and Polhemus and Polhemus (2010). 1. The seventh abdominal segment bears a spine on each side. The spines of the male are lateroventral and triangular in shape, while those of the female are lateral and directed caudad (Fig. 6.312). The color is yellowish brown with obvious dark brown and blackish mottling and frosted areas arranged in a longitudinal pattern. Length of micropterous male: c. 10.5 to 10.6 mm; female: c. 12.5 mm. The male has not been described. ........................................................Bacillometroides mulfordi (Hungerford, 1927) (Bolivia). Syn: Hydrometra mulfordi Hungerford, 1927; Bacillometra mulfordi (Hungerford, 1927). - The seventh abdominal segment lacks paired spine-like processes, although a median protuberance or spine may be present on the eighth abdominal segment (Fig. 6.313). There is no obvious mottling. The maximum length is less than 12 mm. .......................................................................................................................2

Fig. 6.312 Bacillometroides mulfordi female: head in dorsal view (above), apex of the abdomen in dorsal (lower left) and lateral view (lower right). Based on Hungerford and Evans (1934).

Fig. 6.313 Bacillometroides fuallagana: apical abdominal segments of a male (left) and a female (right) in lateral view. Based on Nieser (1970b).

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2. (1) The fourth segment of the antenna is much longer than the second. The abdomen of the female ends in a mid-dorsal spine, while that of the male does not (Fig. 6.313). Length: 8.0 to 10.1 mm. The color is dark to blackish brown with a median grayish-yellow longitudinal stripe, interrupted by fuscous markings. The eyes are reddish. ............................................................Bacillometroides fuallagana (Drake, 1956) (Peru). Syn: Bacillometra fuallagana Drake, 1956. - The second and fourth segments of the antenna are subequal in length, or the fourth appears somewhat shorter (Fig. 6.302). .................................................Bacillometroides woytkowskii (Hungerford, 1935) (Peru). Syn: Bacillometra woytkowskii Hungerford, 1935.

Hebridae Key to the genera in South America Information for the keys was provided by Drake and Harris (1943) and Drake and Chapman (1958a). 1. The antenna obviously consists of four robust segments; the fourth segment is usually as long as the first (Fig. 6.314). The antenna is usually shorter than the maximum width of the pronotum. ..............................................................Merragata Buchanan-White, 1877..p. 267 This genus includes species formerly classifed under the generic name Lipogomphus Berg, 1879, which is no longer considered a valid generic name. - The antenna appears to consist of five segments because the fourth has a constriction in the middle (Fig. 6.315). The antenna is longer than the maximum width of the pronotum. The segments of the antenna are narrow, and the fourth is longer than the first. .....................................................................................Hebrus Curtis, 1879..p. 269

Fig. 6.314 Merragata hebroides: habitus (left), pregenital segment (upper right), and male paramere (lower right). Based on Drake and Cobben (1960a).

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Fig. 6.315 Hebrus elimatus (left to right): habitus, pregenital segment, and a paramere of a male. Based on Drake and Cobben (1943). Key to the species of Merragata in South America Information for the keys was provided by Drake (1917, 1952c), Drake and Harris (1943), Drake and Chapman (1958a), and Estéves and Schnack (1980a). The generic name Lipogomphus Berg, 1879, is now considered a synonym of this genus. 1. The apex of the scutellum is slightly emarginated or obviously forked. ..........2 - The apex of the scutellum is bluntly rounded and not forked. The male lacks a subapical spine on the posterior face of the hind tibia (Fig. 6.314). ....................3 2. (1) The apex of the scutellum is forked. The male bears a subapical spine on the posterior face of the hind tibia (Fig. 6.316). The first segment of the antenna is the shortest and thickest, and the second, third, and fourth are subequal in length. Only the macropterus form has been found. .........................................................................Merragata lacunifera (Berg, 1897) (Bolivia, Paraguay, Argentina, Uruguay, Minas Gerais). Syn: Lipogomphus lacuniferus Berg, 1897. - The apex of the scutellum is narrow and emarginate at the apex. The second segment of the antenna is shorter than the first, the ratios of the four antenna segments being 10:9:14:16. The first two antenna segments are dark red, the third is testaceous, and the fourth is pale. The body and legs are very hairy. Length: c. 2.2 to 2.3 mm. Width across the humeri: 0.9 to 1.0 mm. The color is dark red with lighter reddish markings, a black head, and white markings on the hemielytra. ............................................................Merragata sessoris Drake and Harris, 1943 (Mato Grosso).

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Fig. 6.316 Merragata lacunifera male: hind tibia and tarsus (left) and right paramere (right). Based on Drake and Chapman (1958). 3. (1) The length of macropterous specimens is about 1.55 mm, and the width is about 0.5 mm. The length ratios of antenna segments I to IV are 3:3:3:4. Color: mainly brown but darker laterally. The legs are testaceous. The body has a sparce covering of pale hairs. An apterous form has not been found. ................................................................................Merragata quieta Drake, 1952 (Mexico, Panama, Colombia). - The body and legs are sparcely hairy. The first three segments of the antenna are subequal, while the fourth is thick and fusiform (Fig. 6.314). The body is less than three times as long as its maximum width. Length: greater than 1.75 mm and less than 2.25 mm. ..........................................................................Merragata hebroides White, 1877 (Cape Verde Islands, Canary Islands, North and Central America, West Indies, Hawaii, Colombia, Venezuela, Peru, Ecuador, Bolivia, Paraguay, Uruguay, Argentina, Amazonas). Syn: Merragata foveata Drake, 1917; Merragata slossoni Van Duzee, 1921; Merragata lindbergi Poisson, 1954. Key to the species of adult Hebrus known from South America Information for the keys was provided by Stål (1858), Champion (1897-1901), Drake and Harris (1943), Porter (1952), Drake and Chapman (1953a, 1958a, 1970), and Drake and Cobben (1960a). The male of Hebrus pilosidorsus has not been described. 1. The pronotum is covered by deep, coarse pits, scattered or in irregular lines (Fig. 6.317). ..........................................................................................................2 - The pronotum is not deeply pitted; if pits are present, they are few or not coarse. Facial tubercles are not present. The dorsal surface does not appear densely hairy. ........................................................................................................6

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Fig. 6.317 Habitus of a brachypterous female Hebrus pilosidorsus with the dense coat of hair-like setae removed. Based on Polhemus (1970).

Fig. 6.318 Hebrus plaumanni: habitus without appendages (left) and paramere of a male (right). Based on Porter (1952). 2. (1) The head of the female bears prominent facial tubercles (Fig. 6.317). The dorsal surface is clothed in dense, hair-like setae. Length of female: c. 2.0 mm. The male has not been described. ......................................................................Hebrus pilosidorsus Polhemus, 1970 (Peru). - The head lacks facial tubercles (Fig. 6.318). ....................................................3

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3. (2) The antenna is not slender. The head, pronotum, and elytra lack a short, golden pubescence. The coarse punctures on the pronotum are arranged in irregular rows around the posterior lobes and along the posterior border. The dorsal surface is not clothed in dense, hair-like setae. The apex of the scutellum is forked. There is no white marking at the base of the elytra. The male paramere is strongly recurved near the apex. The head, pronotum, and veins on the hemelytra are mainly fuscous, while the rest of the hemelytra are smoky brown and smoky gray with whitish and fulvous markings (Fig. 6.318). Length: 1.85 to 2.50 mm. ..............................................................................Hebrus plaumanni Porter, 1952 (Rio de Janeiro). - The antenna is slender (Fig. 6.31). The hemielytra are dark fumose or fuscous with other markings, including either one or three white marking at the base of the hemielytra. ......................................................................................................4 4. (3) Length: c. 2.6 mm. The length ratios of antenna segments I to V are 25:15:22:16:20, respectively. An irregular row of large pits separates the anterior and posterior lobes of the pronotum. The ratio of the length to width of the notably wide pronotum is 5:9 (Fig. 6.31). The body, legs, and antennae are finely pubescent. Color: dark fuscous brown with a pearl-white marking from the base of the clavus almost to the apex of the scutellum and four white markings on the otherwise dark fuscous membrane. The head is black with a bluish tinge beside the eyes; it lacks a median furrow. The pronotum bears a slightly flattened gold pubescense. The description is based on macropterous specimens. ...............................................................................Hebrus major Champion, 1898 (Mexico, Colombia). Syn: Hebrus acapulcana Drake and Chapman, 1954. - The length is nearly always shorter than 2.2 mm. .............................................5 5. (4) The head, pronotum, and elytra are covered with a short, golden pubescence. The coarse punctures on the pronotum are arranged in a transverse row along the posterior border of the collar and irregularly between the anterior and posterior lobes of the pronotum, on the posterior lobes, and along the posterior border. There is a pair of especially large pits on the anterior lobe, one on each side of the mid-line. The apex of the scutellum is not forked. There are pearly-white markings at the bases of the hemielytra. The head, pronotum, and veins on the hemelytra are mainly fuscous, while the rest of the hemelytral surface is mainly fulvous. The legs are light yellow. Length: 1.9 to 2.2 mm. ...........................................................Hebrus camposi Drake and Chapman, 1954 (Ecuador). - The pronotum has a sparce covering of pale pubescent setae. The coarse punctures are especially dense between the anterior and posterior lobe of the pronotum, which is blackish with reddish brown on the disc and sometimes on part of the posterior lobe. The scutellum is blackish and has punctures and a rounded and slightly bifid apex (Fig. 6.315). The hemielytra are dark fumose with prominent blackish veins, pale pubescence, and three white spots at the base, one in the closed cell and a pair on each side of the scutellum. There are

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also three whitish spots on the apical half of the membrane. The legs are stramineous with dusky apices of the femora, tibiae, and tarsi. Length of male: 1.6 to 1.8 mm; female: 1.8 to 2.1 mm. ..............................................................Hebrus elimatus Drake and Cobben, 1943 (Netherlands Antilles, assumed to be native to northern South America). 6. (1) The head, pronotum, and scutellum are mainly blackish or dark with a metallic bluish green sheen. The hemielytra are dark with a white, wedgeshaped marking on the clavus and other white markings on the hemielytra. The maximum length barely exceeds 2 mm. ...............................................................7 - The color is not blackish, or the membrane of each hemieltyron displays one pale spot so that two are visible when the insect is at rest, or there are no distinct pale spots at all on the membrane. ........................................................................8

Fig. 6.319 Hebrus spiculus male: genital capsule with short setae removed (left) and two views of the right paramere (right). Based on Polhemus and McKinnon (1983).

Fig. 6.320 Male paramere of Hebrus concinnus. Based on Drake and Cobben (1960a). 7. (6) The ground color is blackish. The hemielytra are blackish with a white wedge-shaped marking on the clavus, a narrow and indistinct white stripe on the corium, and two white spots on each hemielytron, so that three are visible when the insect is at rest. The shape is oblong and rather narrow, with a length of about 2 mm and a width of c. 0.75 mm. The legs are pale. .....................................................................................Hebrus parvulus Stål, 1858 (Rio de Janeiro). Descriptions of this species are sketchy, and it cannot be clearly distinguished from Hebrus consolidus Uhler, 1894, also known by its synonym, Naeogeus consolidus Blatchley, 1926, which has not yet been

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reported from South America, although it is known to range from North America as far south as Panama and Trinidad. Champion (1898) reported that H. parvulus can be distinguished by its longer first antenna segment, the more distinct separation of the two lobes on its pronotum, white streaks between the nerves on its corium, and less conspicuous white spots on the membrane of its hemielytra. Because all of these are rather subjective distinctions, these two nominal species can be considered indistinguishable until better descriptions are available. - The ground color of the head, pronotum, and scutellum is fuscous with a metallic bluish green sheen. The corium and posterior part of the clavus are piceous, and an anterior, wedge-shaped mark on the corium is pearly white. The membrane is brown with one oval, white spot next to the corium. There is a longitudinal midline sulcus on the pronotum. The male paramere has a spur on the inner surface (Fig. 6.319). Length: c. 1.6 to 1.8 mm. Width across humeri: 0.6 to 0.8 mm. ...................................................Hebrus spiculus Polhemus and McKinnon, 1983 (Costa Rica, Panama, Colombia). 8. (6) Length: 1.75 to 2 mm. The third and fifth antenna segments are equal and longer than the others. The scutellum is notched at the apex. Coloration: predominantly coppery, chestnut brown, and reddish. .........................................9 - Length: greater than 2 mm. .............................................................................10 9. (8) The color is generally reddish with the head, pronotum, and scutellum darker than the rest of the dorsum. Length: 1.75 to 2 mm. ..................................................................Hebrus priscus Drake and Harris, 1943 (Panama, Peru, Paraguay, Mato Grosso). - The head, pronotum, and scutellum are not notably darker than the rest of the dorsum. The color is generally dark reddish with patterns on the pronotum and hemielytra. Length: 1.75. ...................................................................Hebrus paulus Drake and Harris, 1943 (Mato Grosso). From the rather poor descriptions of these two species, it is very difficult to find any definitive differences between them. Drake and Harris (1943) failed to give a reason for believing that their specimens from Mato Grosso belong to two rather than a single species. 10. (8) The body and appendages are moderately covered with long hairs. The third segment of the antenna is the longest and 1/3 longer than the first. The apex of the scutellum is deeply notched. Length: 2.4 mm. ...............................................................Hebrus gloriosus Drake and Harris, 1943 (Mato Grosso). - The body and appendages are, at most, sparcely hairy. ..................................11 11. (10) Length: 3.0 mm. The scutellum is deeply impressed and has a prominent median carina and carinae along its lateral and hind margins; its base is wider than its length in the middle. The scutellum converges sharply from its base for a distance of about 2/5 of its total length; for its posterior 3/5, its sides are gradually rounded, and its apex is rounded evenly without a notch. The anterior lobe of the pronotum is strongly and concavely constricted laterally and

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much wider across its humeral angles than at the widest point across the anterior lobe. ...........................................................Hebrus engaeus Drake and Chapman, 1958 (Argentina). - The length is not more than 2.6 mm. ..............................................................12 12. (11) The fourth and fifth antenna segments are subequal, and each is shorter than the third. The color is mainly fuscous or reddish brown dorsally and polished fuscous black with a thin black median stripe ventrally. The head is dark brown with a fuscous apical margin. The pronotum has a depressed rufous margin, and the hemielytra each have a white stripe beginning at the base of the clavus, a longer streak on the subcostal areole, pale margins, and a pale marking near the apex. The hemielytra of some specimens show a trace of another pale marking. There is a silky pubescence on the underside and hemielytra. The male paramere curves about 145º at the apex (Fig. 6.320). Length: 2.25 to 2.50 mm. Width of pronotum: c. 1 mm. ................................................................................Hebrus concinnus Uhler, 1894 (North and Central America, West Indies, Peru). Syn: Naeogeus concinnus Blatchley, 1926. - The fourth antenna segment is shorter than the fifth, or there is only one small white spot on each side near the scutellum. .......................................................13 13. (12) Length: c. 2.25 mm. There is a small white spot at the base of the hemielytra on each side of the scutellum. The right male paramere folds against itself near the apex and bears two setae more than half as long as itself near the folding point (Fig. 6.321). ..........................................................Hebrus limnaeus Drake and Chapman, 1958 (Santa Catarina). - The length is at least 2.4 mm. .........................................................................14

Fig. 6.321 The right male paramere of Hebrus limnaeus. Based on Drake and Chapman (1958a). 14. (13) The fourth antenna segment is only half as long as the fifth; the first is slightly longer than the third and fifth. Length: 2.45 mm. ...........................................................Hebrus hungerfordi Drake and Harris, 1943 (Ecuador).

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- The fourth antenna segment is more than 4/5 as long as the fifth; the first and third are nearly equal. Length: 2.56 mm. ..............................................................Hebrus ecuadoris Drake and Harris, 1943 (Ecuador).

Mesoveliidae Key to the genera in South America Information for the keys was provided by Andersen and Polhemus (1980) and Froeschner (1981) 1. Macropterous specimens (Fig. 6.322). .............................................................2 - Apterous, i.e. wingless specimens (Fig. 6.323). ...............................................3 2. (1) Two dark ocelli are present. The pronotal lobe is not trapezoidal but rather constricted anteriorly. The mesoscutellum is trapezoidal with somewhat rounded sides and separated along its posterior margin from a separate metanotal lobe. The tarsal claws originate at the apices of the tarsi. The forewing has two or three basal cells but lacks an apical cell (Fig. 6.18, 6.322). Length: 2.0 to 4.5 mm. ..............................................................Mesovelia Mulsant and Rey, 1852..p. 278 - Ocelli are absent. The pronotal lobe is trapezoidal. The mesoscutellum is triangular. The origin of the tarsal claws is subapical. The fore-wing has two basal cells and one apical cell. The fourth segment of the antenna is not subdivided. The insertion of the antenna is separated from the eye by a distance not greater than the diameter of the first antenna segment. The femora each have two dark, spinose, subapical setae (Fig. 6.324). Length: 3.5 to 4.2 mm. .................................................................Mesoveloidea Hungerford, 1929..p. 280

Fig. 6.322 Habitus of Mesovelia amoena. Based on Cobben (1960a).

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Fig. 6.323 Cryptovelia terrestris: habitus of an apterous female in dorsal view (upper left) and in lateral view without appendages (lower left), apterous male without appendages in dorsal view (upper right), middle leg of a male (upper middle right) and enlarged middle tarsus with surface setae removed (lower right center), metathoracic scent gland visible in a macerated specimen (lower center), first (lower middle right) and second (lower right) gonocoxa and gonapophysis of the female genitalia and ovipositor in lateral view, and a ripe ovarian egg (upper center). Based on Andersen and Polhemus (1980).

Fig. 6.324 Mesoveloidea williamsi habitus in dorsal view (left), head and thorax in lateral view (upper middle right), and a hemelytron (lower middle right). Based on Alvarez and Roldán (1983).

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Fig. 6.325 Darwinivelia fosteri apterous female: habitus in dorsal view (upper left) and in lateral view without appendages (upper right), enlarged head and thorax in dorsal view (right center) and the head without the rostrum in ventral view showing the longitudinal carinae (right of head and thorax), hind leg (lower left), segments of middle tarsus with surface setae removed (lower center), first gonocoxa and gonopophysis of the female genitalia (lower right), second gonocoxa and gonopophysis (lower right center), and two views of a ripe ovarian egg (upper and middle right). Based on Andersen and Polhemus (1980) and Froeschner (1985). 3. (1) The eyes are vestigial with few ommatidia remaining from the compound eyes. The head is shorter than the thorax and has a transverse carina at its base (Fig. 6.323). Length: 1.2 to 1.3 mm. .............................................................Cryptovelia Andersen and Polhemus, 1980 The only known species in this genus is Cryptovelia terrestris Andersen and Polhemus, 1980, which was found in moist soil litter of secondary forests on the terra firma in Pará. - The compound eyes are distinct and globular (Fig. 6.325). The length is at least about 1.8 mm. ...............................................................................................4

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4. (3) The mesonotum is shorter than the pronotum, transverse, and slightly shorter along the midline than toward the lateral margins. The insertion of the antenna is separated from the eye by about half the diameter of the eye and a distance greater than the diameter of the first antenna segment. On the ventral side of the head, there is a pair of prominent longitudinal carinae posterior to the base of the rostrum (Fig. 6.325). Total length: c. 1.8 mm. The species are marine and found in mangrove swamps. ...............................................Darwinivelia Andersen and Polhemus, 1980..p, 281 - The mesonotum is elongated along the midline, and it is clearly longer than the pronotum. The insertion of the antenna is separated from the eye by about the diameter of the eye or by a distance not greater than the diameter of the first antenna segment (Fig. 6.19, 6.322), best seen in lateral view. The known species vary in length from 2.0 to 4.5 mm. Freshwater species. ..............................................................Mesovelia Mulsant and Rey, 1852..p. 278 Key to the species of adult Mesovelia in South America Information for the key was provided by Drake and Harris (1946), Drake (1949a), Spangler (1990), Nieser and Alkins-Koo (1991), and Moreira et al. (2008). 1. There is a prominent row of black spines on the ventral edge of the middle femur (Figs. 6.18, 6.19, 6.326). ............................................................................2 - The middle femur lacks a prominent row of black spines (Fig. 6.322). ...........4 2. (1) The first genital segment of the male, which is the eighth abdominal segment, lacks a medial spine and a cluster of black spinules. In lateral view, the male paramere resembles a hook with an acute apex directed anteriad (Fig. 6.327). Length: greater than 3.5 mm. .......................................................Mesovelia hambletoni Drake and Harris, 1946 (Ecuador). - The first genital segment of the male bears two tightly-packed clusters of black spinules (Figs. 6.18, 6.19, 6.326). ...............................................................3

Fig. 6.326 Mesovelia halirrhyta male: middle femur (left) and genital segments in ventral view (right). Based on Polhemus (1975).

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3. (2) There are about 25 spines in the ventral row on the middle femur. On the ventral side of the first male genital segment, which is the eighth abdominal segment, there are two well-separated rows of three to five thin spinules (Fig. 6.326). Length: 2.5 to 2.8 mm. .....................................................................Mesovelia halirrhyta Polhemus, 1975 (Central America, Colombia). - There are 8 to 15 spines in the ventral row on the middle femur. On the ventral side of the first male genital segment, there are two dense clusters of thick, black spinules. Length: 3.0 to 4.0 mm. The head is yellowish without brown longitudinal stripes near the midline. The fore and middle femora of the male each bear a ventral row of spines. (Figs. 6.18, 6.19). .............................................................................Mesovelia mulsanti White, 1879 (North and Central America, West Indies, Colombia, Venezuela, Guyana, Peru, Bolivia, Paraguay, Argentina, Amapá, Bahia, Pará, Pernambuco, Amazonas, Rondônia, Mato Grosso, Minas Gerais, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul). Syn: Mesovelia bisignata Uhler, 1884; Mesovelia mulsanti bisignata Uhler, 1884; Mesovelia mulsanti meridionalis Jaczewski, 1930; Mesovelia mulsanti caraiba Jaczewski, 1930.

Fig. 6.327 The male paramere of Mesovelia hambletoni. Based on Spangler (1990). 4. (1) The sixth abdominal segment of the male lacks a posteromedial fringe of black spinules on the ventral side. The color is mainly brown. There is an obvious depression on each side of the dorsal midline on the anterior lobe of the pronotum of winged males; the pronotum has a sinuous posterior margin (Fig. 6.322). Length: 1.8 to 2.5 mm. Males are seldom found. These insects seem to inhabit damp moss rather than open water. ...............................................................................Mesovelia amoena Uhler, 1894 (North and Central America, West Indies, Colombia, Ecuador with Galapagos Islands, Pará, Amazonas, Rondônia, Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo). Syn: Mesovelia douglasensis Hungerford (1924). - The sixth abdominal segment of the male has a posteromedial fringe of black spinules on the ventral side (Fig. 6.328). .............................................................5

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Fig. 6.328 Mesovelia bila (left to right): head and thorax of a female in dorsal view, middle femur of a female, male gonapophysis, and the apex of the male abdomen in ventral view. Based on Jaczewski (1928b). 5. (4) There are no black spinules on the ventral side of the seventh abdominal segment. The color is mainly yellowish. There are no depressions on the anterior lobe of the pronotum of winged males; the pronotum has a straight posterior margin. The sixth abdominal segment has a ventral row of about 20 small black spines near the posterior margin (Fig. 6.33). Length of male: 1.70 to 2.00 mm; female: 2.1 to 2.3 mm. Males are abundant. ................................................................Mesovelia zeteki Harris and Drake, 1941 (Panama, Colombia, Pará, Amazonas). - There are two clusters of black spinules on the ventral side of the seventh abdominal segment (Fig. 6.328). ..............................................................................Mesovelia bila Jaczewski, 1928 (Panama, Argentina, Paraná). Key to the species of adult Mesoveloidea in South America Information for the key was provided by Hungerford (1929e) and Drake (1949a). 1. The ratios of antenna segments I to IV are 26:28:25:33, respectively. Length of male: 2.9 mm. Color: The head and anterior portion of the pronotum are yellow; the rest of the pronotum and scutellum are somewhat shiny, fuscous black with a slight bluish sheen. .....................................................................Mesoveloidea peruviana Drake, 1949 (Peru). - The lengths of the second and third antenna segments are shorter relative to the first and fourth segments (Fig. 6.324). Length: 2.9 mm. Color: The head and anterior portion of the pronotum are yellow; the rest of the pronotum and scutellum are metallic bluish black with a coppery sheen. The hemielytra are smoky with contrasting brown veins. The ventral surface is purplish brown.

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The color of the beak, antennae, and legs is yellow, and the acetabula are paler yellow. ..............................................................Mesoveloidea williamsi Hungerford, 1929 (North and Central America, West Indies, Colombia, Venezuela, Surinam, Ecuador, Peru, Minas Gerais, Rio de Janeiro). Key to the species of adult Darwinivelia in South America Information for the key was provided by Carvalho (1984). 1. The compound eye consists of 10 to 12 ommatidia. The posterior margins of the pronotum and mesonotum are sinuate (Fig. 6.325). The male has not been described. ...............................................Darwinivelia fosteri Andersen and Polhemus, 1980 (Galapagos Islands). - The compound eye consists of 28 to 30 ommatidia. The posterior margins of the pronotum and mesonotum are evenly curved (Fig. 6.329). Length: 1.5 to 1.9 mm. Color: mainly brown. ....................................................................Darwinivelia polhemi Carvalho, 1984 (Pará).

Fig. 6.329 Darwinivelia polhemi: habitus of an apterous female in dorsal view (upper left), head with beak in oblique lateral view (upper center), hind claw (upper right center), hind tarsus (upper right), ventral view of the head and anterior part of the thorax (lower left), pygophore in ventral (lower right center) and lateral view (lower left center), and two views of the male paramere (center and lower right). Based on Carvalho (1984).

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6.7.2 Infraorder Nepomorpha Key to families in South America Information for the key was provided by Brues et al. (1954), Schuh and Slater (1995), and Pereira et al. (2007). Adults and most late instar larvae can be distinguished. 1. The rostrum appears to be on the ventral side of the head; it is very short and broad with no distinct segmentation. The base of the head overlaps the anterior margin of the pronotum. The fore-tarsus consists of a single segment and is modified to form a pala, a broad plate or robust hook without a claw at the end but often with combs of bristles. Nymphs have three scent gland openings on the dorsal surface of the abdomen (Fig. 6.330). .....................................................................................................Corixidae..p. 292 - The rostrum extends from the apex of the head and is distinctly conical or cylindrical with 3 or 4 segments. There are no palae on the fore-legs. The base of the head is inserted in the pronotum. If scent gland openings are present in the abdomen of larvae, they are present on only one segment (Fig. 6.331). ........2

Fig. 6.330 Tenagobia socialis: habitus in dorsal view (middle left), fore-leg of a male (upper left) and a female (lower left), abdomen of a male in dorsal view (lower center), receptaculum seminis (upper center), right parameres (upper right) and left parameres of two males (lower right). Based on Nieser (1977a) and Alvarez and Roldán (1983). 2. (1) A respiratory siphon is present, although, if it is retractile, it may appear very short. The body is either flat and ovoid or cylindrical. Veins are evident on the membranes of the hemielytra. The size is from medium to very large.

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The fore-legs of all known South American species are raptorial, but the characteristic of one African genus is the lack of such fore-legs (Fig. 6.331). ....3 - There is no respiratory siphon. The body is never cylindrical and is small to medium in size (Fig. 6.332). The hind coxae are joined broadly to the pleura, limiting their rotation. ...........................................................................................4

Fig. 6.331 Curicta borellii: habitus in dorsal view (upper left), head and anterior part of the thorax in lateral view (just above center), male operculum (upper right center), apical abdominal segments of three specimens in lateral view (right center to upper middle right), mesosternum and metasternum in ventral view (upper right), fore-femur in anterior view (upper middle left), male genital capsule in dorsal, ventral, and lateral view (lower middle row, left to right), parameres of three different male specimens (lower left), female genitalia in lateral view (lower center), and first female gonapophysis in medial view (lower right). Based on Keffer (1997). 3. (2) The respiratory siphon is not retractile and either consists of two long, slender posterior appendages or is short and strap-like. The hind coxae are short and free to rotate. The hind tibiae are not markedly flattened or modified with swimming hairs, and all tarsi consist of only one segment (Fig. 6.331). All larval instars have relatively short, thick respiratory siphons and fore-legs resembling those of adults (Fig. 6.333). ........................................................................................................Nepidae..p. 356 This family is sometimes split into Nepidae and Ranatridae, but it is maintained here sensu lato.

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- The respiratory siphon is short and retractile, and when retracted, only the apices are evident. The hind coxae are conical and fused with the metapleuron. The middle and hind tarsi consist of two or three segments, and, except in a few cases, the fore-tarsus is, as well. The veins in the membranes of the hemielytra are reticulate (Fig. 6.334). The nymphs lack dorsal abdominal scent gland openings (Fig. 6.335). ............................................................................................Belostomatidae..p. 393

Fig. 6.332 Gelastocoris vandamepompanoni: habitus in dorsal view with the outline of the right side of the head and pronotum above it (left), hemielytron showing locations of the large, pearl-like granules (upper right center), penis (right center), male paramere (lower center), posterior margin of the seventh abdominal segment (lower right center), and apex of the female abdomen in dorsal (upper right) and ventral view (lower right). Based on Boulard and Jauffret (1984).

Fig. 6.333 Curicta bonaerensis: habitus of each larval instar in dorsal view without appendages and its fore-leg above it: instar I (upper left), II (upper center), III (middle left), IV (middle right), and V (lower center). Based on López Ruf et al. (2001).

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Fig.6.334 Belostoma micantulum: habitus in dorsal view (upper left), right hemielytron in dorsal view (upper right), head and part of the pronotum in lateral view (lower left), fore-leg in ventral view (lower right center), and antenna in ventral view (lower right). Based on Alvarez and Roldán (1983).

Fig. 6.335 Lethocerus bruchi: the five larval instars from the first to the fifth (above, left to right and left and right below, respectively). Surrounding the dorsal habitus shown without appendages are a lateral view of the head (below), an antenna (above left), a fore-tarsus (above right) and a hind tarsus (above the antenna and fore-tarsus). Based on Estévez and Pérez Goodwyn (1999), who referred to the species by its synonym: Lethocerus truncatus.

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4. (2) Ocelli are present, or, if they are vestigial or absent, the compound eyes are pedunculate or nearly so, and the head is transverse. The legs are cursorial (Fig. 6.332). ..........................................................................................................5 - Ocelli are lacking, and the compound eyes are sessile. There are fringes of long swimming hairs on the middle or hind legs, except on a few minute species of Pleidae, and one or both of these legs are often flattened (Fig. 6.336). ..........6

Fig. 6.336 Martarega membranacea male: habitus of a brachypterous specimen in dorsal and ventral view (above, left and right), pattern on a hemielytron (left center), penis (lower left), and the male genital capsule in left (lower center) and right lateral view (lower right). Based on Truxal (1949) and Angrisano (1982). 5. (4) The fore-legs are raptorial; the femora are very broad and grooved along their inner edges near the curved tibiae. The antennae are relatively short, incrassate, and concealed in grooves between the eyes and prothorax. The head is markedly transverse, and the eyes are pedunculate or nearly so. The anterior surface of the face is sulcate or carinate. The scutellum is tumid, and the forefemur is incrassate (Fig. 6.332). The larvae closely resemble the adults in body shape and non-genital structures (Fig. 6.337). .............................................................................................Gelastocoridae..p. 435 - The fore-legs are similar to the middle legs and suitable for running. The antennae are long and not completely concealed in dorsal view. The head may be moderately transverse, and the eyes are sessile. The scutellum is flat (Fig. 6.338). ....................................................................................................Ochteridae..p. 457

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Fig. 6.337 Nerthra ranina: habitus without appendages in dorsal view with the antenna at the upper left side and the rostrum in lateral view shown from the left to the lower left of the habitus of Stages I (upper left), II (upper center), III (upper right), IV (lower left), and V (lower right). The claws on the fore-leg of instar V (lower right center) and an adult (lower right) are shown below the Stage V instar. Based on Estéves and Schnack (1978).

Fig. 6.338 Ochterus barberi: habitus in dorsal view (left) and the apical part of the genital capsule (right). Based on Schuh and Slater (1995). 6. (4) In cross section, the dorsum is strongly convex, often tectiform, or, if it is flattened, then the head and pronotum are fused, and the cephalonotal sulcus is not complete. The fore-legs are not raptorial, but those of males are sometimes modified in other ways (Fig. 6.336). ....................................................................7

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- In cross section, the dorsum is flat or only moderately convex, and the head and thorax are never fused. The antenna is fairly long and protrudes beyond the margin of the head, or the fore-legs are obviously raptorial (Fig. 6.339). ...........9

Fig. 6.339 Limnocoris insignis: habitus of a brachypterous female (left), apex of the male abdomen in dorsal view (upper center), apex of the female abdomen in ventral view (upper right), and the right (lower center) and left pseudoparamere (lower right). Based on Nieser and Lopez Ruf (2001).

Fig. 6.340 Neotrephes lanemeloi: habitus of a brachypterous female (left), male left (upper center) and right paramere (lower center), penis in ventral view (upper right), and the seventh abdominal segment of a female in ventral view (lower right). Based on Nieser and Chen (2002). 7. (6) The form is elongate; usually somewhat wedge-shaped. The compound eyes are large. The length is greater then 4 mm. The hind legs are long and oarlike and lack distinct claws dorsally (Fig. 6.336). The head is not fused with the thorax. There is no ventral midline carina on the posterior part of the thorax, but there is a ventral keel on the abdomen. ................................................................................................Notonectidae..p. 464

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- The shape is oval, and the body appears robust. The adults are less than 3 mm long, and the hind legs are not long or oar-like and usually bear two distinct claws dorsally. The head and thorax do not articulate with each other (Fig. 6.340). The anterior abdominal segments have an irregular carina. ...................8

Fig. 6.341 Paraplea puella (above, left to right): habitus in dorsal and lateral view, the ovipositor of a female specimen, and (below, left and right): segments of the fore-tarsus and operculum of the female. Based on Nieser (1975).

Fig. 6.342 Larval instars of Neoplea striola (Fieber, 1844), described as Plea striola Fieber, 1844 (upper row, left to right): dorsal view of larval instar I, II, III, and IV without appendages, and (middle row, left to right), ventral view of larval instar I, II, III, and IV, and (lower row, left to right), larval instar V in ventral, dorsal, and lateral view, the last two without appendages. The presence of this North and Central American species in South America has not been confirmed. Based on Gittelman (1974a).

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8. (7) The head and pronotum are fused dorsally and separated only by a shallow suture; the cephalonotal suture, i.e. the suture between the head and the pronotum, is not straight (Fig. 6.340). In dorsal view, the antenna appears to consist of one or two segments. ..............................................................................................Helotrephidae..p. 504 - The head and pronotum are not fused; the cephalonotal suture is straight. In dorsal view, the antenna appears to consist of three segments (Fig. 6.341). In cross section, the dorsum of adults is always strongly convex, although the larvae tend to be less so (Fig. 6.342). .........................................................................................................Pleidae..p. 509 9. (6) The antennae are short, do not extend beyond the margins of the head, and are seldom visible in dorsal view. The short, thick labium does not extend farther posteriad than the posterior margin of the prosternum. The head is usually transverse with the anterior margin extending only slightly anterior to the compound eyes. The immobile fore-tarsus consists of only one or two segments, bearing one or two tiny claws or lacking claws. The membranes of the hemielytra lack veins (Fig. 6.339). Nymphs have a pair of widely separated dorsal scent gland openings between the second and third abdominal tergites; the general habitus, fore-legs, and antennae of the five larval stages resemble those of the adults (Fig. 6.343). ...................................................................................................Naucoridae..p. 514 - The antennae are long and extend beyond the lateral margins of the head. The slender labium sometimes reaches posteriad to the mesosternum. The head is narrow and elongate with its anterior margin produced well anterior to the eyes. The fore-tarsus is mobile, consists of one to three segments, and bears two robust claws (Fig. 6.344). ...................................................................................10

Fig. 6.343 Pelocoris bipunctulus larval instars: habitus without appendages in dorsal view with the antennae above left and fore-leg above right of instars I to IV (above, left to right) and instar V (below left) with the fore-leg to the right. Based on Lopez-Ruf (1992), who referred to the species as Pelocoris lautus.

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10. (9) The fore-tarsus consists of one segment, while the tarsi on the middle and hind legs consist of two. Only species with macropterous or coleopteroid hemielytra are known. In macropterous forms, the margin separating the corium and membrane is sinuous. There are erect setae on the lateral margins of the head, pronotum, and corium (Fig. 6.344). The labium does not reach posteriad beyond the prosternum. .............................................................................................Potamocoridae..p. 574 - All tarsi consist of three segments, but in some small species, they are fused. The wings range from macropterous to micropterous. In macropterous forms, the margin separating the corium and membrane is straight. There are no erect setae on the lateral margins of the head, pronotum, or corium. The labium reaches at least to the mid-length of the mesosternum. .......................................................................................................Aphelocheiridae This family is known only from the Eastern Hemisphere and is not further considered here.

Fig. 6.344 Potamocoris nieseri female (above, left to right): habitus without appendages in dorsal view, fore and middle leg, and (middle row, left to right): habitus in ventral view with the left legs removed, microscopic sculpture on the coriaceous area of the fifth abdominal sternite, apex of the abdomen in ventral view, and antenna, and (below), the hind leg. Based on van Doesburg (1984).

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Corixidae The key to the subfamilies of adults in South America Information for the key was provided mainly by Hungerford (1927b, 1948), Jaczewski (1927a, b; 1928c), Bachmann (1961a, b; 1962c, f, g), and Froeschner (1981). Only Micronectinae, Heterocorixinae, and Corixinae are known to be represented in South America. 1. The pronotum covers only the anterior part of the scutellum. Ocelli are absent. The antenna consists of three segments. The tarsal claw of the female resembles a spike; that of the male can be folded back into an excavation on the tarsus (Figs. 6.330, 6.345). The posterior margin of the pronotum is broadly but shallowly concave. The subfamily Diaprepocorinae Lundblad, 1928, which is known to occur in Asia, Australia, and islands in between, has never been reported from South America. The presence of two ocelli permits it to be distinguished immediately from Micronectinae. ....................................................................Micronectinae Jaczewski, 1924.p. 294 - The posterior margin of the pronotum is convex or roundly angulate so that the scutellum is completely covered by the pronotum, or rarely, only the posterior tip may be visible (Fig. 6.346). The rostrum has transverse sulcations. The nodal furrow is complete. Vein M on the hemielytra usually curves to meet Cu at or proximal to the nodal furrow. .................................................................2

Fig. 6.345 Tenagobia melini male: antenna (upper left), prothorax in anterior view (upper left center), scutellum (right of prothorax), hemielytra (lower middle left), hind wing (upper center), fore-leg (lower right), apex of the hind leg with claw and terminal spines (lower left), fifth through seventh abdominal segments (center), right (upper right) and left genital lobe (right center), left (above and left of fore-leg) and right lobe of the eighth abdominal segment (lower center). Based on Lundblad (1928a).

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Fig. 6.346 Heterocorixa westermanni male: head in anterior view (upper left), pronotum in dorsal view (upper left center), hemielytra (upper center), pala (upper right center), inner surface of the fore-leg (upper right), metaxyphus (upper middle left), lateral lobe of the pronotum (right of metaxyphus), strigil (lower middle right) and prestrigil (lower right), fourth through seventh abdominal segments in ventral view (lower left to center), right and left paramere in lateral view and left paramere in dorsal view (lower right center, above to below). Based on Lundblad (1928b). 2. (1) The infraocular portion of the genae is very broad, so that the ventral angle of the eye appears much closer to the anterior than the posterior margin of the head in lateral view. The lower margin of the eye is concave. The hypoocular suture arises near a subacute production of the inferior angle of the eye. Vein M is indistinct and runs parallel and very close to Cu (Fig. 6.346). ..............................................................Heterocorixinae Hungerford, 1948..p. 310 - The infraocular portion of the genae is usually not very broad, but if it is, the hypo-ocular suture, if present, arises midway along the ventral margin of the eye. Generally, the ventral angle of the eye appears to be located midway between the anterior and posterior margins of the head in lateral view. Vein M is distinct throughout most of its length, and it usually runs midway between Cu and the costal margin until it fuses with Cu (Fig. 6.347). ...........................................................................Corixinae Enderlein, 1912..p. 320

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Fig. 6.347 Ectemnostegella stridulata: habitus is dorsal view (upper left), foreleg of a male (upper right), head in dorsal view (lower left), apical segments of the male abdomen in dorsal view (lower left center), male genital capsule (lower right center), and apex of a female abdomen in ventral view (lower right). Based on Hungerford (1948). Subfamily Micronectinae Only one micronectine genus is known in the Americas: Tenagobia Bergroth, 1899. Key to the South American species of Tenagobia Information for the key was provided by Hungerford (1948), Nieser (1975, 1977a), and Polhemus and Nieser (1997). The subgenera inserted in parentheses are those of Nieser (1977a); they are usually omitted from more recent publications. 1. The pronotum is truncate laterally and not narrowed at its lateral ends. The posterior margins of the eyes are sinuate. The postocular space at the inner margin of the compound eye is 4 or 5 times as long as an eye facet (Fig. 6.348). The female has a simple receptaculum seminis and usually knife-like bristles on the proepimera. .....................................................................................................2 - The posterior margin of the eye is not sinuate. The pronotum is narrowed at its lateral ends and not truncate, or, if it is truncate, the postocular space at the inner margin of the compound eye is about 2½ times as long as an eye facet (Fig. 6.349). ..........................................................................................................3

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Fig. 6.348 Tenagobia balia female: head and pronotum in dorsal view (upper left), fore-tibia and pala (upper center), prothorax in anterior view (right), and receptaculum seminis (lower left to center). Based on Nieser (1977a).

Fig. 6.349 Tenagobia pulchra male (above, left to right): abdomen in dorsal view with enlarged setae along the margin where they occur and the right (above) and left paramere (below), and female (below, left to right): apical segments of the abdomen in ventral view, seventh and eighth urosternites with enlarged setae from along the margin, gonopodia, and an egg. Based on Bachmann (1983). 2. (1) The postocular space at the inner margin of the compound eye is about 5 times as long as an eye facet (Fig. 6.348). Length of female: 3.0 to 4.5 mm. The proepimera of the female bear knife-like bristles. The color is lighter than chestnut brown. The male has not been described. ............................................................Tenagobia (Baliagobia) balia Nieser, 1977 (Amazonas).

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- The postocular space at the inner margin of the compound eye is about 4 times as long as an eye facet. Length of male: 2.4 to 2.6 mm; female: 2.3 to 2.7 mm. (Fig. 6.350). The color is predominantly chestnut brown with yellow on the head between the eyes, dark costal margins on the hemielytra, and a hyaline stripe on the membrane. The legs are yellowish, but their joints and claws are dark. ..........................…Tenagobia (Baliagobia) castanea Polhemus and Nieser, 1997 (Venezuela).

Fig. 6.350 Tenagobia castanea (left to right): apex of the male abdomen in dorsal view, male left (above) and right (below) parameres, and the female receptaculum seminis. Based on Polhemus and Nieser (1997). 3. (1) At the base of the elytra, the pronotum is truncate. The inner postocular space does not exceed 2½ times the length of an eye facet. The sutures on the vertex converge posterior to the eyes (Fig. 6.349). ..............................................4 - At the base of the elytra, the pronotum is not truncate. The inner postocular space exceeds 2½ times the length of an eye facet, or, if not, then the sutures on the vertex tend to diverge posterior to the eyes (Fig. 6.351). .............................10

Fig. 6.351 Tenagobia peruana (above, left to right): left side of the eighth tergite of a male and two views of the right parameres of two specimens, and (below, left and right, respectively) the right paramere of the male and the receptaculum seminis of a female. Based on Nieser (1977a).

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4. (3) The length is 2.8 to 3.3 mm. A suture is evident on the left membrane. The seventh abdominal segment bears two lateral spines and two bristles. The lower row on the pala bears about 18 to more than 30 bristles, and the inner row bears more than 20 setae (Fig. 6.352). .................................................................5 - The length is 1.8 to 2.3 mm. There is no left membranal suture. The seventh abdominal segment bears three or four short lateral spines and two or three bristles. The lower row on the pala bears 14 to 17 bristles, and the inner row bears 11 to 17 setae (Fig. 6.349). .........................................................................6

Fig. 6.352 The fore-leg of a male Tenagobia fasciata. Based on Nieser (1977a).

Fig. 6.353 Tenagobia schreiberi: head and pronotum of a male in dorsal view (upper left), antenna (middle left), right and left hemielytra of a male (upper center, above and below, respectively), male fore-leg (lower left), apex of the middle tarsus (upper right), apical segments of the male (middle right) and female abdomen (lower right), male genital capsule (center), and right paramere (lower center). Based on Espinola (1975).

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5. (4) There are about 30 bristles in the lower row on the pala. There are two spines forming a basal row on the fore-femur (Fig. 6.352). Length: 2.8 to 3.3 mm. .....................................................Tenagobia (Fasciagobia) fasciata Nieser, 1977 (São Paulo, Rio de Janeiro, Santa Catarina). - There are about 18 bristles in the lower row on the pala. There are no spines forming a basal row on the fore-femur (Fig. 6.353). Length: 3.0 to 3.1 mm. ..............................................Tenagobia (Fasciagobia) schreiberi Espinola, 1975 (Minas Gerais). 6. (4) The seventh abdominal tergite of the male does not bear a patch of small setae in the right anterior corner. The ocular index of the male is not greater then 1.4. See Fig. 6.16, p. 28, to calculate. The sixth segment of the male abdomen bears three spine-like setae on each side. The fore-tarsus of the male bears 5 long, hair-like setae in the upper row, 17 bristle-like setae in the lower row, and 14 in the inner row (Fig. 6.349). Length of male: 2.1 to 2.2 mm; female: 2.17 to 2.25 mm. The female cannot yet be identified with certainty. ................................................Tenagobia (Tenagobia) pulchra Hungerford, 1927 (Bolivia, Argentina). - The seventh abdominal tergite of the male bears a distinct patch of small setae in the right anterior corner. The ocular index of the male is usually about 1.5 or greater (Fig. 6.354). ..............................................................................................7 7. (6) There are more than 20 setae on the left free lobe of the eighth abdominal tergite of the male. The sixth segment of the male abdomen bears four spinelike setae on each side. The pala of the male bears 4 long, hair-like setae in the upper row, 14 to 18 bristle-like setae in the lower row, and 14 or 15 in the inner row (Fig. 6.354). Length of male: 2.1 to 2.2 mm; female: 2.3 mm. The female cannot yet be identified with certainty. .........................................................Tenagobia (Tenagobia) truncata Deay, 1930 (Ecuador, Peru). - There are more than 20 setae on the left free lobe of the eighth abdominal tergite of the male (Fig. 6.355). ............................................................................8

Fig. 6.354 Tenagobia truncata male (left to right): head and pronotum, fore-leg, left posterior apex of the eighth abdominal segment, and left paramere. Based on Hungerford (1948).

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Fig. 6.355 Tenagobia marmorata male (left to right): left and right paramere and the left lobe of the eighth abdominal segment. Based on Nieser (1970c). 8. (7) The greatest width of the pronotum equals about three times the width of the scutellum at its base. The outer apical corner of the fore-femur of the male bears two short, strong setae and one long seta more than half as long as the tibia. The pala of the male bears 4 or 5 long, hair-like setae in the upper row, 12 to 15 bristle-like setae in the lower row, and 11 to 14 setae in the inner row (Fig. 6.356). The general coloration is dark. The length of the male is greater than 2 mm. ........................................................Tenagobia (Tenagobia) testacea Nieser, 1977 (Colombia, Ecuador). - The greatest width of the pronotum equals about twice the width of the scutellum at its base. The length of the male does not exceed about 1.9 mm (Fig. 6.357). ..........................................................................................................9

Fig. 6.356 Tenagobia testacea male (left to right): fore-leg and the left and right parameres. Based on Nieser (1977a).

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9. (8) The small patch of minute setae in the right anterior corner of the seventh abdominal tergite of the male is not round. The scutellum is only slightly longer than the pronotum. The pala of the male bears 5 or 6 long, hair-like setae in the upper row, 16 or 17 bristle-like setae in the lower row, and 14 to 17 setae in the inner row. The right paramere ends in a finger-like process. The free lobe on the left part of the eighth tergite bears 13 setae. The right paramere ends in a finger-like process (Fig. 6.355). The fore-tibia is longer than the pala. Length: 1.9 mm. ...............................................Tenagobia (Tenagobia) marmorata Bergroth, 1899 (Venezuela, Amazonas). - The small patch of minute setae in the right anterior corner of the seventh abdominal tergite of the male is round. The free lobe on the left part of the eighth tergite bears 18 setae. There are four moderately long bristles near the base of the right paramere (Fig. 6.357). The tibia and tarsus of the fore-leg are subequal in length. The pala of the male bears 4 or 5 long, hair-like setae in the upper row, 16 bristle-like setae in the lower row, and 14 or 15 setae in the inner row. Length: 1.8 to 2.0 mm. ............................................Tenagobia (Tenagobia) costaricana Jaczewski, 1930 (Central America). Syn: Tenagobia minuta Deay, 1930.

Fig. 6.357 Tenagobia costaricana male (left to right): right and left male paramere. Based on Deay (1935). 10. (3) There is a ventral row of only two or rarely four spines on the forefemur. The trochanter of the hind leg appears knob-like. The middle legs are usually yellow with a brown band in the basal ¼ of the femur. The ocular index is 0.5 to 0.8 (Fig. 6.351). See Fig. 6.16, p. 28, to calculate. .............................11 - There is a ventral row of three to nine spines on the fore-femur. The trochanter of the hind leg does not appear knob-like. The middle femur lacks a brown band on the basal ¼. The ocular index is 0.7 to 1.6 (Fig. 6.358). Only macropterous specimens have been found. ........................................................17 11. (10) There are four spines forming a basal row on the fore-femur. The spotted pattern on the hemielytra is usually distinct. The left side of the eighth tergite of the male bears very long setae (Fig. 6.351). Length: 2.7 to 2.9 mm. ..................................................Tenagobia (Romanogobia) peruana Nieser, 1977 (Peru).

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- There are only two spines forming a basal row on the fore-femur. The spotted pattern on the hemielytra is usually indistinct (Fig. 6.359). ...............................12

Fig. 6.358 Tenagobia selecta male: head in anterior view partially covered by one fore-leg (upper left) and head in dorsal view (upper center), anterior surface of the prothorax (upper right center), antenna (lower left), inner surface of the pala (lower middle left), right maxilla and right mandible (lower center, above and below, respectively), and left mandible and left maxilla (lower right center, above and below, respectively), hind and middle leg (center, left and right, respectively), claw (between pala and antenna), abdomen in dorsal and ventral view (right center, above and below, respectively), right (upper right) and left paramere (lower right). Based on Hungerford (1948). 12. (11) The ocular index is less than 0.7 (Fig. 6.359). See Fig. 6.16, p. 28, to calculate. .............................................................................................................13 - The ocular index is more than 0.7 (Fig. 6.360). ..............................................15 13. (12) The description is for macropterous specimens only. The posterior leg of the V-shaped mark is not notably broad. Each eye is about 1.4 times as wide as the interocular space (synthlipsis). The left paramere does not bear apical projections (Fig. 6.359). Length: 2.7 to 3.1 mm. .................................Tenagobia (Romanogobia) pseudoromani Nieser, 1970 pars (Surinam, Amazonas, Pará). See Couplet 16 for the brachypterous form. - The descriptions in the next couplet are for brachypterous specimens only. The posterior leg of the V-shaped mark is very broad (Fig. 6.361). ..................14

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Fig. 6.359 Tenagobia pseudoromani male (left to right): base of a hemielytron, left half of the pronotum, and left and right paramere. Based on Nieser (1975).

Fig. 6.360 Tenagobia romani male (left to right): inner surface of the fore-leg, left part of the eighth abdominal segment, and the left and right paramere. Based on Hungerford (1948).

Fig. 6.361 Tenagobia aconita male (above, left to right): male fore-leg, prothorax in anterior view, apex of the abdomen, and the right paramere, and (below, left to right): an enlarged claw on the fore-leg, antenna (below) and the left paramere. Based on Nieser (1977a).

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14. (13) The ratio of the body length to the width is about 2.8:1. The left paramere does not bear long apical projections (Fig. 6.361). Length: c. 2.4 mm. ...................................................Tenagobia (Romanogobia) aconita Nieser, 1977 (Colombia). - The ratio of the body length to the width varies from about 2.2:1 to 2.4:1. The left paramere bears long apical projections (Fig. 6.362). Length: 2.6 to 2.8 mm. ..............................................Tenagobia (Romanogobia) latioculata Nieser, 1970 (Surinam).

Fig. 6.362 Tenagobia latioculata male (left to right): left and right paramere and the left lobe of the eighth abdominal segment. Based on Nieser (1970c). 15. (12) There are about 20 to 22 setae in the lower row on the pala. The middle and hind legs are yellow with distinct dark brown bands and blotches. Each lateral margin of the eighth abdominal segment bears two spine-like setae. The left paramere bears short apical projections (Fig. 6.360). Length of male: 2.8 mm. ...............................................Tenagobia (Romanogobia) romani Lundblad, 1928 (Amazonas). - There are about 13 to 15 setae in the lower row on the pala (Fig. 6.363). .....16

Fig. 6.363 Tenagobia constricta male (left to right): inner surface of the foreleg, left part of the eighth abdominal segment, and the left and right paramere. Based on Hungerford (1948).

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16. (15) The median margin of the clavus is about 2.5 times as long as the scutellum. The right paramere has apical projections. The legs are uniformly yellowish. The lower row on the pala bears 14 setae. The ocular index is 0.78 to 0.85 (Fig. 6.16) Each lateral margin of the eighth abdominal segment bears one spine (Fig. 6.363). Length of male: 2.8 mm; female: 2.7 to 3.0 mm. .................................................Tenagobia (Romanogobia) constricta Deay, 1930 (Ecuador). Syn: Tenagobia (Romanogobia) constricta Nieser, 1977. - For brachypterous specimens only, the median margin of the clavus is about 2.0 times as long as the scutellum. The right paramere lacks apical projections (Fig. 6.359). Length: 2.3 to 3.1 mm. .................................Tenagobia (Romanogobia) pseudoromani Nieser, 1970 pars (Surinam, Amazonas, Pará). See Couplet 13 for the macropterous form. 17. (10) Most specimens are brachypterous. The transparent V-shaped mark on the clavus has a broad posterior part. The femur of the fore-leg has a basal row of seven to nine spines. The inner postocular space is long. The margins of the eighth abdominal segment bear four to five spines and two long bristles (Fig. 6.358). .................................................................................................................18 - Most specimens are macropterous. The posterior part of the transparent Vshaped mark on the clavus is not broad. The femur of the fore-leg has a basal row of three to six spines. If the inner postocular space is long, then the margins of the eighth abdominal segment bear two spines and two long bristles (Fig. 6.364). .................................................................................................................19

Fig. 6.364 Tenagobia schadei male: left posterior apex of the eighth abdominal segment (left), and the left (center) and right paramere (right). Based on Hungerford (1948), who referred to the species as Tenagobia hungerfordi. 18. (17) The eighth tergite is produced into a long, finger-like process (Fig. 6.358). Length of male: 4.2 to 4.6 mm; female: 4.4 to 4.8 mm. ......................................................Tenagobia (Fuscagobia) selecta (White, 1879) (Colombia, Bolivia, Paraguay, Amazonas).

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- The eighth tergite is not produced. Each eye is about 1.25 times the interocular space. There are one terminal and three lateral spine-like setae on each lateral margin of the eighth abdominal segment, and seven to nine spinelike setae on the inner side of the fore-femur (Figs. 6.25, 6.365). Length of male: 3.3 to 3.45 mm; female: 3.4 to 3.75 mm. Bressa and Papeschi (2007) reported the 2n chromosome number of the male to be 30 (XY). .........................................................Tenagobia (Fuscagobia) fuscata (Stål, 1859) (Paraguay, Argentina, Uruguay). Syn: Sigara fuscata Stål, 1859.

Fig. 6.365 Tenagobia fuscata male: antenna (upper left); prothorax in anterior view (upper middle left); hemielytra (lower right); apex of fore-leg with the pala and its claw enlarged to its lower right (center); tarsal claw on the hind leg (lower left); sixth, seventh, and right side of eighth abdominal segment (upper right); left (lower middle left) and right genital lobe (lower center). Based on Lundblad (1928a) and Galliussi (1989). 19. (17) The inner postocular space is at least 3.5 times as long as the diameter of a compound eye facet. The ocular index varies from 0.85 to 1.1 (Fig. 6.16). There are two spines and two long bristles on the margins of the eighth abdominal segment (Fig. 6.364). The pruinose area on the embolium covers a length equalling 0.4 to 0.6 times the length of a hemielytron. ...........................20 - The length of the inner postocular space is no more than 2.5 times the diameter of a facet of the compound eye. The ocular index varies from 1.2 to 1.6 (Fig. 6.16). There are four spines, three long bristles, and one relatively short seta on each margin of the eighth abdominal segment (Fig. 6.366). The pruinose area on the embolium covers a length equalling 0.2 to 0.3 times the length of a hemielytron. ......................................................................................21 20. (19) Proximal to the fold just beyond its mid-length, the left paramere is only slightly expanded. The right paramere has a pointed angle at the apex (Fig. 6.364). The lower row on the pala consists of 12 to 15 bristles; the inner row consists of 13 to 16 setae; the upper row has 4 to 5 long, hair-like setae. The

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receptaculum seminis of the female is shaped like an urn. Length of male: 3.1 to 3.4 mm; female: 3.3 to 3.7 mm. ..............................................Tenagobia (Schadeogobia) schadei Lundblad, 1928 (Ecuador, Colombia, Surinam, Bolivia, Paraguay, Argentina, Uruguay, Pernambuco, Pará, Amazonas, Goiás, Minas Gerais, Mato Grosso). Syn: Tenagobia hungerfordi Deay, 1930. - Proximal to the fold just beyond its mid-length, the left paramere has a conspicuous swelling. The right paramere is broadly rounded at its apex. The lower row on the pala consists of 13 to 14 bristles; the inner row consists of 13 to 15 setae; the upper row has 4 to 5 long, hair-like setae (Fig. 6.345). The receptaculum seminis of the female is elongate. Length of male: about 3.3 to 3.4 mm; female: 3.4 to 3.9 mm. These two nominal species can only be distinguished by the shapes of the parameres, and it appears questionable whether they should be regarded as separate species. ...........................................................................Tenagobia melini Lundblad, 1928 (Amazonas).

Fig. 6.366 Tenagobia spinifera male: left and right paramere and the left lobe on the eighth abdominal segment. Based on Nieser (1970c).

Fig. 6.367 Tenagobia carapachay: head in dorsal view (upper left), habitus in dorsal view (upper left center), two views of the fore-leg (upper and middle right), lateral lobe of the eighth abdominal segment (lower left), and the right (lower center) and left male paramere (lower right). Based on Bachmann (1961).

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21. (19) There are more than 10 long, hair-like setae in the upper row on the pala (Fig. 6.366). ........................................................................................................22 - The upper row on the pala consists of no more than 10 long, hair-like setae (Fig. 6.367). ........................................................................................................23 22. (21) Length: 3.3 to 3.8 mm. There are 20 to 25 bristle-like setae in the lower row on the pala (Fig. 6.366). ...................................................Tenagobia (Incertagobia) spinifera Nieser, 1970 (Venezuela, Colombia). - The total length is shorter than 3 mm. There are no more than 15 bristle-like setae in the lower row on the pala. The toe of the right paramere is bluntly rounded. Its heel forms a right angle and is not protracted (Fig. 6.368). Length of male: 2.4 to 2.8 mm; female: 2.6 to 3.0 mm. ...................................................Tenagobia (Incertagobia) signata (White, 1879) (Venezuela, Surinam, Amazonas). Syn: Sigara signata White, 1879; Sigara socialis var. sobrina White, 1879; Sigara seducta White, 1879; Sigara simulans White, 1879.

Fig. 6.368 Tenagobia signata male (left to right): left and right paramere, left lobe of the eighth abdominal segment, and left and right paramere of a different specimen from Amazonas. Based on Nieser (1970c, 1975). 23. (21) There are 17 stiff bristles in the lower row on the pala, 15 in the inner row, and 8 long setae in the upper row. Ventral from the beak-like process on the right paramere is a small tooth (Fig. 6.369). Length of male: c. 2.3 mm; female: 2.5 to 2.7 mm. ..................................................Tenagobia (Incertagobia) lundbladi Nieser, 1977 (Venezuela). - There are 14 or 15 stiff bristles in the lower row on the pala. The right paramere lacks a small tooth near a beak-like process (Fig. 6.330). ..................24

Fig. 6.369 The left (left) and right paramere (right) of a male Tenagobia lundbladi. Based on Nieser (1977a).

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24. (23) There are 12 bristles in the inner row on the male pala, 12 to 14 in the lower row, and 6 or 7 long setae in the upper row. The posteroventral margin of the right paramere is evenly curved (Fig. 6.367). Length: 2.1 to 2.7 mm. ........................................Tenagobia (Incertagobia) carapachay Bachmann, 1961 (Argentina). - There are 14 or 15 bristles in the inner row on the male pala. The posteroventral margin of the right paramere is angular (Fig. 6.330). ................25 25. (24) The apex of the left paramere is abruptly expanded into a rounded point. The heel of the right paramere is acutely pointed and bears a toe-like process (Fig. 6.330). The right paramere may also have a double row of serrations ventrally. The receptaculum seminis of female specimens is shaped like a dumb-bell. Length of male: 2.25 to 2.75 mm; female: 2.6 to 3.1 mm. ...................................................Tenagobia (Incertagobia) socialis (White, 1879) (Central America, Trinidad, Colombia, Venezuela, Guyana, French Guiana, Surinam, Bolivia, Argentina, Amazonas, Pará Amapá, Pernambuco). Syn: Sigara socialis White, 1879; Tenagobia serrata Deay, 1930. - The apex of the left paramere is not greatly expanded.and bears only a small toe-like process (Fig. 6.370). The receptaculum seminis of the female is shaped like an urn. ..........................................................................................................26

Fig. 6.370 Tenagobia campula: the left (upper left) and right paramere (lower left); the receptaculum seminis (right). Based on Nieser (1977a). 26. (25) The left paramere is hooked (Fig. 6.370). The pala of the male bears 5 to 7, long, hair-like setae in the upper row, 13 to 15 bristle-like setae in the lower row, and 18 or 19 setae in the inner row. Length of male: 2.6 to 2.9 mm; female: 3.2 to 3.5 mm. ...................................................Tenagobia (Incertagobia) campula Nieser, 1977 (Peru). - The left paramere is not hooked (Fig. 6.371). Length of male: 2.6 to 2.9 mm; female: 3.2 to 3.5 mm. ........................................................................................27

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Fig. 6.371 Tenagobia incerta male: left (left) and right paramere (right). Based on Hungerford (1948). 27. (26) The left paramere has a long, narrow dorsal expansion at the apex and a distinctly produced apicoventral edge (Fig. 6.372). The receptaculum seminis of female specimens is very narrow. Length of male: 2.3 to 2.5 mm; female: 2.5 to 2.7 mm. .................................................Tenagobia (Incertagobia) mexicana (Deay, 1930) (North and Central America). Syn: Sigara socialis Uhler, 1894 pars. - The left paramere has a short dorsal expansion at the apex and no more than a vestige of a process on the apicoventral edge (Fig. 6.371). The receptaculum seminis of female specimens is broad. .................................................Tenagobia (Incertagobia) incerta Lundblad, 1928 (Central America, Grenada, Trinidad, Colombia, Venezuela, Surinam, Peru, Bolivia, Paraguay, Argentina, Pernambuco, Ceará, Baia, Paraíba, Pará, Amazonas, Roraima, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo, Santa Catarina, Rio Grande do Sul).

Fig. 6.372 Tenagobia mexicana male (left to right): a pair of hooked and apically serrate setae and the left and right paramere. Based on Nieser (1977a).

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Subfamily Heterocorixinae Only one South American genus is known: Heterocorixa White, 1879. Key to the species and subspecies of Heterocorixa in South America Information for the key was provided by Hungerford (1928a, 1948) and Nieser (1970c, 1975). 1. The claws on the middle leg are longer than the tarsus. The corium bears both slender hairs and short spines. The metaxyphus is usually shorter than the inner line of the hind coxae. The hind femur is pilose on its basal 2/5, and its ventral surface bears many spines. The male has a short prestrigilar comb and a triangular seventh abdominal tergite (Fig. 6.373). ...............................................2 - The claws on the middle leg are equal to or shorter than the tarsus. There are no short spines on the corium. The metaxyphus is as long or longer than the inner line of the hind coxae. Little more than the basal tip of the hind femur is pilose, and there are no more than 12 spines on its ventral side. The male possesses a long prestrigilar comb or lacks one, and its seventh tergite is not produced into a triangular form (Fig. 6.374). .....................................................10

Fig. 6.373 Heterocorixa williamsi male (left to right): tibia and pala, left and right parameres, and the apex of the abdomen in dorsal view. Based on Hungerford (1948).

Fig. 6.374 Heterocorixa anduzei male: fore-tibia and pala (upper left), left (center) and right paramere (right), and apex of the abdomen (lower left). Based on Nieser (1970c).

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2. (1) The head is broadly rounded and longer than the very short pronotal disc, which it embraces (Fig. 6.373). Length: 4.2 to 4.6 mm. ..............................................................Heterocorixa williamsi Hungerford, 1928 (Ecuador). - The pronotal disc is not surrounded by the head, which is not broadly rounded (Fig. 6.375). ..........................................................................................................3 3. (2) The pattern on the hemielytra consists of fine, smooth, and closely set markings (Fig. 6.375). ..........................................................................................4 - The pattern on the hemielytra is coarse (Fig. 6.376). .......................................6

Fig. 6.375 Heterocorixa hesperia female (left to right): pronotum in dorsal view, metaxyphus (above), lateral lobe of the pronotum (below), hemielytra, and pala. Based on Lundblad (1928b).

Fig. 6.376 Heterocorixa lundbladi male (left to right): tibia and pala, left and right parameres, and the apex of the abdomen in dorsal view. Based on Hungerford (1948). 4. (3) The metaxyphus is not more than half the length of the inner line of the hind coxa (Fig. 6.375). The male of the nominate subspecies has not been described. While the male of the Venezuelan subspecies is known, the status of this subspecies is doubtful. ......................................................................Heterocorixa hesperia (White, 1879) (Bolivia, Goiás, Pará, Rio Grande do Norte). A subspecies, H. hesperia venezuelana Hungerford, 1948, is known only from Venezuela and Colombia, but Nieser (1975) noted that the sexual organs of this form hardly differ at all from those of Heterocorixa braziliensis. The taxonomic status of this subspecies is therefore very doubtful.

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- The metaxyphus is longer than half the length of the inner line of the hind coxa (Fig. 6.377). .................................................................................................5

Fig. 6.377 Heterocorixa braziliensis (above, left to right): abdomen of a male in dorsal view, fore-leg of a male, genital capsule above the left paramere, and (middle row, left to right): abdomen of a female in dorsal view, fore-leg of a female, gonopodia of a female, hind femur in ventral view made transparent to show the row of setae across the dorsal side, and (below, left to right): metaxyphus, antenna (above) and the claw on the hind leg (below), basal row of modified setae on the fore-femur of a male above the same row in profile view (upper left center of row), spermatheca and its ducts, and the prestrigil and strigil. Based on Bachmann (1979a). 5. (4) The metaxyphus is at least 2/3 the length of the inner line of the hind coxa (Figs. 6.24, 6.377). Length: 4.6 to 5.6 mm. ...........................................................Heterocorixa braziliensis Hungerford, 1928 (Ecuador, Argentina, Mato Grosso do Sul, and questionable reports from Paraíba, Amazonas, Pernambuco, and Goiás). Syn: Heterocorixa brasiliensis auctt. (misspelled name). When the status of H. hesperia is better known, H. braziliensis may prove to be a junior synonym of that species, and H. hesperia venezuelana Hungerford, 1948, will even more likely prove to be a junior synonym (Nieser, 1975). - The metaxyphus is only slightly more than half the length of the inner line of the hind coxa. The pala of the male is only slightly wider than the fore-tibia at the apex (Fig. 6.378). Length: 4.6 to 5 mm. ..................................................................Heterocorixa hintoni Hungerford, 1948 (Bolivia).

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Fig. 6.378 Heterocorixa hintoni male (left to right): tibia and pala, left and right paramere, and the apex of the abdomen in dorsal view. Based on Hungerford (1948). 6. (3) The seventh abdominal segment of the male is indented (Fig. 6.346). ............................................................Heterocorixa westermanni Lundblad, 1928 (Amazonas?). - There is no indentation on the seventh abdominal segment of the male (Fig. 6.376). ...................................................................................................................7 7. (6) The pala of the male is bluntly rounded at the tip. The metaxyphus is broadly or unevenly rounded at the apex (Fig. 6.376). Length: 5.3 to 5.9 mm. ..............................................................Heterocorixa lundbladi Hungerford, 1948 (Amazonas). - The pala of the male is almost pointed at the apex (Fig. 6.379). The metaxyphus is pointed. .........................................................................................8

Fig. 6.379 Heterocorixa jaczewskii male (left to right): tibia and pala, left and right paramere, and the apex of the abdomen in dorsal view. Based on Hungerford (1948).

314

8. (7) The median lobe of the seventh segment of the male is slightly produced and forms nearly a straight line with the right tergite (Fig. 6.379). Length of male: 6.1 mm; female 7 mm. ................................................................Heterocorixa jaczewskii Lundblad, 1928 (Pernambuco). - The shape of the seventh abdominal segment of the male differs, not forming a nearly straight line with the right tergite (Fig. 6.380, 6.381). Length: 5.8 to 6.8 mm. ..........................................Heterocorixa wrighti Hungerford, 1948……………..9 9. (8) The male has no spines to the left of the strigil and 12 to 14 teeth in the prestrigilar comb (Fig. 6.380). .....................................................Heterocorixa wrighti wrighti Hungerford, 1948 (Ceará). - The male has 1 to 3 spines on the sixth abdominal segment to the left of the strigil and 8 or 9 teeth in the prestrigilar comb (Fig. 6.381). ......................................................Heterocorixa wrighti ollalai Hungerford, 1948 (Amazonas).

Fig. 6.380 Heterocorixa wrighti wrighti male (left to right): inner surface of the fore-tibia and pala, left and right paramere, and apex of the abdomen in dorsal view. Based on Hungerford (1948).

Fig. 6.381 Heterocorixa wrighti ollalai male (left to right): inner surface of the fore-tibia and pala, left and right paramere, and apex of the abdomen in dorsal view. Based on Hungerford (1948).

315

10. (1) The metaxyphus is very long and reaches the apex of the hind coxal projection (Fig. 6.374). .......................................................................................11 - The metaxyphus may be long, but it does not reach the tip of the hind coxal projection (Fig. 6.382). .......................................................................................15 11. (10) The male fore-tibia is greatly produced beyond the joint with the pala (Fig. 6.374). There is one strigil and no prestrigilar comb. Length: c. 4.8 mm. .................................................................Heterocorixa anduzei Hungerford, 1948 (Venezuela, Amazonas). - The fore-tibia of the male is not greatly produced (Fig. 6.383). ......................12

Fig. 6.382 Heterocorixa woytkowskii male: tibia and pala (upper left), left and right paramere (middle and lower left, respectively), apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.383 Heterocorixa genupes male: fore-tibia and tarsus (upper left), apical segments of the abdomen (lower left), and the left (right center) and right paramere (right). Based on Nieser (1970c).

316

12. (11) The pronotum is distinctly rugulose with transverse grooves. The pala of the male is geniculate (Fig. 6.383). Length of male: 4.3 to 4.5 mm; female: 4.3 to 4.8 mm. ........................................................................Heterocorixa genupes Nieser, 1970 (Amazonas). - The pronotum is at most slightly rugulose without transverse grooves. The pala of the male is not geniculate (Fig. 6.384). ..................................................13

Fig. 6.384 Outline of the posterior margins of the head and compound eye of Heterocorixa longixiphus in lateral view. Based on Nieser (1975). 13. (12) The metaxyphus extends beyond the tips of the coxal projections. The posterolateral margin of the head is not strongly sinuous (Fig. 6.384). Length: c. 4.05 mm. ...................................................................Heterocorixa longixiphus Nieser, 1970 (Surinam). - The metaxyphus does not extend beyond the tips of the coxal projections. The posterolateral margin of the head is strongly sinuous (Fig. 6.385). ...................14

Fig. 6.385 Heterocorixa similis (left to right): outline of the posterior margins of the head and compound eye in lateral view, pala, left and right parameres, and apex of the male abdomen. Based on Nieser (1970c).

317

Fig. 6.386 Heterocorixa hungerfordi male (left to right): outline of the posterior margins of the head and compound eye showing the sigmoid outline of the posterior margin and the two parameres. Based on Nieser (1970c, 1975). 14. (13) The right clasper of the male has a drawn-out acute apex (Fig. 6.385). Length of male: 3.6 to 3.8 mm; female: 4.0 to 4.2 mm. ...........................................................................Heterocorixa similis Nieser, 1970

(Surinam, Amazonas).

- The right clasper of the male is blunt at the apex (Fig. 6.386). Length of male: c. 4.8 mm. ..................................................................Heterocorixa hungerfordi Nieser, 1970 (Amazonas). That the two species in this couplet are distinct is subject to doubt. 15. (10) Length: c. 3.0 to 3.2 mm. The posterolateral margin of the head is concave (Fig. 6.387). ..........................................................................Heterocorixa minuta Nieser, 1970 (Amazonas). - Length: greater than 3.5 mm. ..........................................................................16

Fig. 6.387 The apex of the abdomen of a male Heterocorixa minuta. Based on Nieser (1970c). 16. (15) The predominant color is black (Fig. 6.388). .......................................17 - The prodominant colors are light. ...................................................................18

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Fig. 6.388 Heterocorixa nigra male: pronotum in dorsal view (upper left), antenna (middle left), metaxyphus (upper left center), profile of the metathorax in left lateral view (below metaxyphus), meso and metathorax in left lateral view (left center), hemielytra (upper center), pala of a male (upper right center), inner surface of the right fore-leg (upper right), prestigil (upper middle right) and strigil (lower right), fourth through seventh abdominal segments in ventral view (lower left to center), apical part of the penis with its sheath (right center), genital capsule in left lateral view (below penis apex), left paramere in dorsal view (lower center), and left and right paramere in lateral view (above and right of the dorsal view of the left paramere, respectively). Based on Lundblad (1928b). 17. (16) The fore-tibia of the male is carinate. The metaxyphus is longer than the inner line of the hind coxa (Fig. 6.388). Length: 4.1 to 4.9 mm. ....................................................................Heterocorixa nigra Hungerford, 1928 (Argentina, São Paulo, Minas Gerais). Syn: Heterocorixa reinhardti Lundblad, 1928. - The fore-tibia of the male is not carinate. The metaxyphus is subequal to the inner line of the hind coxa (Fig. 6.382). Length: 4.2 to 4.8 mm. ..........................................................Heterocorixa woytkowskii Hungerford, 1948 (Peru). 18. (16) The posterior pruinose area on the embolar groove is considerably longer than that on the claval suture. The anterior tibia of the male lacks a flap but has a carina on its posterior side (Fig. 6.389). Length: 4.2 to 5.25 mm. ........................................................Heterocorixa chapadiensis Hungerford, 1928 (Mato Grosso, Amazonas, Pará). Syn: Heterocorixa Hesperia var. chapadiensis Hungerford, 1928.

319

- The posterior pruinose area on the embolar groove is a little longer than that on the claval suture. The anterior tibia of the male has a long flap resting in the depression in the pala (Fig. 6.390). ....................................................................19

Fig. 6.389 Heterocorixa chapadiensis male: tibia and pala (upper left), left and right paramere (middle and lower left, respectively), apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.390 Heterocorixa boliviensis male: tibia and pala (upper left), left and right paramere (middle and lower left, respectively), apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 19. (18) The spinose dorsal flap on the left apical lobe of the abdomen has a relatively small apical projection (Fig. 6.390). Length: 4.2 to 5.25 mm. ............................................................Heterocorixa boliviensis Hungerford, 1928 (Bolivia, Ecuador, Peru, Surinam, Amazonas, Pará).

320

- The spinose dorsal flap on the left apical lobe of the abdomen has a distinct, finger-like apical projection (Fig. 6.391). Length of male: 4.8 to 5.0 mm; female: 5.1 to 5.3 mm. ................................................................Heterocorixa surinamensis Nieser, 1970 (Trinidad, Surinam).

Fig. 6.391 Heterocorixa surinamensis male (left to right): outline of the posterior part of the head and compound eyes in lateral view, fore-tibia and pala, left and right paramere, and apex of the abdomen in dorsal view. Based on Nieser (1975). Subfamily Corixinae Key to the South American genera Information for this key was provided by Hungerford (1948) and Padilla and Nieser (1994). 1. Small, shiny insects less than 5.6 mm long. The pala of the male is short and triangular, and the tibia is produced over it. Its genitalia display sinestral asymmetry. The apices of the clavi of the females remain within the lines drawn through the costal margins of the hemielytra at the nodal furrows (Fig. 6.392). ........................................................................Trichocorixa Kirkaldy, 1908 p. 325 - The length is more than 5.6 mm, or, if not, the insects are not shiny, or the pala of the male is enlarged enough so that the tibia is not produced over it (Fig. 6.393). ...................................................................................................................2 2. (1) The interocular space is equal to or greater than the width of an eye measured along the posterior margin. The ventral surface of the hind femur is pubescent only on the basal 1/4 or 1/5 of the anterior margin and bears two or more irregular rows of short pegs on the glabrous portion; the pegs at the distal end of the row on the male pala are elongate and crowded into a double row; the basal pegs are elongate and spiniform (Fig. 6.393). .............................................3 - At least one of the features above is lacking. ....................................................4 3. (2) The anterior tibia of the male overlaps the base of the pala. The glabrous ventral portion of the hind femur bears numerous fine hairs interspersed among pegs (Fig. 6.393). .....................................................................Ectemnostega Enderlein, 1912 p. 330

321

- The anterior tibia of the male does not overlap the pala. The glabrous ventral portion of the hind femur bears only pegs (Fig. 6.394). .................................................................Ectemnostegella Lundblad, 1928 p. 330

Fig. 6.392 Trichocorixa milicorum (above, left to right): apical segments of the abdomen of a male in dorsal view, fore-leg of a male, genital capsule (above) and right paramere (below), spermotheca and its ducts, and (middle row, left to right): apical segments of the abdomen of a female in dorsal view, fore-leg of a female, gonopods of a female, strigil, and the claws on the hind leg, and (below, left to right): disc of the pronotum, antenna (above) and the right hemielytron of a female (below), metaxyphus, and the hind femur in ventral (above) and dorsal view (below). Based on Bachmann (1979a). 4. (2) The length of the pruinose area along the claval suture is less than twice the length of the distance between the shining basal apices of the corium and the clavus; the postnodal pruinose area measured from the cubital angle is shorter than or barely equal to the meron. Males lack a strigil or strigilar stalk (Fig. 6.395). .......................................................................Centrocorisa Lundblad, 1928 p. 336 - At least one of the above features is lacking. ....................................................5 5. (4) There is a reticulate pattern on the hemielytra. The face and hemielytra appear hairy. The median portion of the seventh abdominal segment of the females is incised (Fig. 6.396). .................................................................................Sigara Fabricius, 1775..p. 337 - The color pattern may include unbroken light stripes, but there is no conspicuous reticulation (Fig. 6.397). ..................................................................6

322

Fig. 6.393 Ectemnostega quadrata: habitus in dorsal view (lower left) and (above, left to right): dorsal outlines of the head and pronotum, lateral margin of a hemielytron, femur (left center), genital capsule of a male, right paramere of a male (center), male strigil (lower right center), apex of the female abdomen, and the tibia and pala of a male. Based on Hungerford (1948).

Fig. 6.394 Ectemnostegella venturii (above, left to right): dorsal view of head and pronotum of a macropterous specimen and the hemielytron of a female, and (below, left to right): head in lateral view, pronotum of a brachypterous specimen, femur, and apex of the female abdomen in ventral view. Based on Hungerford (1948).

323

Fig. 6.395 Centrocorisa kollari: habitus of a male in dorsal view (upper left), head and pronotum of a male (lower left) and a female (lower right center) in lateral view, fore-leg with the pala of a male (lower left center) and a female (lower right), abdomen of a male (upper right) and a female (middle right) in dorsal view. Based on Alvarez and Roldán (1983).

Fig. 6.396 Sigara yala (upper row, left to right): head of a male in lateral view, disc of the pronotum, inner surface of the fore-leg of a male and dorsal view of its pala, apical part of the fore-leg of a female, metaxyphus, fore-femur in ventral (middle right) and dorsal view, strigil (middle left), male left paramere (right of center), and (below, left to right): male abdomen in dorsal view, genital capsule, and female gonopodia. Based on Bachmann (1979a).

324

Fig. 6.397 Ramphocorixa rotundocephala male: dorsal view of head as shown by Nieser (1969a) and Hungerford (1927c) in his original description of the species (left and left center, respectively), tibia and pala (center), strigil (upper right center), genital capsule (lower right center), and the left (lower right) and right (upper right) paramere. Based on Hungerford (1927c) and Nieser (1969a). 6. (5) The dorsal margin of the male pala is deeply incised, and a strigil is present. In dorsal view, the vertex of the male displays a rounded median prominence (Fig. 6.397). The claws on the palae of both sexes are serrate basally. The pattern on the hemielytra does not appear sharp and is sometimes effaced. The length of the only species reported from South America is about 5 to 6 mm. The width of the head is less than 2 mm. ...................................................................................Ramphocorixa Abbott, 1912 The only known South American species in this genus is Ramphocorixa rotundocephala Hungerford, 1927, from North America, the West Indies, and Venezuela. The generic name is frequently misspelled as Rhamphocorixa. - The dorsal margin of the pala is not deeply incised, or a strigil is absent, or the vertex of the male is rounded and not protruding, or the length is longer than 7 mm or shorter than 4 mm. ....................................................................................7 7. (6) Broadly compact species 5.7 to 6.9 mm long. The upper palmer row of bristles on the male pala is interrupted near the distal end; pegs are inserted in the opening in two of the three species (Fig. 6.398). ..........................................................................Neosigara Lundblad, 1928..p. 350 - Length: greater than 7 mm, or if slightly shorter, then the apical section of the lower peg row consists of hairs rather than elongated pegs (Fig. 6.399). ........................................................................Orocorixa Nieser and Padilla, 1992 The only known South American species in this genus is Orocorixa makrocheira Nieser and Padilla, 1992, from Colombia.

325

Fig. 6.398 Neosigara paramo male (left to right): head in anterior view showing the fovea in the middle, fore-leg, hind femur in ventral view, the sixth and the seventh abdominal segments in dorsal view, and the right paramere in lateral view. Based on Tinerella and Polhemus (2006).

Fig. 6.399 Orocorixa makrocheira male (left to right): fore-leg, sixth and seventh abdominal segments in dorsal view, and the right paramere. Based on Nieser and Padilla (1992). Key to the species of Trichocorixa in South America Information for the key was provided by Jaczewski (1933a, b), Hungerford (1948), Nieser (1970c), and Bachmann (1979a). 1. The width of the synthlipsis exceeds the width of one eye measured along the posterior margin, as viewed from above (Fig. 6.400). .........................................2 - The width of the synthlipsis is usually less than the width of one eye measured along the posterior margin, as viewed from above (Fig. 6.401). .........................3

Fig. 6.400 Trichocorixa beebei male: pronotum in dorsal view (left), hemielytra (upper left center), apex of the abdomen (right center), fore-leg in inner view (right), and the right (lower left center) and left parameres (lower center). Based on Hungerford (1948).

326

Fig. 6.401 Trichocorixa verticalis (left to right): head in anterior view and the male parameres. Based on Hungerford (1948) and Nieser (1975). 2. (1) The length of a pala is equal to the width of a compound eye measured along the ventral margin. There are about 12 pegs forming an arch-like curve on the pala of the male (Fig. 6.400). Length of male: c. 3.0 mm; female: 3.2 mm. ............................................................................Trichocorixa beebei Sailer, 1948 (Galapagos Islands). - The length of the pala is less than 2/3 the width of a compound eye measured along the ventral margin. There are about 18 pegs forming a slightly sinuous curve on the pala of the male (Fig. 6.402). Length of male: 2.8 to 4.9 mm; female: 3.4 to 5.2 mm. .................................................Trichocorixa reticulata (Guérin-Ménéville, 1857) (China, North and Central America, Hawaiian Islands, West Indies, Venezuela, Columbia, Surinam, Peru, Ecuador, including the Galapagos Islands). Syn: Corisa reticulata Guérin-Ménéville, 1857; Corixa wallengreni Stål, 1859; Corixa blackburni White, 1877; Corixa mariae Champion, 1901; Arctocorixa blackburni Kirkaldy, 1910; Trichocorixa mariae Jaczewski, 1927; Trichocorixa blackburni Hungerford, 1928; Trichocorixa wallengreni Lundblad, 1929. 3. (1) Male: The expansion of the anterior lateral angle of the left posterior lobe of the abdomen is small and includes not more than the anterior third of the lateral margin, and the pala is shorter than the tibia and evenly rounded on the dorsal margin. The distance between the compound eyes measured across the vertex is greater than the width of an eye. The pala of the male has a row of about 16 pegs in the form of a wide arch, which does not come close to the dorsal margin. Female: The length of the polished area along the costal margin of each hemielytron anterior to the nodal furrow is equal to or less than the length of the middle tibia (Fig. 6.401). Length of male: 3.6 to 4.6 mm; female: 3.8 to 5.4 mm. Bressa and Papeschi (2007) reported the 2n chromosome number to be 24 (XY). ....................................................................Trichocorixa verticalis (Fieber, 1851) (North and Central America, West Indies, Surinam). Only the subspecies T. verticalis verticalis has been reported from South America. Syn: Corisa verticalis Fieber, 1851; Corisa pygmaea Fieber, 1851; Trichocorixa pygmaea Kirkaldy, 1908; Corixa sellaris Abbott, 1913; Corixa verticalis Abbott, 1913; Trichocorixa verticalis var. sellaris Sailer, 1948.

327

- Male: The expansion of the anterior lateral angle of the left posterior lobe of the abdomen is large and includes the anterior half of the lateral margin, or the pala is as long as the tibia and straight or slightly concave along the dorsal margin. Female: The length of the polished area along the costal margin anterior to the nodal furrow of each hemielytron is greater than the length of the middle tibia (Fig. 6.403). ......................................................................................4

Fig. 6.402 Trichocorixa reticulata male: head in anterior view (lower right), pronota of three individuals in dorsal view (lower left to lower center), hemielytron (upper left), apex of the abdomen (upper right), fore-leg in inner view (middle left), and the right (lower left center) and left paramere (middle right). Based on Hungerford (1948).

Fig. 6.403 Trichocorixa orinocoensis male: pronotum in dorsal view (left), hemielytron (upper center), apex of the abdomen (right), fore-leg in inner view (lower left center), and the right (lower center) and left paramere (lower right center). Based on Hungerford (1948).

328

4. (3) Male: There are only about 11 pegs in the row on the pala, which is obtusely angled (Fig. 6.403). The expansion of the anterior lateral angle of the left posterior lobe of the abdomen is large and includes the anterior half of the lateral margin. The apex of the posterior left lobe is narrowly truncate. Length: c. 4.3 mm. Female: At least along the thickened anterior half of the polished prenodal area on the embolar groove, the costal margin is impressed and thickened laterally to give the impression of an oblique groove originating ventrally and progressing posteriorly to the dorsal margin. Length: 4.3 to 4.5 mm. .........................................................Trichocorixa orinocoensis Hungerford, 1948 (West Indies, Venezuela, Colombia, Surinam, Paraíba, Pará). Syn: Trichocorixa orinocensis Nieser, 1970, misspelled. - Male: The pala has approximately 18 pegs in a single row, which is either slightly curved or sinuous (Fig. 6.404). Female: The costal margin is flattened and blade-like along the polished prenodal area of the embolar groove, and it is sometimes thickened or has a rounded edge on the anterior fourth. If this is not the case, the pale transverse lines on the pronotum are notably thicker than the dark lines. .............................................................................................................5

Fig. 6.404 Trichocorixa mendozana mendozana: pattern on the disc of the pronotum (left) and on a hemielytron (upper center), inner surface of the foretibia and pala of a male (left center), apical segments of the male abdomen in dorsal view (lower right center), left (lower right) and right paramere (middle right), and apical segments of a female abdomen (lower left). Based on Hungerford (1948). 5. (4) Male: The row of approximately 18 pegs on the pala is simply angled and closely approaches the dorsal margin of the pala only at its distal end. In both sexes, the pale transverse lines on the pronotum are notably thicker than the 10 or 11 dark lines (Fig. 6.392). Length of male: 3.8 to 4.5 mm; female: 3.6 to 4.9 mm. ...............................................................Trichocorixa milicorum Bachmann, 1979 (Argentina).

329

- Male: The row of about 18 pegs on the pala is strongly sinuous and reaches or nearly reaches the dorsal margin at its distal end and at a second point at the point of an angle. The expansion of the anterior lateral angle of the left posterior lobe of the abdomen is small and does not include more than the anterior third of the lateral margin; the apex of this lobe is not narrowly truncate and similar to the right lobe. Length: 4.1 to 4.4 mm. Female: The costal margin is flattened and blade-like along the polished prenodal area of the embolar groove, and it is sometimes thickened or has a rounded edge on the anterior fourth (Fig. 6.404). Length: 4.3 to 4.6 mm. In both sexes, the pale transverse bands across the pronotum are about equal to or thinner than the dark bands. ..............................................................Trichocorixa mendozana Jaczewski, 1927 Two subspecies are recognized, as distinguished by the following key. Supplemental key to the subspecies of Trichocorixa mendozana Information for the key was provided by Hungerford (1948). 1. Males: The right paramere is nearly as broad at the apex as in the middle. Females: The width of the synthlipsis is equal to the posterior width of an eye (Fig. 6.404). ...........................................Trichocorixa mendozana mendozana Jaczewski, 1927 (Argentina). 2. Males: The right paramere is less than half as wide at the apex as in the middle. Females: The width of the synthlipsis is greater than the posterior width of an eye (Fig. 6.405). .........................................Trichocorixa mendozana darpomorza Jaczewski, 1933 (Paraíba).

Fig. 6.405 Trichocorixa mendozana darpomorza male (left to right): strigil and the left and right paramere. Based on Hungerford (1948).

330

Key to the species of Ectemnostega in South America Information for the key was provided by Jaczewski (1927c) and Hungerford (1948). The key can be used only for females because the male of Ectemnostega darwini has not been described. 1. The pronotum is crossed by 8 or 9 narrow, broken black lines, which are impressed in the surface and overlap somewhat. The length of the head along the midline is slightly greater than 1/3 of its maximum width. The disc of the pronotum is less than twice its length along the midline (Fig. 6.393). Length: 6.0 to 6.7 mm. ................................................................Ectemnostega quadrata (Signoret, 1885) (Argentina, Chile). Syn: Corixa quadrata Signoret, 1885. - The disc of the pronotum is crossed by 12 to 14 very irregular brown lines, which fuse with each other laterally on the distal half. The length of the head along the midline is slightly less than 1/2 of its maximum width. The disc of the pronotum is more than twice its length along the midline (Fig. 6.406). Length of female: c. 7.8 mm. The male has not been described. ................................................................Ectemnostega darwini Hungerford, 1948 (Argentina).

Fig. 6.406 Ectemnostega darwini female (left to right): dorsal views of the head, pronotum, lateral margin of a hemielytron (above), femur (below), and apex of the abdomen. Based on Hungerford (1948). Key to the species of Ectemnostegella in South America Information for the key was provided by Hungerford (1948). 1. The pruinose area along the claval suture is shorter than the interocular space (Fig. 6.407). ..........................................................................................................2 - The pruinose area along the claval suture is as long as or longer than the interocular space (Fig. 6.408). ..............................................................................4

331

Fig. 6.407 Ectemnostegella lundbladi male (left to right): tibia and pala, left and right parameres, and the apical segments of the abdomen in dorsal view. Based on Hungerford (1948).

Fig. 6.408 Ectemnostegella woytkowskii male: head in dorsal view (upper left), fore-leg (upper right), genital capsule (lower left), and apical segments of the abdomen in dorsal view (lower right). Based on Hungerford (1948). 2. (1) The length of the pruinose area along the claval suture is greater than the width of one compound eye. The male tibia is more than ¾ the length of the pala (Fig. 6.407). Length: 6.3 to 7.3 mm. .........................................................Ectemnostegella lundbladi Hungerford, 1948 (Peru).

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- The length of the pruinose area along the claval suture is less than the width of one compound eye (Fig. 6.409). ...........................................................................3

Fig. 6.409 Ectemnostegella jamesi male: head in dorsal view (upper left), inner surface of the fore-leg (upper right), apical portion of the genital capsule (lower left), and apical segments of the abdomen in dorsal view (lower right). Based on Hungerford (1948). 3. (2) There is an indistinct nodal furrow that divides the embolar groove into two nearly equal halves. The pruinose postnodal area measured along the costal margin is about twice as long as the meron. The pegs on the apical 1/5 of the male pala form two rows (Fig. 6.409). Length: c. 6.5 mm. ..............................................................Ectemnostegella jamesi Hungerford, 1948 (Bolivia). - There is a very distinct nodal furrow that divides the embolar groove into a relatively long anterior half and shorter posterior half. The pruinose postnodal area measured along the costal margin is about the same length as the meron (Fig. 6.410). Length: c. 5.3 to 6.1 mm. ................................................Ectemnostegella tumidacephala Hungerford, 1948 (Peru, Chile). 4. (1) The interocular space (synthlipsis) equals the width of one compound eye (Fig. 6.394). Length of the female: 5.7 to 6.2 mm. ............................................................Ectemnostegella venturii Hungerford, 1948 (Argentina). - The interocular space is greater than the width of one compound eye (Fig. 6.408). ...................................................................................................................5

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Fig. 6.410 Ectemnostegella tumidacephala male: head in dorsal view (upper left), strigil (lower left), fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.411 Ectemnostegella peruana: head in dorsal view (upper left), male foreleg (upper center), femur (upper right), male genital capsule (lower center), apical segments of the male abdomen in dorsal view (lower right), and apex of the female abdomen in ventral view (lower left). Based on Hungerford (1948). 5. (4) The greatest width of the shiny costal margin just anterior to the nodal furrow is equal to the postocular space, measured at the inner angle of the eye. The vertex of the male projects well anterior to the compound eyes and forms an angle slightly greater than 90° at the midline, when observed in dorsal view (Fig. 6.408). Length: 7.4 to 7.8 mm. ....................................................Ectemnostegella woytkowskii Hungerford, 1948 (Peru).

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- The greatest width of the shiny costal margin just anterior to the nodal furrow is equal to 1/2 of the postocular space, measured at the inner angle of the eye (Fig. 6.411). If the hemielytra are mostly shiny, then the head is evenly rounded along the anterior border between the compound eyes. .......................................6 6. (5) The anterior margin of the lateral lobe of the prothorax is nearly straight and longer than its width at the base. The vertex of the head is conically produced in both sexes. The pala of the male is short and broad (Fig. 6.411). Length: 5.9 to 6.8 mm. .............................................................Ectemnostegella peruana Jaczewski, 1933 (Peru). - Both anterior margins of the lateral lobe of the prothorax are tapering and almost as broad at the base as long, or the margins of the lobe are subparallel, extremely rounded, and somewhat truncate. In both sexes, the vertex is not conical, even if it is produced (Fig. 6.411). ..........................................................7

Fig. 6.412 Ectemnostegella pilosafrons male: head in dorsal (upper left) and lateral view (lower left), fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 7. (6) The face of the male is covered with hair-like setae; the face of the female is depressed and appears very hairy. The male lacks a stridular area at the base of the fore-femur (Fig. 6.412). Length: 7.0 to 7.6 mm. ......................................................Ectemnostegella pilosafrons Hungerford, 1948 (Peru). - The male has no more than a few hair-like setae on the face. The male usually has a stridular area at the base of the fore-femur (Fig. 6.413). There is no way yet reported to reliably distinguish the female of E. quechua from that of E. pilosafrons, but in the case of other females, if the face is depressed, it does not appear hairy, but if it appears hairy, it is not depressed. ......................................8

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Fig. 6.413 Ectemnostegella montana male: head in dorsal view (left), fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 8. (7) The femur of the fore-leg is at least 2.5 times as wide as the tibia, and its stridular area is wider than long (Fig. 6.413). There are 5 pegs in the upper distal row of the male pala, and 11 short and 5 long pegs in the lower row. The face of the female appears hairy but is not depressed. Length: 6.5 to 7.7 mm. .............................................................Ectemnostegella montana Lundblad, 1928 (Argentina). - The femur of the fore-leg of the male is no more than twice as wide as the tibia (Fig. 6.414). The face of the female is depressed. ......................................9 9. (8) The stridulatory organ is obviously large. There are 7 pegs in the upper distal row of the male pala, and 13 short and 8 long pegs in the lower row (Fig. 6.414). The hemielytra are entirely shiny except for a slightly rugose area at the base. The face of the female is depressed and hairy; it cannot yet be reliably distinguished from that of E. pilosafrons. Length: 7.1 to 8.9 mm. The specimens were taken in water bodies on open pampas 3,400 meters above sea level. The ground color is blackish, and there are 15 fine, irregular, transverse lines crossing the pronotum. ............................................................Ectemnostegella quechua Bachmann, 1961 (Argentina). - The stridular area is longer than wide (Fig. 6.347). There are 5 pegs in the upper distal row of the male pala, and 12 short and 4 or 5 long pegs in the lower row. The face of the female is depressed but lacks a dense coat of hair-like setae. Length: 6.7 to 7.0 mm. ........................................................Ectemnostegella stridulata Hungerford, 1948 (Peru, Argentina).

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Fig. 6.414 Ectemnostegella quechua: head in dorsal view (left), fore-leg of male (upper left center), metaxyphus (left center), strigil (upper center), outline of the lateral lobes of the prothorax of a male (lower center) and a female (lower right center), the apex of the male abdomen (right), and the male paramere (lower left center). Based on Bachmann (1961a). Key to the species of Centrocorisa in South America Information for the key was provided by Nieser (1969a). 1. The right paramere of the male has a thick knob. The left male paramere is broadly expanded at the apex. There are no tufts of hair-like setae on the corium (Fig. 6.415). Length: c. 7 to 8 mm. ............................................................Centrocorisa nigripennis (Fabricius, 1803) (North America, West Indies, Venezuela). Syn: Sigara nigripennis Fabricius, 1803; Corisae cubae Guérin-Menéville, 1843; Corisa irrorata Herrich-Schäffer, 1853; Corisa nigripennis Stål, 1868; Corisa cubae Uhler, 1894; Arctocorisa antiguensis Torre-Bueno, 1923. - The knob of the right paramere of the male is fairly small. The apex of the left male paramere is only slightly expanded (Fig. 6.395). There is a tuft or patch of hair-like setae on the corium of the female near the nodal furrow and a stripe of long setae on the clavus near the distal third of the claval suture. Length: c. 6.5 to 7.5 mm. ........................................................................Centrocorisa kollari (Fieber, 1851) (North and Central America, West Indies, Colombia, Ecuador, Peru, Bolivia, Paraguay, Argentina, Pernambuco, and Minas Gerais). Syn: Corisa kollari Fieber, 1851; Centrocorisa kollarii Hungerford, 1948.

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Fig. 6.415 Centrocorisa nigripennis male: tibia and pala (left), left (lower right) and right (upper right) paramere. Based on Nieser (1969a). Key to the species of Sigara in South America Information for the key was provided by Hungerford (1928b, 1948), (Jaczewski 1931a, b), Bachmann (1960, 1961a, 1966c. 1979a, 1986), and Nieser (1993b). One species known from Argentina belongs to the subgenus Aphelosigara. All others from South America belong to the subgenus Tropocorixa, found south of the tropics in the subtropical or temperate climate zones. Unfortunately, the females of some species cannot be distinguished using this key. 1. The hemielytra are partially hyaline. The apex of the tarsus of the middle leg does not extend beyond the apex of the hemielytra. The pala of the male has a basal carina. The sperm duct is relatively short and thick (Fig. 6.416). Length of male: 6.4 to 6.7 mm; female: 6.0 to 6.2 mm. ......................................................Sigara (Aphelosigara) tucma Bachmann, 1961 (Argentina). - The hemielytra are opaque. The sperm duct is long and thin (Fig. 6.417). ................................................................................subgenus Tropocorixa...........2 2. (1) The right male paramere is narrowed by a preapical invagination and then widened to an irregularly blunted apex. The space between the eyes across the vertex is wider than a compound eye. The infraocular width of the genae at the level of the hypo-ocular suture is clearly less than the diameter of the middle femur. The median length of the pronotum is about 1.5 times the length of the head. The pronotum is dark with six or seven regularly spaced yellow transverse stripes. The male pala has a continuous, arching row of

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approximately 25 pegs. The strigil is semicircular with about six irregularly arranged combs (Fig. 6.418). Length of male: c. 5.5 mm. The female has not been described. .............................................................................Sigara forciceps (Spinola, 1852) (Chile, Argentina?). Syn: Corixa forciceps Spinola, 1852; Corisa forciceps Spinola, 1852 (spelling change); Arctocorisa forciceps (Spinola, 1852); Arctocorixa forciceps (Spinola, 1852); Sigara rehi Jaczewski, 1931; Sigara argentiniensis chichinal Bachmann, 1960. Bachmann (1962) noted that the authors of major 20th century revisions of the South American Sigara species had not examined any of Spinola’s type specimens and based their descriptions of his species on specimens probably belonging to other species, most frequently a new species Bachmann named Sigara platensis. He declared Sigara forciceps to be a species inquirenda. In 1986, he expressed the opinion that Sigara argentiniensis chichinal and Sigara rehi are both junior synonyms of Sigara forciceps, but did not seem to formally propose a change in the nomenclature. Accepting his proposal as a formal change, this key assumes descriptions of its synonyms in the major revisions to be the only available descriptions of Sigara forciceps since the original description by Spinola was published in 1852. These descriptions are also sketchy due to a lack of undamaged specimens. - The right paramere of the male lacks a pre-apical invagination, or it is not irregularly blunt at the apex (Fig. 6.419). If not, then the infraocular width of the genae at the level of the hypo-ocular suture is very wide, equaling the diameter of the middle femur. ..............................................................................3

Fig. 6.416 Sigara tucma male: head (upper left), metaxyphus (lower left), foreleg showing the stridulatory structure (upper center), the pala of a different specimen (loower left center), apex of the abdomen (upper right), and right paramere (lower right). Based on Bachmann (1961a).

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Fig. 6.417 Sigara denseconscripta male: head in dorsal view (upper left), foreleg (upper right), genital capsule (lower left), and apex of the abdomen in dorsal view (lower right). Based on Hungerford (1948).

Fig. 6.418 Sigara forciceps male (left to right): pala, strigil, and right paramere. Based on Jaczewski (1931a), who described it under his synonym, Sigara rehi. 3. (2) The interocular space is approximately equal to the width of one eye (Fig. 6.417). ...................................................................................................................4 - The interocular space is clearly narrower than the width of one eye (Fig. 6.419). .................................................................................................................11 4. (3) The metaxyphus is obviously broader than long (Fig. 6.417). ...................5 - The metaxyphus is as long or longer than broad (Fig. 6.420). .........................7 5. (4) The claval suture has a pruinose area that is obviously longer than the post-nodal pruinose area. The length of the head is greater than the synthlipsis (Fig. 6.421). Length: 6.3 to 7.4 mm. Bressa and Papeschi (2007) reported the 2n chromosome number to be 24 (XY). ..........................................................................Sigara rubyae (Hungerford, 1928) (Argentina). Syn: Arctocorixa rubyi Hungerford, 1928; Sigara rubyi (Hungerford, 1928), error in Latin grammar.

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- The pruinose area on the claval suture is nearly equal to the post-nodal pruinose area (Fig. 6.417). ...................................................................................6

Fig. 6.419 Sigara platensis male: head (left), fore-leg (upper center), genital capsule (right), sheath for penis on the genital capsule and a right paramere (lower left center, above and below, respectively), and apex of the abdomen in dorsal view (lower center). Based on Hungerford (1948), who referred to it by its synonym, Sigara forciceps.

Fig. 6.420 Sigara trimaculata male: head in dorsal view (left), inner surface of the fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

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6. (5) The pala of the male is broadest at its base (Fig. 6.417). Length 6.7 to 8.0 mm. While the size may permit a probable identification, there is still no reliable way to distinguish the females of this species from those of the next. Bressa and Papeschi (2007) found the 2n chromosome number to be 24 (XY). .................................................................Sigara denseconscripta (Breddin, 1897) (Paraguay, Argentina, Uruguay, Bolivia). Syn: Corixa denseconscripta Breddin, 1897; Arctocorisa denseconscripta (Breddin, 1897); Arctocorixa denseconscripta (Breddin, 1897). - The pala of the male is uniform in width throughout most of its length (Fig. 6.422). Length 5.9 to 6.7 mm. .........................................................................Sigara schadei (Hungerford, 1928) (Paraguay, Argentina, Uruguay, Chile). Syn: Arctocorixa rubyi var. schadei Hungerford, 1928.

Fig. 6.421 Sigara rubyae male: head in dorsal view (upper left), fore-leg (upper right), genital capsule (lower left), and apical segments of the abdomen in dorsal view (lower right). Based on Hungerford (1948). 7. (4) Length: 7.1 to 8.2 mm. The middle femur is spinose. The pegs on the male pala form an even row, linear at the base and curving evenly apically (Fig. 6.420). ....................................................................Sigara trimaculata (Le Guillou, 1841) (Chile, Argentina, Rio Grande do Sul). Syn: Corixa trimaculata Le Guillou, 1841; Corisa maculata Guillon (alternate spelling by Fieber in 1851); Corixa maculata (spelling change); Arctocorixa maculata (catalogued by Kirkaldy and Torre-Bueno in 1909); Arctocorixa fazi Hungerford, 1928. - Length: about 5 to 6 mm. The middle femur is not spinose or armed only with small spines (Fig. 6.421). .....................................................................................8

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Fig. 6.422 Sigara schadei male: head in dorsal view (upper left), inner surface of the fore-leg (upper right), genital capsule (middle left) with another view of the penis sheath (lower left), and apical segments of the abdomen in dorsal view (lower right). Based on Hungerford (1948).

Fig. 6.423 Sigara femoridens male (left to right): fore-leg, apical segments of the abdomen in dorsal view, and genital capsule. Based on Hungerford (1948). 8. (7) The pruinose area on the claval suture is nearly equal to the post-nodal pruinose area. The ventral margin of the male fore-femur has a small projection (Fig. 6.423). Length: 6.3 to 6.7 mm. ......................................................................Sigara femoridens Hungerford, 1939 (Bolivia). - The pruinose area on the claval suture is longer than that on the post-nodal region. There is no small projection on the ventral margin of the male forefemur (Fig. 6.424). ...............................................................................................9

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Fig. 6.424 Sigara boliviensis male (left to right): fore-leg, apical segments of the abdomen in dorsal view, and genital capsule. Based on Hungerford (1948). 9. (8) The fore-femur of the male lacks a stridular area. Any small indentations near the midline of the apical abdominal tergites are simple and without medial protuberances (Fig. 6.424). Length: c. 5.5 mm. ......................................................................Sigara boliviensis Hungerford, 1939 (Bolivia). - There is a stridular area on the fore-femur of the male (Fig. 6.425). ..............10 10. (9) The posterolateral corners of the apical abdominal segments of the male are drawn out into slender spines (Fig. 6.396). Length of the male: 5.4 to 5.9 mm; female: 5.8 to 6.1 mm. ..................................................................................Sigara yala Bachmann, 1979 (Argentina). - The posterolateral corners of the apical abdominal segments of the male are triangular and somewhat wedge-shaped (Fig. 6.425). Length: c. 5 mm. .................................................................................Sigara czakii Jaczewski, 1927 (Paraná). Syn: Arctocorixa czakii (Jaczewski, 1927).

Fig. 6.425 Sigara czakii male: head in dorsal view (left), pala (upper left center), metaxyphus (lower left center), three views of the right paramere (upper center and right center), left paramere (lower right center), and apex of the abdomen in dorsal view (right). Based on Hungerford (1948).

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11. (3) The infraocular width of the genae at the level of the hypo-ocular suture, which is the short suture ventral to and ending at the margin of the compound eye, is as great as the diameter of the middle femur (Fig. 6.419). The right male paramere is narrowed by a preapical invagination. The seventh abdominal segment of the male is somewhat trapeziodal and bears a group of long setae. The metaxyphus is extremely short and wide. The right paramere of the male appears variable in shape because the distal lobe is thin and transparent, making it difficult to see in microscopic preparations. Bressa and Papeschi (2007) found the 2n chromosome number to be 24 (XY). The species in this couplet and several in subsequent couplets have been revised several times by Bachmann (1960, 1961a, 1966c. 1979a, 1985) but are still not clearly defined. ...........................................................................Sigara platensis Bachmann, 1962 (Argentina, Paraguay, Uruguay, Santa Catarina, Rio Grande do Sul). Syn: Sigara forciceps auctt. (nec Spinola, 1852). See Couplet 2. - The infraocular width of the genae at the level of the hypo-ocular suture is less than the diameter of the middle femur (Fig. 6.426). ..........................................12

Fig. 6.426 Sigara denseconscriptoidea male: head in dorsal view (upper left), inner surface of the fore-leg (upper center), genital capsule (lower left), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 12. (11) The metaxyphus is as long or longer than broad (Fig. 6.426). .............13 - The metaxyphus is obviously broader than long (Fig. 6.427). .......................15 13. (12) The vertex of the male is triangularly produced (Fig. 6.426). Length: 5.7 to 6.1 mm. ....................................................Sigara denseconscriptoidea (Hungerford, 1928) (Argentina, São Paulo, Espirito Santo). Syn: Arctocorixa denseconscriptoidea Hungerford, 1928; Sigara bahiensis Lundblad, 1929. - The vertex of the male is not produced (Fig. 6.427). There are six dark bands across the pronotal disc. .....................................................................................14

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Fig. 6.427 Sigara termasensis male: head in dorsal view (upper left), fore-leg (upper right), genital capsule (lower left), and apical segments of the abdomen in dorsal view (lower right). Based on Hungerford (1948).

Fig. 6.428 Sigara argentiniensis male: head in dorsal view (upper left), inner surface of the fore-leg (upper center), tibia and pala in dorsal view (center), genital capsule (lower left), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.429 Sigara vuriloche male (left to right): outline of head and eye in lateral view, strigil, and a paramere. Based on Bachmann (1959).

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14. (13) Length of male: c. 5.5 mm; female: c. 5.8 to 6.3 mm. The strigil is oval (Fig. 6.428). .................................................................Sigara argentiniensis Hungerford, 1948 (Argentina). - Length of male: c. 6.0 to 6.6 mm; female: c. 6.3 to 7.0 mm. There are six dark bands across the pronotal disc. The strigil is large and round (Fig. 6.429). Other than size, there is little to distinguish these two nominal species. ..........................................................................Sigara vuriloche Bachmann, 1960 (Argentina). Syn: Sigara argentiniensis vuriloche Bachmann, 1960. 15. (12) The disc of the pronotum is reduced laterally (Fig. 6.427). Length: 5.7 to 5.9 mm. ..................................................................Sigara termasensis (Hungerford, 1928) (Chile). Syn: Arctocorixa termasensis Hungerford, 1928. - The disc of the pronotum is not reduced laterally (Fig. 6.430). ......................16 16. (15) The abdomen of the male lacks a strigil (Fig. 6.430). Length: 4.2 to 4.7 mm. ...................................................................Sigara jensenhaarupi Jaczewski, 1927 (Argentina). Syn: Sigara jensen-haarupi Jaczewski, 1927. - The abdomen of the male has a strigil (Fig. 6.431). .......................................17

Fig. 6.430 Sigara jensenhaarupi male (left to right): head in dorsal view, pala (above), genital capsule (below), apical segments of the abdomen in dorsal view. Based on Hungerford (1948).

Fig. 6.431 Sigara roberti male (left to right): head in dorsal view, pala (above), metaxyphus (below), two views of the right paramere, and apex of the abdomen in dorsal view. Based on Hungerford (1948).

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17. (16) The fore-femur of the male lacks a stridular area (Fig. 6.431). ............18 - The fore-femur of the male has a stridular area (Fig. 6.432). .........................20

Fig. 6.432 Sigara dita male: head in dorsal view (left), inner surface of the foreleg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.433 Sigara hungerfordi male: head in dorsal view (left), inner surface of the fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 18. (17) The post-nodal pruinose area is shorter than that of the claval suture. The male fovea is poorly defined (Fig. 6.431). Length: 4.5 to 5.0 mm. .............................................................................Sigara roberti Hungerford, 1948 (São Paulo). - The post-nodal pruinose area is as long or slightly longer than the similar area along the claval suture. The male fovea is clearly defined (Fig. 6.433). ..........19

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19. (18) The male has a fringe of long, hair-like setae on the dorsolateral margin of the fovea (Fig. 6.433). Length: 5.8 to 6.25 mm. .......................................................................Sigara hungerfordi Jaczewski, 1927 (Argentina, Paraná, São Paulo, Santa Catarina). - The male lacks hair-like setae on the dorsolateral margin of the fovea (Fig. 6.434). Length: c. 7.1 mm. .....................................................................Sigara braziliensis Hungerford, 1948 (Rio de Janeiro).

Fig. 6.434 Sigara braziliensis male: head in dorsal view (left), inner surface of the fore-leg (upper center), genital capsule (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948).

Fig. 6.435 Sigara santiagiensis male: head in dorsal view (left), fore-leg (upper center), genital capsule (right), and apical segments of the abdomen in dorsal view (lower center). Based on Hungerford (1948).

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20. (17) The right margin of the male seventh abdominal segment is not angulate or produced (Fig. 6.435). ....................................................................................21 - The right margin of the male seventh abdominal segment is angulate or produced (Fig. 6.436). ........................................................................................22 21. (20) The median lobe of the seventh segment of the male abdomen has a fringe of hair-like setae (Fig. 6.432). Length: 4.2 to 5.4 mm. ....................................................................................Sigara dita Jaczewski, 1927 (Argentina, Paraná, São Paulo, Santa Catarina). Syn: Arctocorixa dita (Jaczewski, 1927). - The median lobe of the seventh segment of the male abdomen does not have a fringe of hair-like setae (Fig. 6.435). Length: 5.8 to 6.2 mm. ................................................................Sigara santiagiensis (Hungerford, 1928) (Chile, Argentina, Falkland Islands alias Malvinas). Syn: Arctocorixa santiagiensis Hungerford, 1928.

Fig. 6.436 Sigara egbertae male: head in dorsal view (upper left), inner surface of the fore-leg (upper center), genital capsule (lower left), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). 22. (20) The right margin of the male seventh abdominal segment forms two angles (Fig. 6.436). Length: c. 5.9 mm. ..........................................................................Sigara egbertae Hungerford, 1948 (Argentina, Uruguay, Paraguay, Bolivia, Paraná, Minas Gerais). - The right margin of the male seventh abdominal segment does not form two angles (Fig. 6.437). Length 5.1 to 6.3 mm. ......................................................................Sigara chrostowskii Jaczewski, 1927 (Argentina, Paraguay, Uruguay, São Paulo, Paraná, Santa Catarina. Rio Grande do Sul). Syn: Arctocorixa chrostowskii (Jaczewski, 1927); Arctocorixa chrostowskii var. brachypala Hungerford, 1928; Arctocorixa chrostowskii var. townsendi Hungerford, 1928; Sigara townsendi (Hungerford, 1928); Arctocorixa hosfordi Hungerford, 1928; Sigara hosfordae (Hungerford, 1928); Sigara (Tropocorixa) brachypala Hungerford, 1948.

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Fig. 6.437 Sigara chrostowskii (male): head in dorsal view (left), fore-leg (upper center), genital capsule with another view of the apex of the paramere above it (lower center), and apical segments of the abdomen in dorsal view (right). Based on Hungerford (1948). Key to the species of adult male Neosigara in South America Information for the key was provided by Hungerford (1948), Nieser and Padilla (1992), Padilla and Nieser (1994), and Tinerella and Polhemus (2006). 1. The asymmetrical modifications of the abdomen are sinestral, or, if dextral, then the strigil is absent or nearly absent (Fig. 6.438). ........................................2 - The asymmetrical modifications of the abdomen are dextral. The strigil is well developed (Fig. 6.439). .................................................................................3

Fig. 6.438 Neosigara griffini male (left to right): fore-leg, genital capsule, and apical abdominal segments. Based on Hungerford (1948).

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Fig. 6.439 Neosigara columbiensis male: habitus (above) and (below, left to right): fore-leg, apical segments of the abdomen in dorsal view, ventral surface of the femur (below), and genital capsule (above). Based on Hungerford (1948). 2. (1) A strigil is absent. The right paramere is broad apically (Fig. 6.438). ........................................................................Neosigara griffini (Kirkaldy, 1899) (Ecuador, Peru). Syn: Corixa (Callicorixa) Griffini Kirkaldy, 1899; Sigara griffini (Kirkaldy, 1899); Corixa williamsi Hungerford, 1928. - The strigil is well developed. The right paramere is narrow apically (Fig. 6.440). Length: 4.5 to 6.0 mm. ..........................................................Neosigara aristera Nieser and Padilla, 1992 (Colombia).

Fig. 6.440: Neosigara aristera male (left to right): fore-leg, sixth and seventh abdominal segments, and the right paramere. Based on Nieser and Padilla (1992).

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3. (1) The deep, broad, and densely pilose frontal fovea reaches midway between the eyes (Fig. 6.439). ..............................................................................4 - The shallow, sparcely pilose fovea reaches only to the base of the eyes. The median lobe of the seventh abdominal segment has a sinuate posterior margin (Fig. 6.441). ..........................................................................................................5

Fig. 6.441 Neosigara murilloi male (left to right): fore-leg, apical segments of the abdomen in dorsal view, genital capsule (above), and ventral surface of the hind femur (below). Based on Hungerford (1948). 4. (3) The median lobe of the seventh abdominal segment is bilobate with the left part broad and evenly rounded caudally. The glabrous ventral surface of the hind femur bears 20 to 25 spines (Fig. 6.439). Length: 5.1 to 6.4 mm. The female reaches 6.7 mm. ................................................................Neosigara columbiensis Lundblad, 1928 (Colombia). - The median lobe of the seventh abdominal segment is not divided into separate lobes, and the left part is acutely pointed at the apex. The glabrous ventral surface of the hind femur bears 3 or 4 tiny spines (Fig. 6.398). ..................................................Neosigara paramo Tinerella and Polhemus, 2006 (Colombia).

Fig. 6.442 Neosigara sterea male (left to right): fore-leg, sixth and seventh abdominal segments, and right paramere. Based on Nieser and Padilla (1992).

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Fig. 6.443 Neosigara akanthinomeros male (left to right): fore-leg, sixth and seventh abdominal segments in dorsal view, and right paramere. Based on Padilla Gil and Nieser (1994). 5. (3) The glabrous portion of the hind femur bears about 5 spinules (Fig. 6.441). Length: 4.6 to 6.0 mm. The female reaches 6.9 mm. .....................................................................Neosigara murilloi Hungerford, 1948 (Colombia). - The glabrous portion of the hind femur bears more than 15 spinules (Fig. 6.442). ...................................................................................................................6 6. (5) The glabrous portion of the hind femur bears about 16 spinules. The right paramere is 0.75 mm long. The left side of the median lobe of the seventh abdominal tergite of the males is somewhat rounded rather than acutely pointed (Fig. 6.442). Length: 4.8 to 6.9 mm. .............................................................Neosigara sterea Nieser and Padilla, 1992 (Colombia). - The glabrous portion of the hind femur bears about 25 spinules. The right paramere is 0.5 mm long (Fig. 6.443). Length: 4.4 to 6.1 mm. .............................................Neosigara akanthinomeros Padilla and Nieser, 1994 (Colombia). Key to the species of known fifth instar Neosigara and Orocorixa larvae in South America Information for the key was provided by Padilla and Nieser (1994). 1. The pala is twice as long as the fore-tibia. The mesonotum has a medioposterior pilose band. The scent glands on the fourth and fifth tergites are not clearly evident (Fig. 6.444). ...................................................Orocorixa makrocheira Nieser and Padilla, 1992 (Colombia).

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- The pala is only 1.4 to 1.6 times as long as the fore-tibia. Pilosity on the mesonotum is restricted to the anterior half. The scent glands on the fourth and fifth tergites are heart-shaped and clearly evident (Fig. 6.445). ..........................2

Fig. 6.444 Orocorixa makrocheira (left to right): egg, fifth larval instar of an individual that will be capable of flight as an adult and one that will not. Based on Padilla and Nieser (1994).

Fig. 6.445 Neosigara aristera (left to right): egg, head of fifth larval instar in anterior view, habitus of fifth larval instar without appendages, and hind femur of the fifth instar. Based on Padilla and Nieser (1994). 2. (1) The ventral glabrous portion of the hind femur bears at least 20 spinules. The length of the body measured along its mid-line is about 2.4 to 2.9 times the maximum width (Fig. 6.445). ...............................................................................3 - The ventral glabrous portion of the hind femur bears no more than 17 spinules. The median length of the body is about 2.0 to 2.1 times the maximum width (Fig. 6.446). ..........................................................................................................4 3. (2) The ventral glabrous portion of the hind femur bears 20 to 25 spinules, which are arranged in a regular pattern (Fig. 6.445). ..........................................................Neosigara aristera Nieser and Padilla, 1992 (Colombia). - The ventral glabrous portion of the hind femur bears more than 25 spinules, which are arranged irregularly (Fig. 6.447). .............................................Neosigara akanthinomeros Padilla and Nieser, 1994 (Colombia).

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Fig. 6.446 Neosigara sterea (left to right): egg, habitus of fifth larval instar in dorsal view, mesothorax and metathorax, and hind femur of the fifth instar. Based on Padilla and Nieser (1994). 4. (2) The ventral glabrous portion of the hind femur bears 15 to 17 spinules. The abdomen is very wide (Fig. 6.446). .............................................................Neosigara sterea Nieser and Padilla, 1992 (Colombia). - The ventral glabrous portion of the hind femur bears no more than 10 spinules. In dorsal view, the anterior margin of the vertex of the head appears evenly rounded (Fig. 6.448). ............................................................................................5

Fig. 6.447 Neosigara akanthinomeros fifth larval instar (left to right): head in anterior view, habitus without appendages in dorsal view, and hind femur. Based on Padilla and Nieser (1994).

Fig. 6.448 Neosigara columbiensis: egg (left) and habitus without appendages in dorsal view (right). Based on Padilla and Nieser (1994).

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Fig. 6.449 Neosigara murilloi: egg (left) and habitus without appendages in dorsal view (right). Based on Padilla and Nieser (1994). 5. (4) The ventral glabrous portion of the hind femur bears about 10 spinules. The wing buds are aligned parallel to the body axis (Fig. 6.448). ................................................................Neosigara columbiensis Lundblad, 1928 (Colombia). - The ventral glabrous portion of the hind femur bears no more than 5 spinules. The wing buds diverge posteriad (Fig. 6.449). .....................................................................Neosigara murilloi Hungerford, 1948 (Colombia).

Nepidae Key to the genera of Nepidae The information for the key was provided by De Carlo (1964a), Lansbury (1974a), Nieser (1975), and Froeschner (1981). 1. The body is flattened, and the parasternites of the abdomen are visible. The head is distinctly narrower than the pronotum and partly concealed by its anteriolateral angles (Fig. 6.450). .........................................................................2 - The body is subcylindrical and lacks visible parasternites. The head is not partially concealed by the pronotum; the head and pronotum are subequal in width (Fig. 6.451). ................................................................................................3 2. (1) The body is elongate; its ratio of length to width is 5:1 to 6:1 (Fig. 6.450). ........................................................................................Curicta Stål, 1861..p. 359 - The body is ovoid; its ratio of length to width is not more than 3:1. The respiratory tube is broad and strap-like, distinguishing this genus from the Holarctic genus Nepa Linnaeus (Fig. 6.452). ...........................................................................Telmatotrephes Stål, 1854..p. 368

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Fig. 6.450 Curicta tibialis male: habitus in dorsal view (upper left), fore-leg in anterior view (lower left), male operculum (upper right), and parameres (lower right). Based on Keffer (1997).

Fig. 6.451 Ranatra (Amphischizops) compressicollis: habitus in dorsal view (upper middle left to center), head and thorax in dorsal (upper left) and lateral view (lower middle left), ventral view of metathorax (upper right), and apex of the abdomen with the respiratory siphon in dorsal (lower center to right) and lateral view (lower left to center). Based on Montandon (1898).

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3. (1) The eyes are set on lateral extensions of the head, which are not concealed by dense patches of hairs (Fig. 6.451). Length: 43 mm to the end of the genitalia; 64 mm with appendages. ........................................................... Ranatra (Amphischizops) Montandon, 1903 Regarded by some authors as a genus, Amphischizops is treated here as a subgenus of Ranatra. It is represented in South America by Ranatra (Amphischizops) compressicollis (Montandon, 1898), known only from Venezuela. Lansbury (1974b) reduced Amphischizops to a subgenus of Ranatra, but his opinion has not been universally accepted (Nieser, 1975). The species appears in the key to Ranatra. - The eyes are not set on lateral extensions of the head (Fig. 6.453). ..............................................................................Ranatra Fabricius, 1790..p. 370 All South American species of Ranatra, other than R. (Amphischizops) compressicollis, should key out here.

Fig. 6.452 Telmatotrephes grandicollis: male (upper left) and female antenna (upper center); male head and pronotum in dorsal (upper middle left) and lateral view (lower middle left); male thorax in ventral view (above center, below female antenna); visible part of male abdomen in ventral view (upper right); male fore-leg (lower left); male genital capsule in dorsal (lower right center), ventral (lower right), and lateral view (lower center); female operculum (lower left center, right of head and thorax in lateral view). Based on Lansbury (1972).

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Fig. 6.453 Ranatra heydeni male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946). Key to the species of Curicta in South America Information for the key was provided by Kuitert (1949a), De Carlo (1951a, 1960a, 1967a, b), Nieser (1975), and Keffer (1997, 1998, 1999). The species Curicta longimanus and C. lenti cannot presently be identified with certainty. Few specimens have been described, and Keffer (1997) was unable to locate a sufficient number to examine to elucidate their status in the latest review of this genus. In the key, C. longimanus is shown as a probable junior synonym of C. volxemi, while C. lenti appears in a couplet together with C. montei, from which it presently cannot be distinguished with certainty. 1. The sulcus of the fore-femur bears one tooth on its anteroventral edge (Fig. 6.454). ...................................................................................................................2 - The sulcus of the fore-femur bears two teeth, one at each end (Fig. 6.455). ....4 2. (1) The single tooth on the fore-femur is truncate and not prominent. The pronutum lacks carinae. The posterior width of the pronotum is subequal to its median length (Fig. 6.454). Length of male: 19.5 mm. The female has not been described. ................................................................................Curicta bilobata Kuitert, 1949 (Espirito Santo). - The single tooth on the fore-femur is prominent and pointed at the apex (Fig. 6.456). ...................................................................................................................3

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Fig. 6.454 Curicta bilobata male: habitus without appendages in dorsal view (upper left), fore-leg in anterior view (lower left), distal fifth of the fore-femur in dorsal view showing the tubercles in the sulcus (lower right), operculum (upper right), and a paramere (middle right). Based on Keffer (1997).

Fig. 6.455 Curicta peruviana (left to right): antenna, outline of the pronotum, apical segments of the fore-leg, and paramere. Based on De Carlo (1960a).

Fig. 6.456 Curicta pelleranoi male: habitus (upper left), fore-femur in anterior (lower left) and posterior view (lower right), posterolateral metasternal process (center), operculum (upper right), and parameres (middle right). Based on Keffer (1997).

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3. (2) Distal to the prominent tooth, the fore-femur is only moderately arched. The total body length is only about 13 to 15.5 mm long. The respiratory tube is 8 mm long (Fig. 6.456). ..........................................................................Curicta pelleranoi De Carlo, 1951 (Argentina, São Paulo). - Distal to the prominent tooth, the fore-femur is remarkably arched (Fig. 6.457). Length: 13 to 16 mm. Siphon length: 6.2 to 9.0 mm. ...........................................................................Curicta bonaerensis (Berg, 1879) (Argentina, Uruguay). Syn: Helotenthes bonäerensis Berg, 1879; Nepa bonaerensis von Ferrari, 1888; Nepoidea falloui Martin, 1898; Curicta falloui Montandon, 1903.

Fig. 6.457 Curicta bonaerensis male: habitus in dorsal view (upper left), foreleg in anterior view (lower left), ventral view of the apical part of the fore-femur (lower right), operculum (upper right), posterolateral metasternal process (right center), and parameres of three specimens (middle right). Based on Keffer (1997). 4. (1) On the dorsal side of the last abdominal segment of the female, there are two well-developed carinae arched dorsomediad to form a dome-like structure. The teeth on the fore-femur are located close to its mid-length: c. 47% of the distance from the base to the apex (Fig. 6.458). Length of female: 16.1 mm. Siphon length: 8.5 mm. Color: dark brown with yellowish markings. Only one specimen, a female, has been described. ..........................................................................Curicta johnpolhemi Keffer, 1999 (Minas Gerais). - The posterior abdominal segment lacks two carinae on the dorsal surface. Two Central American species have carinae on the dorsal side of the last abdominal segment, but these are not oriented to form a dome-like structure (Fig. 6.455). The length is usually at least 17 mm. .............................................5

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Fig. 6.458 Curicta johnpolhemi female: habitus (upper left), fore-femur (right center), and the apical segment of the abdomen in dorsal (lower left) and lateral view (right). Based on Keffer (1999) 5. (4) The teeth on the fore-femur are located in the proximal half, and the distance from the base of the fore-femur to the apex of the anteroventral tooth is no more than 45% of the length of the fore-femur. The length of the respiratory appendages is about 75% of the total length. The process on the preapical antenna segment is cone-shaped with a rounded apex (Fig. 6.455). Length: 22 to 26.5 mm. Color: grayish black with tibiae yellowish brown mottled with reddish brown. There is a broad, dark band around the middle of the fore-tibia. .............................................................................Curicta peruviana Kuitert, 1949 (Peru). - The teeth on the fore-femur are usually located in the distal half, and the distance from the base of the fore-femur to the apex of the anteroventral tooth is more than 45% of the length of the fore-femur (Fig. 6.459). ...............................6 6. (5) The posterior half of the pronotum is remarkably swollen (Fig. 6.459). The parasternum ends in a broad process, which extends posteriad beyond the posterior border of the operculum (Fig. 6.459). ..............................................................................Curicta grandis De Carlo, 1951 (Surinam, Paraguay, Bolivia, Argentina). Syn: Curicta paraguayensis De Carlo, 1951; Curicta borellii De Carlo, 1951 (nec Montandon). - The posterior half of the pronotum is not notably swollen. The parasternum ends in a finger-like or thumb-like process, which sometimes extends posteriad beyond the posterior border of the operculum (Fig. 6.460). ................................7

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7. (6) The lateral margin of the sulcus in the fore-femur has a distinct distal notch. The teeth in the sulcus of the fore-femur are obviously large (Fig. 6.450). Length: 26 to 32 mm. ................................................................................Curicta tibialis (Martin, 1898) (Paraguay, Uruguay, Rio Grande do Sul). Syn: Nepoidea tibialis Martin, 1898; Curicta suspecta Montandon, 1903; Curicta beckeri De Carlo, 1956. - A distal notch in the lateral margin of the sulcus in the fore-femur is absent or present only distally (Fig. 6.460). ........................................................................8

Fig. 6.459 Curicta grandis: habitus in dorsal view (upper middle left), head and prothorax in lateral view (lower middle left), fore-leg in anterior view (center below habitus), antennae of three specimens (upper left), male operculum (upper center), apical abdominal segment in ventral (upper right) and lateral view (middle right), and the male parameres of six different specimens (lower row). Based on Keffer (1997).

8. (7) There are no teeth on the sulcus of the fore-femur. Along the mid-line of the vertex of the head, there is always a prominent carina (Fig. 6.460). .............9 - Teeth are present on the sulcus of the fore-femur (Fig. 6.461). ......................10

Fig. 6.460 Curicta carinata (left to right): antenna, outline of the pronotum, apical segments of the fore-leg, and paramere. Based on De Carlo (1960a).

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Fig. 6.461 Curicta granulosa (left to right): antenna, outline of the pronotum, apical segments of the fore-leg, and paramere. Based on De Carlo (1960a), who referred to it as Curicta venezolana.

Fig. 6.462 Curicta decarloi: habitus in dorsal view (upper left), fore-leg in anterior view (lower left), ventrolateral view of the apex of the fore-femur (lower right), and apex of the posteriormost abdominal segment in lateral view (upper right). Based on Keffer (1997). 9. (8) The annulus at the midlength of the fore-tibia is dark. The length of the respiratory appendages is less than about 70% of the total length. There is a pair of well-developed, elevated, longitudinal carinae on the mesosternum. The dorsal surface is covered with granulations, especially dense on the pronotal ridges and margins of the hemielytra. The process on the preapical segment of the antenna is long and finger-like (Fig. 6.460). The sternite on the distal segment of the abdomen ends in finger-like processes. Length of male: 17.5 to 21 mm; female: 17 to 25 mm. Length of respitatory tube: 11.5 to 17 mm. ................................................................................Curicta carinata Kuitert, 1949 (Central America, Trinidad, Colombia, Ecuador, Peru, Bolivia, Paraguay, Argentina). Syn: Curicta dilatata De Carlo, 1951. - The insect is dark, and the annulus at the mid-length of the fore-tibia is faint. The length of the respiratory appendages is more than about 80% of the total length. The carinae on the mesosternum are not obviously elevated. The dorsal surface is not covered with obvious granulations (Fig. 6.462). The sternite on

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the distal segment of the abdomen ends in inflated processes. Length of female: c. 25.5 mm. Length of respitatory tube: c. 22 mm. The male has not been described. .................................................................................Curicta decarloi Keffer, 1997 (Pernambuco). 10. (8) Dense dark granulations cover the pronotum and are especially dense on the ridges and humeral lobes (Fig. 6.461). .........................................................11 - The pronotum is not densely covered by dark granulations (Fig. 6.463). ......12 11. (10) The carina on the vertex is usually strongly developed. The fore-tibia usually has a dark annulus at mid-length, which is occasionally vague, especially in females. On the pronotum, the longitudinal ridges are usually carinate, and the sulci, deep. There is usually a pair of broad, parallel, longitudinal, tomentose bands on the mesosternum (Fig. 6.461). Length of the male: 18 to 22 mm; female: 20 to 25 mm. Length of siphon: 12 to 20 mm. ..........................................................................Curicta granulosa De Carlo, 1951 (Colombia, Venezuela, Peru, Paraguay, Bolivia, Pará, Amazonas, Paraiba, Bahia, Goiás, Minas Gerais, São Paulo, Mato Grosso, Mato Grosso do Sul, Rondônia, Acre). Syn: Curicta venezolana De Carlo, 1966; Curicta intermedia Roback and Nieser, 1974 (nec Martin). - A carina on the vertex is sometimes visible but never strongly developed. The fore-tibia usually lacks a dark annulus at mid-length, but if one is visible, it is interrupted and does not completely surround the tibia. On the pronotum, the longitudinal ridges are usually subcarinate or undeveloped, and the sulci, shallow. There is usually a pair of oblique tomentose bands on the mesosternum, which arise where the posterior margin of the mesosternum joins the metacoxae and extend only half the length of the mesosternum (Fig. 6.331). Length of the male: 21 to 26 mm; female: 24 to 30 mm. ..........................................................................Curicta borellii Montandon, 1903 (Argentina, Paraguay, Acre). Syn: Curicta riggii De Carlo, 1951; Curicta dureti De Carlo, 1951; Curicta carinata De Carlo, 1951 (nec Kuitert); Curicta bachmanni De Carlo, 1956; Curicta sanmartini De Carlo, 1956. 12. (10) Males are 24 to 27 mm long but usually longer than 26 mm. Females are 24.5 to 30.5 mm long but usually longer than 29 mm. The legs appear very long, so that the length of the fore-femur always exceeds the length of the pronotum measured along the lateral margin by more than 1 mm (Fig. 6.463). .......................................................................Curicta volxemi (Montandon, 1895) (Colombia, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Rio Grande do Sul; Mato Grosso do Sul). Syn: Nepoidea volxemi Montandon, 1895; Curicta schoutendeni Montandon, 1909; Curicta brasiliensis De Carlo, 1951; Curicta femoralis Roback and Nieser, 1974; probably also Curicta longimanus De Carlo, 1951. - Males are equal to or shorter than 26 mm, and females are equal to or shorter than 27 mm. The legs are not noticeably long, so the length of the fore-femur only exceeds the length of the pronotum measured along the lateral margin by less than 1 mm (Fig. 6.464). ...............................................................................13

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Fig. 6.463 Curicta volxemi: habitus in dorsal view (upper left), fore-leg in anterior view (lower left), apical abdominal segment in ventral (upper right) and lateral view (lower right), and the male parameres of seven different specimens (center, top to bottom). Based on Keffer (1997).

Fig. 6.464 Curicta intermedia male: habitus in dorsal view without the breathing tube (upper left), ventral view of the posterolateral metasternal process (lower left), fore-leg in anterior view (lower center), and apical abdominal segment in lateral view (right). Based on Keffer (1997). 13. (12) At the posterior end of each parasternum of the last abdominal segment is a finger-like but not thumb-like process. The vertex has a well-developed carina (Fig. 6.464). Length of male: c. 19.7 mm; female presumed to belong to this species: 21.8 mm. Length of respiratory tube of presumed female specimen:

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16 mm. The species is known from two specimens, although one of them, the female, was only presumptively assigned to this species. ..........................................................................Curicta intermedia (Martin, 1898) (Colombia). Syn: Nepoidea intermedia Martin, 1898. - At the posterior end of the parasternum of the last abdominal segment is a large, thumb-like process. The vertex usually lacks a well-developed carina. The paramere is hooked almost 180° near the apex (Fig. 6.465). ......................14

Fig. 6.465 Curicta montei (above, left to right): antenna, outline of the pronotum in dorsal view, fore-leg, and paramere, and (below, left to right): antenna of a different specimen somewhat enlarged, posterior margins of the pronotum and metasternum, and a different view of a paramere. The upper row is based on De Carlo (1960a), and the lower row on Nieser (1975), who referred to the species as Curicta doesburgi. 14. (13) Length of male: 18 to 23 mm; female: 20 to 27 mm. Length of siphon: 14 to 24 mm. The color is mainly dark brown with only the apical 2/3 of the anterior tibiae light yellow. There are three tooth-like plates along the posterior margin of the metasternum (Fig. 6.465). .............................................................................. Curicta montei De Carlo, 1960 (Venezuela, Surinam, Guyana, French Guiana, Maranhão, Ceará, Pernambuco, Minas Gerais). Syn: Curicta doesburgi De Carlo, 1967. - Length of male: c. 25 mm; female: 25 to 25.5 mm. Length of siphon: 19.5 mm. Aside from the slightly greater length of the male, existing information does not include any single feature that can be used to positively distinguish the two species in this couplet. ...................................................................................Curicta lenti De Carlo, 1951 (Rio de Janeiro). The two species in this couplet may be synonyms (Keffer, 1997), and if this is so, the valid name would be C. lenti. A better description of C. lenti is urgently needed, and studies of a much larger series of specimens will be necessary to determine the degree of intraspecific variability of both nominal species.

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Key to the species of Telmatotrephes in South America Information for the key was provided by Kuitert (1949a), De Carlo (1956), and Lansbury (1972). 1. The total body length, excluding the siphon, is more than 30 mm. The posterolateral lobes of the metasternum extend beyond the hind margin of the second sternite. The lateral margins of the posterior third of the hemielytra are not concave (Fig. 6.466). Length: 30 to 31 mm. Maximum body width: 14.5 mm. Siphon length: c. 6 mm. A description of a female is not available. ..................................................................Telmatotrephes sculpticollis Stål, 1854 (Colombia). - The total body length, excluding the siphon, is less than 28 mm. The posterolateral lobes of the metasternum do not extend beyond the hind margin of the second sternite (Fig. 6.467). ...........................................................................2

Fig. 6.466 Telmatotrephes sculpticollis male: antenna (upper left), head and pronotum in dorsal (middle left) and lateral view (upper center), prothorax in ventral view (left of center), meso and metathorax in ventral view (right of center), operculum (lower middle right), fore-coxa (upper middle right), fore-leg (upper right), genital capsule in lateral view (lower left), paramere (lower center). Based on Lansbury (1972). 2. (1) The siphon is less than half as long as the operculum. The maximum body width is less than half the length (Fig. 6.468). Length: 25 mm. ...............................…...........................Telmatotrephes carvalhoi De Carlo, 1956 (Rio de Janeiro).

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- The siphon is as long as the operculum. The maximum body width is about half the length (Fig. 6.467). ..................................................................................3 3. The length is less than 23 mm. The interocular space is much less than three times the width of an eye (Fig. 6.467). Only the female has been described. ........................................................Telmatotrephes ecuadorensis Lansbury, 1972 (Ecuador). - The length is 25 to 27 mm. The interocular space is about three times the width of an eye. The lateral margin of the posterior third of the hemielytra is concave. The body and legs bear numerous clusters of small hairs (Fig. 6.452). ..............................................................Telmatotrephes grandicollis Kuitert, 1949 (Peru, Amazonas). Syn: Telmatotrephes brasiliensis De Carlo, 1956.

Fig. 6.467 Telmatotrephes ecuadorensis female: antenna (lower left center), head and pronotum in dorsal (upper left) and lateral view (lower left), thorax in ventral view (upper left center), fore-leg (right center), operculum (upper right), fore-coxa (lower right). Based on Lansbury (1972).

Fig. 6.468 Telmatotrephes carvalhoi male (left to right): habitus without appendages in dorsal view, fore-leg, and apical part of abdomen in ventral view. Based on De Carlo (1956).

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Key to the species of Ranatra in South America Information for the key was provided by De Carlo (1946, 1950a, 1963b, 1964a, 1973), Kuitert (1949b), Drake (1949b), Nieser (1975), and Nieser and Burmeister (1998). The key lacks information on several species, as noted, because the descriptions were poor or specimens of both sexes are not available. There is not enough information about the males of Ranatra signoreti, R. oliveiracesari, R. unidentata, R. ecuadoriensis, and R. mixta or the females of R. brasiliensis, R. ornitheia, R. sattleri, and R. lenti to include them in the key. 1. The eyes are set on lateral extensions of the head, which are not concealed by dense patches of hairs (Fig. 6.451). Length of male: 37 mm; female: 43 mm. .................................Ranatra (Amphischizops) compressicollis Montandon, 1898 (Venezuela). This species was placed in a separate genus by Montandon (1903), but Lansbury (1974b) returned it to Ranatra. The taxonomic position of Amphischizops remains to find a consensus. - The eyes are not set on lateral extensions of the head (Fig. 6.469), or, if they are, the extensions are obscured by dense, hairy patches (see R. macrophthalma, Couplets 25 and 40), and the male is usually longer. ...........................................2 2. (1) The second segment of the antenna has a relatively small lateral process, which is, at most, barely longer than half the length of the third segment and sometimes reduced to a barely detectable vestige (Fig. 6.469). ...........................3 - The second segment of the antenna has a long lateral process, which is nearly equal in length to the third antennal segment (Fig. 6.470). ..................................9

Fig. 6.469 Ranatra segrega male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

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Fig. 6.470 Ranatra tuberculifrons male (left to right): antenna, fore-leg, and paramere. Based on De Carlo (1967c). 3. (2) The third antennal segment is well developed and clearly longer than the second. The apical portion of the fore-femur is abruptly and strongly arched (Fig. 6.471). Length of female: 42.5 mm. The femora lack dark bands. ...............................................................................Ranatra montei De Carlo, 1946 (Minas Gerais). Described from a single specimen, catalog No. 47173 in the Museo Argentino de Ciencias Naturales. - The third antennal segment is subequal to or shorter than the second (Fig. 6.469). ...................................................................................................................4

Fig. 6.471 Ranatra montei male: habitus in dorsal view (middle), antenna (upper left), and fore-tibia and tarsus (lower left). Based on De Carlo (1946). 4. (3) The apical portion of the fore-femur is moderately arched. The length of the second antenna segment between its junctions with the first and third is somewhat more than half the length of the third segment and subequal to its lateral process. The width of an eye is about 1.4 times the width of the head between the eyes.

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The male paramere has a short subapical tooth that is wide at its base (Fig. 6.472). The ratio of the hind femur of the female to the total length is about 20:45, and the middle femur nearly reaches the apex of the abdomen. The female operculum extends well beyond the apex of the abdomen. Length of male: c. 40 mm; female: c. 45 mm. The respiratory tubes of the specimens described measured 36 mm in the male and 39 mm in the female. The middle and hind legs display distinct pale and dark rings. .........................................................................Ranatra robusta Montandon, 1905 (São Paulo, Amazonas, Pará). - The apical portion of the fore-femur is no more than slightly and evenly arched (Fig. 6.469). The proportions of the antennal segments are different. ................5

Fig. 6.472 Ranatra robusta male: habitus in dorsal view (middle), antenna (upper left), fore-tibia and tarsus (below). Based on De Carlo (1946).

Fig. 6.473 Ranatra wagneri: antenna of a male with the second and third segments separate and distinct (upper left) and another with those segments fused (upper left center), habitus in lateral view (middle left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

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5. (4) The second and third antennal segments appear completely fused or only slightly separated, leaving scarcely a trace of the lateral process on the second segment (Fig. 6.473). Body length of male: 29 to 29.5 mm; female: 33 to 35.5 mm. Length of respiratory siphon of male: 14 to 14.5 mm; female: 14.5 to 15 mm. ........................................................................Ranatra wagneri Hungerford, 1929 (Argentina, Santa Catarina, Mato Grosso). - The second segment of the antenna has a lateral process not at all fused with the third segment (Fig. 6.469). .............................................................................6

Fig. 6.474 Ranatra brevicauda male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

Fig. 6.475 Ranatra machrisi (upper row, left to right): antenna of female, head and anterior part of the thorax of a female in lateral view, male paramere, and (upper middle row, left and right): fore-leg and apex of the paramere of a male enlarged, and (lower middle row, left and right): tibia and tarsus folded against the femur and the genital capsule of a male, and (lower row, left and right): genital operculum of a female and the sclerites supporting the male genital capsule. Based on Nieser and Burmeister (1998).

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6. (5) The process on the second antenna segment is barely more than a swelling, and the second segment, measured along its main axis, is longer than the third (Fig. 6.474). ....................................................................Ranatra brevicauda Montandon, 1905 (Argentina, Mato Grosso). - The process on the second antenna segment is well developed but significantly shorter than the third segment (Fig. 6.469). .........................................................7 7. (6) The process on the second antenna segment is well developed and digitiform but only slightly longer than half of the third segment. The basal part of the second segment is shorter than the third segment, but the combined length of the basal part of the second segment and its lateral process is somewhat greater than the length of the third (Fig. 6.475). Length: 37 to 41 mm; length of siphon: 25.5 to 27.0 mm. ......................................................Ranatra machrisi Nieser and Burmeister, 1998 (Goiás). - The process on the second antenna segment is bluntly conical, narrowing toward its rounded apex, and the length of the second segment measured along its main axis is subequal to the length of the third (Fig. 6.469). ..........................8

Fig. 6.476 Ranatra flokata male (upper row, left to right): antenna, head and anterior part of the thorax in lateral view, paramere, and (middle row, left and right): fore-leg and apex of the paramere enlarged, and (lower row, left to right): tibia and tarsus folded against the femur, sclerites supporting the genital capsule, and the genital capsule. Based on Nieser and Burmeister (1998). 8. (7) The siphon does not bear a strong fringe of setae, and setae at the distal end of the siphon, if any, are not twice as long as the width of the siphon. The male paramere is not strongly sigmoid (Fig. 6.469). .........................................................................Ranatra segrega Montandon, 1905 (Uruguay, Argentina).

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- The siphon is short and bears a strong fringe of setae, which increase in length caudad until the caudalmost setae are at least twice as long as the width of the siphon. The male paramere is strongly sigmoid (Fig. 6.476). .........................................................Ranatra flokata Nieser and Burmeister, 1998 (Amazonas). 9. (2) There is a tubercle in the interocular space (Fig. 6.477). The respiratory tube is shorter than the abdomen, except perhaps in the case of R. subinermis, for which this feature has not been described. ...................................................10 - There is no tubercle in the interocular space (Fig. 6.478). .............................12 10. (9) Total length: about 24 mm. Color: Medium to light brown without annulations on the legs. The paramere has a small subapical indentation (Fig. 6.477). ...............................................................................Ranatra weberi De Carlo, 1970 (Amazonas). - Total length: greater than 37 mm. ...................................................................11 11. (10) The interocular tubercle is indistinct. The hind femur of the female does not quite reach the base of the genital operculum. Length: c. 44 mm. Color: light brown without annulations on the legs. ....................................................................Ranatra subinermis Montandon, 1907 (French Guiana, Amazonas). - The interocular tubercle is very distinct. The paramere of the male curves more than 90º rather evenly in its apical 1/3 (Fig. 6.470). The hind femur of the female reaches to about the midpoint of the genital operculum. Length: 37 to 48 mm. Color: Medium to light brown without annulations on the legs. ...............................................................Ranatra tuberculifrons Montandon, 1907 (Surinam, Amazonas).

Fig. 6.477 Ranatra weberi male: head and anterior part of the pronotum in lateral view (left) and paramere (right). Based on Nieser (1975). 12. (9) The dorsal part of the abdomen is pinkish. The respiratory tube is slightly greater than 75% of the body length (Fig. 6.22). Each eye is about 1.2 times as wide as the distance between the eyes. Length: 40 mm. Only the female has been described. .................................................................................Ranatra unidentata Stål, 1861 (Minas Gerais).

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- The abdomen is yellowish or brownish, or, if it is somewhat reddish, then its size is greater, or the length of its respiratory tube is significantly shorter or longer than 75% of the body length (Fig. 6.478). ..............................................13

Fig. 6.478 Ranatra neivai male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946). 13. (12) The lateral process on the second antenna segment is almost as long as the third segment but only about half as wide, and it has a slight constriction at about its mid-length and a slight widening again toward the apex. The portion of the second segment between the borders of the first and third is only slightly longer than half the length of the third. The width of an eye is only slightly greater than the distance between the eyes. The subapical tooth on the male paramere is small (Fig. 6.478). The female genital operculum is long, narrow, only slightly curved, and visible well beyond the apex of the abdomen. Length of male: c. 35 to 36 mm; respiratory tube length: c. 24 mm. Length of female: c. 37 to 38 mm; respiratory tube length: 25 to 27 mm. ................................................................................Ranatra neivai De Carlo, 1946 (São Paulo, Paraná). - The lateral process on the second antenna segment is nearly parallel-sided for most of its length, or it is clearly more than half as wide as the third segment, or the width of an eye is at least about 1.1 times the width of the head between the eyes (Fig. 6.479). If in doubt, the total length or length of the respiratory tube is different from that reported in the first part of this couplet. ...............................14 14. (13) The operculum reaches more than 0.1 mm beyond the apex of the abdomen, and 10 or 11 small dorsal teeth on the operculum are visible beyond the apex of the abdomen (Fig. 6.480). Each eye is 1.1 to 1.2 times the interocular space. The respiratory tube folded back over the dorsum almost reaches the apex of the scutellum. The hind femur of the female almost reaches or just reaches the base of the operculum. Length: 37 to 38 mm. Color: dark brown with lighter legs, which are indistinctly banded and have dark anterior coxae. The male has not been described. .............................................................................Ranatra mixta Montandon, 1907 (Trinidad, French Guiana, Surinam, Pará).

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- The length is at least about 40 or less than 35 mm, or, if not, then fewer than 10 dorsal teeth on the operculum are visible beyond the apex of the abdomen (Fig. 6.479), or the transverse width of one eye is about 1 1/3 times or greater or less than 1 1/10 times the interocular width. .......................................................15

Fig. 6.479 Ranatra obscura: habitus in dorsal view (middle), antenna (upper left), and fore-tibia and tarsus (lower left). Based on De Carlo (1946).

Fig. 6.480 Ranatra mixta: apex of the female abdomen in lateral view. Based on Nieser (1975).

Fig. 6.481 Ranatra ecuadoriensis female: antenna (left) and fore-leg (right). Based on De Carlo (1964a).

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15. (14) The ground color is dark or very dark brown. The hind femur of the male does not reach or just reaches the base of the operculum, while that of the female does not reach that far (Fig. 6.479). .......................................................16 - The color is light brown, yellow, gray, or black, or, if it is dark brown, the hind femur of the female reaches the base of the operculum, or the operculum does not reach more than 0.1 mm beyond the apex of the abdomen; if in doubt, the number of teeth on the operculum that reach beyond the apex of the abdomen is greater or less than three. Otherwise, the hind femur of the male does not reach or reaches well beyond the base of the operculum, or the subapical tooth of the male paramere is sharply pointed (Fig. 6.481). ..............17 16. (15) Length of female: 31 to 33 mm; male: 29 to 32 mm. Color: dark brown; the legs are lighter and have dark bands. The hind femur of the male just reaches the base of the operculum, while that of the females does not reach that far. The apex of the operculum reaches more than 0.1 mm beyond the apex of the abdomen, and three of the dorsal teeth reach beyond the apex. The male paramere has a blunt subapical tooth, and the pronotum is about 3 mm wide (Fig. 6.479). .........................................................................Ranatra obscura Montandon, 1907 (Surinam, French Guiana, Guyana, Pará). Syn.: Ranatra annulipes White, 1879 (nec Stål, 1854). - Length of female: c. 47 mm; male: c. 40 mm. Color: uniformly very dark brown. The respiratory tube of the male is about 35 mm long, and that of the female is about 41 mm. The transverse width of an eye is about 1 1/3 times the length of the interocular space. The posterior part of the pronotum is about 3.3 mm wide. The fore-femur is about 12.6 mm long, and the large tooth on the femur is located c. 7.5 mm from the base. A very sketchy original description and lack of illustrations makes this a very difficult species to identify, and a better description is greatly needed. That the hind femur does not reach the base of the posteriormost abdominal segment is an assumption of later authors who reviewed the original description. ..............................................................................Ranatra attenuata Kuitert, 1949 (Ecuador). 17. (15) The general color of the female is uniformly brownish black with blackish legs. Length of female: c. 33 mm. The length of the respiratory tube is 37 mm. The lateral process of the second segment of the antenna and the third segment are long and parallel (Fig. 6.481). The male has not been described. ...................................................................Ranatra ecuadoriensis De Carlo, 1950 (Ecuador). Syn: Specimens from Ecuador identified by De Carlo (1946) as Ranatra annulipes Stål, 1854, actually belong to this species, which will remain difficult to identify until better descriptions, including that of the male, are available. - The general color is not darker than dark brown, or the legs are lighter than the body, with or without darker bands or mottling. In case of doubt, the ratio of the body length to the length of the respiratory tube is different from 33:37. .........18

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18. (17) The hind femur of the male extends posterior to the anterior margin of the posteriormost abdominal segment, but the hind femur of the female just reaches the anterior margin. The transverse width of an eye is about 1 1/3 times the length of the interocular space. The width of the posterior part of the pronotum is c. 4.1 mm. The genital operculum of the female extends well posterior to the apex of the abdomen. Length of male: 40 to 44 mm; female: 42 to 47 mm. Length of respiratory tube of the male: 32 to 37 mm; female: 32 to 39 mm. Color: testaceous to dark brown. The description of this species was cursory and lacked illustrations, so identification will be difficult until a better description is provided. ..........................................................................Ranatra hungerfordi Kuitert, 1949 (Paraguay). - The length is shorter than 40 mm, or, if the length is greater, the hind femora are longer or shorter, or another part of the description does not fit. .................19 19. (18) The paramere of the male is strongly curved and lacks a distinct preapical tooth (Fig. 6.482). The hind femur does not reach the base of the operculum; the apex of the operculum reaches more than 0.1 mm beyond the apex of the abdomen; either five or six of the dorsal teeth on the operculum extend beyond the apex. The pronotum is not narrower than about 3 mm. ......20 - The paramere of the male bears a distinct preapical tooth. The hind femur reaches the base of the operculum or beyond, or, if it does not, then the apex of the operculum fails to extend more than 0.1 mm beyond the apex of the abdomen, or fewer than five or more than six of the dorsal teeth on the operculum extend beyond the apex (Fig. 6.483). ...............................................22

Fig. 6.482 Ranatra doesburgi: paramere of a male (left) and the apex of a female abdomen showing the ovipositor (right). Based on Nieser (1975).

Fig. 6.483 Ranatra magna (left to right): antenna, fore-leg, and male paramere. Based on De Carlo (1964a). 20. (19) The length is 23 to 25 mm. There is no indication of a subapical tooth on the paramere. The respiratory tube folded over the dorsum does not reach the

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pronotal pits. The operculum of the female does not reach beyond the apex of the abdomen. The width of an eye is about 1.2 times the interocular space (Fig. 6.484). Color: medium to dark brown with lighter, unbanded legs. ........................................................................Ranatra sarmientoi De Carlo, 1967 (Colombia, Surinam, Venezuela). Syn: Ranatra ameghinoi De Carlo, 1970. - The length is at least 35 mm. ..........................................................................21

Fig. 6.484 Ranatra sarmientoi: habitus (left) and male paramere (right). Based on Nieser (1975). 21. (10) There is at most a faint indication of a subapical tooth on the paramere (Fig. 6.485). The respiratory tube of the female folded over the dorsum reaches almost to the midlength of the scutellum. About five of the dorsal teeth on the operculum extend beyond the apex of the abdomen. Length of male: 35.5 to 37 mm; female: 38 to 40 mm. Color: dark brown, the legs are lighter with indistinct darker bands. .............................................................................Ranatra moderata Kuitert, 1949 (Pará, Amazonas). - There is no indication of a subapical tooth on the paramere. The respiratory tube folded over the dorsum reaches to the pronotal pits. About six of the dorsal teeth on the operculum are visible posterior to the apex of the abdomen (Fig. 6.482). Length of male: 32 to 33 mm; female: 35 to 37 mm. Color: dark brown; the legs are lighter and have dark bands. .........................................................................Ranatra doesburgi De Carlo, 1963 (Colombia, Surinam, Pará, Amazonas). Syn.: Ranatra usingeri De Carlo, 1970.

Fig. 6.485 Male paramere of Ranatra moderata. Based on Nieser (1975).

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22. (19) The body length of the female is greater than 40 mm, and that of the male is at least 38 mm. .......................................................................................23 - Males and females are shorter than 40 mm. ....................................................29 23. (22) Length of male: at least 46 to 53 mm; female: 49 to 58 mm. The transverse width of an eye is 1.5 times the interocular space. Color: light brown with darker brown along the margins of the hemielytra, pronotum, and forefemur; the fore-femur and coxae are mottled, and the legs may have indistinct bands (Fig. 6.483). ..................................................................................Ranatra magna Kuitert, 1949 (Amazonas). Syn: Ranatra carvalhoi De Carlo 1954. - Length of male: less than 44 mm; female: no more than 50 mm. The lateral prolongation of the second antenna segment extends apicad more than half of the length of the third segment (Fig. 6.486). ......................................................24 24. (23) Length of male: 38 to 40 mm; female: 40.5 to 43 mm. ........................25 - Length of male: 41 to 44 mm. The pronotum is densely coated by somewhat clavate bristles. The known females reach a length of 46 to 50 mm. ................26

Fig. 6.486 Ranatra macrophthalma male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

Fig. 6.487 Ranatra travassosi female: antenna (left) and fore-leg (right). Based on De Carlo (1964a). 25. (24) The respiratory tube of the male is almost as long as he body. The apical portion of the male fore-femur is only slightly curved at the tip. The hind femur of the female reaches about to the midlength of the operculum (Fig.

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6.486). Specimens may be confused with Ranatra heydeni (Couplet 27), but the operculum of the female reaches well beyond the apex of the abdomen, while that of R. heydeni reaches only slightly beyond. Length: about 42 mm. Color: dark brown; the legs are light brown with well defined dark bands. ...........................................Ranatra macrophthalma Herrich-Schaffer, 1849 pars (Trinidad, Colombia, Surinam, French Guiana, Bolivia, Paraguay?, Amazonas, Pará, Minas Gerais). Syn: Ranatra surinamensis De Carlo, 1963. Only unusually large specimens will key out here. Specimens of normal size will key out below. - The respiratory tube is only about 3/4 as long as the body. Length: about 40.5 mm. The bands on the femora are somewhat faint. The tooth on the fore-femur is located somewhat proximal to the apical third (Fig. 6.487). .........................................................................Ranatra travassosi De Carlo, 1950 (São Paulo). 26. (24) The respiratory tube is only about half as long as the body. Length of male: 41.5 to 44 mm. Length of respiratory tube: c. 21 mm. The transverse length of one eye is slightly more than 0.75 times the interocular width. Color: light testaceous with a much darker prothorax, mesosternum, and metasternum. The female has not been described. .....................................................................Ranatra dolichodentata Kuitert, 1949 (São Paulo). - The respiratory tube is at least about 2/3 of the body length (Fig. 6.453). .....27 27. (26) The respiratory tube of the male is longer than the total body length. The legs are yellowish and unbanded (Fig. 6.453). Some specimens may be confused with Ranatra macrophthalma (Couplet 25), but the operculum of the female reaches only slightly beyond the apex of the abdomen, while that of R. macrophthalma reaches well beyond. Length of male: 43 to 44 mm; female: 46 to 50 mm. .........................................................................Ranatra heydeni Montandon, 1909 (Argentina, Minas Gerais). - The respiratory tube is less than 80% as long as the body. .............................28 28. (27) The respiratory tube is only about 2/3 of the body length, but that of the type may have been broken. The male paramere is widest near its mid-length (Fig. 6.488). The legs are brown and unbanded. Length of male: 44 mm. ..............................................................................Ranatra sattleri De Carlo, 1967 (Trinidad, Pará). Only the male has been described. Nieser (1975) cites some confusion about the holotype, which was labelled differently from the way it was reported by De Carlo (1967a). The similarity of this species, known only from the type male, and R. heydeni is so great that no reliable way of distinguishing the two can be found in the descriptions. A synonymy seems likely. - The respiratory tube is more than 3/4 of the body length. Length of male: about 42 mm. .................................................................................Ranatra cruzi De Carlo, 1950 (Espirito Santo). Only the male has been described.

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Fig. 6.488 Ranatra sattleri male (left to right): antenna, fore-leg, and the male paramere. Based on De Carlo (1967a). 29. (22) The median length of the pronotum is 1.05 to 1.1 times as long as the fore-femur. The respiratory tube is less than 2/3 of the total length. The subapical tooth on the male paramere is small and acute (Fig. 6.489). Length: 31 to 36 mm. Color: mainly dark brown with mottled legs on which the dark brown patches do not usually form bands. ......................................................................Ranatra curtafemorata Kuitert, 1949 (Amazonas). - The median length of the pronotum is no more than 0.8 to 0.9 times as long as the fore-femur (Fig. 6.490). ................................................................................30

Fig. 6.489 Ranatra curtafemorata male (left to right): antenna, fore-leg, and paramere. Based on De Carlo (1964a). 30. (29) The respiratory tube is clearly longer than the body (Fig. 6.488). ........31 - The respiratory tube is slightly longer than, equal to, or shorter than the total body length (Fig. 6.491). ....................................................................................36 31. (30) The interocular space is no more than about 0.8 times as wide as each eye (Fig. 6.490). ..................................................................................................32 - The interocular space is equal to or greater than 0.9 times the width of an eye and is most often greater (Fig. 6.492). ................................................................33

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32. (31) The metaxyphus has a low carina between the metacoxae (Fig. 6.490). Length: 26 to 29.5 mm. Color: medium to dark brown; legs lighter and only vaguely banded. ........................................................Ranatra rabida White, 1879 (Paraguay, Uruguay, Argentina, Amazonas, Mato Grosso do Sul). While Nieser (1975) stated that the respiratory tube is always longer than the body, DeCarlo (1964a) reported that it is sometimes the same length or slightly shorter. - The metaxyphus does not have a low carina between the metacoxae (Fig. 6.493). Length: 28 to 33 mm. Color of legs: uniform yellowish with no notable markings. ........................................................................Ranatra horvathi Montandon, 1910 (São Paulo). The similarity or the two species in this couplet leads to the suspicion that the names are synonymous.

Fig. 6.490 Ranatra rabida: habitus in dorsal view (middle left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

Fig. 6.491 Ranatra oliveiracesari male: habitus in dorsal view (middle), antenna (upper left), and fore-tibia and tarsus (lower left). Based on De Carlo (1946).

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Fig. 6.492 Ranatra costalimai male (left to right): antenna, fore-leg, and paramere. Based on De Carlo (1954a).

Fig. 6.493 Ranatra horvathi: habitus in dorsal view (middle), antenna (upper left), and fore-tibia and tarsus (lower left). Based on De Carlo (1946). 33. (31) The male paramere curves sharply slightly more than 90º well proximal to the insertion of the preapical tooth (Fig. 6.492). ............................................34 - The male paramere curves evenly much greater than 90º, just proximal to the preapical tooth (Fig. 6.494). Length of male: always less than 30 mm. ...........35 34. (31) The process on the second segment of the antenna is nearly parallel sided (Fig. 6.492). The male is about 31.5 mm long. The hind femur does not extend as far as the apex of the abdomen. The female remains undescribed. ........................................................................Ranatra costalimai De Carlo, 1954 (Minas Gerais). The two species in this couplet appear very similar. - The process on the second antenna segment tapers slightly to the apex (Fig. 6.495). Length of male: 29 to 31 mm; female: 32 to 34 mm. Male respiratory tube: 33 to 36 mm; female: 38 to 40 mm. Maximum eye width of male: 0.95 to 1.05 mm; female: 1.1 to 1.2 mm. Interocular space of male: 0.9 to 1.0; female: 1.0 to 1.1 mm. The metaxyphus extends 0.5 mm ventral to the margin of the hind coxa. The color is light brown with slightly darker areas. ...............................................................Ranatra zeteki Drake and De Carlo, 1953 (West Indies, Panama, Colombia, Venezuela).

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Fig. 6.494 Ranatra signoreti: habitus in dorsal view (lower left), antenna (upper left), fore-tibia and tarsus (lower right center), and paramere (right). Based on De Carlo (1946).

Fig. 6.495 Ranatra zeteki (left to right): antenna, paramere, and its apex. Based on Nieser (1969).

Fig. 6.496 Ranatra parvula: habitus in dorsal view (lower right) and parameres (upper left). Based on Nieser (1975). 35. (33) The hind femur does not reach to the base of the operculum (Fig. 6.494). Length: 28 to 29 mm. Color: medium to dark brown; legs lighter with indistinct dark bands. The male is unknown. .......................................................................Ranatra signoreti Montandon, 1907 (Venezuela, Bolivia, Paraguay, Argentina, Amazonas, Rio Grande do Sul).

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- The hind femur reaches at least to the base of the operculum (Fig. 6.496). Length: 24 to 26 mm. Color: the body and legs are light to medium brown. ................................................................................Ranatra parvula Kuitert, 1949 (Colombia, Pará, Amazonas). 36. (30) The respiratory tube is covered by long hairs for its entire length, and denser groups of these hairs are present on the basal third (Fig. 6.491). ...........37 - The respiratory tube is not covered by long hairs for its entire length (Fig. 6.497). .................................................................................................................38

Fig. 6.497 Ranatra sjostedti male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

Fig. 6.498 Ranatra lanei male: habitus in dorsal view (middle left), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946). 37. (36) The respiratory tube is only about 2/3 as long as the body length (Fig. 6.491). The female holotype is 31 mm long. Color: uniform chestnut brown. The male has not been described. ...................................................................Ranatra oliveiracesari De Carlo, 1946 (Mato Grosso).

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- The respiratory tube is at least almost 5/6 as long as the body (Fig. 6.498). The female is about 37 mm long. The middle and hind tibiae bear long hairs for their entire lengths. Length of male: 30 to 33.5 mm. Color: light yellow with no dark bands. .................................................................................Ranatra lanei De Carlo, 1946 (Rio de Janeiro).

Fig. 6.499 Ranatra annulipes: habitus in dorsal view (middle left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946). 38. (36) Each eye is about 0.8 mm, equal to or slightly narrower than the interocular space (Fig. 6.497). Length of male: 27.7 to 29 mm; female: 28 to 33 mm. Color: uniform yellowish without dark bands. ........................................................................Ranatra sjostedti Montandon, 1911 (Argentina, Uruguay). - Each eye is generally equal to or wider than the interocular space (Fig. 6.499). In case of doubt, the color is not uniformly yellowish. ......................................39

Fig. 6.500 Ranatra siolii: habitus (left) and a male paramere (right). Based on De Carlo (1970) and Nieser (1975).

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39. (38) The middle and hind femora are clearly marked with three dark and three light bands. The respiratory tube is about equal to the body in length (Fig. 6.499). .................................................................................................................40 - The middle and hind femora are only vaguely banded or completely lack dark bands (Fig. 6.500). ..............................................................................................42 40. (39) The abdomen has prominent dark markings. The eyes are about 1 mm wide and equal to the interocular space at the midpoint between the eyes (Fig. 6.499). Length of males and females: 30 to 31 mm. ..................................................................................Ranatra annulipes Stål, 1854 (Uruguay, Rio de Janeiro, Rio Grande do Sul). - The eyes are wider than the interocular space (Fig. 6.500). ...........................41 41. (40) The total length is about 25 to 27 mm. The body length is subequal to or slightly shorter than the length of the respiratory appendices. The genital operculum of the female does not extend beyond the apex of the abdomen. The lateral margin of the posterior part of the abdomen bears two kinds of teeth, which contact the free end of the male genital operculum (Fig. 6.501). ........................................................................Ranatra camposi Montandon, 1907 (Ecuador). - The total length is 35 to 42 mm. The body length is subequal to or slightly longer than the length of the respiratory tube. The genital operculum of the female clearly extends beyond the apex of the abdomen. The lateral margin of the posterior part of the abdomen does not bear any teeth that contact the free end of the male genital operculum (Fig. 6.486). ...........................................Ranatra macrophthalma Herrich-Schaffer, 1853 pars (Trinidad, Surinam, French Guiana, Bolivia, Paraguay?, Pará). Syn: Ranatra surinamensis De Carlo, 1963. See Couplet 25.

Fig. 6.501 Ranatra camposi male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

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Fig. 6.502 Ranatra chagasi male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946). 42. (39) Length of male: 28 mm; female: 27 mm. The width of the pronotum is 2 mm. The color is light yellowish brown or grey-brown with no trace of bands on the legs. The respiratory filaments are 2/3 to 4/7 of the total length. The hind femur of the male reaches to about halfway between the base and apex of the operculum. The subapical tooth on the male paramere is blunt (Fig. 6.500). .................................................................................Ranatra siolii De Carlo, 1970 (Pará, Amazonas). - If the length is about 27 to 28 mm, and the width of the pronotum is 2 mm, then the color is medium or dark brown, or the respiratory filaments are longer (Fig. 6.502). ........................................................................................................43

Fig. 6.503 Ranatra brasiliensis male: habitus in dorsal view (left to center), antenna (upper left), fore-tibia and tarsus (lower left), and paramere (right). Based on De Carlo (1946).

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43. (42) The width of an eye is subequal to or as much as 1.2 times the interocular space. The male paramere has a subapical tooth that is no longer than its width at its base (Fig. 6.502). Length of male and female: 35 to 35.5 mm. Color: dark chestnut brown. .............................................................................Ranatra chagasi De Carlo, 1946 (São Paulo, Paraná). - Each eye is about 1.2 to more than 1.6 times as wide as the interocular space (Fig. 6.503). ........................................................................................................44 44. (43) Each eye is at least 1.6 times as wide as the interocular space. The subapical tooth of the paramere is small and narrow, at least 1.5 times as long as its width at the base. The width of the pronotum is about 3.4 mm. The female genital operculum extends well beyond the apex of the abdomen. Length: 35 to 36 mm. Length of respiratory tubules: 26 to 27 mm. Color: medium brown with similarly colored, unbanded legs (Fig. 6.503). .......................................................................Ranatra brasiliensis De Carlo, 1946 (Amazonas?). - Each eye is about 1.2 to 1.4 times as wide as the interocular space (Fig. 6.504). .................................................................................................................45

Fig. 6.504 Ranatra mediana: paramere of a male (left) and the apex of a female abdomen showing the ovipositor (right). Based on Nieser (1975). 45. (44) Each eye is about 1.4 times as wide as the interocular space. The length of the male is approximately 28 to 30 mm. ........................................................46 - Each eye is 1.2 to 1.3 times as wide as the interocular space (Fig. 6.505). ....47 46. (45) The subapical tooth of the male paramere is relatively large and sharply pointed (Fig. 6.504). Length of the male: 28 to 30 mm; female: 31 to 33 mm. Color: dark brown, usually with a lighter venter; legs lighter with very faint bands. ........................................................................Ranatra mediana Montandon, 1910 (Trinidad, Surinam, French Guiana, Guyana, Pará). Syn: Ranatra williamsi Kuitert, 1949. - The subapical tooth of the paramere is relatively small and rounded (Fig. 6.506). Length: c. 29 mm. The pronotum is 2.8 mm wide. Color: medium brown with lighter, unbanded legs. The female has not been described. ...............................................................................Ranatra ornitheia Nieser, 1975 (Guyana).

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47. (45) The respiratory tube is slightly longer than 4/5 of the body length. The subapical tooth of the male paramere is small and blunt (Fig. 6.505). Length of the male: 27 to 28 mm; female: 33 mm. Color: medium brown; legs lighter and only vaguely banded. .......................................................................Ranatra adelomorpha Kuitert, 1949 (Surinam). - The respiratory tube is slightly shorter than 4/5 of the body length. Length of the male: 28 mm. The femora are completely unbanded. The third segment of the antenna is evenly curved (Fig. 6.507). The female has not been described. ..................................................................................Ranatra lenti De Carlo, 1950 (Espirito Santo, Rio de Janeiro, Minas Gerais). The reported differences between this species and R. adelomorpha are minor, raising doubts about its validity.

Fig. 6.505 Ranatra adelomorpha: habitus (left) and one paramere of a male and the apex of the other (right). Based on Nieser (1975).

Fig. 6.506 Male parameres of Ranatra ornitheia. Based on Nieser (1975).

Fig. 6.507 Antenna of Ranatra lenti. Based on De Carlo (1964a).

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Belostomatidae Key to the South American genera Information for the key was provided by Torre Bueno (1907), Cummings (1933), Lauck and Menke (1961), De Carlo (1966a), and Nieser (1975). 1. There is a suture-like fold delimited by a suture on each side that subdivides the fifth and sixth sternites; it may be concealed by silky setae. Spiracles are located on or adjacent to the mesal margins of the ventral laterotergites (Fig. 6.508). ...................................................................................................................2 - There is no suture-like fold but only a single sulcus subdividing the fifth and sixth sternites. The spiracles are located near the centers of the ventral laterotergites (Fig. 6.509). .................................................Belostomatinae..........3

Fig. 6.508 Horvathinia pelocoroides: head in dorsal (upper left) and lateral view (lower left), habitus (left center), sternites showing the location of the pilosity in black (right center), and male genitalia in dorsal, ventral (upper right, left and right, respectively), and lateral view (lower right). Based on Schnack and Estéves (2005). 2. (1) The mesal margins of the laterotergites contact the genital plate near its apex. The second and third segments of the antenna sometimes bear angular, digitiform projections. The tibia and tarsus of the hind leg are compressed, thin, and much more dilated than the middle tibia and tarsus. The fore-tarsus appears to consist of two segments with one well-developed claw. The head is not conically produced. The rostrum is short and stout (Fig. 6.510). The tarsal claws are prominent on all larval instars (Fig. 6.335). .......................................................Lethocerinae....Lethocerus Meyr, 1853..p. 395 - The mesal margins of the laterotergites contact the genital plate near its basal angle. The third segment of the antenna bears a large expanded and flattened basal lobe. The tibiae and tarsi of the middle and hind legs are similarly compressed and narrow but not broadly dilated. The fore-tarsus appears to consist of one segment, and both claws are vestigial (Fig. 6.508). What may be

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the only South American species in this genus has been found to inhabit wetlands. ........................................Horvathiniinae....Horvathinia Montandon, 1911..p. 403

Fig. 6.509 Weberiella rhomboides: habitus in dorsal view (left), profile of the right side of the genitalia in dorsal (upper center) and ventral view (right), genitalia in lateral view (lower center). Based on Nieser (1975).

Fig. 6.510 Lethocerus delpontei: habitus in dorsal view (below), ventral surface of the hind leg (upper left), male genitalia in lateral view (middle left), and shape of the apical abdominal segment in ventral view (upper right). Based on Cummings (1933) and Nieser (1975). 3. (1) The fore-tarsus consists of one segment. The lateral margins of the abdomen are interrupted at the borders between the segments (Fig. 6.509). Length: about 21 mm. ......................................................................................Weberiella De Carlo, 1966 This is a monotypic genus including only Weberiella rhomboides (Menke, 1965) from French Guiana and Amazonas.

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- The fore-tarsus consists of two segments. The lateral margins of the abdomen form an uninterrupted curve (Fig. 6.511). ............................................................4 4. (3) The membrane is well developed, being wider than the clavus at its widest point. Most of its cells form long, narrow rectangles of equal length (Fig. 6.511). ............................................................................Belostoma Latreille, 1807..p. 404 - The membrane on the hemielytra is reduced, so that its greatest width is equal to or less than the maximum width of the clavus. Its cells are usually small and feebly developed, often forming short rectangles of variable length. There is a strong mid-ventral keel on the metasternum. ....................................................................................................Abedus Stål, 1862 Known from California to Panama; it is just beyond the geographical range of this volume.

Fig. 6.511 Belostoma porteri: head in dorsal view (left) and dorsal view of the habitus. Based on De Carlo (1942). Key to the species of Lethocerus reported from South America Information for the key was provided by Menke (1962), De Carlo (1931a, 1962b, 1964b) Pérez Goodwyn (1997, 2006), and Ribeiro (2005). Menke (1979) successfully petitioned the Intenational Commission on Zoological Nomenclature to preserve the name Lethocerus. 1. The inner apical angle on the underside of the hind tibia forms a sharp point (Fig. 6.512). ..........................................................................................................2 - The inner apical angle on underside of posterior tibia does not form a sharp point; it is typically broadly rounded (Fig. 6.513). ..............................................4

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Fig. 6.512 Lethocerus grandis: tarsus and claw showing the location and numbering of the tarsal segments (upper left), the sharply pointed internal projection on the ventral side at the apex of the hind tibia (lower left), the aedeagus in lateral (upper center) and ventral view (lower center), and the paramere in lateral and ventral view (right). Based on Peréz Goodwyn (2006).

Fig. 6.513 Lethocerus jimenezasuai: diagrammatic dorsal view of a head showing divergent inner margins and oblique posterior margins of the compound eyes (left), aedeagus in lateral (upper left center) and ventral view (lower left center), paramere in lateral (upper right center) and ventral view (upper right), lateral view of the female gonopophysis (lower right center), and diagrammatic ventral view of the apex of the hind tibia showing its rounded inner angle (lower right). Based on Peréz Goodwyn (2006). 2. (1) The lateral margins of the pronotum are broadly foliaceous, that is, very thin and expanded. The width of the fore-femur is at least 1.3 times the width of the hind femur; and the fore-femur is not pointed at its apex. The claw on the fore-leg is longer than the combined length of the tarsal segments. The anterior lobe of the pronotum is fuscous with a pair of dark median lines on the anterior third and a pair of conspicuous brown oval spots. The margins of the hind tibiae are parallel. The dorsal margins of the compound eyes are broadly rounded, and their inner margins are divergent; their posterior margins lie along a

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straight line. The male paramere does not appear expanded at the apex in dorsal or ventral view, and the hook is large and curved (Fig. 6.514). Length of male: 73 to 80 mm; female: 79 to 88 mm. Width: 28 to 33 mm. Color: dark with a patchy pattern. .................................................................Lethocerus camposi (Montandon, 1900) (Ecuador). Syn: Amorgius camposi Montandon, 1901. - The lateral portions of the pronotum are not broadly foliaceous. The forefemur is only 1.1 to 1.2 times as wide as the hind femur, and it has a pointed apex, which is visible when the leg is folded. The hook on the male genitalia is large and curved (Fig. 6.515). ..............................................................................3

Fig. 6.514 Lethocerus camposi: diagrammatic dorsal view of a head showing divergent inner margins and straight posterior margins of the compound eyes (left), prosternal keel (left center), aedeagus in lateral (upper right center) and ventral view (lower right center), and paramere in lateral (upper right) and ventral view (lower right). Based on Peréz Goodwyn (2006).

Fig. 6.515 Lethocerus maximus (left to right): aedeagus in lateral (above) and ventral view (below); paramere in lateral (above) and ventral view (below); female gonopophysis. Based on Peréz Goodwyn (2006). 3. (2) The fore and hind femora are roughly equal in length, or the fore-femur is slightly longer. There is a variable, median chestnut marking on the anterior part of the head. The inner apical angle on the underside of the hind tibia forms a sharp point, and the claw on the fore-tarsus is longer than the combined length

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of the two tarsal segments, measured along their outer margins. The male paramere appears somewhat widened in dorsal and ventral view; the hook is large and curved (Fig. 6.512). Length of male: 81 to 92 mm; female: 95 to 102 mm; width: 31 to 38 mm. The color is light with variable patches of darker pigment. ......................................................................Lethocerus grandis (Linnaeus, 1758) (Colombia, Paraguay? Argentina? Mato Grosso, São Paulo, Rio de Janeiro, Paraná, Minas Gerais). Syn: Nepa grandis Linnaeus, 1758; Belostoma grandis (Linnaeus, 1758) Latreille, 1809; Belostomum grande Burmeister, 1835; Belostoma grande Blanchard, 1840; Belostoma annulipes sensu HerrichSchäffer, 1845; Iliastus grandis Gistel, 1848; Lethocerus grandis = annulipes Herrich-Schäffer, 1849; Lethocerus largus Cummings, 1933. - The fore-femur is shorter than the hind femur. The length is greater than 85 mm, and the claw on the fore-tarsus is equal to or shorter than the combined length of the two tarsal segments. The male parameres appear very much widened in dorsal and ventral view (Fig. 6.515). Length of male: 90 to 102 mm; female: 100 to 110 mm; width: 33 to 41 mm. The overall color is dark with a dark longitudinal stripe between the compound eyes. There are pale stripes on the pronotum and pale foveae. The middle and hind femora each have three dark bands. .......................................................................Lethocerus maximus De Carlo, 1938 (West Indies, Venezuela, Colombia, Guyana, French Guiana, Surinam, Peru, Bolivia, Paraguay, Argentina, Pará, Mato Grosso, Mato Grosso do Sul, Rio de Janeiro, Rio Grande do Sul, Espirito Santo). Syn: Lethocerus paraensis Lanzer de Souza, 1991, Belostoma grande Amyot and Audinet Serville, 1843; Lethocerus grandis (partim) misidentified by De Carlo (1930). 4. (1) The hind femur is longer than the fore-femur (Fig. 6.516). .......................5 - The fore-femur is longer than the hind femur (Fig. 6.517). ..............................6 5. (4) The mesosternum has one dark mid-ventral stripe. There is no carina between the eyes. The fore-claw is longer than the combined length of the tarsal segments. There are three conspicuous dark rings on the hind femur and middle and hind tibiae. The compound eye inner margins are parallel, and the posterior margins are oblique to each other. The hind femur is 1.05 to 1.07 times as wide as the hind tibia. The parameres do not appear enlarged in dorsal or ventral view, and the hook is large and curved (Fig. 6.516). Length of male: 48 to 58 mm; female: 62 to 64 mm. Maximum width of male: 19 to 24 mm; female: 24 to 26 mm. Coloration: densely spotted. .........................................................Lethocerus medius (Guérin-Méneville, 1857) (North and Central America; one old record from Venezuela). Syn: Belostoma medium Guérin-Méneville, 1857; Belostoma curtum Guérin-Méneville, 1857; Belostoma Signoreti Dufour, 1863 (partim); Belostoma annulipes Uhler, 1876 (nec Herrich-Schäffer, 1845); Amorgius annulipes Torre Bueno, 1906 (nec Herrich-Schäffer, 1845); Lethocerus curtus Kirkaldy and Torre Bueno, 1908; Lethocerus annulipes Van Duzee, 1917; Lethocerus delpontei Cummings, 1933 (partim).

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- The mesosternum has three dark longitudinal stripes, which may be faint in pale individuals. There are faint and diffuse dark markings on the middle and hind legs. The inner margins of the compound eyes are divergent, and their posterior margins are oblique to each other. There is a carina between the compound eyes. The hind femur is about 1.10 times as wide as the hind tibia. The paramere appears widened in dorsal and ventral view, and the hooks are large and curved (Fig. 6.513). Length of male: 59 to 65 mm; female: 62 to 69 mm. Width: 23 to 26 mm. Color: mainly dark with a dark marking between the eyes. ................................................................Lethocerus jimenezasuai De Carlo, 1957 (Ecuador).

Fig. 6.516 Lethocerus medius male (left to right): aedeagus in lateral and ventral view; paramere in lateral and ventral view. Based on Peréz Goodwyn (2006).

Fig. 6.517 Lethocerus bruchi male (above, left to right): aedeagus in lateral and ventral view; paramere in lateral and ventral view; and (below) gonopophyses of a female in lateral view. Based on Peréz Goodwyn (2006). 6. (4) The posterior tip of the metaxyphus is truncate and straight in the middle, not sharply pointed. The inner margins of the compound eyes are almost parallel, and their posterior margins form a straight line. There is a carina between the compound eyes. The aedeagus does not appear widened at the apex in ventral view, but the paramere is considerably widened in lateral view, and each appears almost rectangular. The hook is considerably reduced (Fig. 6.516).

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The larval instars are shown in Figure 6.335. Length of male: 69 to 78 mm; female: 64 to 79 mm. Maximum width: 25 to 30 mm. ..........................................................................Lethocerus bruchi De Carlo, 1931 (Argentina, Uruguay, Paraguay, São Paulo, Paraná, Minas Gerais) Syn: Lethocerus truncatus Cummings, 1933. - The metaxyphus has a sharply pointed posterior apex. The parameres do not appear rectangular (Fig. 6.518). ...........................................................................7

Fig. 6.518 Lethocerus annulipes: habitus (left), male genitalia in lateral view (lower right), and shape of the apical abdominal segment in ventral view (upper right). Based on Cummings (1933) and Nieser (1975). 7. (6) There are fuscous stripes on the third through sixth parasternites, and sometimes on the seventh, as well. The paramere is only slightly widened in dorsal and ventral view, and the hook is large and curved (Fig. 6.518). Length of male: 54 to 69 mm; female: 58 to 75 mm. Width: 19 to 27 mm. ......................................................Lethocerus annulipes (Herrich-Schäffer, 1845) (North and Central America, West Indies, Colombia, Ecuador, Venezuela, Guyana, Surinam, French Guiana, Paraguay, Uruguay, Argentina, Mato Grosso, Minas Gerais, Goiás, São Paulo, Rio de Janeiro, Santa Catarina, Rio Grande do Sul). Syn: Belostoma annulipes Herrich-Schäffer, 1845; Belostoma ruficeps Dufour, 1863; Belostoma Signoreti Dufour, 1863; Amorgius annulipes (HerrichSchäffer, 1846) Montandon, 1909; Belostoma annulipes var. mayri Montandon, 1896; Belostoma mayri Kirkaldy and Torre Bueno, 1908. - There are no fuscous stripes on the underside of the abdomen (Fig. 6.519). ...8 8. (7) The posterior borders of the compound eyes are aligned along one straight line (Fig. 6.519). ...................................................................................................9 - The posterior borders of the compound eyes are oblique to each other (Fig. 6.520). .................................................................................................................10

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9. (8) The hind wings are dark. The tarsal claw is longer than the total length of the tarsal segments of the fore-legs. The male paramere appears only slightly widened in dorsal and ventral view; the hook is large and curved (Fig. 6.521). Length of male: 63 to 71 mm, female: 64 to 72 mm. Width: 24 to 30 mm. Color: light chestnut brown with little trace of spots and sometimes with a reddish abdomen. ..........................................................................Lethocerus mazzai De Carlo, 1962 (Peru, Bolivia, Argentina). - The hind wings are pale. The abdomen has a fuscous margin. The anterior tarsi are white. The paramere is only slightly widened in dorsal and ventral view; the hook is very small and only slightly curved (Fig. 6.519). Length of male: 77 to 82 mm; female: 69 to 72 mm. Width of male: 30 to 32 mm; female: 27 to 28 mm. .......................................................................Lethocerus dilatus Cummings, 1933 (Peru, Venezuela, Surinam, Bolivia, Santa Catarina).

Fig. 6.519 Lethocerus dilatus: diagrammatic dorsal view of a head showing parallel inner margins and straight posterior margins of the compound eyes (left), aedeagus in lateral (left center) and ventral view (upper right center), paramere in lateral (upper right) and ventral view (lower right), and lateral view of a female gonopophysis (lower center). Based on Peréz Goodwyn (2006). 10. (8) The exterior and interior margins of the hind tibia are curved so that they run approximately parallel. The abdomen lacks a fuscous margin. The anterior tarsi are not white. The paramere is widened in dorsal and ventral view; the hook is large and curved (Fig. 6.510). Length of male: 51 to 62 mm; female: 61 to 70 mm. Width: 20 to 28 mm. The species is found at elevations as high as 2000 m. ......................................................................Lethocerus delpontei De Carlo, 1933 (North and Central America, West Indies, Venezuela, Ecuador, Surinam, Paraguay, Argentina, Goiás, Mato Grosso, São Paulo, Rio de Janeiro, Paraná, Santa Catarina).

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- The exterior margin of the hind tibia is nearly straight, while the inner margin is somewhat curved, causing the width of the tibia to vary along its length. The fore-claw is longer than the combined length of the tarsal segments. The head has a mid-dorsal carina. The apex of the aedeagus appears greatly enlarged in dorsal and ventral view. The male paramere is slightly widened in dorsal and ventral view; the hook is large and curved (Fig. 6.520). Length of male: 58 to 67 mm; female: 64 to 70 mm. Width: 22.5 to 28 mm. Color: light chestnut with irregular spotting. ..................................................................Lethocerus melloleitaoi De Carlo, 1933 (Argentina, Paraguay, Uruguay, Colombia? São Paulo, Rio de Janeiro, Santa Catarina, Minas Gerais, Mato Grosso do Sul, Mato Grosso). Syn: Belostoma annulipes Mayr, 1871 (partim); Lethocerus mello-leitãoi De Carlo, 1933. Ribeiro (2005) reported that this species cannot always be clearly distinguished from L. delpontei, but that the variation in width along the hind tibiae is a fairly reliable characteristic for distinguishing it.

Fig. 6.520 Lethocerus melloleitaoi (above, left to right): antenna, ventral view of the apex of a male and a female abdomen, male genitalia in lateral view, and (below, left to right): lateral view of the female gonapophysis, paramere in lateral and ventral view, and the aedeagus in ventral view. Based on Peréz Goodwyn (2006).

Fig. 6.521 Lethocerus mazzai (left to right): the male genitalia in ventral and lateral view and the female gonapophyses. Based on De Carlo (1964b) and Pérez Goodwyn (2006).

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Key to the species of Horvathinia reported from South America Information for the key is provided by De Carlo (1930, 1938a, 1958) and Schnack and Estéves (2005), who speculated that Horvathinia includes only one species but continued to provisionally recognize H. lenti because they had not examined the type specimen. The differences described by De Carlo (1958) and used in the key below certainly do not support his assumption that H. lenti is a distinct species. 1. The angle formed between the midline and the posterior margin of the metasternal plate is usually less than 70º. The anterolateral corner of the prosternal carina is sometimes broadly rounded (Fig. 6.508). Length: 25 to 30 mm. Maximum width: 13.5 to 15.5 mm. ..........................................................Horvathinia pelocoroides Montandon, 1911 (Argentina, Minas Gerais, Rio Grande do Sul). Syn: Horvathinia doellojuradoi De Carlo, 1930; Horvathinia pelleranoi De Carlo, 1930; Horvathinia meyeri De Carlo, 1938; Horvathinia castilloi De Carlo, 1938; Horvathinia schubarti De Carlo, 1957; Horvathinia bollei De Carlo, 1938; Horvathinia forsteri Lauck, 1958; Horvathinia argemii Schnack, 1972. - The angle formed at the midline of the posterior margin of the metasternal plate is usually greater than than 70º. The anterolateral corner of the prosternal carina is usually bluntly angled (Fig. 6.522). Length: c. 29 mm. Maximum width: c. 16 to 16.5 mm. Only a female has been described. ............................................................................Horvathinia lenti De Carlo, 1958 (São Paulo).

Fig. 6.522 Horvathinia lenti female (left to right): posterior part of the membrane, the posterior margin of the metasternal plate, and the anterior part of the prosternal carina. Based on De Carlo (1958).

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Key to the species of Belostoma reported from South America Information for the key was provided by De Carlo (1935a, 1938b, 1960b), Lauck (1962, 1963, 1964), Schnack (1973a, b), Bachmann (1962h), Nieser (1975), Lanzer-de-Souza (1996), Ribeiro (2005), Ribeiro and Alecrim (2008), and Ribeiro and Estévez (2009). The literature includes much confusion of one species with one of more others, and even relatively recent publications must be used with caution. Future taxonomic revisions can be expected. Belostoma minor (Palisot de Beauvois, 1805) is omitted from the key, relying on the opinion of Ribeiro (2007) that a report of this North American species from Brazil is based on a misidentification. 1. The pilosity of the ventral side of the abdomen reaches the genital operculum and covers the entire connexivum (Fig. 6.523). ...................................................2 - The pilosity on the ventral side does not reach the genital operculum, or, if it does, it does not completely cover the connexivum (Fig. 6.524). ......................13 2. (1) Pilosity covers at least part of the sternites (Fig. 6.525). ............................3 - Pilosity does not extend onto the sternites. .......................................................9 3. (2) The eyes are triangular with straight inner margins (Fig. 6.525). Length: 19 to 30 mm. .........................................................................................................4 - The eyes are globular with convex inner margins (Fig. 6.523). .......................7

Fig. 6.523 Belostoma noualhieri male (above, left to right), the prosternal keel in lateral view, ventral view of the extent of the pilosity on one side of the abdomen, and the ventral diverticulum of the penis in dorsal and ventral view, and (below), the ventral diverticulum of the penis in lateral view. Based on Ribeiro and Estévez (2009). 4. (3) The pilosity covers the entire ventral surface of the abdomen and about 2/3 of the genital operculum. Length: c. 28 to 30 mm. The color is chestnut brown. ...................................................................................................................5 - The sternites are only partially covered by pilosity (Fig. 6.525). The length does not usually exceed 28 mm. ...........................................................................6

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Fig. 6.524 Belostoma pygmeum (above, left to right): head in dorsal and lateral view, habitus in dorsal view, the extent of the ventral pilosity on one side of the abdomen marked in black, and (below, left to right): the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007).

Fig. 6.525 Habitus of Belostoma costalimai in dorsal view (left) and the extent of the pilosity on the ventral surface of one side of the abdomen marked in black (right). Based on De Carlo (1960b) and Nieser (1975).

Fig. 6.526 Belostoma fittkaui: habitus in dorsal view without appendages (left) and head in dorsal view (right). Based on De Carlo (1966).

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Fig. 6.527 Belostoma sayagoi: habitus in dorsal view without appendages (left), head in dorsal view (center), and penis in dorsal view (right). Based on De Carlo (1966).

Fig. 6.528 Belostoma hirsutum male: penis in dorsal (upper left), ventral (upper right), and lateral view (lower center). Based on Roback and Nieser (1974).

Fig. 6.529 Belostoma testaceopallidum male (left to right), habitus in dorsal view and the membranes of two specimens. Based on De Carlo (1963c). 5. (4) The body of the male is obviously flattened. There is a longitudinal carina between the eyes (Fig. 6.526). Length: c. 29 to 30 mm. Maximal body width: 15.0 to 15.2 mm. Color: very uniform dark chestnut brown. The female has not been described. ..........................................................................Belostoma fittkaui De Carlo, 1966 (Amazonas). This species was described from only two male specimens.

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- The pronotum is somewhat convex dorsally. There is no longitudinal carina between the eyes (Fig. 6.527). Length: c. 28 mm. Maximum width: c. 12.8 mm. Color: uniform chestnut brown. The female has not been described. .........................................................................Belostoma sayagoi De Carlo, 1966 (Pará). This species was described based on only one male specimen. 6. (4) Maximum width: 12 to 14.2 mm. Length: 19 to 28 mm. Color: dark brown with yellowish patches (Fig. 6.525). .....................................................................Belostoma costalimai De Carlo, 1938 (Colombia, Surinam, Roraima, Pará, Tocantins, Mato Grosso, Goiás, Minas Gerais, Espírito Santo, São Paulo, Rio de Janeiro). Syn: Belostoma truxali De Carlo, 1960. - Maximum width: 11 to 12 mm. Length: 25 to 28 mm. In lateral view, the penis arcs evenly ventrad (Fig. 6.528). Color: brown with a median longitudinal stripe on the pronotum and scutellum, which is sometimes indistinct. The legs are yellowish with brown markings, which sometimes coalesce to form bands. The ventral surface of the thorax is yellow with brown spots, and the ventral surface of the abdomen is variegated brown and yellow. ........................................................Belostoma hirsutum Roback and Nieser, 1974 (Colombia). 7. (3) The maximum width of the body is approximately half of the length (Fig. 6.529). The first segment of the rostrum is distinctly shorter than the second. Length: 28.9 to 33.4 mm. ...........................................................Belostoma testaceopallidum Latreille, 1807 (Guadalupe, Argentina, Mato Grosso, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina, Bahia?). Syn: Belostoma pallidum Gistel, 1848; possibly Zaitha stolli Mayr, 1863 (nec Amyot and Serville); probably Zaitha margineguttata Dafour, 1863; probably Zaitha carbonaria Dafour, 1863; Belostoma grandicollum De Carlo, 1934; Belostoma longirostrum De Carlo, 1934; Belostoma stolli De Carlo, 1950 (nec Amyot and Serville). - The maximum width of the body is less than half its length (Fig. 6.530). .......8

Fig. 6.530 Belostoma ribeiroi: habitus in dorsal view (left) and the penis in dorsal (upper right), ventral (middle right), and lateral view (lower right). Based on De Carlo (1935a) and Lauck (1963).

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8. (7) The first segment of the beak is longer than the second. The pilosity on the ventral side of the abdomen covers the entire connexivum and extends onto the genital operculum and the sternites (Fig. 6.523). Length: 20 to 21.5 mm; maximum width: 9.5 to 10.5 mm, less than half its length. .................................................................Belostoma noualhieri Montandon, 1903 (Rio Grande do Sul, Santa Catarina, São Paulo). - The first segment of the beak is about ¾ as long as the second. The length is slightly greater than twice the maximum width (Fig. 6.530). Length: 27.5 to 32.5 mm. .........................................................................Belostoma ribeiroi De Carlo, 1933 (Santa Catarina, Rio de Janeiro, Minas Gerais, Distrito Federal, Goiás, Mato Grosso). Syn: Belostoma dufouri De Carlo, 1933; Belostoma lundbladi De Carlo, 1963.

Fig. 6.531 Habitus of Belostoma elegans in dorsal view (below) and (above, left to right): the penis in dorsal, ventral, and lateral view. Based on De Carlo (1930) and Lauck (1964).

Fig. 6.532 Belostoma angustum male (left to right): penis in dorsal, ventral, and lateral view. Based on Lauck (1964). 9. (2) The body length is less than 30 mm (Fig. 6.20). ......................................10 - The body is considerably broadened and longer than 30 mm. ........................12

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10. (9) The body is ovoid, and the width of the scutellum is less than half of the hemielytral width (Fig. 6.20). Length: 20 to 23 mm. ...................................................................Belostoma bifoveolatum Spinola, 1852 (Argentina, Chile). Syn: Zaitha bifoveolata Dafour, 1863. - The body is elongate, so the width of the scutellum is approximately half as wide as the hemielytra (Fig. 6.531). ...................................................................11

Fig. 6.533 The left half of the head and anterior part of the pronotum of Belostoma dilatatum in dorsal view (left), and the penis in dorsal (upper center), ventral (upper right), and lateral view (lower right). Based on Lauck (1963) and Ribeiro (2005). 11. (10) The interocular space equals 1.2 to 1.3 times the width of a compound eye. The length of the head anterior to the eyes is subequal to the distance between the compound eyes (Fig. 6.531). Length: 20 to 23 mm. Color: light yellowish brown to greenish gray. .............................................................................Belostoma elegans (Mayr, 1871) (Peru, Bolivia, Argentina, Uruguay, Paraguay, Rio Grande do Norte, Rio Grande do Sul). Syn: Zaitha elegans Mayr, 1871. - The interocular space equals 1.4 to 1.6 times the width of a compound eye. The length of the head anterior to the eyes is less than the distance between the compound eyes. The apex of the penis is wide (Fig. 6.532). Length: 21 to 24.5 mm. Color: mainly dark brown. ...........................................................................Belostoma angustum Lauck, 1964 (Bolivia). 12. (9) The second segment of the beak is longer than the first. The compound eyes are much longer than wide (Fig. 6.532). Length: 42 to 53 mm. .......................................................................Belostoma dilatatum (Dufour, 1863) (Bolivia, Paraguay, Argentina, Uruguay, Pará, Paraíba, Mato Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo, Rio de Janeiro, Rio Grande do Sul, Paraná). Syn: Zaitha dilatata Dufour, 1863; probably Belostoma ripicolum Lanzer-de-Souza, 1992. - The first segment of the beak is longer than the second (Fig. 6.534). .....................................................................Belostoma martini (Montandon, 1899) (Argentina, Uruguay).

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Fig. 6.534 The penis of Belostoma martini in lateral view. Based on De Carlo (1960b). 13. (1) The pronotum and hemielytra are covered by fine, golden brown scalelike setae. The pronotum and scutellum have a distinct yellowish median carina. The body is generally elongate (Fig. 6.535). Length: 33 to 36 mm; width at base of pronotum: 12 to 13 mm; maximum width across hemielytra: 14.5 to 15.2 mm. .........................................................Belostoma aurivillianum (Montandon, 1899) (Colombia, Venezuela, Surinam, Bolivia, Paraguay, Acre, Roraima, Pará, Mato Grosso, Goiás, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Rio Grande do Sul). Syn: Zaitha aurivilliana Montandon, 1899. - There are no golden brown scales on the pronotum and hemielytra. There is no distinct yellowish median carina on the pronotum and scutellum, or if a faint carina is present, the body shape is oval or broadly oval (Fig. 6.536). ..............14

Fig. 6.535 Belostoma aurivillianum: habitus (left) and the penis in dorsal (upper right center), ventral (upper right), and lateral view (lower right). Based on Nieser (1975). 14. (13) The eye is more or less triangular with nearly straight outer margins (Fig. 6.536). ........................................................................................................15 - The eye appears globose with rounded outer margins (Fig. 6.537). ...............18

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Fig. 6.536 Belostoma bosqi male (left to right): habitus in dorsal view and the penis in dorsal and ventral view. Based on Nieser (1975).

Fig. 6.537 Belostoma gestroi: habitus in dorsal view (upper left) and penis in dorsal (center), ventral (right), and lateral view (lower left). Based on Nieser (1975).

Fig. 6.538 The head of Belostoma discretum in dorsal view. Based on Nieser (1975).

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Fig. 6.539 Belostoma martinezi male (left to right): penis in dorsal, ventral, and lateral view. Based on Lauck (1964). 15. (14) The second segment of the beak is longer than the first. The synthlipsis is greater than the width of an eye (Fig. 6.538). Length: 22.5 to 27.5 mm. ..................................................................Belostoma discretum Montandon, 1903 (Peru, Bolivia, Paraguay, Argentina, Pará, Amazonas, Rondônia, Acre, Goiás, Mato Grosso, Mato Grosso do Sul, São Paulo, Rio de Janeiro). - The first segment of the beak is longer than the second. ................................16 16. (15) The lateral margins of the body are greatly inflated and angular. There are abrupt expansions of the dorsal arms of the phallobase (Fig. 6.539). Length: 21 to 23.5 mm. .......................................................................Belostoma martinezi De Carlo, 1957 (Bolivia, Amazonas). - The lateral margins of the body are not greatly inflated or angular, and the general shape is elliptical. Abrupt expansions of the dorsal arms of the phallobase are absent (Fig. 6.536). .....................................................................17

Fig. 6.540 Belostoma bergi male: habitus in dorsal view (upper left), prosternal keel of two different specimens (upper right), penis of two different specimens shown above and below each other in dorsal (lower left), ventral (center), and lateral view (lower right). Based on Ribeiro (2007).

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17. (16) There is a longitudinal median carina on the head between the compound eyes (Fig. 6.536). The length of the first segment of the beak is about 1.25 times the length of the second. Length: 20.5 to 22 mm. .............................................................................Belostoma bosqi De Carlo, 1932 (Venezuela, Surinam, Peru, Bolivia, Paraguay, Argentina, Amazonas, Rondônia, Goiás, Minas Gerais, Mato Grosso, Mato Grosso do Sul). Syn: Belostoma bergi De Carlo, 1930 (nec Montandon, 1899) - There is no longitudinal median carina on the head between the compound eyes (Fig. 6.540). Length: 20 to 24 mm. .......................................................................Belostoma bergi (Montandon, 1899) (Peru, Uruguay, Paraguay, Argentina, São Paulo, Rio Grande do Sul). Syn: Zaitha anura Distant, 1891; Zaitha bergi Montandon, 1899; Zaitha boops Berg, 1879; Belostoma asiaticum De Carlo, 1930 (nec Mayr, 1863). 18. (14) The total length is not less than 28 mm. The first segment of the beak is equal to or longer than the second. .....................................................................19 - The total length does not exceed 24 mm. ........................................................40 19. (18) There is a prominent carina on the vertex. The scutellum does not reach the nodal line, that is, a line connecting the lateralmost points along the nodal furrow (Fig. 6.541). ............................................................................................20 - The vertex lacks a median carina, and the scutellum usually reaches the nodal line (Fig. 6.542). .................................................................................................21

Fig. 6.541 Belostoma stollii (left to right): habitus and the penis in dorsal, ventral, and lateral view. Based on Nieser (1975). 20. (19) The base of the clypeus is located anterior to the ocular line, that is, the line drawn between the anteriormost corners of the compound eyes (Fig. 6.537). Length: 39 to 45 mm. The ground color varies from yellowish brown to dark chocolate. .......................................................................Belostoma gestroi Montandon, 1903 (Surinam, Peru, Bolivia, Paraguay, Argentina, Amazonas, Mato Grosso).

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- The clypeus has its base at or posterior to the ocular line. The disc of the pronotum often bears a dense mat of short, scale-like hairs, but sometimes the hairs are sparce (Fig. 6.541). Length: 31 to 42 mm. Color: predominantly dark or grayish brown, sometimes with yellow mottling and dark lateral markings on the scutellum. The legs are banded, sometimes faintly. .........................................................Belostoma stollii (Amyot and Serville, 1843) (Guyana Surinam, French Guiana, Venezuela, Peru, Bolivia, Pará, Amazonas, Mato Grosso, Minas Gerais, Rio de Janeiro). Syn: Zaitha stollii Amyot and Serville, 1843; Belostoma brasiliensis De Carlo, 1950; Belostoma planum Lauck, 1963; Belostoma stolii auctt. 21. (19) There is a distinct apicoventral protuberance on the ventral diverticulum of the male genitalia (Fig. 6.542). ......................................................................22 - The apicoventral protuberance on the ventral diverticulum of the male genitalia is only moderately developed (Fig. 6.543). ........................................................31

Fig. 6.542 Penis of Belostoma foveolatum in ventral view. Based on Ribeiro (2005).

Fig. 6.543 Belostoma venezuelae male (left to right): head in dorsal view and penis in dorsal, ventral, and lateral view. Based on Lauck (1962). 22. (21) The first rostral segment is only slightly longer than the second, its length usually being shorter than 1.1 times the length of the second. The color is uniform dark brown. ...........................................................................................23

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- The first rostral segment is at least 1.3 times as long as the second (Fig. 6.544). .................................................................................................................24

Fig. 6.544 Belostoma bordoni male (left to right): head in dorsal view and the penis in dorsal and lateral view. Based on De Carlo (1966a).

Fig. 6.545 Belostoma doesburgi male: head in dorsal view (left) and the penis in lateral view (right). Based on De Carlo (1966a). 23. (22) The interocular space of the female is about 1.3 to 1.4 times the width of one compound eye. The penis is widest in its apical half (Fig. 6.542). Length of the female: 38 to 39 mm; maximum width: 16.0 to 17.2 mm. .......................................................................Belostoma foveolatum (Mayr, 1863) (French Guiana, Paraguay, Argentina, São Paulo, Rio de Janeiro, Minas Gerais, Mato Grosso do Sul, Ceará, Pará). Syn: Zaitha foveolata Mayr, 1863. - The interocular space of the male is no more than 1.25 times the width of one eye (Fig. 6.545). Length: 38 to 40 mm; maximum width: 16 to 18.5 mm. ......................................................................Belostoma doesburgi De Carlo, 1966 (Surinam). Sketchy descriptions of these species make it difficult to distinguish them reliably, if they are actually distinct. 24. (22) The part of the head anterior to the eyes in dorsal view is less than 1.3 times as long as the interocular space (Fig. 6.544). ...........................................25 - The part of the head anterior to the eyes in dorsal view is 1.3 times as long as the interocular space or more (Fig. 6.546). ........................................................28

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Fig. 6.546 Belostoma malkini: habitus (upper left), head in dorsal view (lower left), and penis in dorsal (upper center), ventral (upper right), and lateral view (lower right). Based on Nieser (1975). 25. (24) The first segment of the beak is about 1.65 times as long as the second segment. The widest point along the penis in dorsal view forms an abrupt angular bend (Fig. 6.544). Length: 36.5 to 40 mm. Maximum width: 15 to 16 mm. Color: uniform brown to dark brown. .........................................................................Belostoma bordoni De Carlo, 1966 (Venezuela). - The first segment of the beak is never much longer than 1.3 times the length of the second and usually shorter. ...........................................................................26 26. (25) In dorsal view, the narrowest point of the penis, just apical to the base, is about ½ the greatest width, located just proximal to the apex. The space between the eyes is about 3 mm, and the total width of the head, including the eyes, is about 7.8 mm (Fig. 6.547). The length of the head, measured along the midline, is about 6.5 mm. Length: 40 to 42 mm. Maximum width: c. 18 mm. Color: light brown. ..........................................................................Belostoma sattleri De Carlo, 1966 (Amazonas). - In dorsal view, the narrowest point of the penis, slightly apical from the base, is much more than ½ its greatest width, located just proximal from the apex (Fig. 6.548). ........................................................................................................27

Fig. 6.547 Belostoma sattleri male (left to right): head in dorsal view and the penis in dorsal and lateral view. Based on De Carlo (1966a).

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27. (26) The anterior width of the vertex is less than 1.3 times the width of an eye. In dorsal view, the head anterior to the eyes is about 1.1 times the interocular space (Fig. 6.511). Length: 37 to 44 mm. ...........................................................................Belostoma porteri De Carlo, 1942 (Panama, Venezuela, Peru). - The anterior width of the vertex is at least 1.35 times the width of a compound eye. The part of the head anterior to the eyes in dorsal view is 1.2 times as much as the interocular space, or more (Fig. 6.548). Length: 40 to 43 mm. ........................................................................Belostoma harrisi Lauck, 1962 pars (Guyana, Surinam, Pará). Also see Couplet 30.

Fig. 6.548 Belostoma harrisi: habitus in dorsal view (upper left), outline of the head in dorsal view (upper right), and the penis in dorsal (lower left), ventral (lower center), and lateral view (lower right). Based on Nieser (1975). 28. (24) The length is usually greater than 45 mm. The part of the head anterior to the eyes in dorsal view is greater than the interocular space but less than 1.5 times that space (Fig. 6.549). .............................................................................29 - The length is almost always shorter than 45 mm. The part of the head anterior to the eyes in dorsal view is no more than 1.4 times as much as the interocular space. The dorsal arms of the male genitalia are slender (Fig. 6.546). .............30 29. (28) The length of the head anterior to the eyes in dorsal view is 1.35 times as much as the interocular space, or more. The dorsal arms of the male genitalia are broad, and a ventral, preapical, tooth-like process is evident on the penis in lateral view. In dorsal or ventral view, the width narrows proximal to the widest point just proximal to the apex but increases to another wide point near the base (Fig. 6.549). Length: 42 to 51 mm. Color: usually a uniform dark brown but occasionally olive brown suffused with creamy yellow. ..........................................................................Belostoma dentatum (Mayr, 1863) (Venezuela, Peru, Bolivia, Paraguay, Uruguay, Argentina, Pará, Piauí, Amazonas, Rondônia, Acre, Mato Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo, Rio de Janeiro, Rio Grande do Sul). Syn: Zaitha dentata Mayr,

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1863; Zaitha eumorpha Dufour, 1863 pars?; Zaitha mayri Berg, 1884?; probably also encompasses the forms major and mayri of Montandon, 1903. - The part of the head anterior to the eyes in dorsal view is about 1.31 times as much as the interocular space. In dorsal or ventral view, the penis narrows gradually but evenly from the subapical location of its maximum width (Fig. 6.550). Length: 46 to 50 mm. Color: uniform brown. ...................................................................Belostoma peruvianum De Carlo, 1966 (Peru).

Fig. 6.549 Belostoma dentatum: habitus in dorsal view (upper left), extent of the ventral pilosity on one side of the abdomen (upper right), and the penis in dorsal, ventral, and lateral view (below, left to right). Based on De Carlo (1935a) and Nieser (1975).

Fig. 6.550 Belostoma peruvianum male (left to right): head in dorsal view and the penis in dorsal and lateral view. Based on De Carlo (1966a). 30. (28) The shape is elongate elliptical with a length:width ratio of 2.3:1 to 2.4:1 (Fig. 6.546). Length: 39 to 45 mm. ..............................................................................Belostoma malkini Lauck, 1962 (Trinidad, Venezuela, Guyana, Bolivia). The two names in this couplet are very probably synonyms. Lauck (1962) and Nieser (1975) did not agree about which specimens should be assigned to which species, and no definitive differences could be found between them. The general body shape, considered by Nieser (1975) to constitute a genuine difference, can be very variable among populations of a given species, and the lack of a difference between the male genitalia strongly suggests that they are conspecific forms.

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- The shape is broad and ovoid with a length:width ratio of 2.1:1 to 2.2:1 (Fig. 6.548). Length: 40 to 43 mm. ........................................................................Belostoma harrisi Lauck, 1962 pars (Guyana, Surinam, Pará). Also see Couplet 27. 31. (21) The pilosity on the connexivum extends along the margins of the genital operculum (Fig. 6.551). Length: 34.5 to 35.5 mm. .....................................................................Belostoma cummingsi De Carlo 1935 (Argentina, Uruguay, Minas Gerais, São Paulo, Santa Catarina, Rio Grande do Sul). Syn: Belostoma foveolatum De Carlo, 1930 (nec Mayr, 1863), Belostoma cachoeirinhensis Lanzer-de-Souza, 1996. - The pilosity on the connexivum extends only slightly posterior to the penultimate segment (Fig. 6.552). ......................................................................32

Fig. 6.551 Belostoma cummingsi (above, left to right): beak in lateral view, habitus in dorsal view, one side of the abdomen showing the pilosity on the connexivum, and (below, left to right): penis in dorsal, ventral, and lateral view, and the head in dorsal view. Based on De Carlo (1935a) and Lauck (1962).

Fig. 6.552 Belostoma oxyurum male (above, left to right), habitus without appendages in dorsal view and the head in dorsal and lateral view, and (below, left to right): one side of the abdomen in ventral view showing the extent of the pilose area, and the ventral diverticulum of the penis in dorsal, ventral, and lateral view. Based on Ribeiro and Estévez (2009).

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32. (31) The length of the head anterior to the compound eyes is greater than the interocular width (Fig. 6.553). ...........................................................................33 - The length of the head anterior to the compound eyes is equal to or less than the interocular width (Fig. 6.552). ......................................................................35 33. (32) On the head, there are distinct lateral foveae, which form grooves at the anterior corners of the compound eyes. The length of the head anterior to the compound eyes is equal to or greater than the width between the anterior corners of the eyes (Fig. 6.553). Length: 27.5 to 33.0 mm. ..........................................................Belostoma anurum (Herrich-Schäffer, 1848) (Mexico, Central America, West Indies, Guyana?, Chile, Argentina, Paraguay, Maranhão, Ceará, Alagoas, Rio Grande do Norte, Bahia, Minas Gerais, Espírito Santo, São Paulo, Rio de Janeiro, Santa Catarina). Syn: Diplonychus anurus Herrich-Schäffer, 1848; Zaitha anurus (Herrich-Schäffer, 1848) in Dufour, 1863; probably Zaitha stollii Dufour, 1863 (nec Amyot and Serville); probably Zaitha subspinosa Dufour, 1863 (nec Palisot de Bouvois); Belostoma boscii Montandon, 1903 pars (nec Le Peletier de Saint-Fergeau and Serville); Belostoma dallasi Schnack, 1973 (nec De Carlo). - The lateral foveae on the head are only moderately developed. The part of the head anterior to the compound eyes is slightly longer than the width between the anterior corners of the eyes (Fig. 6.554). ...........................................................34

Fig. 6.553 Belostoma anurum: left side of the head with compound eye and anterior portion of the prothorax in dorsal view (left) and penis in ventral view (right). Based on Ribeiro (2005). 34. (33) In lateral view, the length of the head anterior to the compound eyes and the length of the eye are equal (Fig. 6.554). Length: 25 to 27.5 mm. .......................................................................Belostoma ellipticum Latreille, 1833 (Ecuador, Peru, Chile). Syn: Zaitha anurus Mayr, 1871 pars; Zaitha boops Dufour, 1863 pars; Zaitha carbonaria Dufour, 1863 possibly; Belostoma asiaticum Montandon, 1903 (nec Mayr, 1863) pars; Belostoma impavidum Torre Bueno.

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- In lateral view, the length of the head anterior to the compound eyes is shorter than the length of the eye (Fig. 6.555). Length: 33.5 to 35 mm. ..........................................................................Belostoma columbiae Lauck, 1962 (Colombia).

Fig. 6.554 Belostoma ellipticum (left to right): head in dorsal view and the male genitalia in dorsal, ventral, and lateral view. Based on Lauck (1962).

Fig. 6.555 Belostoma columbiae (left to right): head in dorsal view and the male genitalia in dorsal, ventral, and lateral view. Based on Lauck (1962).

Fig. 6.556 Belostoma pseudoguianae male (left to right): penis in dorsal, ventral, and lateral view. Based on Roback and Nieser (1974). 35. (32) The following combination of characters is diagnostic: The first and second segments of the rostrum are subequal in length; the interocular space is equal to 1.4 to 1.5 times the width of a compound eye; the anterior interocular width is equal to about 1.2 times the length of the head anterior to the eyes; the length is about 2.2 to 2.3 times the maximum width; the length is 29 to 32 mm. The dorsal margin of the penis is nearly straight with an abrupt obtuse angle formed beyond the mid-length (Fig. 6.556). Color: mainly light brown or yellowish brown. ..............................................Belostoma pseudoguianae Roback and Nieser, 1974 (Colombia).

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- The first two segments of the rostrum are not subequal in length, or, if they are, then the interocular space is equal to 1.2 to 1.3 times the width of a compound eye, or the length is about 2.1 times the maximum width (Fig. 6.552). .............36 36. (35) The base of the clypeus is located at or proximal to the ocular line, that is, the line drawn between the anteriormost corners of the compound eyes (Fig. 6.537). .................................................................................................................37 - The base of the clypeus is at least slightly anterior to the ocular line (Fig. 6.543). .................................................................................................................38 37. (36) The first two segments of the beak are subequal. The base of the clypeus is only slightly anterior to the ocular line (Fig. 6.543). Length: 27 to 31.5 mm. .........................................................................Belostoma venezuelae Lauck, 1962 (Venezuela, Ecuador). - The first segment of the beak is 1.2 times the length of the second. The body is widest near its mid-length (Fig. 6.557). Length: 25 to 27.5 mm. .........................................................................Belostoma asiaticum (Mayr, 1863) (Ecuador). Syn: Zaitha asiatica Mayr, 1863; Zaitha anura Distant, 1891 (nec Herrich-Schäffer, 1848); Zaitha boops Campos, 1919. This species was initially reported from Borneo, a mistake apparently caused by a missing label.

Fig. 6.557 Belostoma asiaticum (left to right): habitus in dorsal view, head in dorsal view (lower left center), penis in dorsal, ventral, and lateral view. Based on De Carlo (1930) and Lauck (1962).

Fig. 6.558 Habitus of Belostoma elongatum in dorsal view (left), and penis in dorsal (upper right), ventral (middle right), and lateral view (lower right). Based on Lauck (1962) and Ribeiro (2007).

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38. (36) The body is elongate, 2.5 times as long as it is wide (Fig. 6.558). Length: 33 to 41 mm. Color: generally brown with yellow markings. The ventral surface of the thorax is dark chocolate brown or nearly black with brown or yellow patches. .................................................................Belostoma elongatum Montandon, 1908 (North and Central America, West Indies, Colombia, Peru, Bolivia, Paraguay, Argentina, Uruguay, Rondônia, Minas Gerais, Mato Grosso, Mato Grosso do Sul). Syn: Belostoma boscii De Carlo, 1930 (nec Herrich-Schäffer, 1848). - The body is ovate, not more than 2.25 times as long as wide (Fig. 6.559). ....39 39. (38) The first two segments of the beak are subequal in length (Fig. 6.559). Length: 29.5 to 35.5 mm. Color: mainly dark chocolate brown, either uniform or with a mottled pattern. .............................................................................Belostoma guianae Lauck, 1962 (Guyana, Surinam). - The first segment of the beak is obviously longer than the second (Fig. 6.560). Length: 31.8 to 34.6 mm. Color: mainly brown. ...........................................................................Belostoma dallasi De Carlo, 1930 (Argentina, Ceará, Minas Gerais, Goiás). Syn: Zaitha anurus Berg, 1879 (nec Herrich-Schäffer, 1848) probably pars; Belostoma boscii Kirkaldy and Bueno, 1909 (nec Herrich-Schäffer, 1848); Belostoma bosqi De Carlo, 1960 (nec De Carlo, 1932).

Fig. 6.559 Belostoma guianae male (left to right): habitus and penis in dorsal, ventral, and lateral view. Based on Nieser (1975).

Fig. 6.560 Belostoma dallasi (left to right): head in dorsal view and the penis in dorsal, ventral, and lateral view. Based on Lauck (1962).

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40. (18) There is an obvious thickening along the dorsolateral margins of the ventral diverticulum of the male genitalia (Fig. 6.524). The length does not exceed about 17 mm. ..........................................................................................41 - No thickening is evident along the dorsolateral margins of the ventral diverticulum of the male genitalia, or the length is equal to or greater than c. 18 mm (Fig. 6.561). .................................................................................................43

Fig. 6.561 Belostoma denticolle (above, left to right): habitus in dorsal view and head in dorsal and lateral view, and (below, left to right): male genitalia in dorsal view, area covered by fine setae on the lateral surface of the abdomen, and male genitalia in ventral and lateral view. Based on Estévez and Polhemus (2001).

Fig. 6.562 Belostoma lariversi (above, left to right): head in dorsal and lateral view, habitus in dorsal view, the extent of the ventral pilosity on one side of the abdomen marked in black, and (below, left to right): the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007).

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41. (40) The length of the first segment of the rostrum is 0.88 to 0.95 the length of the second. The interocular space is 1.5 to 1.7 times the width of an eye (Fig. 6.524). Length: 10 to 13 mm. Color: yellowish brown to light brown. .......................................................................Belostoma pygmeum (Dufour, 1863) (Paraguay, Peru, Bolivia, Amazonas). - The length of the first segment of the rostrum is 0.80 to 0.87 the length of the second (Fig. 6.562). ............................................................................................42 42. (41) The prosternal carina is not prominent. In ventral view, the length of the ventral diverticulum of the phalosoma is three times its length, and the width of its dorsal arms is less than that of the phalosoma at its base (Fig. 6.562). Length: 15 to 17 mm. Color: yellowish brown. .........................................................................Belostoma lariversi DeCarlo, 1960 (Peru). - The prosternal carina is prominent. In ventral view, the length of the ventral diverticulum of the phalosoma is about 2.5 times its length, and the width of its dorsal arms equals that of the phalosoma at its base (Fig. 6.563). Length: 11.5 to 14 mm. Color: brown with yellow mottling. ...................................................Belostoma nicaeum Estéves and Polhemus, 2007 (Amazonas).

Fig. 6.563 Belostoma nicaeum (above, left to right): head in dorsal and lateral view, habitus in dorsal view, the extent of the ventral pilosity on one side of the abdomen marked in black, and (below, left to right): the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007). 43. (40) The posterior margin of the clypeus is located along the imaginary line drawn between the inner anterior corners of the compound eyes. The coloration is almost uniformly brownish black on the dorsal side and dark brown ventrally, but there are yellow or blackish markings on the membranes of the hemielytra, which display a network of veins. There are yellow markings on the last three connexival segments, and there are yellow bands on both sides of the predominantly blackish legs. There is a faint median, longitudinal carina on the vertex, but it does not reach the posterior margin of the head. The prosternal

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keel is very narrow and has a rounded apex. The first segment of the beak is somewhat shorter than the second. Length: c. 18 to 19.5 mm. Maximum width: 9.5 to 10 mm. ....................................................................Belostoma bachmanni De Carlo, 1957 (Paraguay). - The posterior margin of the clypeus is usually located anterior or posterior to the imaginary line drawn between the inner anterior corners of the compound eyes. In case of doubt, the coloration is mainly brownish, the legs are not obviously banded, or there are no contrasting yellow and black marks on the apical segments of the abdomen. Only a few species reach 18 mm in length, and these are either narrower or their prosternal keel is wider and more prominent (Fig. 6.564). ........................................................................................................44 44. (43) The ventral diverticulum of the male genitalia is flat, spatulate, or discshaped (Fig. 6.564). ............................................................................................45 - The ventral diverticulum of the male genitalia is not at all flattened (Fig. 6.565). .................................................................................................................54

Fig. 6.564 Belostoma orbiculatum (above, left to right): habitus in dorsal view with the area on the lateral surface of the abdomen covered by fine setae shown to the lower right, the head in dorsal and lateral view, and (below, left to right): male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2001).

Fig. 6.565 Belostoma bicavum (left to right): habitus and penis in dorsal, ventral, and in lateral view with the apex pointing left. Based on Nieser (1975).

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45. (44) The space between the inner anterior corners of the compound eyes equals about 1.25 times the width of a compound eye. The ventral diverticulum is spatulate (Fig. 6.564). .....................................................................................46 - The space between the inner anterior corners of the compound eyes equals about 1.5 times the width of a compound eye. The ventral diverticulum is somewhat circular in shape (Fig. 6.566). ...........................................................50

Fig. 6.566 Belostoma sanctulum male (above, left to right), habitus without appendages in dorsal view and the head in dorsal and lateral view, and (below, left to right): one side of the abdomen in ventral view showing the extent of the pilose area, and the ventral diverticulum of the penis in dorsal, ventral, and lateral view. Based on Ribeiro and Estévez (2009). 46. (45) The prosternal keel is simple and rounded at its apex. .........................47 - The prosternal keel is acute at its apex, and it obviously projects anteriad. ...48 47. (46) The prosternal keel is prominent and rounded. The arms of the phalosoma are shorter than its posterior part (Fig. 6.564). Length of male: 13.4 to 14.6 mm; female: 13.5 to 14.7 mm. Color: brown with few discernible markings. .............................................Belostoma orbiculatum Estévez and Polhemus, 2001 (Argentina, Uruguay, Rio de Janeiro, Santa Catarina). - The prosternal keel is not prominent and simply rounded. The arms of the phalosoma are longer than its posterior part (Fig. 6.567). Length: c. 11.0 mm. Maximum width: 5 to 6 mm. Color: very dark with alternating light and dark bands on the dorsal part of the connexivum. ......................................................Belostoma loprettoai Estévez and Reyes, 2009 (Argentina). 48. (46) The prosternal keel appears low in lateral view, with its height 1/3 of its length or less. The maxillary plate is compressed (Fig. 6.568). Length of male: 11 to 12 mm; female: 11.5 to 12.0 mm. Color: brown with few discernible markings. ...................................................Belostoma retusum Estévez and Polhemus, 2001 (Paraguay, Argentina).

428

- The prosternal keel appears high in lateral view, with its height 2/3 or more of its length. The maxillary plate is not compressed (Fig. 6.569). ........................49

Fig. 6.567 Belostoma loprettoai male (above, left to right): habitus without appendages in dorsal view and the head in dorsal and lateral view, and (below, left to right): the extent of the pilosity on the underside of the abdomen and the ventral diverticulum of the penis in dorsal, ventral, and lateral view. Based on Ribeiro and Estéves (2009).

Fig. 6.568 Belostoma retusum (above, left to right): habitus in dorsal view with the area on the lateral surface of the abdomen covered by fine setae shown to the lower right, head in dorsal and lateral view, and (below, left to right): male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2001). 49. (48) The suture joining the anteclypeus and maxillary plate is slightly shorter than the anteclypeal suture. The ventral diverticulum is spatulate (Fig. 6.561). Length of male and female: 12 to 14 mm. Color: Light or dark brown, usually with brownish yellow markings. ..................................................................Belostoma denticolle Montandon, 1903 (Colombia, Guyana, French Guiana, Surinam, Venezuela, Amapá, Pará, Amazonas).

429

- The suture joining the anteclypeus and maxillary plate and the anteclypeal suture are about equal in length (Fig. 6.569). Length of male: 13 to 15 mm; female: 14.5 to 16.0 mm. Color: uniform light brown. ...............................................Belostoma amazonum Estévez and Polhemus, 2001 (Venezuela, Amazonas, Bahia, Santa Catarina).

Fig. 6.569 Belostoma amazonum (above, left to right): habitus in dorsal view with the area on the lateral surface of the abdomen covered by fine setae shown to the lower right, head in dorsal and lateral view, and (below, left to right): male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2001). 50. (45) About half of the genital operculum is covered by pilosity, and the pilose area on the ventral surface of the lateral parts of the abdominal segments covers more than half of the entire area. The anterior anteocular space is about 1.5 times the width of an eye. The ventral diverticulum is almost circular (Fig. 6.26). Length: 16 to 17 mm. ........................................................................Belostoma oxyurum (Dufour, 1863) (Uruguay, Argentina, Paraná, Rio Grande do Sul). Syn: Zaitha oxyura Dufour, 1863. - Less than half of the genital operculum is covered by pilosity. The wings do not completely cover the abdomen (Fig. 6.566). ................................................51 51. (50) The anteocular length and interocular width are approximately equal. The length of the pronotum measured along the dorsal mid-line is slightly less than half of its maximum width due to a fold in the middle of its anterior margin (Fig. 6.566). Length of males and females: 14 to 19 mm. Width of male: 6.6 to 7.8 mm; female: c. 8.5 mm. .................................................................Belostoma sanctulum Montandon, 1903 (Argentina, Espírito Santo, Rio de Janeiro, São Paulo, Santa Catarina). Syn: Belostoma amici Piza-Jr., 1975 (pars). - The anteocular length is shorter than the interocular width. Measured along the dorsal mid-line, the pronotum is longer than half of its maximum width, and there is no fold in the middle of its anterior margin (Fig. 6.570). ......................52

430

Fig. 6.570 Belostoma horvathi male (above, left to right), habitus without appendages in dorsal view and the head in dorsal and lateral view, and (below, left to right): one side of the abdomen in ventral view showing the extent of the pilose area, and the ventral diverticulum of the penis in dorsal, ventral, and lateral view. Based on Ribeiro and Estévez (2009). 52. (51) The apex of the prosternal keel is obtuse, truncate, or occasionally tuberculated. The pilosity covering the abdomen clearly does not reach the abdominal suture. The dorsal arms of the phallotheca of the male genitalia are parallel or convergent (Fig. 6.570). The second segment of the beak is slightly longer than the first. Length: 14 to 19 mm. ....................................................................Belostoma horvathi Montandon, 1903 (Argentina, Bahia, Goiás, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina, Rio Grande do Sul). Syn: Belostoma oxyurum Nieser and de Melo, 1997 (nec Dufour, 1863); Belostoma sanctulum Ribeiro, Nessimian, and Mendonça, 1998 pars (nec Montandon, 1903). - The apex of the prosternal keel has a slightly rounded, nearly acute apex. The first segment of the beak is equal to or shorter than the second (Fig. 6.571). The pilosity covering the abdomen almost reaches the abdominal suture. The dorsal arms of the phallotheca of the male genitalia are divergent. ...................53 53. (52) The margin of the prosternal keel displays distinct angles along its curve. The dorsal arms of the ventral diverticulum are slender (Fig. 6.571). The first segment of the rostrum is subequal to the second. Length: 15 to 20 mm. ...............................................................Belostoma candidulum Montandon, 1903 (Argentina, Uruguay, Goiás, Minas Gerais, São Paulo, Rio de Janeiro, Santa Catarina, Rio Grande do Sul). Syn: Belostoma horvathi De Carlo, 1938 (nec Montandon, 1903); Belostoma machrisi Menke and Lauck, 1962; Belostoma amici Piza-Jr., 1975 (pars); Belostoma sanctulum Ribeiro, Nessimian, and Mendonça, 1998 pars (nec Montandon, 1903).

431

- The margin of the prosternal keel has smooth, even curves. The dorsal arms of the ventral diverticulum are enlarged, so that in dorsal veiw, they completely cover the sides of the diverticulum itself (Fig. 6.572). Length: 15.0 to 17.5 mm. Maximum width: c. 6.4 to 7.0 mm. .................................................Belostoma carajaensis Ribeiro and Estévez, 2009 (Pará).

Fig. 6.571 Belostoma candidulum: lateral profile of the prosternal ridge (left) and the penis in lateral view (right). Based on Ribeiro (2005).

Fig. 6.572 Belostoma carajaensis male (left to right): prosternal keel in lateral view and the ventral diverticulum of the penis in dorsal, ventral, and lateral view. Based on Ribeiro and Estéves (2009). 54. (44) The interocular space is nearly twice the width of one eye (Fig. 6.565). The prosternal keel is triangular with an acute apex. .........................................55 - The interocular space is no greater than 1.5 times the width of one eye. The prosternal keel is semicircular (Fig. 6.573). Most South American specimens are shorter than 16 mm. ......................................................................................56 55. (54) The first two rostral segments are subequal. The anteocular and interocular distances are subequal (Fig. 6.574). ........................................................................Belostoma triangulum Lauck, 1964 (Peru, Mato Grosso). - The first rostral segment is shorter than the second. The length of the head anterior to the eyes is about 1.25 times the space between the inner anterior corners of the eye (Fig. 6.565). Length: 19 to 20 mm. Color: light to medium brown, often mottled with yellow. The legs are chestnut brown with yellowish bands and patches. .............................................................................Belostoma bicavum Lauck, 1964 (Amazonas). Nieser (1975) considers Belostoma parvoculum Lauck, 1964, which was described from one specimen, to be a synonym of B. bicavum.

432

Fig. 6.573 Belostoma estevezae: habitus without appendages in dorsal view (left), prosternal keel (upper center), and penis in dorsal (upper right), ventral (lower right), and lateral view (lower center). Based on Ribeiro and Alecrim (2008).

Fig. 6.574 Belostoma triangulum male (left to right): penis in dorsal, ventral, and lateral view. Based on Lauck (1964). 56. (54) The length of the first segment of the beak is 0.80 to 0.87 the length of the second (Fig. 6.575). ..................................................................................57 - The length of the first segment of the beak is 0.88 to 0.95 the length of the second (Fig. 6.576). ............................................................................................58

Fig. 6.575 Belostoma minusculum (above, left to right): head in dorsal and lateral view, habitus in dorsal view, the extent of the ventral pilosity on one side of the abdomen marked in black, and (below, left to right): the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007).

433

Fig. 6.576 Belostoma plebejum (above, left to right): head in dorsal and lateral view, habitus in dorsal view, and (below, left to right): the extent of the ventral pilosity on one side of the abdomen marked in black and the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007).

Fig. 6.577 Belostoma parvum (above, left to right): head in dorsal and lateral view, habitus in dorsal view, the extent of the ventral pilosity on one side of the abdomen marked in black, and (below, left to right): the male genitalia in dorsal, ventral, and lateral view. Based on Estévez and Polhemus (2007). 57. (56) The laterodorsal margins of the ventral diverticulum of the penis are prominent and well defined; the dorsal arms of the diverticulum converge gradually (Fig. 6.573). Length: c. 11.3 mm. The female has not been described. ....................................................Belostoma estevezae Ribeiro and Alecrim, 2008 (Rondônia).

434

- The laterodorsal margins of the ventral diverticulum of the penis are inconspicuous and poorly defined; the dorsal arms of the diverticulum are narrow and convergent (Fig. 6.575). Length: 11.5 to 14 mm. Color: dark brown. .....................................................................Belostoma minusculum (Uhler, 1884) (Central America, Trinidad, Venezuela). Syn: Zaitha minuscula Uhler, 1884; Belostoma drakei De Carlo, 1950. 58. (56) The prosternal carina narrows and projects slightly anteriad (Fig. 6.577). Total length: 9.5 to 11.1 mm. ....................................................Belostoma parvum Estévez and Polhemus, 2007 (Colombia, Venezuela, Guyana, Surinam, Pará). - The prosternal carina is rounded and does not project anteriad (Fig. 6.576). The total length is at least 10.1 mm and frequently exceeds 11 mm. .................59 59. (58) The lateromedian section of the laterodorsal margins of the ventral diverticulum display an accentuated curvature, giving the diverticulum an elliptical shape. The ventro-apical portion of the diverticulum bears a prominent protuberance (Fig. 6.576). Length: 13 to 17 mm. .............................................................................Belostoma plebejum (Stål, 1860) (Venezuela, Peru, Bolivia, Paraguay, Argentina, Uruguay, Bahia, Pará, Amazonas, Mato Grosso, Goiás, Minas Gerais, Espírito Santo, São Paulo, Rio de Janeiro, Santa Catarina). Syn: Zaitha plebeja Stål, 1860; Zaitha maculosa Dafour, 1863; Zaitha limbata Defour, 1863; Zaitha adusta Dufour, 1863; Zaitha difficilis Dufour, 1863; probably Zaitha micantula Dufour, 1863 (nec Stål); Belostoma amici Piza-Jr., 1975 pars (some of the paratypes). - The lateromedian section of the laterodorsal margins of the ventral diverticulum show a slight curvature, making the diverticulum narrowly elliptical in shape. The ventro-apical portion of the diverticulum bears only a vestigial protuberance (Fig. 6.578). Length: 10 to 13 mm. ...............................60

Fig. 6.578 Belostoma nessimiani male: habitus without appendages in dorsal view with the position of the wings somewhat distorted (left), outline of the prosternal carina in lateral view (upper right center), and penis in dorsal (upper right), ventral (middle right), and lateral view (lower right). Based on Ribeiro and Alecrim (2008).

435

60. (59) The median and dorsal depression of the ventral diverticulum of the male genitalia is well developed, and its dorsal arms diverge (Figs. 6.27, 6.334). Length: 10.4 to 13.3 mm. Maximum width: 5.0 to 6.3 mm. Color: dark brown. ........................................................................Belostoma micantulum (Stål, 1858) (Colombia, Venezuela, Guyana, Surinam, Bolivia, Paraguay, Uruguay, Argentina, Amapá, Ceará, Tocantins, Pará, Amazonas, Mato Grosso, Mato Grosso do Sul, Goiás, Minas Gerais, Rio de Janeiro, Rio Grande do Sul). Syn: Zaitha micantula Stål, 1858; Zaitha zelotypus White, 1879; Belostoma husseyi De Carlo, 1960; Belostoma apache De Carlo, 1960 (nec Kirkaldy, 1908). - The median and dorsal depression of the ventral diverticulum of the male genitalia is poorly developed, and its dorsal arms abruptly converge (Fig. 6.578). Length: 10.1 to 11.8 mm. ..................................................Belostoma nessimiani Ribeiro and Alecrim, 2008 (Pará, Amazonas).

Gelastocoridae The toad bugs usually inhabit the littoral zone of water bodies, where their form, coloration, and coating of aufwuchs provide them with camouflage. They are amphibious, generally hopping on moist ground near water or on wetlands but also entering shallow water to avoid predators. Some species of Nerthra, however, seem to have adopted terrestrial habits and are often found far from water. Some may burrow in the sediment. Most species are active mainly during the daytime, relying on their camouflage for protection. Their hopping movements and general shape cause them to be confused with small toads just after metamorphosis, which may be an additional aid for survival. Key to subfamilies and genera of adults in South America The information for the key was provided by Melin (1929) and Todd (1955). 1. The fore-tarsus is free to articulate and not fused to the tibia. Adults bear two well-developed tarsal claws on the forelegs. The rostrum arises from the apex of the head; it is stout and recurved posteriorly (Fig. 6.579). Larvae closely resemble adults (Fig. 6.580). ...........................................................................Subfamily Gelastocorinae..p. 437 The only genus in the Western Hemisphere is Gelastocoris Kirkaldy, 1897. - The fore-tarsus is fused to the tibia and cannot articulate. Adults bear one well-developed tarsal claw on each foreleg. The rostrum appears to arise from the ventral side of the head; it is slender and projects ventrally or anteriorly (Fig. 6.581). Larval instars have been described (Fig. 6.337). ..................................................................................Subfamily Nerthrinae..p. 447 The only genus in the Western Hemisphere is Nerthra Say, 1832.

436

Fig. 6.579 Gelastocoris willneri: habitus of a male (upper left) and a female in dorsal view (lower left), male genital capsule (upper right), and the right part of the ovipositor (lower right). Based on De Carlo (1960c).

Fig. 6.580 Habitus of a larval Gelastocoris major. Based on Melin (1929), who referred to it by its synonym, Gelastocoris andinus peruensis.

437

Fig. 6.581 Nerthra praecipua female: habitus in dorsal view (below) and (above, left to right): head in dorsal view, trochanter of the fore-leg showing the black anterior denticles, and the apex of the abdomen in dorsal, ventral, and lateral view. Based on Todd (1957b) and Polhemus and Cekalovic (1998). Subfamily Gelastocorinae Key to the species of Gelastocoris reported from South America The information for the key was provided by Martin (1928), Melin (1929), Todd (1955; 1957b, c; 1961a, b), De Carlo (1960c), Nieser (1977b), Schnack and Estéves (1979b), and Estéves and Schnack (1977, 1980b). There has been some controversy about whether several species described by DeCarlo (1954b) are valid, with differing opinions being proposed by Todd (1955) and Schnack and Estévez (1979b). Melin (1929) believed that Gelastocoris angulatus should be placed in a separate genus, Montandonius, but Todd (1955) disagreed. While there are differences between G. angulatus and the other species of the genus, they still do not seem sufficient for the taxon Montandonius to merit more than subgeneric status, if it should be considered valid at all. For the sake of completeness, the key includes two species that have been reported from South America but probably do not occur there at all (Todd, 1955). Both sexes of some species may not yet be identifiable with certainty. Due to disagreements among taxonomists in the past resulting in conflicting opinions, the identities of individual species may be difficult to determine. 1. Along the posterior border of the pronota of both males and females, there are six to eight longitudinal carinae (Fig. 6.582). The ventral apex of the head is truncate. The dorsal surface is rough, and a few bright, pearl-like structures are present on the hemielytra. .....................................................................................2

438

- No carinae are present on the basal part of the pronotum of either sex. The ventral apex of the head is acutely pointed (Fig. 6.583). .....................................5

Fig. 6.582 Gelastocoris angulatus male (left to right): pronotum in dorsal view; genitalia; apex of the abdomen in ventral view; outline of the habitus in dorsal view showing the claval suture. Based on Todd (1955) and De Carlo (1960c).

Fig. 6.583 Gelastocoris hungerfordi male: pronotum in dorsal view (left) and genitalia (right). Based on Todd (1955).

Fig. 6.584 Gelastocoris bridarollii: habitus in dorsal view (left) and the right part of the ovipositor (right). Based on De Carlo (1960c).

439

Fig. 6.585 Gelastocoris mansosotoi female: habitus in dorsal view (left) and the right part of the ovipositor (right). Based on De Carlo (1960c). 2. (1) The claval suture is present (Fig. 6.582). ...................................................3 - The claval suture is absent (Fig. 6.584). ...........................................................4 3. (2) The anterior margin of the pronotum is strongly convex (Fig. 6.582). The hemielytral sutures are petiolate. Length of male: 7.4 to 8.8; female: 8.1 to 10.6 mm. .....................................................................Gelastocoris angulatus (Melin, 1929) Range of the species, sensu Todd, 1955: Guyana, Surinam, Bolivia, Paraguay, Mato Grosso, Amazonas, Pará, Minas Gerais, Rio de Janeiro, Bahia. Range sensu Schnack and Estéves, 1979b: Amazonas, Pará. Syn. Montandonius angulatus Melin, 1929; Gelastocoris flavus Martin, 1929 auctt. - The anterior margin of the female pronotum is only slightly convex. The hemielytral sutures are not petiolate, and sometimes the entire suture is obsolete (Fig. 6.585). Length: 9.4 mm. The male has not been described. .....................................................Gelastocoris mansosotoi (De Carlo, 1954) pars (Bolivia, Amazonas). Syn: Montandonius mansosotoi De Carlo, 1954. Todd (1955) reduced this specific name to a junior synonym of G. angulatus. Schnack and Estéves (1979b) disagreed. 4. (2) The posterior margin of the pronotum has eight short longitudinal carinas. The hemielytra cover the lateral parts of the posteriormost sternites (Fig. 6.579). Length: 9.8 mm. ...................................................................Gelastocoris willneri (De Carlo, 1954) (Paraguay, Mato Grosso). Syn: Montandonius willneri De Carlo, 1954. Todd (1955) reduced this species to a junior synonym of G. angulatus. Schnack and Estéves (1979b) disagreed.

440

- The posterior margin of the pronotum has six short longitudinal carinas. The hemielytra do not cover the lateral parts of the posteriormost sternites (Fig. 6.584). Length: 9.8 mm. ...............................................................Gelastocoris bridarollii (De Carlo, 1954) (Argentina). Syn: Montandonius bridarolli De Carlo, 1954. Todd (1955) reduced this species to a junior synonym of G. angulatus. Schnack and Estéves (1979b) disagreed.

Fig. 6.586 Gelastocoris major male: pronotum in dorsal view (left) and male genitalia (right). Based on Todd (1955). 5. (1) The length of the membrane on the hemielytron is subequal to or less than the width of the hind femur. The male genitalia have a bilobed pan (Figs. 6.579, 6.586). .....................................................................Gelastocoris major Montandon, 1910 (Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Chile, Brazil). Syn: Galgulus flavus Campos, 1919; Gelastocoris duplicatus Martin, 1928; Gelastocoris andinus Melin, 1929; Gelastocoris andinus peruensis Melin, 1929; Gelastocoris incasiana Poisson, 1954. - The length of the membrane on a hemielytron is more than twice the width of the hind femur, or, if not, the pan of the male genitalia is not bilobed (Fig. 6.587). ...................................................................................................................6

Fig. 6.587 Gelastocoris oculatus male: habitus without appendages in dorsal (left) and with legs on one side in ventral view (right). Based on Todd (1955).

441

Fig. 6.588 Gelastocoris amazonensis male: pronotum in dorsal view (left) and genitalia (right). Based on Todd (1955). 6. (5) The pronotal disc is slightly elevated. The tumescence of the right paramere of the male is not adnate to the base of the paramere, and the keel hook is not recurved back to the base of the pan. A secondary keel hood is prominent, or, if not, then the pan of the male genitalia is very thin, elongate, narrow, and twisted apically, turned completely over (Fig. 6.588). ....................7 - The pronotal disc is very prominently elevated, and the male genitalia are not as above (Figs. 6.17, 6.587). ................................................................................8

Fig. 6.589 Gelastocoris quadrimaculatus: the black and pinkish yellow thorax and abdomen in dorsal view (upper left), lateral margins of the thoraces of two males (lower left), membrane of a hemielytron (lower center), and the male genital capsules of four different specimens (right). Based on De Carlo (1960c) and Nieser (1972).

442

7. (6) The anterolateral margin of the pronotum does not have a triangular projection and forms an obtuse angle with the mediolateral margin. The posterolateral margin is dentate. The fringe of the keel hood of the male genitalia is bent at the apex toward the midline and is obliquely concave to the right margin of the keel apex. A secondary keel hood is prominent (Fig. 6.583). Length of male: 5.9 to 6.5 mm; female: 6.2 to 7.4 mm. ....................................................................Gelastocoris hungerfordi Melin, 1929 (Mexico, Central America, Colombia, Venezuela, Acre). Syn: Gelastocoris cucullatus Martin, 1929. - The anterolateral margin of the pronotum has a triangular projection and forms a right angle with the mediolateral margin. The pan of the male genitalia is very thin, elongate, narrow, and twisted apically, turning completely over (Fig. 6.588). Length of male: 7.9 to 8.2; female: 7.5 to 8.6 mm. ...................................................................Gelastocoris amazonensis Melin, 1929 (Amazonas).

Fig. 6.590 Gelastocoris fuscus: pronotum (left) and male genitalia (right). Based on Todd (1955). 8. (6) The anterolateral margin of the pronotum has a triangular projection elevated above the plane of the pronotum where it borders on the head; the projection may be rounded off (Fig. 6.587). ........................................................9 - The anterolateral margin of the pronotum usually has a triangular projection, but it is not oriented vertically enough to elevate it above the plane of the anterior border with the head (Fig. 6.589). ........................................................10 9. (8) The keel hood of the male genitalia is located on the right side of the keel (Fig. 6.590). Length of male: 7.3 to 8.2 mm; female: 8.4 to 9.5 mm. ............................................................................Gelastocoris fuscus Martin, 1928 (Ecuador, Peru, Surinam, Chile, Paraguay, Bolivia, Amazonas). Syn: Gelastocoris martinezi De Carlo, 1954; Gelastocoris problematicus Poisson, 1954.

443

- The keel hood is either located on the left side or covers the entire apex. The pan is more or less thickened and elongate, not distinctly in a plane oblique to that of the genital capsule (Figs. 6.17, 6.587). Length of male: 5.7 to 6.5 mm; female: 6.6 to 7.5 mm. ..................................................................Gelastocoris oculatus (Fabricius, 1798) (North America, Colombia, Argentina?). Only one subspecies, Gelastocoris oculatus variegatus (Guérin-Méneville, 1844), has been reported to occur in South America, perhaps as far south as Argentina (Champion, 1901). 10. (8) The dorsal surface of the head, pronotum, and hemielytra is rough and pustulated; the hemielytra bear a few large, light bluish, pearl-like structures. The head at the base of the eyes is enlarged. The lateral margins of the pronotum bear only a few minute teeth. Length: c. 6.3 to 6.5 mm. The width of the head, including the eyes, is c. 2.7 to 2.8 mm. The embolium is slightly inflated along its lateral margin at the widest point of the body, which is about 3.7 to 3.9 mm (Fig. 6.332). The general color is yellow to light brown. ...............................Gelastocoris vandamepompanoni Boulard and Jauffret, 1984 (Pará). - The dorsal surfaces of the head, pronotum, and hemielytra are not notably rough and pustulated, and any brightly-colored granules present are small, not pearl-like, or not bluish (Fig. 6.589). .................................................................11 11. (10) The dorsal coloration is black with pinkish yellow markings covering the head, lateral parts of the pronotum, small markings on the scutellum, and extreme edges of the hemielytra. The ventral side of the abdomen is black at the base with the sutures between the apical sternites yellow or pinkish. The internal lateral tuberculated elevations on the pronotum are separated from each other but not greatly elevated (Fig. 6.589). The large, colored granules are greenish. Length: 6.7 to 8.1 mm. ......................................Gelastocoris quadrimaculatus (Guérin-Méneville, 1844) (Peru, Bolivia, Argentina). Syn: Galgulus quadrimaculatus Guérin-Méneville, 1844. - The general coloration is brownish or yellowish with variable patterns, sometimes including a longitudinal black stripe on the pronotum. ....................12

Fig. 6.591 Gelastocoris decarloi male (left to right): pronotum in dorsal view, scutellum, a hemielytron showing the locations of the colored granules, and apex of the abdomen in ventral view. Based on Estéves and Schnack (1977).

444

12. (11) The anterolateral margin of the pronotum diverges only slightly posteriad and forms an angle slightly more than 90º with the strongly divergent anterior margin of the posterolateral expansion of the pronotum, which is denticulated along its lateral margin. The scutellum has long, narrow anterolateral extensions. The lateral external elevations of the pronotum are broadened anteriorly and bear denticles (Fig. 6.591). Length: 6.9 to 7.6 mm. ..................................................Gelastocoris decarloi Estéves and Schnack, 1977 (Santa Catarina). - The anterolateral margin of the pronotum forms an angle more than 100º with the moderately divergent anterior margin of the posterolateral expansion of the pronotum (Fig. 6.592). .......................................................................................13 13. (12) The median sulcus of the pronotum is clearly delimited, even if fairly shallow (Fig. 6.592). ..........................................................................................14 - The median sulcus of the pronotum is faintly delimited or incomplete (Fig. 6.593). .................................................................................................................16

Fig. 6.592 Gelastocoris flavus: pronotum in dorsal view (left) and the male genitalia (right). Based on illustrations labelled Gelastocoris nebulosus by Todd (1955).

Fig. 6.593 Gelastocoris vicinus male: pronotum in dorsal view (left) and genitalia (right). Based on Todd (1955).

445

14. (13) The posterolateral margin of the pronotum appears truncate and finely denticulated. There are two granules between the hemielytral sutures. The tumescence on the right paramere is adnate to the base of the paramere. The keel hook is recurved completely back over the base of the pan (Fig. 6.592). Length of male: 5.5 to 8.0 mm; female: 6.4 to 8.8 mm. ........................................................Gelastocoris flavus (Guérin-Méneville, 1835) (Peru, Ecuador, Venezuela, Bolivia, Paraguay, Uruguay, Argentina, Amazonas, Pará, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, São Paulo, Santa Catarina?). Syn: Galgulus flavus Guérin-Méneville, 1835; Galgulus nebulosus Guérin-Méneville, 1844; Gelastocoris stali Torre-Bueno, 1909; Gelastocoris quadrimaculatus Martin, 1929 (nec Guérin-Méneville, 1835); Gelastocoris vianai De Carlo, 1954. Probable synonyms include Gelastocoris quadrimaculatus Martin, 1928 (nec Guérin-Méneville, 1844), and Gelastocoris apureensis Melin, 1929. In addition, several species were placed in synonymy by Todd (1955) but reinstated as valid by Schnack and Estéves (1979b) and are found in this key. - The posterolateral margin on the pronotum is broadly rounded at the anterior corner and bears variable sized teeth (Fig. 6.594). .............................................15

Fig. 6.594 Gelastocoris paraguayensis (left to right): lateral outline of the thorax of a male (lower left) and a female (upper left), the seventh abdominal segment (center), and the genital capsule (right). Based on De Carlo (1960c). 15. (14) The anterior and anterolateral edges of the pronotum are straight (Fig. 6.594). Length: 7 mm. ...........................................................Gelastocoris paraguayensis De Carlo, 1954 (Paraguay, Mato Grosso). - The anterior edge of the pronotum is concave, and the anterolateral edge is straight anteriorly and indented posteriorly (Fig. 6.595). Length: 7.7 mm. .................................................................Gelastocoris bolivianus De Carlo, 1954 (Bolivia).

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16. (13) The front edge of the head is smooth. There are no blister-like granules between the hemielytral sutures. The posterior margin of the seventh abdominal sternite of the male is sinuous with a deep excavation along the midline (Fig. 6.596). Length: 6.8 to 7.5. .....................................................................Gelastocoris monrosi De Carlo, 1959 (Santa Catarina, São Paulo). - The front edge of the head is quite rough. The tumescence of the right paramere is not adnate to the base of the paramere. The keel hook is not recurved back to the base of the pan, which is flattened and wider than long. It lies in a plane oblique to the horizontal plane of the genital capsule (Fig. 6.593). Length of male: 6.2 to 7.1 mm; female: 7.1 to 8.0 mm. .....................................................................Gelastocoris vicinus Champion, 1901 (Champion reported the species from the southern and western United States to Brazil, but Todd [1955] doubted that the species actually occurs anywhere in South America).

Fig. 6.595 Gelastocoris bolivianus: left side of the habitus showing the location of the granules on the hemielytra (lower left), lateral outline of the thorax of a male (upper left) and a female (middle left), membrane of the hemielytra (upper center), and the genital capsule (right). Based on De Carlo (1960c).

Fig. 6.596 The seventh abdominal sternite of a male Gelastocoris monrosi. Based on Nieser (1972).

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Subfamily Nerthrinae Key to the species of Nerthra reported from South America Information for the key was provided by Todd (1955), Schnack and Estévez (1979b), and Estévez and Schnack (1980c, d). There are many synonyms for the Nerthra species in South America. Synonyms for the generic name include Mononyx Laporte, 1833; Phintius Stål, 1861; Matinus Stål, 1861; Peltopterus Guérin-Méneville, 1843; and Scylaecus Stål, 1861. These names appear frequently in the older literature. 1. Ocelli are absent. The hemielytron has a membrane, which is reduced or entirely coriaceous. The urosternites of the female are symmetrical or nearly so (Fig. 6.597). ..........................................................................................................2 - Ocelli are present. The membrane is well developed (Fig. 6.598). .................5

Fig. 6.597 Habitus of Nerthra rugosa without appendages. Based on Todd (1955).

Fig. 6.598 Nerthra gaucha: outline of the anterior margin of the head between the compound eyes (left), pronotum of a male (left center), scutellum (center), hemielytron (lower right center), right paramere of the male enlarged (upper center to right center), apices of the abdomens of a male in ventral view (upper right) and a female (lower right). Based on Estéves and Schnack (1980d).

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Fig. 6.599 Nerthra americana: right paramere of a male (left) and habitus (right). Based on Todd (1955). 2. (1) The margins of the pronotum are not differentiated (Fig. 6.597). Length of female: 6.0 to 7.1 mm. ..........................................................................Nerthra rugosa (Desjardins, 1837) (Australia, Mauritius, New Guinea, Florida, Central America, Brazil). Syn: Naucoris rugosa Desjardins, 1837. - The margins of the pronotum are differentiated (Fig. 6.599). ..........................3 3. (2) The head bears supra-apical tubercles and has a concave excavation at the apex. There are two lateral longitudinal and one apical elevation on the scutellum. The hemielytra have both longitudinal and transverse carinae. The connexivum protrudes considerably and is serrate. The trochanter of the foreleg bears two black denticles at the anterior end (Fig. 6.581). Length of female: 8.2 mm. ................................................................................Nerthra praecipua Todd, 1957 (Chile). - The head lacks supra-apical tubercles and is pointed at the apex. There are no elevations on the scutellum. The hemielytra lack carinae (Fig. 6.599). ..............4 4. (3) The length exceeds 8 mm. The last three sternites of the male are small. The right side of the seventh sternite is spatulate and partially covered by the sixth sternite. The male paramere is simple, sickle-shaped, and has spiniform bristles (Fig. 6.600). .................................................................................Nerthra williamsi Todd, 1955 (São Paulo). - Length: less than 7.5 mm. The last three sternites of the male are large. The right side of seventh sternite is completely covered by the sixth. The male paramere bears a large dorsomedial process and clavate bristles (Fig. 6.599). Length of male: 6.3 to 6.6 mm: female: 6.6 to 7.1 mm. ..................................................................Nerthra americana (Montandon, 1905) (Espirito Santo, São Paulo).

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Fig. 6.600 Right paramere of a male Nerthra williamsi. Based on Todd (1955). 5. (1) The edge of the anterior dilation of the fore-femur nearly forms a right angle with the posterior side of the femur, and the length is shorter than about 6 mm (Fig. 6.601). ...................................................................................................6 - The edge of the anterior dilation of the fore-femur forms an acute angle with the posterior side of the femur, or the length exceeds 6 mm (Fig. 6.602). ..........7

Fig. 6.601 Nerthra raptoria (left to right): apical part of the fore-leg, right paramere of the male, and apical abdominal sternites of a female. Based on Todd (1955).

Fig. 6.602 Nerthra fuscipes (left to right): anterior margin of the head, apical segments of the male abdomen, right paramere of a male, and apex of the abdomen of a female in ventral view. Based on Todd (1955). 6. (5) The anterior basal angle of the anterior dilation of the fore-femur is more or less rounded. There is an elevation on the head between the eyes and a tumescence on either side of the emargination on the ventral posterior margin of

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the last abdominal segment of the female (Fig. 6.601). Length of male: 5.0 to 5.5 mm: female: 5.4 to 5.5 mm. ..........................................................................Nerthra raptoria (Fabricius, 1803) (Mexico, Central America, Colombia, Venezuela, French Guiana, Surinam, Argentina, Pará, Amazonas, Mato Grosso) - The anterior basal angle of the anterior dilation of the fore-femur is pointed. There is no tumescence on either side of the emargination on the ventral posterior margin of the last abdominal segment of the female. The hemielytral membrane does not extend as far as the apex if the abdomen (Fig. 6.603). Length of female: 4.2 mm. .....................................................................................Nerthra buenoi Todd, 1955 (Pernambuco, Goiás).

Fig. 6.603 Nerthra buenoi: apical part of the fore-leg (left) and apical abdominal segments of a female in ventral view. Based on Todd (1955). 7. (5) The eighth abdominal sternite of the male is twice as long as the ninth sternite. The ovipositor of the female is small and as wide as long (Fig. 6.602). Length of male: 8.2 to 10 mm; female: 8.8 to 11 mm. ............................................................Nerthra fuscipes (Guérin-Méneville, 1843) (Numerous from Mexico to Colombia; one isolated individual has been reported from Rio de Janeiro). - The eighth abdominal sternite of the male is almost subequal to or shorter than the ninth. The ovipositor of the female of most species is longer than wide, (Fig. 6.604). ..........................................................................................................8 8. (7) The scutellum bears prominent coats or patches of black bristles. .............9 - No patches of black bristles are on the scutellum. ..........................................14 9. (8) There are narrow longitudinal bands of bristles on the scutellum. ...........10 - The bristles on the scutellum do not form bands and are usually in oval patches or form a more extensive coating of the surface. ..................................12 10. (9) The posterior part of the abdomen is not strongly dilated laterally (Fig. 6.604). Length: 7.2 to 7.7 mm. .....................................................................Nerthra unicornis (Melin, 1929) pars (French Guiana, Surinam, Paraguay, Pará, Rio Grande do Sul). - The posterior part of the abdomen is strongly dilated laterally (Fig. 6.605). ...11

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Fig. 6.604 Nerthra unicornis male (left to right): anterior margin of the head, apical portion of the paramere, and apex of the abdomen in ventral view. Based on Todd (1955).

Fig. 6.605 Nerthra montandoni (left to right): enlarged apical portion of the male paramere and ventral views of the apices of the male and female abdomen in ventral view. Based on Todd (1955).

Fig. 6.606 Nerthra peruviana (left to right): apical portion of the paramere and apex of the male and female abdomen in ventral view. Based on Todd (1955). 11. (10) The bristles on the scutellum are short and clavate (Fig. 6.605). .........................................................................Nerthra montandoni (Melin, 1929) (Trinidad, Venezuela, Colombia). - The bristles on the scutellum are long and scarcely or not at all clavate (Fig. 6.606). ...................................................................Nerthra peruviana (Montandon, 1905) (Peru, Argentina, Ecuador, Bolivia). Syn: Mononyx peruviana Montandon, 1905. 12. (9) The apex of the head is projected to a point; the tubercles at its apex are fused basally. The urosternites of the female are asymmetrical (Fig. 6.604). .....................................................................Nerthra unicornis (Melin, 1929) pars (French Guiana, Surinam, Paraguay, Pará, Rio Grande do Sul). - The apex of the head is concavely excavated. The urosternites of the female are symmetrical (Fig. 6.598). .............................................................................13

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13. (12) The anterior margin of the head projects relatively slightly in the middle, and the lateral parts of this margin are generally concave (Fig. 6.14). The larvae have been described (Fig. 6.337). ........................................................Nerthra ranina (Herrich-Schäffer, 1853) pars (Colombia, Argentina, Chile, Peru, Paraguay, Bolivia, Santa Catarina, Minas Gerais, Pará, São Paulo). Syn: Mononyx melini Poisson, 1954; Mononyx melini forma siviae Poisson, 1954. Also see Couplet 17. - The anterior margin of the head projects considerably beyond its lateral margins, leaving the lateral parts generally convex (Fig. 6.598). Length: 7.2 to 8.4 mm. Maximum width: 4.4 to 6.0 mm. The bristles on the scutellum cover the entire area. Deficiencies in the original description will make this species difficult to identify. ...........................................................Nerthra gaucha Estéves and Schnack, 1980 (Rio Grande do Sul).

Fig. 6.607 Nerthra amplicollis (left to right): right paramere of a male and the apical abdominal segments of a male and a female in ventral view. Based on Todd (1955).

Fig. 6.608 Nerthra parvula male: apical part of the right paramere (left) and apical abdominal segments of a male in ventral view (right). Based on Todd (1955). 14. (8) The apex of the head is not projected to a point but rather concavely excavated. The urosternites of the female are symmetrical (Fig. 6.607). .........15 - The apex of the head projects to a point and terminates in one or more apical tubercles (Fig. 6.608). ........................................................................................19 15. (14) The apical half of the male paramere is simply sword or sickle-shaped (Fig. 6.607). For female specimens, compare the descriptions of the known females in the three following couplets. .............................................................16 - The apical half of the male paramere is bifurcate, or it bears a dorsomedial process, which is much smaller than the rest of the paramere (Fig. 6.609). ......18

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Fig. 6.609 Nerthra ecuadorensis (left to right): portion of the right paramere of the male genitalia and apex of the male and the female abdomen in ventral view. Based on Todd (1955). 16. (15) The apical half of the male paramere is bifurcate (Fig. 6.607). The left ovipositor lobe of the female is not twisted at its apex to cover the apex of the right lobe. The apex of the head is broadly and concavely excavated with a short, broad tubercle on each side of the excavation. Lateral tubercles are indistinct. Length 10.8 to 14.7 mm. ..............................................................................Nerthra amplicollis (Stål, 1854) (Costa Rica, Colombia, Venezuela, Ecuador, Peru, Bolivia). Syn: Mononyx amplicollis Stål, 1854. - The apical half of the male paramere is not bifurcate but has a dorsomedial process, much smaller than the main part of the paramere (Fig. 6.608). ...........17

Fig. 6.610 Apical abdominal segments of a female Nerthra ater in ventral view. Based on Todd (1955), who referred to it by its synonym, Nerthra rudis. 17. (16) The lateral margins of the male pronotum are rounded and convex outward (Fig. 6.14). The last visible abdominal sternite of the female has an emargination that is prolonged as a long groove curving to the right, forming a lobe or flap. ........................................................Nerthra ranina (Herrich-Schäffer, 1853) pars (Colombia, Argentina, Chile, Peru, Paraguay, Bolivia, Santa Catarina, Minas Gerais, Pará, São Paulo). Syn: Mononyx melini Poisson, 1954; Mononyx melini forma siviae Poisson, 1954. Also see Couplet 13. - The lateral margins of the male pronotum are straight or slightly sinuous. The male paramere curves more than 90º in its basal half (Fig. 6.608). The urosternites of the female are symmetrical. Length of male: 5.8 to 6.1 mm; female: 7.0 to 7.5 mm. Also see Couplet 21. ....................................................................Nerthra parvula (Signoret, 1864) pars (Chile).

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18. (15) The dorsomedial process of the male paramere is cylindrical or flattened in a plane that is radial to the longitudinal axis of the paramere (Fig. 6.609). There are only about two tubercles on the anterior margin of the head. The left ovipositor lobe of the female is not twisted at its apex to cover the apex of the right lobe. Length: 8.1 to 10.2 mm. .......................................................................Nerthra ecuadorensis (Melin, 1929) (Panama, Colombia, Ecuador, Peru). Syn: Mononyx amplicollis var. ecuadorensis Melin, 1929. - The dorsomedial process of the male paramere is flattened in a plane tangential to the longitudinal axis of the paramere. The left ovipositor lobe of the female is twisted to the right at its apex to cover the apex of the right lobe (Fig. 6.610). Length of female: 8.9 to 10.6 mm. Color: yellowish brown to black with a small yellowish marking at the posterior corner or each connexival segment and sometimes a darker color on the scutellum. ......................................................................................Nerthra ater (Melin, 1929) (Mexico, Colombia, Ecuador, Bolivia). Syn: Mononyx ater Melin, 1929; Mononyx fuscipes var. rudis Melin, 1929; Nerthra rudis (Melin, 1929). 19. (14) Superapical tubercles are usually present but may be small and inconspicuous or even absent, especially in the male. .......................................20 - Superapical tubercles are completely absent. ..................................................22 20. (19) The posteriormost abdominal sternite of the female lacks obvious depressions or tumescences and is almost symmetrical. The male has not been described. .............................................................................Nerthra lata (Montandon, 1899) (Ecuador). Syn: Mononyx latus Montandon, 1899; Mononyx latus Torre-Bueno, 1906. - The posteriormost abdominal sternite of the female has obvious depressions or tumescences and is clearly asymmetrical. ..........................................................21 21. (20) One superapical tubercle is usually present along the anterior margin of the head, but it may be inconspicuous. The apical half of the male paramere is simple and more or less sword or sickle-shaped. The urosternites of the female are symmetrical. Length of male: 5.8 to 6.1 mm; female: 7.0 to 7.5 mm. Also see Couplet 17. ....................................................................Nerthra parvula (Signoret, 1864) pars (Chile). - Superapical tubercles are present and distinct along the anterior margin of the head of the male, which bears a total of five tubercles. The female may lack apical tubercles or bear only one. The apical abdominal segments are relatively symmetrical (Fig. 6.611). Length 7.5 to 8.2 mm. ...................................................................Nerthra quinquedentata (Melin, 1929) (Argentina, Chile, Santa Catarina). 22. (19) The posteriormost abdominal sternite of the male is very large, about 2/5 as wide as the fifth sternite (Fig. 6.612). The anterior part of the emargination on the posteriormost visible abdominal sternite of the female forms an acute angle. ..........................................................................................23

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- The posteriormost abdominal sternite of the male is only about 1/4 as wide as the fifth sternite. The anterior part of the emargination on the last visible abdominal sternite of the female is rounded (Fig. 6.613). .................................24

Fig. 6.611 Apical segments of a male Nerthra quinquedentata. Based on Todd (1955).

Fig. 6.612 Nerthra terrestris (left to right): right paramere of the male, apical segments of the abdomen of a male and a female in ventral view. Based on Todd (1955).

Fig. 6.613 Nerthra nepaeformis: anterior margin of the head (left), apical segments of the male abdomen in ventral view (upper center), right paramere of a male (lower left to center), and apex of the abdomen of a female in ventral view (right). Based on Todd (1955). 23. (22) The lateral margins of the pronotum converge noticeably anteriorally and distinctly project beyond the base of the embolium at the level of the

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transverse furrow. The right paramere of the male curves more than 90º near its base (Fig. 6.612). Length: 8.2 to 8.8 mm. .............................................................................Nerthra terrestris (Kevan, 1948) (Trinidad, Guyana, French Guiana, Surinam, Colombia, Ecuador, Peru, Bolivia, Venezuela, Amazonas, Goiás, Mato Grosso do Sul, São Paulo). Syn: Mononyx bipunctatus Melin, 1929 preoccupied; Mononyx terrestris Kevan, 1948; Mononyx titschacki Poisson, 1954. - The lateral margins of the pronotum are more or less parallel and project only slightly beyond the base of the embolium at the level of the transverse furrow. The apex of the male right paramere is nearly straight but is excavated slightly on the ventral side (Fig. 6.614). ...............................................................................Nerthra borealis (Melin, 1929) (One specimen has been reported from Brazil without information on location). This species may very well be synonymous with N. terrestris.

Fig. 6.614 Apex of the right paramere of a male Nerthra borealis in lateral view. Based on Todd (1955). 24. (22) The basal third of the embolium is dilated, and its edges are rounded. The body is flattened. The greatest width of the male seventh abdominal sternite is less than half the width of the posterior margin of the fourth abdominal sternite. The apex of the right paramere of the male is knob-like (Fig. 6.613). ...................................................................Nerthra nepaeformis (Fabricius, 1775) (Mexico, Colombia, Chile, Argentina, Uruguay, Bahia, Rio de Janeiro, São Paulo, Santa Catarina). Syn: Naucoris nepaeformis Fabricius, 1775; Mononyx bipunctatus Stål, 1854. - The basal third of the embolium is narrowed, not rounded, and usually slightly sinuous. The body is not notably flattened. The greatest width of the seventh abdominal sternite of the male is approximately equal to 1/2 the width of the posterior margin of the fourth abdominal sternite (Fig. 6.615 ................................................................................Nerthra tenebrosa Todd, 1955 (Mexico, Belize, Panama, Colombia, Ecuador, Bolivia).

Fig. 6.615 Nerthra tenebrosa (left to right): right paramere of a male and apices of a male and a female abdomen in ventral view (right). Based on Todd (1955).

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Ochteridae There are only three genera presently recognized in this family: Megochterus Jaczewski, 1934, is confined to the Eastern Hemisphere, while Ochterus Latreille, 1807, and Ocyochterus Drake and Gomez-Menor, 1954, occur in South America. The ochterids are inhabitants of wetlands and moist littoral habitats. They fly away when disturbed and are therefore difficult to catch. Practically nothing is known with certainty about their ecology. Because of the dearth of information about these insects and the probability that the ranges of the species are greatly underestimated, the geographical area encompassed by the keys is somewhat increased. Key to the genera of adults in South America Information for the key was provided by Drake (1952d) and Drake and Gomez-Menor (1954). 1. The pronotal and hemielytral margins are widely expanded. The anterior margins of the eyes are obliquely and acutely rounded. The ocelli are widely separated (Fig. 6.616). Length: estimated to be about 6.5 to 7.5 mm. .........................................................Ocyochterus Drake and Gomez-Menor, 1954 The only species described is Ocyochterus victor (Bolivar, 1879), known from Ecuador. It is described as generally black with a violet tinge with yellow on marginal and transverse posterior lines, round spots above the eyes, and on the exterior, posterior margin. It also appears in the literature under its synonyms: Pelogonus victor Bolivar, 1879; Pelogonus splendidulus Montandon, 1898; Ochterus victor Torre Bueno, 1909; Ochterus splendidulus Schell, 1943. - The pronotal and hemielytral margins are not widely expanded. The anterior margins of the eyes are not almost acutely rounded. The ocelli are rather close together (Fig. 6.617). Length: less than 7 mm. ..............................................................................Ochterus Latreille, 1807..p. 458

Fig. 6.616 Ocyochterus victor: habitus in dorsal view (left) and the head in anterior view (right). Based on Drake and Gomez-Menor, 1954.

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Fig. 6.617 Habitus of Ochterus perbosci. Based on Nieser (1975). Key to the adult Ochterus species in the Western Hemisphere Information for the key was provided by Schell (1943), Kormilev and DeCarlo (1952), Drake (1958b), and Nieser (1975). The key is fully applicable only to males. 1. The anterior angles of the pronotum have pointed projections. There is a series of yellow spots along the exterior margin of the hemielytra (Fig. 6.617). Length: 6.5 mm. .........................................................Ochterus perbosci (Guérin-Méneville, 1843) (Texas, Mexico, West Indies, Central America, Surinam, Peru, Ecuador, Paraguay, Amazonas, Minas Gerais). Syn: Pelogonus perbosci GuérinMéneville, 1843. - The anterior angles of the pronotum do not bear projections (Fig. 6.618). ......2

Fig. 6.618 Habitus of Ochterus aeneifrons surinamensis. Based on Nieser (1975).

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2. (1) The frons is metallic green. .........................................................................3 - The frons is black or very dark. ........................................................................5

Fig. 6.619 The genital capsule of Ochterus hungerfordi. Based on Schell (1943). 3. (2) The labrum and rostrum are brown. The apex of the genital capsule is narrow and pointed, and it almost reaches the apex of the right paramere (Fig. 6.619). Length: c. 4 mm. .........................................................................Ochterus hungerfordi Schell, 1943 (Cuba; not known from South America). - The labrum and rostrum are light yellow. .........................................................4 4. (3) The right paramere of the male bears a retrorse prominence on the inner margin (Fig. 6.620). Length: 4.25 to 5.5 mm. .................................................................Ochterus viridifrons (Champion, 1901). (Guatamala to southern Panama; not yet reported from South America). - The right paramere of the male is club-shaped (Fig. 6.621). Length: 5.1 to 5.2 mm. ...............................................................Ochterus acutangulus (Champion, 1901). (Mexico, Panama).

Fig. 6.620 The genital capsule of Ochterus viridifrons. Based on Schell (1943).

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Fig. 6.621 The genital capsule of Ochterus acutangulus. (1943).

Based on Schell

5. (2) The entire dorsum is light brown. The right paramere of the male widens preapically (Fig. 6.622). Length: 3.3 mm. ..................................................................................Ochterus parvus Schell, 1943 (Ecuador). - At least part of the dorsum is black. ..................................................................6

Fig. 6.622 The genital capsule of Ochterus parvus. Based on Schell (1943).

Fig. 6.623 The genital capsule of Ochterus flaviclavus. Based on Schell (1943). 6. (5) The clavus is lemon yellow. The right paramere extends well beyond the apex of the genital capsule and has an extended process beyond its mid-point (Fig. 6.623). Length: 3.5 mm. ..........................................................................Ochterus flaviclavus Barber, 1913 (Florida).

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- The clavus is not yellow. ...................................................................................7 7. (6) The posterior half of the pronotum and hemielytra, except the membrane, are ferruginous, at least to a large part. ................................................................8 - The posterior half of the pronotum is not ferruginous. .....................................9 8. (7) There is a fine median carina on the vertex of the head. The coloration is variable. Length of male: 4.5 to 4.7 mm; female: 5.2 to 5.5 mm. ....................................................Ochterus foersteri Kormilev and DeCarlo, 1952 (Paraguay). - There is no median carina on the vertex of the head. The right paramere has blunt preapical teeth (Fig. 6.624) Length: 4.8 mm. .....................................................................Ochterus brunneus Hungerford, 1927 (Bolivia).

Fig. 6.624 The genital capsule of Ochterus brunneus. Based on Schell (1943). 9. (7) The lateral margin of the pronotum has an almost spherical, yellow spot near the anterior angle. The paramere extends well beyond the apex of the genital capsule (Fig. 6.625). Length: c. 6.5 mm. ........................................................................Ochterus americanus (Uhler, 1884) (North America, Cuba). - The lateral margins of the pronotum are broadly pale. ...................................10

Fig. 6.625 The genital capsule of Ochterus americanus. Based on Schell (1943).

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10. (9) The right paramere of the male has a distinct hook at the apex, which might not be visible from all sides (Fig. 6.626). ................................................11 - The right paramere lacks a hook at the tip (Fig. 6.627). .................................12 11. (10) There are bluish gray markings on a vivid brown background on the head behind the eyes, the disc of the pronotum, near the lateral margin of the pronotum, at the base of the scutellum, at the base of the clavus, and elsewhere (Fig. 6.618). Length: 3.5 to 5 mm. .................................................................Ochterus aeneifrons (Champion, 1901). (Mexico to Panama, West Indies, Surinam, Ecuador, Amazonas, Minas Gerais). Syn: Pelogonus aeneifrons Champion, 1901. Only the subspecies Ochterus aeneifrons surinamensis Neiser, 1975, is known from continental South America. It can be distinguished from the nominate subspecies by its entirely brownish posterior half of the pronotum, which is blackish except for the margin in O. a. aeneifrons. - There are at most faint gray markings on the dorsum (Fig. 6.626). Length: 5.1 to 5.6 mm. ...........................................................................Ochterus tenebrosus Nieser, 1975 (Surinam, Amazonas).

Fig. 6.626 Habitus of Ochterus tenebrosus. Based on Nieser (1975).

Fig. 6.627 The genital capsule of Ochterus bidentatus. Based on Schell (1943).

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12. (10) The right paramere of the male has two small teeth on the inner side (Fig. 6.627). Length: 4.7 mm. ............................................................................Ochterus bidentatus Schell, 1943 (Peru). - The right paramere lacks distinct teeth on the inner side (Fig. 6.628). ...........13 13. (12) The right paramere of the male has an enlarged tip and a series of very fine teeth on the outer side (Fig. 6.338). Length: 5 mm. ................................................................................Ochterus barberi Schell, 1943 (Arizona). - The right paramere does not bear a series of fine teeth (Fig. 6.628). .............14

Fig. 6.628 The genital capsule of Ochterus banksii. Based on Schell (1943). 14. (13) The right paramere of the male bears a heart-shaped lobe on the outer margin (Fig. 6.628). Length: 4 mm. ................................................................................Ochterus banksii Barber, 1913 (Eastern U.S.A.). - The right paramere does not bear a heart-shaped lobe on the outer margin (Fig. 6.629). Length: 4.1 mm. ..........................................................................Ochterus manni Hungerford, 1927 (Costa Rica, Peru, Bolivia).

Fig. 6.629 The genital capsule of Ochterus manni. Based on Schell (1943).

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Notonectidae Key to the South American genera The information used for the key was provided by Truxal (1953) and Nieser (1975). 1. The hemielytral commisure has a hair-lined pit at the anterior end. The ventral keel on the abdomen extends to the last segment of all known South American species. The male genital capsule is closed posteriorly. Males possess a stridular protuberance on the fore-tibia. Females have long gonapophyses. The anterior tarsi of the males of all South American species consist of two segments (Fig. 6.630). ........................Subfamily Anisopinae....................Buenoa Kirkaldy, 1904..p. 466 (This is an exclusively New World genus belonging to a nearly cosmopolitan subfamily. It encompasses many South American species). - The hemielytral commissure lacks a hair-lined pit at the anterior end (Fig. 6.631). .............................................................Subfamily Notonectinae. .........................2

Fig. 6.630 Buenoa paranensis male (above, left to right): outline of the anterior part of the body, third rostral segment of a male in anterior view, inner surface of the fore-leg, and the left (lower center) and right genital clasper (center) and the spine on the outer sinestral margin of the seventh abdominal segment (lower right). Based on Truxal (1953).

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Fig. 6.631 Notonecta virescens (above, left to right): habitus of a male with a color pattern originally ascribed to the form known by the synonym, N. bicirca; male genital capsules originally ascribed to N. bicirca, N. virescens, and N. bicircoidea; and (below, left to right): female habitus with color patterns originally ascribed to N. bicircoidea and N. virescens; middle leg with a typically rounded trochanter. Based on Hungerford (1933).

Fig. 6.632 Martarega mcateei: head and thorax in dorsal view (upper left); head of a male (lower left) and a female in lateral view (lower left center); hemielytron of a macropterous male (upper center), brachypterous male (center), and brachypterous female (upper right); male gonapophyses (lower right center); female genitalia (lower right). Based on Jaczewski (1928d).

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2. (1) The middle tibia lacks a pointed preapical protuberance, and the antenna of all South American species consists of four segments (Fig. 6.632). The antennae of some Old World species have three segments. ......................................Tribe Nychina..................Martarega White, 1879..p. 488 - The middle femur has a pointed protuberance before the apex, and the antenna always consists of four segments (Fig. 6.631). ........................................................................................Tribe Notonectini...........3 3. (2) The anteriolateral margins of the prothorax are not foveate (Fig. 6.631). ...........................................................................Notonecta Linnaeus, 1758..p. 493 - The anteriolateral margins of the prothorax are foveate (Fig. 6.633). ..............................................................................Enithares Spinola, 1837..p. 503

Fig. 6.633 The left lateral (left) and right lateral view (right) of the male genital capsule of Enithares braziliensis. Based on Brooks (1953). Key to the species Buenoa in South America The information for the keys was provided by Truxal (1953, 1957), Nieser (1970d, 1975), Argrisano (1982), Padilla and Nieser, (1992), Nieser et al. (1997), and Padilla-Gil (2002). Diagnostic features of many females have not been reported, so the key can be used with a reasonable degree of reliability only for males, although it can be used to tentatively identify females of a few species. 1. The synthlipsis of the male is one half or more of the anterior width of the vertex (Fig. 6.634). ...............................................................................................2 - The synthlipsis of the male is less than one half of the anterior width of the vertex (Fig. 6.635). .............................................................................................17 2. (1) The rostral prong is shorter than or the same length as the third rostral segment. The pronotum has very little sculpture and is not tricarinate. The length of the fore-femur is about twice its width at the apex (Fig. 6.636). ..........3 - The rostral prong is longer than the third rostral segment (Fig. 6.634). ...........6 3. (2) The fore-femur of the male lacks a stridulatory area (Fig. 6.636). .............4 - The fore-femur of the male has a stridulatory area (Fig. 6.637). ......................5

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Fig. 6.634 Buenoa serrana: head and pronotum in dorsal view (left), rostrum (beak) of a male in lateral view (upper center), male parameres (right center), penis (upper right), spine on the seventh sternite (middle right), fore-femur of a male (lower center), and valve of an ovipositor showing four coarse apical teeth and a row of smaller teeth ventral the coarse ones (lower right). Based on Angrisano (1982a).

Fig. 6.635 Buenoa konta male (left to right): head in dorsal view, lateral view of the rostrum (beak), fore-leg, and an enlargement of the tibial comb. Based on Nieser and Pelli (1994).

Fig. 6.636 Buenoa machrisi male (left to right): the tylus and rostrum (beak), enlarged left stridulatory comb on the fore-tibia, left fore-leg, and inner surface of the hind trochanter. Based on Truxal (1957).

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Fig. 6.637 Buenoa ida male: rostrum and tylus in left lateral view (left), inner surface of the left fore-leg with an enlargement of the tibial stridulatory comb in the area between the femur and tarsi (right). Based on Truxal (1953). 4. (3) The head of the female, measured along the midline, is about 0.8 times the length of the pronotum. There are 15 to 25 teeth in the tibial comb on the foreleg (Fig. 6.638). Length of male: 9.2 to 9.9 mm; female: 8.7 to 9.9 mm. ............................................Buenoa oreia Nieser, Melo, Pelli, and Barbosa, 1997 (Minas Gerais). - The heads of the male and the female are about 0.5 times the length of the pronotum. The tibial comb on the fore-tibia consists of 14 to 16 stout teeth (Fig. 6.636). Length of male: 8.8 to 9.0 mm; female: 8.3 to 8.9 mm. Maximum width of male: 2.4 to 2.6 mm; female: 2.5 to 2.7 mm. ................................................................................Buenoa machrisi Truxal, 1957 (Goiás). 5. (3) The heads of males and females are about 2/3 as long as the pronotum. There are about 4 to 6 weak ridges in the stridulatory area on the fore-femur and about 24 denticles in the stridulatory comb on the fore-tibia (Fig. 6.639). Length of macropterous male: c. 5.9 mm; macropterous female: 5.9 to 6.2 mm. ......................................................................Buenoa koina Nieser and Pelli, 1994 (Minas Gerais). - The head of the female, measured along the midline, is less than half as long as the pronotum. There are about 25 teeth in the tibial comb on the fore-leg (Fig. 6.637). Length of male: 9.1 to 9.7 mm; female: 9.2 to 10.0 mm. Maximum width of male: 2.7 to 2.9 mm; female: 3.1 to 3.3 mm. ..............................................................................Buenoa ida Kirkaldy, 1904 pars (Mexico, Guatamala, Uruguay).

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Fig. 6.638 Buenoa oreia: rostrum of a male in lateral view (upper left), inner surface of the left fore-leg of a male with enlargements of the tibial stridulatory combs of two specimens in the area between the femur and tarsi (center), the tarsus of the male fore-leg in posterior view (right center to lower center), left paramere of a male (upper right), spine from the left-posterior edge of the seventh abdominal tergite of a male (upper middle right), hind trochanter of a female showing the stridular area on its inner surface (lower middle right), and the female ovipositor (lower right). Based on Truxal (1953).

Fig. 6.639 Buenoa koina male (left to right): lateral view of the rostrum (beak), fore-leg, and an enlargement of the tibial comb. Based on Nieser and Pelli (1994). 6. (2) The base of the rostral prong is located laterally or near the distal end of the third rostral segment (Fig. 6.634). ..................................................................7 - The base of the rostral prong is not located laterally or near the distal end of the third rostral segment (Fig. 6.640). ................................................................12 7. (6) Length of male: at least 7.0 mm. The female is at least 7.1 mm. ..............8 - The length is less than 7.0 mm. .........................................................................9

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Fig. 6.640 Buenoa pallens male (left to right): enlargement of the rostral prong, a somewhat varying rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargements of two slightly different tibial stridulatory combs. Based on Truxal (1953). 8. (7) The fore-femur of the male has an obvious prominence on the anterior surface of the distal third (Fig. 6.634). Length of male: 7.0 to 7.8 mm; female: 7.1 to 7.8 mm. ............................................................................Buenoa serrana Angrisano, 1982 (Argentina). - The fore-femur of the male does not have a prominence on the anterior surface of the distal third, or if there is a slight elevation, it is barely discernible. The posterior margin of the rostral prong is almost straight (Fig. 6.641). Length of male: 7.1 to 7.7 mm; female: 7.2 to 8.2 mm. .......................................................................Buenoa crassipes (Champion, 1901) (Guatamala, Costa Rica, Ecuador, Paraná, Santa Catarina). Syn: Anisops crassipes Champion, 1901.

Fig. 6.641 Buenoa crassipes male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, enlargement of the tibial stridulatory comb, and the spine at the left posterior corner of the seventh abdominal segment. Based on Truxal (1953).

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9. (7) The head is at least half the length of the pronotum measured along the median longitudinal axis, and it is narrower than the humeral width of the pronotum. The pronotum is not tricarinate. A stridulatory area is present on the inner surface of the fore-femur (Fig. 6.642). Length of male: 5.5 to 5.9 mm; female: 5.5 to 6.4 mm. Maximum width of male: 1.4 to 1.7 mm; female: 1.4 to 1.8 mm. Color: mainly black but usually with a white or testaceous head, anterolateral part of the pronotum, ventral surface of the thorax, legs, ventral keel on the abdomen, and posterior abdominal segment. .........................................................................Buenoa macrotrichia Truxal, 1953 (Peru). - The head is about 1/3 or less the length of the pronotum measured along the median longitudinal axis (Fig. 6.643). ...............................................................10

Fig. 6.642 Buenoa macrotrichia male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953).

Fig. 6.643 Buenoa rostra male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and an enlargement of the tibial stridulatory comb. Based on Truxal (1953).

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Fig. 6.644 Buenoa pallipes male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and an enlargement of the tibial stridulatory comb. Based on Truxal (1953). 10. (9) The rostrum is robust with the posterior margin of the rostral prong distinctly sinuate. The tylus is only slightly inflated (Fig. 6.643). Length of male: 5.7 to 6.5 mm; female: 5.8 to 6.3 mm. The head, pronotum, ventral surface of the thorax, and limbs are white to testaceous. The dorsal surface of the metathorax is yellowish to dark brown. The abdomen is mainly black with a yellowish white dorsum. The color pattern is variable. .............................................................................Buenoa rostra Truxal, 1953 pars (Trinidad, Venezuela). Also see Couplet 28. - The rostrum is not robust, and the posterior margin of the rostral prong is almost straight. The tylus is distinctly inflated (Fig. 6.644). ............................11 11. (10) In dorsal view, the head is about six times as wide as the anterior width of the vertex, and the vertex appears slightly indented (Fig. 6.644). Length of male: 5.5 to 6.3 mm; female: 5.6 to 6.5 mm. Maximum width of male: 1.5 to 2.0 mm; female: 1.6 to 2.0 mm. Color: white with variable black markings. ...........................................................................Buenoa pallipes (Fabricius, 1803) (Hawaii, Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Paraguay). Syn: Notonecta pallipes Fabricius, 1803; Anisops pallipes Stål, 1868. - In dorsal view, the head is about five times as wide as the anterior width of the vertex, and the vertex appears protuberant (Fig. 6.645). Length of male: 5.8 to 6.6 mm; female: 5.7 to 7.0 mm. Maximum width of male: 1.5 to 1.8 mm; female: 1.5 to 2.1 mm. The head and pronotum are white or yellowish white. The ventral surface of the thorax and the legs are light brown to testaceous. The scutellum is white, sometimes with black at its posterior apex. The abdomen is black with white or testaceous dorsal markings and ventral keel. The color pattern shows considerable variation. ................................................................................Buenoa absidata Truxal, 1953 (Peru).

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Fig. 6.645 Buenoa absidata male: rostrum and tylus in left lateral view (left), inner surface of the left fore-leg with an enlargement of the tibial stridulatory comb in the area between the femur and tarsi (right). Based on Truxal (1953).

Fig. 6.646 Buenoa femoralis male: rostrum and tylus in left lateral view (upper left), inner surface of the left fore-leg with an enlargement of the tibial stridulatory comb in the area between the femur and tarsi (center), and the spine at the left posterior corner of the seventh abdominal segment (right). Based on Truxal (1953). 12. (6) The posterior margin of the tibia bears more than 40 setae in the ventral row. The greatest width of the head is 6 times the anterior width of the vertex (Fig. 6.640). Length of male: 5.5 to 6.9 mm; female: 5.9 to 7.2 mm. Maximum width of male: 1.4 to 1.6 mm; female: 1.7 to 2.1 mm. The head, thorax, and legs are white; darker specimens have more extensive testaceous markings on these parts. The scutellum is usually entirely black with a violate tinge. The dorsal surfaces of the metathorax and the abdomen are black, except for a testaceous ventral keel and sometimes the apical segments of the abdomen. ..........................................................................Buenoa pallens (Champion, 1901) (Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Chile, Amazonas, Minas Gerais). Syn: Anisops pallens Champion, 1901.

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- The posterior margin of the tibia bears 40 or fewer setae in the ventral row (Fig. 6.646). .................................................................................................................13 13. (12) The length exceeds 7.75 mm. The posterior margin of the hind femur bears fewer than 30 large setae in the ventral row (Fig. 6.646). ........................14 - The length is less than 7.75 mm. The posterior margin of the hind femur bears 30 or more small setae in the ventral row (Fig. 6.647). .....................................16

Fig. 6.647 Buenoa tarsalis male (left to right): rostrum and tylus in left lateral view, inner surfaces of the middle tarsus and the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953).

Fig. 6.648 Buenoa antigone antigone: rostrum and tylus in left lateral view (left), inner surface of the left fore-leg of a male with an enlargement of the tibial stridulatory comb in the area between the tibia and tarsi (right), and the ovipositor valve of a female (upper center). Based on Truxal (1953).

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14. (13) The greatest width of the head is about seven times that of the anterior vertex; the head is at least half the length of the pronotum, measured along the median longitudinal axis (Fig. 6.646). Length of male: 8.3 to 8.7 mm; female: 8.7 to 8.9 mm. .............................................................................Buenoa femoralis (Fieber, 1851) (Peru, Paraná). Nieser et al. (1997) surmised that the specimens with labels stating “Porto Rico,” were found near the town of that name along the Paraná River in Brazil and not on the island of Puerto Rico. Syn: Anisops femoralis Fieber, 1851. - The greatest width of the head is six times or less that of the anterior vertex; the head is 1/3 the length of the pronotum measured along the median longitudinal axis (Fig. 6.648). ............................................................................15 15. (14) The pronotum is tricarinate. The fore-femur is three times as long as its width at the apex or longer. The tibial comb consists of 40 to 50 teeth (Fig. 6.648). Length of male: 7.8 to 9.0 mm; female: 8.2 to 9.8 mm. ..........................................................................Buenoa antigone (Kirkaldy, 1899) (Mexico, Central America, West Indies, Ecuador, Peru, Bolivia, Paraguay, Argentina, Rio Grande do Norte, São Paulo, Santa Catarina). Syn: Anisops antigone Kirkaldy, 1899. Two subspecies have been described: B. antigone antigone is the only one reported from South America. B. antigone carinata (Champion, 1901) is known only from North and Central America and is now considered to be a distinct species. - The pronotum has very little sculpture and is not tricarinate. The fore-femur is twice its width at the apex. There are about 25 and always fewer than 30 teeth in the tibial comb (Fig. 6.637). Length of male: 9.1 to 9.7 mm; female: 9.2 to 10.0 mm. Maximum width of male: 2.7 to 2.9 mm; female: 3.1 to 3.3 mm. ..............................................................................Buenoa ida Kirkaldy, 1904 pars (Mexica, Guatamala, Uruguay). 16. (13) The pronotum is distinctly tricarinate. The fore-femur is narrowed at its apex (Fig. 6.647). Length of male: 6.2 to 7.2 mm; female: 6.3 to 8.1 mm. Maximum width of male: 1.6 to 2.0 mm; female: 2.0 to 2.6 mm. Color: white to testaceous, with the head, pronotum, ventral side of the thorax, and legs usually white but sometimes with brown or black on the legs and ventral side of the thorax. The abdomen is black with white on the ventral keel and parts of the dorsum. ..................................................................................Buenoa tarsalis Truxal, 1953 (Pará, Ceará, Pernambuco, Rio Grande do Norte, Parhaíba, Minas Gerais, Rio de Janeiro). - The pronotum has very little trace of coarse sculpture, and it is not tricarinate. The fore-femur is widened at its apex (Fig. 6.649). Length of male: 6.1 to 6.9 mm; female: 6.5 to 7.1 mm. Maximum width of male: 1.7 to 1.9 mm; female: 1.9 to 2.1 mm. Color: variable from pale testaceous to black, sometimes with white legs and white ventral thoracic surfaces. .............................................................................Buenoa nitida Truxal, 1953 pars (Peru, Amazonas, Espirito Santo).

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Fig. 6.649 Buenoa nitida male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953). 17. (1) A stridulatory area is present on the inner surface of the fore-femur (Fig. 6.635). .................................................................................................................18 - There is no stridulatory area on the inner surface of the fore-femur (Fig. 6.650). .................................................................................................................29

Fig. 6.650 Buenoa salutis (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg of a male, and enlargement of the male tibial stridulatory comb. Based on Nieser (1975). 18. (17) The tylus is flat and bears setae, and the stridulatory area on the foretibia consists of about 19 to 22 sclerotized ridges or denticles (Fig. 6.651). .....19 - The tylus is not flat and hairy (Fig. 6.652). ....................................................20

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19. (18) Length of male: 5.9 to 6.0 mm; female: 5.9 to 6.1 mm. The tylus is flat with short setae at the base and long, hair-like setae along the lateral margins (Fig. 6.651). The stridulatory area consists of 19 to 22 sclerotized ridges. ...................................................................................Buenoa fittkaui Nieser, 1970 (Goiás). - Length of brachypterous male: 3.9 to 4.3 mm; brachypterous female: 4.2 to 4.6 mm. The tylus is flat with a few hair-like setae on the anterior surface. The lengths of the rostral prong and third rostral segment are approximately equal (Fig. 6.635). The fore-femur lacks a stridulatory area, and the stridulatory comb on the fore-tibia consists of about 22 denticles. ......................................................................Buenoa konta Nieser and Pelli, 1994 (Minas Gerais).

Fig. 6.651 Buenoa fittkaui male (left to right): rostrum and tylus in left lateral view, enlargement of the tibial stridulatory comb, and inner surface of the left fore-leg. Based on Nieser (1970d). 20. (18) The fore-femur is obviously narrowed at the apex; its length is more than three times the width at its apex (Fig. 6.652). ............................................21 - The fore-femur is widened or only slightly narrowed at the apex; its length is three times or less than the width at its apex (Fig. 6.653). .................................25 21. (20) The rostral prong is equal to or shorter than the third rostral segment (Fig. 6.652). The pronotum is tricarinate. The synthlipsis is less than 1/3 of the anterior width of the vertex. There are four short, peg-like setae at the apex of the fore-tibia inserted on the interior surface. The stridulatory area consists of six to nine sclerotized ridges. Length of male: 5.3 to 6.1 mm; female: 5.5 to 6.2 mm. Maximum width of male: 1.3 to 1.5 mm; female: 1.3 to 1.7 mm. Color: mainly white, except for extensive black ventral markings and yellow dorsal markings on the abdomen. ..................................................................................Buenoa gracilis Truxal, 1953 (Mexico, Central America, West Indies, Colombia, Peru, Venezuela). - The rostral prong is longer than the third rostral segment (Fig. 6.654). .........22

478

Fig. 6.652 Buenoa gracilis male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, group of peg-like setae on the inner surface of the fore-tibia near the apex, and enlargement of the tibial stridulatory comb. Based on Truxal (1953).

Fig. 6.653 Buenoa doesburgi male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and an enlargement of the tibial stridulatory comb. Based on Nieser (1975). 22. (21) The pronotum is distinctly tricarinate (Fig. 6.654). Length of male: 5.3 to 6.1 mm; female: 5.9 to 6.5 mm. Maximum width of male: 1.4 to 1.8 mm; female: 1.6 to 2.0 mm. The head, pronotum, ventral surface of the thorax, and legs are white or testaceous, except for the middle part of the pronotum, which is occasionally orange. The scutellum is orange, sometimes with black or brown markings. The humeral area of each hemielytron is sometimes rufescent, and the abdomen is light brown or black with testaceous areas along the ventral keel, on the connexivium, and on the two apical segments. ..............................................................................Buenoa communis Truxal, 1953 (Bolivia, Surinam, Pará, Paraíba).

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- The pronotum has little notable coarse sculpture; it is not tricarinate (Fig. 6.655). The stridulatory area on the fore-femur is subtriangular. .....................23

Fig. 6.654 Buenoa communis male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953).

Fig. 6.655 Buenoa triangularis male (left to right): the tylus and rostrum (beak), the left stridulatory comb on the tibia enlarged, and the left fore-leg. Based on Truxal (1957). 23. (22) The humeral width of the pronotum is more than twice its median length. The stridulatory area on the fore-femur consists of 5 to 7 sclerotized ridges opposing 15 to 25 teeth in the stridulatory comb on the fore-tibia (Fig. 6.655). Length of male: 6.3 to 6.8 mm; female: 6.6 to 7.0 mm. Greatest width of male: 1.3 to 2.0 mm; female: 1.9 to 2.0 mm. The general color is variable from testaceous to blackish violet or occasionally almost entirely yellowish white except for the black abdomen. ...........................................................................Buenoa triangularis Truxal, 1957 (Goiás).

480

- The width of the pronotum is considerably less than twice its median length. The stridulatory comb on the tibia bears from about 25 to as many as 38 teeth (Fig. 6.656). ........................................................................................................24 24. (23) The width of the pronotum is about 1 2/3 times its median length. The comb on the fore-tibia bears about 25 setaceous teeth (Fig. 6.630). Length: 6.0 to 6.3 mm. Color: yellowish white with dark eyes and black markings on both the dorsal and ventral surfaces of the abdomen. .......................................................................Buenoa paranensis Jaczewski, 1928 (Paraná, Goiás, Minas Gerais). The two species in this couplet are suspiciously similar, but determining synonymy will be difficult because the type specimens of Jaczewski have apparently been destroyed (Truxal, 1953). - The width of the pronotum is from about 1 1/2 to 1 4/7 of its median length. The comb on the fore-tibia consists of about 33 to 38 teeth, the apical ones larger than the proximal ones. The posterior margin of the tibia bears fewer than 40 setae in the ventral row. The greatest width of the head is 6½ times the anterior width of the vertex (Fig. 6.656). Length of male: 5.2 to 6.0 mm; female: 5.6 to 6.3 mm. Maximum width of male: 1.4 to 1.9 mm; female: 1.6 to 1.9 mm. The head, anterior part of the pronotum, ventral surface of the thorax, and legs are white or testaceous. The abdomen is usually black. The rest of the insect is variable and may be black or pale. ...............................................................................Buenoa mutabilis Truxal, 1953 (West Indies, Venezuela, Guyana, Peru, Paraguay, Goiás, Minas Gerais).

Fig. 6.656 Buenoa mutabilis male (left to right): apex of the rostrum, somewhat varying rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953). 25. (20) The head is equal to or slightly wider than the humeral width of the pronotum, which is not tricarinate. .....................................................................26 - The head is narrower than the humeral width of the pronotum. .....................27

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26. (25) The tibial comb of the male consists of 31 to 33 teeth (Fig. 6.649). Length of male: 6.1 to 6.9 mm; female: 6.5 to 7.1 mm. Maximum width of male: 1.7 to 1.9 mm; female: 1.9 to 2.1 mm. Color: variable from pale testaceous to black, sometimes with white legs and ventral thoracic surfaces. .............................................................................Buenoa nitida Truxal, 1953 pars (Peru, Amazonas, Espirito Santo). - The tibial comb of the male consists of 24 to 27 teeth (Fig. 6.653). Length of male: 5.8 to 6.1 mm; female: 6.0 to 6.3 mm. ..............................................................................Buenoa doesburgi Nieser, 1968 (Surinam). 27. (25) The rostral prong forms a distinct anterolateral angle (Fig. 6.21). The synthlipsis of the male is 0.4 times the anterior width of the vertex. The forefemur is only slightly widened at the apex. The stridular area on the fore-femur of the male is triangular and consists of 17 to 21 pegs. The stridular area on the tibia consists of 18 to 21 pegs. Length of male: 5.5 to 6.8 mm; female: 5.7 to 7.3 mm. The maximum width is 1.4 to 2.1 mm. ......................................................Buenoa cucunubensis Padilla and Nieser, 1992 (Colombia). - The rostral prong is almost straight, or its posterior margin is sinuate (Fig. 6.657). .................................................................................................................28

Fig. 6.657 Buenoa platycnemis male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and an enlargement of the tibial stridulatory comb. Based on Truxal (1953). 28. (27) The rostrum is robust, and the posterior margin of the rostral prong is distinctly sinuate. The tylus is only slightly inflated. The pronotum appears tricarinate in posterior view (Fig. 6.643). Length of male: 5.7 to 6.5 mm; female: 5.8 to 6.3 mm. Maximum width of male and female: 1.6 to 1.9 mm. Color: mainly sordid to yellowish white with testaceous markings and sometimes dark brown on the dorsal surface of the metanotum. The abdomen is black except for the yellowish white ventral keel and parts of the connexivum. .............................................................................Buenoa rostra Truxal, 1953 pars (Trinidad, Venezuela). Also see Couplet 10.

482

- The rostrum is not robust, and the posterior margin of the rostral prong is almost straight. The rostrum is distinctly tricarinate (Fig. 6.657). Length of male: 4.5 to 5.4 mm; female: 5.0 to 5.5 mm. Maximum width of male: 1.3 to 1.7 mm; female: 1.4 to 1.7 mm. Color: mainly white or testaceous with brown or black markings, usually including a mainly black abdomen. .........................................................................Buenoa platycnemis (Fieber, 1851) (North and Central America, West Indies, Colombia, Venezuela, Peru, Argentina, Maranhão, Goiás). Syn: Anisops platycnemis Fieber, 1851. 29. (17) The greatest width of the head is five times or less than the anterior width of the vertex. If slightly more (B. fuscipennis), then the synthlipsis is less than 1/4 the anterior width of the vertex (Fig. 6.650). .......................................30 - The greatest width of the head is more than five times the anterior width of the vertex. The synthlipsis is 1/4 or more of the anterior width of the vertex. .......34 30. (29) The rostral prong is equal to or longer than the third rostral segment (Fig. 6.650). ........................................................................................................31 - The rostral prong is shorter than the third rostral segment (Fig. 6.658). ........33

Fig. 6.658 Buenoa unguis male (left to right): rostrum and tylus in left lateral view, outline of the fore-tarsus, inner surface of the left fore-leg, and enlargement of the male tibial stridulatory comb. Based on Truxal (1953). 31. (30) The eyes are not holoptic. The greatest width of the head is more than 3 times the anterior width of the vertex (Fig. 6.650). Length of male: 3.3 to 3.7 mm; female: 3.7 to 4.4 mm. Maximum width of male: 1.0 to 1.2 mm; female: 1.1 to 1.3 mm. Color: white to pale testaceous, except for the mainly black or brown abdomen and often a pair of light brown to black longitudinal stripes on the dorsolateral surfaces of the metathorax. The hemielytra are hyaline, sometimes with a light brown to black marking at the apex of the corium. ................................................................................Buenoa salutis Kirkaldy, 1904 (Colombia, Venezuela, Guyana, French Guiana, Surinam, Bolivia, Paraguay, Argentina, Amazonas, Pará, Ceará, Paraíba, Pernambuco, Mato Grosso, Goiás, Minas Gerais, São Paulo, Paraná, Rio Grande do Sul). Syn: Buenoa mallochi Jaczewski, 1929.

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- The eyes of the male are usually holoptic (Fig. 6.659). The length is at least 4.4 mm. ...............................................................................................................32

Fig. 6.659 Buenoa amnigenus (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg of a male, enlargement of the male tibial stridulatory comb (above), and ovipositor valve of a female (below). Based on Truxal (1953).

Fig. 6.660 Buenoa amnigenopsis male: rostrum and tylus in left lateral view (left), inner surface of the left fore-leg (right center and lower center), enlargement of the tibial stridulatory comb (upper left center), and another view of the fore-femur (right). Based on Nieser (1975). 32. (31) The apex of the fore-femur is not notched (Fig. 6.659). The ventral row along the posterior margin of the hind femur bears 18 to 25 spine-like setae. The greatest width of head is less than three times the anterior width of the vertex. Brachypterous forms are known. Length of male: 4.7 to 5.4 mm; female: 4.8 to 5.9 mm. Maximum width of male: 1.2 to 1.4 mm; female: 1.3 to 2.6 mm. Color: mainly white with parts of the head, thorax, and dorsal surface of the abdomen testaceous; the legs are testaceous or brown. The abdomen is mainly black ventrally. ...........................................................................Buenoa amnigenus (White, 1879) (Guyana, Surinam, Peru, Bolivia, Paraguay, Argentina, Amazonas, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Goiás, Mato Grosso, Mato Grosso do Sul, Paraná). Syn: Anisops amnigenus White, 1879.

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- The apex of the hind femur is notched (Fig. 6.660). The ventral row along the posterior margin of the hind femur bears 12 to 14 spine-like setae. The greatest width of the head is more than 3.8 times the anterior width of the vertex. Length of male: 5.1 to 5.4 mm; female: 5.3 to 5.5 mm. ........................................................................Buenoa amnigenopsis Nieser, 1975 33. (30) The inner tarsal claw on the fore-leg is shield-like at the base (Fig.6.658). The synthlipsis is only about 1/5 the anterior width of the vertex. Length of male: 5.9 to 7.1 mm; female: 5.8 to 7.1 mm. Maximum width of male: 1.4 to 1.8; female: 1.6 to 1.9 mm. Color: white to pale testaceous, except sometimes for anterolateral brown or black markings at the anterolateral corners of the scutellum and the mainly brown or black abdomen, which is yellowish white or testaceous only on the anterodorsal surface, keel, and parts of the connexivum. ....................................................................................Buenoa unguis Truxal, 1953 (Peru, Bolivia, Paraguay, Argentina, Pará, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Minas Gerais, Rio de Janeiro). - The inner tarsal claw on the fore-leg gradually tapers from base to apex. The synthlipsis is about 1/2 the anterior width of the vertex (Fig.6.661). Length of male: 6.8 to 7.3 mm; female: 7.0 to 7.3 mm. Maximum width of male: 1.8 to 2.1 mm; female: 1.9 to 2.1 mm. The head, dorsal and some of the ventral surface of the thorax, legs, and parts of the abdomen, including the mid-ventral keel and posterior margins of the segments, are testaceous, while much of the ventral surface of the abdomen is black. ............................................................................Buenoa fuscipennis (Berg, 1879) (Venezuela, Bolivia, Paraguay, Uruguay, Argentina, Chile, Paraná, Santa Catarina, Mato Grosso). Syn: Anisops fuscipennis Berg, 1879; Anisops naias Kirkaldy, 1899; Buenoa naias (Kirkaldy, 1899) auctt; Buenoa dentipes Jaczewski, 1928.

Fig. 6.661 Buenoa fuscipennis male (left to right): rostrum and tylus in left lateral view, inner surfaces of the middle tarsus and the left fore-leg, enlargement of the tibial stridulatory comb, left (below) and right paramere (above), and the spine at the left posterior corner of the seventh abdominal segment. Based on Truxal (1953).

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34. (29) The tylus is greatly excavated and has a short anteromedial ridge (Fig. 6.662). Length of male: c. 4.5 to 4.6 mm; maximum width of male: c. 1.3 mm. The female has not been described. The color is sordid white with orange or brown ventral markings on the thorax and a black abdomen with orange on the anterior parts of the connexiva. The hemielytra are transparent, except for a violet-black band crossing the humeri and extending along the margins for about 1/3 of the length, as well as a large black spot near the apex of the corium. The two anterior pairs of legs are testaceous or violet black, and the hind legs are yellowish white. ................................................................................Buenoa excavata Truxal, 1953 (Peru). - The tylus is broadly rounded and not excavated (Fig. 6.663). ........................35

Fig. 6.662 Buenoa excavata male (left to right): rostrum and tylus in left lateral view, enlargement of the tibial stridulatory comb, inner surface of the left foreleg, and the spine at the left posterior corner of the seventh abdominal segment. Based on Truxal (1953).

Fig. 6.663 Buenoa oculata male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, enlargement of the tibial stridulatory comb, and the left (below) and right clasper (above). Based on Truxal (1953).

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35. (34) The synthlipsis is 1/15 of the anterior width of the vertex. The pronotum is not tricarinate. The first tarsal segment of the middle leg is emarginate on the inner surface (Fig. 6.663). Length of male: 4.7 to 5.2 mm; female: 4.8 to 5.4 mm. Maximum width of male: 1.1 to 1.3 mm; female: 1.2 to 1.4 mm. Color: mainly white with orange markings sometimes on the middle and posterior parts of the pronotum and the entire scutellum. The abdomen is yellowish white with a central blackish area dorsally and black ventrally, except for the white mid-ventral keel, parts of the connexivum, and at least one of the two apical segments. ..................................................................................Buenoa oculata Truxal, 1953 (Peru). - The synthlipsis is about 1/5 or more of the anterior width of the vertex. The first tarsal segment of the middle leg is straight on its inner surface. ................36

Fig. 6.664 Buenoa incompta male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Truxal (1953). 36. (35) The pronotum is tricarinate. The tibial comb is somewhat narrowed in the middle (Fig. 6.664). Length of male: 5.0 to 5.2 mm; female: 5.3 to 5.8 mm. Maximum width of male: 1.3 to 1.4 mm; female: 1.3 to 1.5 mm. The head and pronotum are white, while the ventral surface of the thorax and legs vary from white to brown. The scutellum is white, sometimes with brown anterolateral markings. The abdomen is yellowish white with black markings dorsally and black ventrally with white on the keel and connexivum. ...............................................................................Buenoa incompta Truxal, 1953 (Bolivia, Surinam, Amazonas, Goiás, Paraíba). - The pronotum lacks a carina. ..........................................................................37 37. (36) The apex of the fore-tibia projects to a sharp point (Fig. 6.665). Length of male: 4.5 to 5.1 mm; female: 5.3 to 5.7 mm. Color: pale yellowish to light brown dorsally with a tinge of orange. ....................................................................................Buenoa truxali Nieser, 1968 (Trinidad, Surinam, Amazonas). - The apex of the fore-tibia projects to a blunt end (Fig. 6.665). ......................38

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Fig. 6.665 Buenoa truxali: rostrum and tylus in left lateral view (left), inner surface of the left fore-leg of a male, enlargement of the prominence near the apex of the tibia, and an enlarged stridulatory comb. Based on Nieser (1968, 1975). 38. (37) Color: variable, but usually sordid white to blackish violet with a black dorsal surface of the thorax and abdomen. The scutellum is usually blackish with a testaceous apex. There is a long row or short setae running most of the length of the hind femur (Fig. 6.666). The hemielytra are hyaline with blackish violet on the posterior 1/3. The color may vary to sordid white with black only on the abdomen and posterior third of the hemielytra. Length of male: 5.0 to 5.5 mm; female: 4.9 to 5.7 mm. Maximum width of male: 1.6 to 1.8 mm; female: 1.7 to 1.9 mm. ...................................................................................Buenoa tibialis Truxal, 1957 (Goiás). - Color: dark brown to blackish with yellowish markings. The row or short setae on the inner surface of the hind femur runs only for about half its length (Fig. 6.667).Length of male: 4.5 to 5.1 mm; female: 5.0 to 5.5 mm. ..................................................................................Buenoa fasciata Nieser, 1975 (Surinam).

Fig. 6.666 Buenoa tibialis male (left to right): the tylus and rostrum (beak), the enlarged left stridulatory comb on the tibia, and the left fore-leg. Based on Truxal (1957).

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Fig. 6.667 Buenoa fasciata male (left to right): rostrum and tylus in left lateral view, inner surface of the left fore-leg, and enlargement of the tibial stridulatory comb. Based on Nieser (1975). Key to the species of brachypterous male Martarega in South America Information for the key was provided by Truxal (1949), Bachmann (1961c), and Nieser (1975). Females of all species cannot yet be identified, but notes on known females are included in the key. 1. The hemielytron is opaque pale yellow with a brown costal margin and two opaque, nearly parallel-sided longitudinal stripes, one whitish along or near the anterior margin, and the other white and brown (Fig. 6.668). Length of male: c. 4.6 to 4.9 mm; female: c. 5.7 to 6.0 mm. Maximum width of male: 1.3 to 1.5 mm; female: 1.5 to 1.6 mm. Color: mainly pale yellow with red eyes. Most specimens described were brachypterous. .......................................Martarega pacifica Manzano, Nieser, and Caicedo 1995 (Colombia). - Each hemielytron lacks a pair of nearly parallel-sided opaque stripes; if opaque, longitudinal stripes are present, the posterior pair has an obvious expansion dorsad (Fig. 6.669). Most specimens are larger or smaller than the size range shown above. .......................................................................................2 2. (1) Males do not exceed 4.3 mm in length. The pronotum is no wider than 1.3 mm. The opaque median longitudinal stripe on the hemielytra forks just proximal to the apex (Fig. 6.669). ........................................................................3 - Males are at least 4.9 mm long. If there is an opaque median longitudinal stripe on the hemielytra, it does not fork at the apex (Fig. 6.670). ......................4 3. (2) The ventral surface of the middle trochanter is smooth (Fig. 6.336). Length of male: 4.1 to 4.2 mm; female: 4.5 to 4.6 mm. Width of pronotum: 1.1 to 1.4 mm. Color: whitish or yellowish with bluish brown eyes. ...................................................................Martarega membranacea White, 1879 (Guyana, Surinam, Colombia, Ecuador, Bolivia, Argentina, Amazonas, Pará, Goiás, Minas Gerais).

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- The ventral surface of the middle trochanter has a median nodule (Fig. 6.669). The hemielytral process does not extend posteriad as far as the apex of the membrane. The opaque stripe on the hemielytron has a rounded rather than acutely angled inner margin at its fork. Length of male: 4.1 to 4.2 mm; female: 4.6 to 4.7 mm. Width of pronotum: 1.1 to 1.4 mm. ................................................................................Martarega chinai Hynes, 1948 (Trinidad, Colombia, Venezuela, Bolivia, Argentina, Pará, Amazonas).

Fig. 6.668 Martarega pacifica: the left hemielytron (left) and the male genital capsule (right). Based on Manzano et al. (1995).

Fig. 6.669 Martarega chinai male (left to right): pattern on a hemielytron, genital capsule in left and right lateral view, and penis. Based on Truxal (1949) and Angrisano (1982a). 4. (2) The lateral margin of the hind trochanter is sharply concave at its distal end. The emarginated costal margins of the female hemielytra bear teeth (Fig. 6.670). Hair-like setae on the mid-ventral keel are found on both the anteroventral and lateral surfaces. Length of male: 5.1 to 5.6 mm; female: 6.5 to 6.6 mm. Width of pronotum: 1.4 to 1.65 mm. Color: whitish with reddish eyes and light yellowish legs. ......................................................................Martarega hungerfordi Truxal, 1949 (Guyana, Surinam, Pará, Amazonas).

490

- The lateral margin of the hind trochanter is not sharply concave at its distal end (Fig. 6.671). The emarginated costal margins of the female hemielytra lack teeth. Hair-like setae on the mid-ventral keel are found only on its lateral margins. ................................................................................................................5

Fig. 6.670 Martarega hungerfordi (left to right): middle trochanter, hemielytron of a female, and left and right views of the genital capsule. Based on Truxal (1949) and Nieser (1975). 5. (4) The hemielytra are mainly hyaline with a single opaque longitudinal stripe, which is not located in the middle. The left paramere is three times as long as wide (Fig. 6.671). Length of brachypterous male: 5.7 to 5.9 mm; brachypterous female: 6.4 to 6.5 mm. Length of macropterous male: 5.8 to 5.9 mm; macropterous female: 6.5 to 6.6 mm. Width of pronotum of brachypterous specimens: 1.4 to 1.6 mm; macropterous specimens: 1.6 to 1.8 mm. Color: pale yellowish with bluish brown eyes. ..........................................................................Martarega gonostyla Truxal, 1949 (Surinam, Bolivia, Pará, Amazonas, Mato Grosso). - The hemielytra are not mainly hyaline with a single opaque longitudinal stripe. The left paramere of the male genital capsule is shorter than three times its width (Fig. 6.672). ...........................................................................................6

Fig. 6.671 Martarega gonostyla male (left to right): middle trochanter, pattern on a hemielytron, and left and right views of the genital capsule. Based on Truxal (1949).

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Fig. 6.672 Martarega williamsi male (left to right): pattern on a hemielytron of a male and the genital capsule in left and right lateral view. Based on Truxal (1949). 6. (5) The outer lateral margin of the middle trochanter bears one or two groups of short bristles ventrally. Each hemielytron has a short, opaque, median stripe at the base (Fig. 6.672). Length of brachypterous male: 5.3 to 5.4 mm; brachypterous female: 6.0 to 6.1 mm. Length of macropterous male: 5.9 to 6.0 mm; macropterous female: 6.4 to 6.5 mm. Width of pronotum of brachypterous specimens: 1.5 to 1.7 mm; macropterous specimens: 1.7 to 1.8 mm. ..........................................................................Martarega williamsi Truxal, 1949 (Panama, Peru, Ecuador, Amazonas). - The outer lateral margin of the middle trochanter does not bear a group of short bristles ventrally. Neither hemielytron has a short, opaque, median stripe at the base (Fig. 6.673). ........................................................................................7

Fig. 6.673 Martarega bentoi male (left to right): middle trochanter, pattern on a hemielytron, left and right views of the genital capsule. Based on Truxal (1949).

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7. (6) The trochanter and femur of the middle leg lack nodule-like structures; the middle trochanter bears a patch of yellow hairs on the ventral surface. There is a narrow median stripe on each hemielytron and a second short, shiny stripe at the base of the hemielytral process. The penis ends with a broad elongation with three processes along its rim (Fig. 6.674). Length of male: 5.0 to 5.4 mm; female: 5.6 to 5.8 mm. Width of pronotum: 1.2 to 1.4 mm. Color: light yellowish or whitish dorsally with reddish brown or dark brown eyes. .......................................................................Martarega brasiliensis Truxal, 1949 (Surinam, Peru, Pará, Ceará, Pernambuco). - The middle trochanter of the male bears a nodule (Fig. 6.675). North and Central American species also bear such structures on the middle femora. ........8

Fig. 6.674 Martarega brasiliensis male (left to right): genital capsule in left and right lateral view and the penis. Based on Truxal (1949) and Angrisano (1982a). 8. (7) The apex of the metaxyphus is narrowly rounded. The width of a compound eye is more than three times the ocular commisure. The length of the membrane of the apterous form is 1.5 times that of the hemielytral process (Fig. 6.632). Length of the male: c. 8.0 mm; female: c. 8.2 mm. Width of pronotum: c. 2.1 mm. .......................................................................Martarega mcateei Jaczewski, 1928 (Argentina, Paraná). - The apex of the metaxyphus is medium or broadly rounded. The width of a compound eye is less than three times the ocular commisure. The length of the membrane is at least twice that of the hemielytral process (Fig. 6.673). .............9 9. (8) The longitudinal hyaline stripe along the midline of the male hemielytra is narrow, so at its midlength, it is about 1/5 the width of the shiny costal area of each hemielytron. The genital capsule forms a long, even curve along its longest side, best seen in right lateral view (Fig. 6.675). The basal exterior margin of the female hind femur is emarginate. The membrane is about three times the length of the hemielytral process. Length of the male: 5.2 to 5.5 mm; female: 5.8 to 5.9 mm. Width of pronotum: 1.2 to 1.5 mm. ....................................................................Martarega uruguayensis (Berg, 1883) (Paraguay, Bolivia, Argentina, Uruguay, Pernambuco, Minas Gerais, Rio de Janeiro). Syn: Signoretiella uruguayensis Berg, 1883.

493

- The longitudinal hyaline stripe along the midline of the male hemielytra is wide, so at its midlength, it is about 2/3 the width of the shiny costal area. The basal exterior margin of the female hind femur is not emarginate. The membrane is about twice the length of the hemielytral process (Fig. 6.673). Length of the male: 5.2 to 5.3 mm; female: 5.7 to 5.8 mm. Width of pronotum: 1.3 to 1.4 mm. ...............................................................................Martarega bentoi Truxal, 1949 (Argentina, Pernambuco).

Fig. 6.675 Martarega uruguayensis male (left to right): genital capsule in left and right lateral view and penis. Based on Truxal (1949) and Angrisano (1982a). Key to the species of Notonecta in South America Information for the key was provided by Hungerford (1930c, 1933), De Carlo (1935b), Angrisano (1982b, c), Bachmann (1963c, d, 1966d), Polhemus (1993b), Mazzucconi (2000), and Mazzucconi and Bachmann (2004). 1. Hair-like setae are present only on the lateral parts of the fourth abdominal sternite, leaving the median keel bare (Fig. 6.676). Length: 13 to 14 mm. Color: usually red and black, with males usually darker. The anterior half of the prothorax, head, and legs are yellow with two black stripes on the fore-femur and one stripe along each middle trochanter and femur. The scutellum is black, and the clavus and corium are orange red or red. ......................................................Notonecta mexicana Amyot and Serville, 1843 (Mexico, Colombia). Syn: Notonecta klugi Fieber, 1851 pars; Notonecta macrocephala Hungerford, 1925. Two “varieties” were described: Notonecta mexicana mexicana Amyot and Serville, 1843, known throughout the range, and the smaller Notonecta mexicana creaseri Hungerford, 1932, known only from the state of Nuevo Leon, Mexico. - Hair-like setae are present on the median keel of the fourth abdominal sternite (Fig. 6.677). ..........................................................................................................2

494

Fig. 6.676 Notonecta mexicana male: habitus (upper left), ventral view of the abdomen (lower left), and the left side of the genital capsule (right). Based on Hungerford (1933). 2. (1) The last abdominal sternite of the female is large and constricted just before the apex. The male genital capsule has a finger-like prolongation. Total length: c. 16 mm. Color: almost entirely chocolate brown to black with the scutellum and venter always black and the head and limbs shiny brown or black (Fig. 6.677). ..................................................................................Notonecta nigra Fieber, 1851 (Brazil, but Hungerford found it to be morphologically identical to Notonecta glauca Linnaeus, 1758, a species widespread in the Eastern Hemisphere). - The last abdominal sternite of the female lacks a pre-apical constriction. The male lacks a digitate prolongation of the genital capsule (Fig. 6.678). ................3

Fig. 6.677 Notonecta nigra male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933).

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3. (2) The lateral margins of the pronotum are divergent and embrace the compound eyes, which are flattened and do not recede from the anterior margin of the vertex. In dorsal view, the synthlipsis equals about half the anterior margin of the vertex. The scutellum is obviously broader than long (Fig. 6.678). The male bears a stout tubercle at the base of the fore-trochanter and a very short, broad hook. .................................................................................................4 - The lateral margins of the pronotum do not embrace the compound eyes. The scutellum is not obviously broader than long. The male lacks a stout tubercle at the base of the fore-trochanter but has a short, broad hook on its anterior surface (Fig. 6.679). ..........................................................................................................5

Fig. 6.678 Habitus of a male Notonecta colombiana. Based on Hungerford (1933).

Fig. 6.679 Notonecta unifasciata: habitus of a male (upper left); male genital capsules of Notonecta unifasciata unifasciata, N. u. angulata, N. u. cochisiana, and N. u. andersoni (lower left to upper right); middle leg with a typically angulate trochanter (lower right). Only N. u. angulata has been reported from South America. Based on Hungerford (1933).

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4. (3) Length: 12.0 to 12.6 mm. There are reportedly 2 color forms, but only the blackish form has been thoroughly described. The scutellum of this form is black, and the hemielytra are brown and black with a light yellow streak along the claval suture and a transverse spot on the proximal third of the corium (Fig. 6.678). The face, legs, and connexivum are usually yellow or greenish yellow. ...............................................................Notonecta colombiana Hungerford, 1933 (Colombia). - Length: 10.8 to 11.4 mm. The color is entirely pale yellow. There is a thickened protuberance on the keel of the male genital capsule. ......................................................................Notonecta ochrothoe Kirkaldy, 1897 (Colombia). Syn: Notonecta shooteri var. ochrothoe Kirkaldy, 1897. 5. (3) The trochanter of the middle leg is angular. ...............................................6 - The trochanter of the middle leg is rounded. ....................................................8

Fig. 6.680 Genital capsule of Notonecta bifasciata in lateral view. Based on Bachmann (1971).

Fig. 6.681 Notonecta confusa: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933). 6. (5) The trochanter of the middle leg is strongly angular. The color pattern includes the scutellum and one band across the dorsum posterior to the clavus (Fig. 6.679). ......................................................Notonecta unifasciata Guérin-Meneville, 1857 (North and South America?). Syn: Notonecta americana Kirkaldy, 1897 pars; Notonecta indica Kirkaldy, 1900 pars; Notonecta undulata Champion, 1901

497

pars. Reference to this species in South America is confined to the labels for two specimens showing “S. Amer.” as the origin without any more information on the location. Four subspecies have been described: Notonecta unifasciata unifasciata Guérin-Meneville, 1857; Notonecta unifasciata andersoni Hungerford, 1933; Notonecta unifasciata cochisiana Hungerford, 1933; Notonecta unifasciata angulata Hungerford, 1933. Although Hungerford (1933) discussed the two South American specimens with his discription of N. u. angulata, he leaves the question of their identity open. The occurrence of this species in South America therefore remains dubious. - The trochanter of the middle leg is weakly angular. .........................................7 7. (6) Length: c. 8.5 to 9.3 mm. The scutellum is black, and the hemielytra are dark with a pale stripe on the basal 2/3 of the clavus. The inner margin bordering the scutellum is entirely brown. There is a pale triangular spot on the corium bordering the clavocorial suture. There is a transverse orange spot and a larger irregular spot above it on the embolium. The apex of the membrane is pale. The hemielytra are covered with silvery setae that are narrowly spatulate. The left paramere of the male has a sparse coat of short spines (Fig. 6.680). ..........................................................................Notonecta bifasciata Guérin, 1844 (Argentina, Uruguay). - Length: c. 12 mm. The type specimen is pale yellow, but Hungerford speculated that most specimens would have a black and white pattern. The branches of the male paramere are broad (Fig. 6.681). ......................................................................Notonecta confusa Hungerford, 1930 (South America without a specific location; the range possibly includes Cuba.). 8. (5) The ratio of the synthlipsis to the posterior border of a compound eye is between 3.9:9 and 4.4:9, and the length varies from 8.4 to 9.6 mm. The color pattern ranges from entirely pale yellow to variable with black on the scutellum and brown and smoky stripes laterally (Fig. 6.631). .....................................................................Notonecta virescens Blanchard, 1852 (Chile). Syn: Notonecta bicirca Hungerford, 1926, Notonecta bicircoidea Hungerford, 1928. - The ratio of the synthlipsis to the posterior border of a compound eye is less than 2:5 or more than 3:5 (Fig. 6.682), or the length is about 11 mm or more. ...9

Fig. 6.682 Notonecta pulchra male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933).

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9. (8) The synthlipsis is less than 1/3 as long as the posterior margin of a compound eye (Fig. 6.682). ...............................................................................10 - The synthlipsis is more than 1/3 as long as the posterior margin of a compound eye (Fig. 6.683). ...............................................................................12

Fig. 6.683 Notonecta vereertbruggheni male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933). 10. (9) Length: c. 7.2 mm. Width of pronotum: c. 2.0 mm. The head, pronotum, legs, and basal 1/3 of the hemielytra are yellow. The scutellum and distal 2/3 of the membrane are black with the apices of the membrane pale. The black areas on the hemielytra are interrupted by two small, irregular orange spots about equal in size. The width of an eye at the base is equal to the vertex, which is more than four times the synthlipsis (Fig. 6.684). ..................................................................Notonecta hungerfordi Polhemus, 1993 (Colombia). Syn: Notonecta minuta Hungerford, 1926 (nec Gmelin, 1790). - The length is at least about 7.5 mm, and the width of the pronotum, at least 2.1 mm. The black scutellum usually lacks tan markings along the lateral margins (Fig. 6.682). ........................................................................................................11

Fig. 6.684 Notonecta hungerfordi male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933).

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Fig. 6.685 Notonecta disturbata male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933). 11. (10) The hemielytra are black with two pale spots at the base and transverse orange markings at the apex of the corium (Fig. 6.682). The hemielytra are covered with silvery setae. Length: 7.5 to 9.0 mm. Width of pronotum: 2.1 to 3.1 mm. The scutellum is either entirely black, or it has a pair of tan markings, each at the lateral margin. ......................................................................Notonecta pulchra Hungerford, 1926 (Paraguay, Argentina, Amazonas, Minas Gerais). - The hemielytra are mainly whitish, sometimes with brownish black at the apex of the clavus and black on the distal half of the corium and basal 2/3 of the membrane (Fig. 6.685). The scutellum is either entirely pale, or it is black with a pair of tan markings, each along a lateral margin. Length: 8.0 to 8.4 mm. Width of pronotum: 2.8 to 2.9 mm. ..................................................................Notonecta disturbata Hungerford, 1926 (Argentina, São Paulo, Rio de Janeiro, Goiás, Minas Gerais, Mato Grosso). 12. (9) The length is usually at least 11 mm, and the width is usually 4.0 to 4.2 mm. The color pattern is not exclusively black and white (Fig. 6.686). ...........13 - The length is usually less than 10.5 mm, or the color pattern is exclusively black and white or black and luteous (Fig. 6.687). ............................................14

Fig. 6.686 Notonecta fazi male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933).

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Fig. 6.687 Notonecta indica: habitus of two females with somewhat different color patterns (left) and the left side of the male genital capsule (right). Based on Hungerford (1933). 13. (12) The synthlipsis is more than 3/5 as long as the posterior margin of a compound eye. Length: 11 to 12 mm. Width of pronotum: c. 4.2 mm. The head and dorsum are black, yellow, and tan (Fig. 6.683). Ventrally, the colors are black and yellow. ......................................................Notonecta vereertbruggheni Hungerford, 1928 (Argentina). - The synthlipsis is less than 3/5 as long as the posterior margin of a compound eye (Fig. 6.686). Length: c. 12 mm. Width of pronotum: c. 4 mm. The head and dorsum are typically black, yellow, orange, and tan arranged in a somewhat variable pattern. Ventrally, the colors are black and yellowish brown. ............................................................................Notonecta fazi Hungerford, 1930 (Chile, Peru). Syn: Notonecta americana Kirkaldy, 1897 pars from Chile; Notonecta virescens Reed, 1901 pars, misidentified. 14. (12) The coloration is exclusively black and white, but some specimens may be entirely luteous (Fig. 6.687). The color pattern is variable. The posterior margin of the male paramere and the sixth visible abdominal sternite of the female are not deeply notched. ............................................................................Notonecta indica Linnaeus, 1771 (North and Central America, West Indies, Colombia, Brazil). Syn: Notonecta americana Fabricius, 1775; Notonecta undulata Say, 1832 pars; Notonecta variabilis var. scutellaris Fieber, 1851 (misidentified specimens from Puerto Rico); Notonecta pallipes Lethierry, 1881 (three misidentified West Indian specimens); Notonecta howardii Bueno, 1905. The species may also encompass all South American varieties of the doubtful species, Notonecta variabilis Fieber, 1851. Hungerford (1933) was unable to find any feature for distinguishing them.

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- The pattern includes colors besides black and white, with at least a pair of tan or yellowish spots but usually with distinct orange, brown, horn-colored, yellow, or tan markings in addition to an extensive black pattern (Fig. 6.688). .............15 15. (14) The posterior margin of the male paramere and the sixth visible abdominal sternite of the female are deeply notched. The synthlipsis is less than half the width of a compound eye along its posterior margin (Fig. 6.688). .......16 - The posterior margin of the male paramere and the sixth visible abdominal sternite of the female are not deeply notched. The synthlipsis is more than half the width of a compound eye along its posterior margin (Fig. 6.689). ..............17

Fig. 6.688 Notonecta inca (left to right): male genital capsule, penis in dorsal and lateral view, left male paramere (above), fifth and sixth abdominal segments of a female in ventral view, the sixth and seventh abdominal sternites and laterotergites of a female, and the eighth abdominal sternite of a female. When counting the abdominal segments, the first is not visible, so the first visible segment is counted as the second, the third as the second, as so forth. Based on Mazzucconi (2000).

Fig. 6.689 Notonecta polystolisma male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933). 16. (15) The length of the male paramere is about 2.5 times its greatest width or less. The seventh, that is, the sixth visible abdominal sternite of the female is obviously concave along the middle of its posterior margin (Fig. 6.688). Total length: 8.1 to 10.4 mm. There are gaps in the description of this species, and judging from the illustrations of the two nominal species in this couplet, it is not fully convincing that they are distinct. ..........................................................................Notonecta inca Mazzucconi, 2000 (Peru).

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- The length of the male paramere is about 3 times its greatest width. The seventh, that is, the sixth visible abdominal sternite of the female is straight or only slightly concave across the middle of its posterior margin. The ratio of the width of the vertex to the synthlipsis is 15:8. The left clasper is acutely pointed at the apex (Fig. 6.690). Length: 9.3 to 10.3 mm. Width of pronotum: 3.4 to 3.5 mm. The scutellum is black, and the hemielytra are dark with a tan or horncolored oblique marking along the base of the clavocorial line. There is an oblique orange line along the posterior margin of the embolium and a dull orange spot on the corium. The membrane is mainly brown. ..................................................................Notonecta peruviana Hungerford, 1933 (Peru, Argentina). Syn: Notonecta huincamoreni Bachmann, 1962.

Fig. 6.690 Notonecta peruviana (left to right): male genital capsule, penis in lateral view, left male paramere, fifth and sixth abdominal segments of a female in ventral view, the sixth and seventh abdominal sternites and laterotergites of a female, and the eighth abdominal sternite of a female. When counting the abdominal segments, the first is not visible, so the first visible segment is counted as the second, the third as the second, as so forth. Based on Mazzucconi (2000), who reported that the illustration by Hungerford (1933) is inaccurate. 17. (15) The typical color is white with a transverse black marking across the clavus enclosing two tan spots. The scutellum is black with the lateral margins lined by short pale stripes. The color may also be lighter, and the pattern is variable. Length: 8.7 to 9.6 mm. Width of pronotum: 2.8 to 3.2 mm. The anterior width of the vertex is three times the synthlipsis, which is much less than half the width of a compound eye measured along its posterior margin (Fig. 6.691). ................................................................................Notonecta sellata Fieber, 1851 (Argentina, Paraguay, Uruguay, southern Brazil). Syn: Notonecta variabilis Kirkaldy, 1897 (nec Fieber); Notonecta bifasciata Hungerford, 1926 (nec Guérin). - The typical color is black with horn-colored or brownish yellow on the anterior half of the pronotum, propleura, head and legs, which are sometimes also tinged with green. The dominant ventral color is black or brown, sometimes with a metaxyphus and connexivum colored like the legs. The scutellum is black, unless the insect is teneral. The color of the hemielytra is

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black or bluish black with a tan forked streak along the basal half of the claval commissure and two tan spots or a tan stripe across the corium at the apex of the clavus. The apex of the membrane is usually pale. Length: 9.0 to 9.6 mm. Width of pronotum: 3.1 to 3.5 mm. The anterior width of the vertex is three times or more than the synthlipsis, which is much less than half the width of a compound eye measured along its posterior margin (Fig. 6.689). ......................................................................Notonecta polystolisma Fieber, 1851 (Argentina, Paraná, São Paulo). Syn: Notonecta polystolisma var. guttata Fieber, 1851; Notonecta polystolisma var. bipunctata Fieber, 1851; Notonecta polystolisma var. fasciata Fieber, 1851; Notonecta polystolisma var. immaculata Fieber, 1851; Notonecta polystolisma var. spatulata Hungerford, 1926; Notonecta bifasciata Kirkaldy, 1904 (listed as a synonym in error).

Fig. 6.691 Notonecta sellata male: habitus (left) and the left side of the genital capsule (right). Based on Hungerford (1933). Key to the species of Enithares in South America Information for the key was provided by Brooks (1953). The females of the South American species, both placed in the subgenus Enitharoides Brooks, 1953, cannot be distinguished. 1. There is a rather long posterior process on the male genital capsule (Fig. 6.692). Length: 13 to 14 mm. The predominant colors are dark brown and black, but the vertex and pronotum are testaceous. The black coloration of the abdomen can be seen through hyaline areas on the hemielytra. .......................................................................Enithares duidaensis Brooks, 1953 (Venezuela). - The posterior part of the male genital capsule ends in an acute angle without any trace of a process (Fig. 6.633). .......................................................................Enithares braziliensis Spinola, 1837 (Brazil).

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Fig. 6.692 The left lateral (left) and right lateral view (right) of the male genital capsule of Enithares duidaensis. Based on Brooks (1953).

Helotrephidae Key to the genera of adults in South America Information for the key was provided by Froeschner (1981) and Nieser (1975). 1. The anterior margin of the pronotum has two deep, angular emarginations separated by a broad, convex, central lobe. The eyes are globose. The labium does not reach the hind coxae (Fig. 6.693). ..........................................................................................Neotrephes China, 1936 - The anterior margin of the pronotum lacks emarginations and is nearly straight. The eyes are not globose. The labium reaches between the hind coxae (Fig. 6.694). .........................................................................................Paratrephes China, 1940 The only species in this genus is Paratrephes hintoni China, 1940, known from Colombia, French Guiana, and Surinam. It is no more than 2 mm long, and its coloration varies from yellowish brown to blackish.

Fig. 6.693 Neotrephes variegatus: habitus of a macropterous male (upper left) and a brachypterous female (lower left), dorsal (upper center) and ventral paramere of a male (center), penis in ventral view (right), and seventh abdominal segment of a female in ventral view (lower center). Based on Nieser and Chen (2002).

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Fig. 6.694 Paratrephes hintoni (left to right): the right half of the head and prothorax in dorsal view, a lateral view of the head and prothorax, a paramere, and the habitus in dorsal view. Based on Nieser (1975). Key to the species of adult Neotrephes in South America Information for the key was provided by China (1936, 1940) and Nieser and Chen (2002). Brachypterous individuals are apparently predominant. 1. The total length is more than 2.2 mm. The narrow part of the ventral paramere is not long (Fig. 6.695), and the female bears no distinct sensory costal lobe. ......................................................................................................................2 - The total length is not more than 2.2 mm, or, if it is slightly longer, then there is a long, narrow part of the ventral paramere (Fig. 6.696), and there is a distinct sensory costal lobe on the female. ........................................................................4

Fig. 6.695 Neotrephes latus male (left to right): dorsal (above) and ventral paramere (below) and the penis in ventral view. Based on Nieser and Chen (2002). 2. (1) The dorsal paramere is straight, long, and narrow. The length of the brachypterous male is c. 2.2 to 2.3 mm, which is only 1.25 times its width (Fig. 6.695). The color is mainly dark brown and black with yellowish and reddish yellow spots. ................................................................Neotrephes latus Nieser and Chen, 2002 (Minas Gerais). - The dorsal paramere is either curved, or it is short and thick. The length is about 1.5 times the width (Fig. 3.697). ................................................................3

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Fig. 6.696 Neotrephes minutus (left to right): dorsal and ventral paramere of a male, penis in ventral view, and seventh abdominal segment of a female in ventral view. Based on Nieser and Chen (2002). 3. (2) The apical part of the ventral paramere of the male is narrow and long. The right hemielytron of the female bears a distinct sensory costal lobe (Fig. 6.697). Length: 2.3 to 2.5 mm. The hemielytra are entirely dark. ............................................................................Neotrephes usingeri China, 1936 (Argentina, Santa Catarina). - The narrow apical part of the male paramere is short. The right hemielytron of the female does not bear a distinct sensory costal lobe (Fig. 6.698). Length: 2.2 to 2.4 mm. The hemielytra have yellow or reddish yellow markings. There is no transverse yellow band on the posterior part of the pronotum; only the humeri are yellow. There is only one vague, large rusty brown spots on the hemielytra. .........................................................................Neotrephes jaczewskii China, 1940 (Minas Gerais, Rio de Janeiro). 4. (1) The total length is 1.6 to 1.7 mm. ...............................................................5 - The total length is 1.8 to 2.2 mm. ................................................................….6 5. (4) The apical part of the male paramere is not narrow. The seventh abdominal sternite of the female has a median caudal process that is narrow, long, and shallowly incised at the apex (Fig. 6.696). Length: 1.65 to 1.8 mm. Only brachypterous specimens have been described. ............................................................Neotrephes minutus Nieser and Chen, 2002 (Minas Gerais). - The apical part of the male paramere is long and narrow. The seventh abdominal sternite of the female has a median caudal process that is rounded at the apex (Fig. 6.699). Length: 1.6 to 1.8 mm. ......................................................Neotrephes transversus Nieser and Chen, 2002 (Minas Gerais).

Fig. 6.697 The dorsal (above) and ventral paramere (below) of a male Neotrephes usingeri. Based on Nieser and Chen (2002).

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Fig. 6.698 Neotrephes jaczewskii Habitus in dorsal view showing the pattern of black, ferrugineous brown, and yellow at the posterolateral corners of the pronotum (right); dorsal (upper left) and ventral paramere (lower left) of a male. Based on China (1940) and Nieser and Chen (2002).

Fig. 6.699 Neotrephes transversus male (left to right): dorsal and ventral paramere of a male, penis in ventral view, and seventh abdominal segment of a female in ventral view. Based on Nieser and Chen (2002). 6. (4) The length is at least 2.0 mm. The dorsal paramere of the male tapers to the apex. The female does not have a sensory costal lobe (Fig. 6.693). Length: 2.0 to 2.2 mm. ......................................................Neotrephes variegatus Nieser and Chen, 2002 (Minas Gerais). - The length is less than 2.0 mm, or, if it is slightly longer, then the dorsal paramere of the male is truncate or slightly enlarged at the apex, and the female bears a sensory costal lobe (Fig. 6.700). ..............................................................7 7. (6) There is a transverse reddish yellow band on the posterior part of the pronotum, which expands only slightly anteriad along the midline. There are several distinct yellow spots on the hemielytra. The female has a sensory costal lobe. The dorsal paramere is enlarged at the apex. The ventral paramere remains nearly the same width throughout the entire basal half (Fig. 6.700). ........................................................................Neotrephes plaumanni China, 1940 (Rio de Janeiro, São Paulo, Minas Gerais).

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- There is a transverse reddish yellow band on the posterior part of the pronotum, which expands anteriad along the midline to reach the cephalonotal suture (Fig. 6.701). ...............................................................................................8

Fig. 6.700 Neotrephes plaumanni (above, left to right): the dorsal (above) and ventral paramere (below) of a male, habitus without appendages in dorsal and lateral view, and (below, left to right): antenna, beak, apex of a female hemielytron showing the sensory lobe, penis, and male abdomen in ventral view. Based on China (1940). 8. (7) The dorsal paramere is strongly curved. The basal part of the seventh abdominal sternite of the female is not enlarged and is barely visible in lateral view (Fig. 6.340). Length: 1.85 to 2.00 mm. ........................................................Neotrephes lanemeloi Nieser and Chen, 2002 (Argentina, Minas Gerais). Syn: Neotrephes jaczewskii China, 1940, pars, misidentified by Nieser and Lane de Melo (1997). - The dorsal paramere is not strongly curved. The basal part of the seventh abdominal sternite of the female is greatly expanded and appears as a ventral hump in lateral view (Fig. 6.701). Length: 1.90 to 2.05 mm. ..........................................................Neotrephes fragosus Nieser and Chen, 2002 (Minas Gerais).

Fig. 6.701 Neotrephes fragosus (left to right): dorsal (above) and ventral paramere (below) of a male, penis in ventral view, and seventh abdominal segment of a female in ventral view. Based on Nieser and Chen (2002).

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Pleidae The family is represented by only two genera in South America, which can be distinguished by the following key. Key to the genera of Pleidae in South America Information for the key was provided by Bachmann (1968, 1979c) and Nieser (1969b, 1975). 1. The fore-tarsus consists of two segments (Fig. 6.341). Length: 1.5 to 1.7 mm. .............................................................................Paraplea Esaki and China, 1928 This genus is represented by only one species in South America, Paraplea puella (Barber, 1923), which ranges from southern North America through Central America and the West Indies to Venezuela and Surinam. Dorsally, it is yellowish with brown patches, and adults are about 1.6 mm long. - The fore-tarsus consists of three segments (Fig. 6.702). The larval instars have been described (Fig. 6.342). A few species in this genus which are not found in South America have only two anterior tarsal segments. ...............................................................................Neoplea Esaki and China, 1928

Fig. 6.702 Neoplea borellii (above, left to right): habitus in dorsal and lateral view; male genital capsule and right paramere, and (middle left to center): fore, middle, and hind tarsi; eighth abdominal segment of a male, and (below, left to right): seventh urosternite of a male, right valve of the ovipositor, and seventh urosternite of a female. Based on Bachmann (1968).

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Key to the species of adult Neoplea in South America Information for the key was provided by Drake and Chapman (1953b), Bachmann (1968, 1979c), Roback and Nieser (1974), and Nieser (1975). 1. The second through the sixth sternites each bear a carina. The posterior margin of the pronotum is narrowly and moderately excavated along the midline (Fig. 6.702). Length: c. 3.25 mm. ......................................................Neoplea (Guazuplea) borellii (Kirkaldy, 1899) (Argentina, Santa Catarina?). - Only the second through the fifth sternites bear carinae; the sixth does not. If the posterior margin of the pronotum is excavated at all, the concave section is very broad or very shallow (Fig. 6.703). ..............................................................2

Fig. 6.703 Neoplea absona (above, left to right): habitus in dorsal and lateral view, male genital capsule and right paramere, and (below, left to right): the eighth segment and the seventh urosternite of a male, the seventh urosternite of a female, and the right valve of the ovipositor. Based on Bachmann (1968).

Fig. 6.704 Neoplea maculosa (above, left to right): habitus in dorsal and lateral view, male genital capsule and right paramere, and (below, left to right): the eighth segment and the seventh urosternite of a male, the seventh urosternite of a female, and the right valve of the ovipositor. Based on Bachmann (1968).

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2. (1) Viewed dorsally, the sides of the pronotum are straight and nearly parallel. The pronotum is no wider than the hemielytra are high (Fig. 6.703). .................3 - Viewed dorsally, the sides of the pronotum are sinuous. The pronotum is as wide or wider than the hemielytra are high. The scutellum is large, its sides are sinuous and its posterior apex is sharply acute (Fig. 6.704). ...............................4

Fig. 6.705 Neoplea lingula (above, left to right): habitus without appendages in dorsal view, eighth abdominal segment of a male, male genital capsule without the aedeagus, and (below, left to right): habitus without appendages in lateral view, operculum of a female, and ovipositor. Based on Roback and Nieser (1974).

Fig. 6.706 Neoplea tenuistyla (above, left to right): habitus without appendages in dorsal view, eighth abdominal segment of a male, the male genital capsule without the aedeagus, right paramere, and (below, left to right): habitus without appendages in lateral view, operculum of the female, and ovipositor. Based on Roback and Nieser (1974). 3. (2) The color is uniformly light. The scutellum is small with straight sides; the total length of the insect is about 4½ times the length of the scutellum, and the pronotum is about 1½ times as long as the scutellum. The claval suture is usually lacking (Fig. 6.703). Length: about 1.5 mm. ........................................Neoplea (Neoplea) absona (Drake and Chapman, 1953) (Argentina, Amazonas). Syn: Plea absona Drake and Chapman, 1953.

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- The color is yellowish with indistinct and irregular light brown markings; the punctures covering the body are not marked with brown. The pronotum is about 1.2 to 1.3 times as long as the scutellum (Fig. 6.705). Length: 1.55 to 1.80 mm. .............................................Neoplea (Neoplea) lingula Roback and Nieser, 1974 (Colombia). 4. (2) The length and maximum width of the scutellum are subequal (Fig. 6.703). The length is approximately 1.6 to 2.1 mm. ...........................................5 - The scutellum is obviously wider than long (Fig. 6.706). ................................7 5. (4) There are dense, fine punctures over most of the dorsal surface, except near the center of the pronotum and on the scutellum. The punctures are marked with dark pigment. The posterior half of the hemielytra, and rarely the anterior half, as well, has dark markings. The underside is mainly blackish. Length: c. 1.75 mm. The identity of this species is doubtful because it was described in a publication that was overlooked for more than 50 years, and the description is sketchy, at best. ........................................................Neoplea (Neoplea) semipicta (Horváth, 1918) (Colombia, Bolivia, Paraguay, Argentina, Minas Gerais). Syn: Plea semipicta Horváth, 1918. - The maximum width is clearly greater than half of the length. In dorsal view, the posterior lobes of the pronotum appear wider than the hemielytra, which are parallel-sided. The right paramere of the male is not slender (Fig. 6.704). ........6

Fig. 6.707 Neoplea gauchita (above, left to right): habitus in dorsal and lateral view, male genital capsule and right paramere, and (below, left to right): the eighth segment and the seventh urosternite of a male, the seventh urosternite of a female, and the right valve of the ovipositor. Based on Bachmann (1968). 6. (5) The length is very variable but generally approximates 2 mm or slightly more. The color is typically a mottled brown. The total length is about four times the scutellum length. In dorsal view, the posterior lobes of the pronotum appear elongated, their maximum width being at the midpoint or slightly posterior to the midpoint of their length (Fig. 6.704). .............................................................Neoplea (Neoplea) maculosa (Berg, 1879) (Surinam, Colombia, Peru, Bolivia, Paraguay, Argentina, Pará, Minas Gerais).

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- Length: c. 1.6 to 1.7 mm. The total length is 3.50 to 3.75 times the scutellum length. In dorsal view, the posterior lobes of the pronotum appear only slightly elongated, their maximum width being at a point 1/4 of the length or less from the posterior end. The last visible sternite of the female has a very distinct spot (Fig. 6.707). ........................................................Neoplea (Neoplea) gauchita Bachmann, 1968 (Bolivia, Argentina, Mato Grosso). 7. (4) The length is clearly greater than 1.6 times the maximum width. The sides of the hemielytra are parallel anteriorly. The right paramere of the male is very slender (Fig. 6.706). Length: 1.95 to 2.1 mm. The color is yellowish with indistinct and irregular light brown markings; the punctures covering the body are marked with brown. .........................................Neoplea (Neoplea) tenuistyla Roback and Nieser, 1974 (Colombia). - The length is equal to about 1.6 times the maximum width or less. The sides of the hemielytra are not parallel anteriorly (Fig. 6.708). ....................................8

Fig. 6.708 Neoplea globoidea (left to right): habitus in dorsal and lateral view and the parameres. Based on Nieser (1975). 8. (7) The length is about 1.6 times the maximum width. Viewed dorsally, the posterior lobes of the pronotum appear much wider than the hemielytra, which are widest at the base and narrow in a continuous curve posteriad (Fig. 6.708). Length: 1.6 mm. Color: light brown with white markings dorsally, dark brown ventrally. ............................................................Neoplea (Neoplea) globoidea Nieser, 1975 (Surinam). - The maximum width is equal to or slightly less than half of the length. Viewed dorsally, the posterior lobes of the pronotum are as wide as the hemielytra or very slightly wider. The hemielytra widen in a regular curve, their greatest width being at a point between 1/2 and 3/4 of the total length. The total length is between 3 2/3 and 3 3/4 times the scutellum length. The last visible sternite of the female has a faint spot (Fig. 6.709). Length: 2 mm. ....................................Neoplea (Neoplea) argentina (Drake and Chapman, 1953) (Argentina). Syn: Plea argentina Drake and Chapman, 1953.

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Fig. 6.709 Neoplea argentina (above, left to right): habitus in dorsal and lateral view, male genital capsule and right paramere, and (below, left to right): the eighth segment and the seventh urosternite of a male, the seventh urosternite of a female, and the right valve of the ovipositor. Based on Bachmann (1968).

Naucoridae Key to the subfamilies Information for the key was provided by Usinger (1941) and Nieser (1975). 1. The anterior margin of the pronotum is deeply emarginate behind the interocular space (Fig. 6.710). ..............................................................................2 - The anterior margin of the pronotum is straight or very slightly concave behind the interocular space (Fig. 6.711). ............................................................4 2. (1) The insertion of the rostrum is well posterior to the apical margin of the head. The head has a plate-like process anterior to the eyes. .........................................................................................................Cheirochelinae Found in Eastern Asia, this subfamily is not known to occur in South America. - The rostrum is inserted in the anterior margin of the head (Fig. 6.710). ..........3 3. (2) Plate-like extensions of the propleurae, which are contiguous at the midline, completely cover the posterior portion of the prosternum. The ventral part of the abdomen is densely pubescent, but the pubescence is interrupted around the spiracles and along transverse rows of punctures behind them. Only macropterous forms are known (Fig. 6.710). Nearctic and Neotropical. ..................................................................................................Ambrysinae..p. 518 - The prosternum is completely exposed and separated from the pleurae by simple sutures. There is no pubescence on the ventral side of the abdomen; a dish-like area is visible near each spiracle. The species are dimorphic, with the brachypterous form being more common (Fig. 6.712). Neotropical. ............................................................................................Cryphocricinae..p. 533

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Fig. 6.710 Carvalhoiella beckeri female: habitus in dorsal view (left), fore-leg (upper center), abdomen in ventral view showing the extent of the pilosity (upper right), and the tibia and tarsus of the hind leg (lower right). Based on De Carlo (1963d).

Fig. 6.711 Ctenipocoris spinipes: habitus in dorsal view (left) and fore-leg (right). Based on Alvarez and Roldán (1983). 4. (1) The anterior margin of the head is bent downward and backward, leaving the insertion of the rostrum well behind the actual anterior margin of the head. The hypostomal region is extremely short. The middle and hind femora each bear two longitudinal rows of conspicuous bristles on the ventral face in addition to the two rows of short bristles along the posterior or inner face. The fore-tarsi consist of two segments and bear two claws, which are often inconspicuous. ............................................................................................................Laccocorinae Formerly, four South American species were included in the predominantly Old World genus, Heleocoris Stål, 1876, which belongs in this subfamily. They

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have since been transferred to the genus Ctenipocoris Montandon, 1897, which is included in the subfamily Naucorinae. Therefore, at the present time, no laccocorine species are known from South America. - The anterior margin of the head is only slightly curved downward or backward. The hypostomal region is moderately long. The middle and hind femora lack distinct additional rows of bristles (Fig. 6.711). ..............................5

Fig. 6.712 Cryphocricos barozzii: habitus with the antenna shown just above the head (upper left), fore-leg (middle left), penis in ventral (lower left center) and lateral view (lower left), ventral view of the apex of the abdomen of a male (upper right) and a female (lower right center), urotergites of a male (lower right), and female genitalia (lower center). Based on López Ruf (1991). 5. (4) The inner eye margins diverge anteriorly. The meso and metasterna bear prominent longitudinal carinae, which are broad and foviate or otherwise excavated in the middle. The body is broadly oval and somewhat flattened. The fore-tarsi each consist of one segment and sometimes bear a single minute claw. The known late instar larvae are almost round (Fig. 6.713). Neotropical. ................................................................................................Limnocorinae..p.538 - The inner eye margins converge anteriorly. The meso and metasterna do not bear prominent longitudinal carinae, or their carinae are small, thin and platelike. The body is strongly convex and robust (Fig. 6.711). The known larvae are somewhat elongate and become wider across the abdomen in later instars (Fig. 6.714). Cosmopolitan. ...................................................................................................Naucorinae..p. 567

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Fig. 6.713 Limnocoris maculiceps: habitus of a macropterous adult (upper left) and a fifth larval instar (upper right), labrum (lower left), apex of the male abdomen in dorsal view with the segments extended apart and the genital capsule to the right (lower left center), apex of the female abdomen in ventral view (lower right), hind tibia with the stridulatory ridge in dorsal view (lower right center), right pseudoparamere (right of center), and left pseudoparamere in medial and dorsal view (right center above tibia, above and below, respectively). Based on Nieser and Lopez Ruf (2001).

Fig. 6.714 Pelocoris binotulatus nigriculus larval instars: habitus without appendages in dorsal view with the antennae above left and fore-leg above right of instars I to III (above, left to right) and instars IV and V (below, left and right, respectively). Based on Lopez-Ruf (1992a).

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Subfamily Ambrysinae Key to the Neotropical genera in the subfamily Ambrysinae Information for the key was provided by De Carlo (1963d), Nieser (1975), and Nieser et al. (1999). The classification of the genera and subgenera within the Ambrysinae has been unsettled in recent decades. The system used herein generally follows that of Nieser et al., 1999. 1. A coat of fine and dense setae covers the ventral surface of the abdomen all the way to the lateral margins (Fig. 6.715). ....................................................................................Ambrysus Stål, 1862..p. 519 - The pilose and setose area on the ventral surface of the abdomen is separated from the lateral margins by bare, shiny stripes (Fig. 6.716). ...............................2

Fig. 6.715 Ambrysus stali: head and prothorax in dorsal (left) and ventral view (left center), spines at the apex of the hind tibia (upper left of center), process on the seventh abdominal segment of a male (upper right of center), male parameres and penis (lower center), apex of the female abdomen in ventral view (upper right center), and outlines of the subgenital plates of two varying female specimens (right). Based on Nieser (1975). 2. (1) There is a deep, medial excision along the anterior margin of the pronotum. The punctures on the dorsal surface of the body contain distinct fine granules. .................................................................................Cataractocoris Usinger, 1941 This genus includes two known species confined to Mexico and Central America and is not further considered.

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- The anterior margin of the pronotum has only a shallow concavity in the middle (Fig. 6.716). If there are fine granules in the punctures on the dorsal surface of the body, they are indistinct. As far as known, the three species in this genus are confined to South America. .....................................................................Carvalhoiella De Carlo, 1963..p. 532

Fig. 6.716 Carvalhoiella stysi female: habitus in dorsal view (left) and apex of the abdomen in ventral view (right). Based on Nieser et al. (1999). Key to the known Neotropical subgenera of the genus Ambrysus Information for the key was provided by Nieser (1975), López Ruf (1996a), and Nieser et al. (1999). All species, regardless of subfamily, are included in the key to the species below this key. 1. There are at least four transverse rows of spines on the distal part of the hind tibia. The eyes are sharply angulate at their lateral angles (Fig. 6.717). The length is at least 10 mm. ..........................................................................................Picrops La Rivers, 1952 There is one species found only in northern South America. - There are no more than three transverse rows of spines on the distal part of the hind tibia. The lateral angles of the eyes are rounded (Fig. 6.718). ....................2 2. (1) The posterolateral angles of the pronotum are always obliquely truncate and feebly emarginated, producing sharp angles at the posterolateral corners. The right hemielytron of the brachypterous form has its membrane reduced to a

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narrow strip (Fig. 6.718). The claval suture is obsolete, and there is no apical transverse emboliar suture. Macropterous forms have never been found. All specimens are longer than 11 mm. .......................................................................................Melloiella De Carlo, 1935 Two species are known, both confined to the region of southern Brazil and northern Argentina. - The posterolateral angles of the pronotum are rounded. Most specimens are macropterous, but even brachypterous forms have hemielytra with welldeveloped membranes, easily distinguishable at least from the corium (Fig. 6.719). ...................................................................................................................3 3. (2) The hind wing lacks a costal cell. ......................................................................................Acyttarsus La Rivers, 1965 Only one species in this genus is known; it inhabits only North America. - The hind wing has a costal cell. ........................................................................4 4. (3) The prosternum is fused with the propleura medially. The general shape is elongate (Fig. 6.719). ........................................................................................Syncollis La Rivers, 1965 Most of the species are found in southern North America and Central America, with a few native to northern South America and one known in southern Brazil. - The middle posterior part of the prosternum appears to pass beneath the propleura, and it is not fused with them (Fig. 6.720). ................................................................................................Ambrysus Stål, 1862 This is by far the largest subgenus, represented by about 60 species ranging from North America to northern Argentina.

Fig. 6.717 Habitus of a male Ambrysus usingeri. Based on La Rivers and Nieser (1952).

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Fig. 6.718 Habitus of a male Ambrysus lamprus. Based on Nieser et al. (1999).

Fig. 6.719 The head and prothorax of Ambrysus teutonius in ventral view, showing the fused propleural plates. Based on Nieser (1975).

Fig. 6.720 Ambrysus kolla: habitus (left) and fore-leg (right). Based on López Ruf (2004).

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Key to the species of adults in the genus Ambrysus in South America Information for the key was provided by Montandon (1909b), La Rivers (1951, 1962a. 1970a), De Carlo (1950b, 1968d), La Rivers and Nieser (1972, 1975), Nieser et al. (1999), and López Ruf (2004, 2007a, b). Melloiella was considered to be a genus by some authors in the past, but it is treated as a subgenus here. 1. There are at least four transverse rows of spines on the distal part of the hind tibia. The eyes are sharply angulate at their lateral angles. The posterolateral angles of the pronotum form sharp angles. The prosternum and propleurae are firmly fused. The male lacks a genital process but bears a short lateral process on the sixth tergite (Fig. 6.717). The length is at least 10 mm. ........................................................Ambrysus (Picrops) usingeri La Rivers, 1952 (Guyana, French Guiana, Surinam, Pará, Amazonas, Mato Grosso). Syn: Ambrysus fittkaui De Carlo, 1966. - There are no more than three transverse rows of spines on the distal part of the hind tibia. The lateral angles of the eyes are rounded (Fig. 6.715). ....................2 2. (1) The posterolateral angles of the pronotum are always obviously obliquely truncate and feebly emarginate, producing sharp angles at the corners of the truncation. The right hemielytron of the brachypterous form has its membrane reduced to a narrow strip (Fig. 6.718). The claval suture is obsolete, and there is no apical transverse emboliar suture. Macropterous forms have never been found. All specimens are longer than 11 mm. .....................................................3 - The posterolateral angles of the pronotum are rounded, weakly and briefly truncate, or form sharp right angles. Most specimens are macropterous, but even brachypterous forms have hemielytra with well-developed membranes, easily distinguishable at least from the corium (Fig. 6.721). ..........................................4

Fig. 6.721 Habitus of Ambrysus truncaticollis in dorsal view. Based on Usinger (1941).

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3. (2) The length is approximately 12 mm. The color is mottled with yellow and brown markings (Fig. 6.718). ...............................Ambrysus (Melloiella) lamprus Nieser, Pelli, and Melo, 1999 (Minas Gerais). - The length is approximately 15 mm. Except for the embolium, the hemielytra are uniformly dark (Fig. 6.721), .............................................Ambrysus (Melloiella) truncaticollis De Carlo, 1935 (Argentina, Rio Grande do Sul, Santa Catarina, São Paulo). 4. (2) The coloration of the head, pronotum, and abdomen is predominantly brown. The lateral margins of the pronotum are not crenulated. The surfaces of the head and pronotum are finely and very densely granulated. The shape is elongate oval. The anterior corners of the pronotum form right angles. The length is shorter than 10 mm but longer than 7 mm. ............................................5 - The coloration is predominately yellowish, mottled, or variegated, or the head and pronotum are densely punctured and not granulated, or the length is not between 7 and 10 mm. ..........................................................................................6 5. (4) The length of the head is not greater than its width between the compound eyes. The embolium is mainly yellowish. Length: c. 9 mm; maximum width: c. 5 mm. .............................................Ambrysus (Ambrysus) colombicus Montandon, 1909 (Colombia). - The length of the head is greater than its width between the compound eyes. Most of the embolium is brownish. Length: 7.5 to 8.0 mm; maximum width: 4.2 to 4.5 mm. ..................................................Ambrysus (Ambrysus) horvathi Montandon, 1909 (Peru). 6. (4) The female subgenital plate has three protuberances along its posterior margin. One is median and two are lateral. At least some of the posterolateral corners of the laterotergites are drawn out into spines (Fig. 6.722). The size range is approximately 7.5 to 11 mm. ..................................................................7 - The female genital plate is not produced into three protuberances along the posterior margin, and if one is present, it is notched along the midline (Fig. 6.723). The male genital process does not have an “exaggerated dog’s head outline.” A better description or an illustration was not provided in the original descriptions of this group. ....................................................................................8 7. (6) The posterior margin of the female subgenital plate forms three points, one median and two lateral. The medial point is not notched along its midline. The male genital process has an “exaggerated dog’s head outline,” which has not been illustrated. The ratio of the distance between the eyes across the vertex to the synthlipsis equals 13:19. Length: c. 9 mm; Maximum width: c. 5 mm. Color: mottled brown and yellow. Only macropterous specimens have been described. The original description is sketchy, at best, but the features of the male and female genitalia are supposed to be distinctive within the genus. ....................................................Ambrysus (Ambrysus) tricuspis La Rivers, 1974 (Colombia).

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- The protuberances along the posterior margin of the female subgenital plate are broadly rounded, making the outline sinuous. The medial protuberance extends farther posteriad than the lateral ones. The dorsal surface of the head is smooth but has micropunctures. The outline of the head and pronotum together is strongly triangular with nearly straight sides and broadly rounded angles (Fig. 6.722). Length: 8 to 9 mm. Only the female was described. .......................................................Ambrysus (Ambrysus) scolius La Rivers, 1970 (Trinidad, Venezuela, Guyana).

Fig. 6.722 Habitus of Ambrysus (Ambrysus) scolius in dorsal view. Based on La Rivers (1970). 8. (6) The prosternum is fused with the propleura medially (Fig. 6.719). ...........9 - The middle posterior part of the prosternum appears to pass beneath the propleurae, and it is not fused with them (Fig. 6.715). ......................................11 9. (8) The dorsal surface is uniform yellow or yellowish brown, and the ventral color is yellow without conspicuous mottling. The posterolateral angles of the pronotum form sharp right angles. The posterior part of the pronotum has distinctive longitudinal grooves. Total length: 9 to 10.3 mm. Maximum width: 5 to 6 mm. .........................................................Ambrysus (Syncollis) planus La Rivers, 1951 (Ecuador, Peru). Two subspecies have been described. All known South American specimens belong to Ambrysus planus planus La Rivers, 1951; Ambrysus planus plax La Rivers, 1967, has been found only in Costa Rica. - The dorsal surface usually displays contrasting colors. The posterolateral angles of the pronotum do not form sharp right angles. The posterior part of the pronotum is not grooved longitudinally. ............................................................10

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10. (9) The dorsum has two weakly contrasting colors, with the color of the head and pronotum lighter or redder than the rest. The ventral coloration is variable but usually yellow or yellowish brown, sometimes with the abdomen distinctly darker than the thorax. The posterolateral corners of the pronotum are well rounded (Fig. 6.719). Length: 9 to 10 mm. Maximum width: 5 to 6 mm. .....................................................Ambrysus (Syncollis) teutonius La Rivers, 1951 (Colombia, Rio Grande do Sul, Santa Catarina, Minas Gerais, Mato Grosso). - The dorsal coloration of the male is shiny chestnut brown with two dark longitudinal stripes between the eyes; it is yellow ventrally (Fig. 6.723). Length: 11 mm. The female has not been described. ....................................................Ambrysus (Syncollis) partridgei De Carlo, 1968 (Amazonas).

Fig. 6.723 Habitus of Ambrysus partridgei in dorsal view. Based on De Carlo (1968d). 11. (8) Course punctures are concentrated on the posterior part of the head, and small punctures may be concentrated in the depression bordering the inner margins of the eyes of at least one species. The posterolateral corners of the connexival segments are obviously acuminate, especially on the posterior segments. The length varies from about 9 to 13 mm, and the maximum width is between about 5 and 7 mm. ................................................................................12 - The punctures on the head are not exclusively on the posterior part, or the posterolateral corners of the connexival laterotergites form pointed angles but are not strongly acuminate, or the length is shorter than 9 mm or longer than 13 mm. .....................................................................................................................14

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12. (11) Large punctures are abundant in the middle of the posterior part of the head and scattered sparcely on the rest of the surface together with a few small punctures. Small punctures are greatly concentrated in the depression bordering the inner margins of the eyes. The pronotum is covered by coarse punctures, and its margin is very weakly and finely crenulated. Length: c. 12.5 to 13.5 mm; maximum width: c. 7.1 to 7.2 mm. Color: mainly fulvous but with a brown labrum, scutellum, and underside; ochraceous connexiva narrowly bordered with fuscous; yellowish fore-legs; testaceous posterior part of the pronotum; and black compound eyes. ........................................................Ambrysus (Ambrysus) fossatus Usinger, 1946 (Colombia, Ecuador, Peru). - There are punctures only on the posterior part of the head. If the length is as long as 12.5 to 13 mm, the maximum width is no more than c. 6.5 mm. ..........13 13. (12) The lateral borders of the pronotum are strongly crenulated. The dorsal surface is shiny. The anterolateral corners of the pronotum are sharply pointed. Length: c. 10 to 11 mm. Maximum width: c. 5 to 6 mm. ...............................................Ambrysus (Ambrysus) crenulatus Montandon, 1897 (Colombia, Ecuador). - The lateral borders of the pronotum are, at most, very weakly crenulated. The dorsal surface is matt. The anterolateral corners of the pronotum are not usually acuminate at the apex. Length: 11 to 13 mm. Maximum width: c. 6 to 6.5 mm. ..............................................Ambrysus (Ambrysus) peruvianus Montandon, 1909 (Peru). 14. (11) The length is greater than 10 mm. ........................................................15 - The length is no more than 10 mm. .................................................................20 15. (14) At the base of the operculum, the median part of the posterior margin of the sixth abdominal segment of the female lacks a tooth (Fig. 6.724). The male cannot yet be identified. .................................................Ambrysus (Ambrysus) gemignanii De Carlo, 1950 (Argentina). - At the base of the operculum, the median part of the posterior margin of the sixth abdominal segment of the female bears a tooth (Fig. 6.725). ...................16

Fig. 6.724 Ambrysus gemignanii female (left to right): antenna, paramere, and operculum. Based on De Carlo (1950b).

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Fig. 6.725 Ambrysus fucatus (left to right): antenna, parameres of two specimens, and genital operculum of a female. Based on De Carlo (1950b). 16. (15) Length: 14.1 to 15.5 mm. ......................................................................17 - The length is less than 14 mm. ........................................................................18 17. (16) The corion of the egg is covered mainly by rectangular and triangular fields, each with 2 to 5 pores. The lateral margins of the pronotum are obviously curved. The length of the fore-femur divided by the width is about 1.41. The apex of the abdomen is broadly rounded (Fig. 6.726). Total length: 14.2 to 15.0 mm. ..................................................Ambrysus (Ambrysus) calilegua López Ruf, 2007 (Argentina). - The corion of the egg is covered mainly by polygonal fields, each with 6 to11 pores. The lateral margins of the pronotum are relatively straight, giving the pronotum a triangular shape. The length of the fore-femur divided by the width is about 1.12. The apex of the abdomen tapers, making it somewhat pointed (Fig. 6.720). Total length: 14.3 to 15.5 mm. .........................................................Ambrysus (Ambrysus) kolla López Ruf, 2004 (Argentina).

Fig. 6.726 Ambrysus calilegua female: habitus without appendages (left) and the fore-leg in posterior view (right). Based on López Ruf (2007b).

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18. (16) The ratio length:width is c. 3:2 (Fig. 6.725). Length: c. 13 mm. Maximum width: c. 8.5 mm. ...............................................................Ambrysus (Ambrysus) fucatus Berg, 1879 (Argentina). - The ratio length:width is 9:5. ..........................................................................19 19. (18) The anterolateral angles of the pronotum and the posterolateral angles of the connexival segments are acuminate. The head is slightly shorter than the pronotum, measured along the mid-line. The widest point across the female genital operculum (plate) is more than three times the distance across the excavation at the apex (Fig. 6.727). Length: 13.0 to 14.4 mm. .......................................................Ambrysus (Ambrysus) bergi Montandon, 1897 (Argentina). - Neither the anterolateral angles of the pronotum nor the posterolateral angles of the connexivum are sharply acuminate. The head is slightly longer than the pronotum, measured along the mid-line. The widest point across the female genital operculum (plate) is less than three times the distance across the excavation at the apex (Fig. 6.728). Length: 11.5 to 13.2 mm. Posterior width of the pronotum: 5.96 to 6.27 mm. ...............................................Ambrysus (Ambrysus) ochraceus Montandon, 1909 (Argentina, Bolivia).

Fig. 6.727 Ambrysus bergi (left to right): antenna, extent of the yellowish area along the lateral margin of the prosternum, parameres of two specimens, and genital operculum of a female. Based on De Carlo (1950b).

Fig. 6.728 Ambrysus ochraceus (left to right): antenna, extent of the yellowish area along the lateral margin of the prosternum, paramere, and genital operculum of a female. Based on De Carlo (1950b).

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20. (14) The posterolateral angles of the laterotergites are obtuse, with a slight degree of spine formation evident only on the fifth segment, i.e., the fourth visible segment. Length: 9.4 mm. Color: brown mottled with yellowish. The female has not been described. ..............................................Ambrysus (Ambrysus) obscuratus Montandon, 1898 (Pernambuco, Minas Gerais). - The posterolateral angles of both the fourth and fifth laterotergites bear distinct spines (Fig. 6.729). ................................................................................21 21. (20) The hemielytron lacks a clavus and is reduced, leaving a broad part of the conexivum and apex of the abdomen uncovered (Fig. 6.729). Length: 8.0 to 8.6 mm. Color: yellowish to light brown variegated with dark brown. ...........................................................Ambrysus (Ambrysus) siolii De Carlo, 1966 (Pará). - The hemielytron has a clavus; all but a narrow part of the conexiva and the apex of the abdomen are covered by the hemielytra. .........................................22

Fig. 6.729 Habitus of Ambrysus siolii. Based on Nieser (1975). 22. (21) Length: 9.1 to 9.8 mm. The female subgenital plate is trisinuate along its apical margin; the lateral sinuosities are angulate, and the median sinuosity is formed by an extended convexity between the two lateral ones. The male genital process is blunt, wide, and rather straight along the inner margin and rounded along the outer edge but not extending laterally to form a salient (Fig. 6.715). Color: brown mottled with yellowish. ...........................................................Ambrysus (Syncollis) stali La Rivers, 1962 (Trinidad, Surinam, French Guiana, Argentina, Pará, Amazonas). Syn: Ambrysus bourquini De Carlo, 1968. Nieser et al. (1999) suggested that this taxon might actually be a subspecies of Ambrysus obscuratus. - The length is less than 9.0 mm. .......................................................................23

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Fig. 6.730 Ambrysus bifidus habitus (below), and (above, left to right): the male genital process on the seventh abdominal tergite of three varying specimens, penis and parameres, apex of the female abdomen in ventral view, and outline of the female subgenital plate. Based on Nieser (1975). 23. (22) The subgenital plate of the female is quadrisinuate at the apex; the lateral sinuosities are bluntly angulate, and the median sinuosity is formed by two close, elevated angles. The genital process of the male is blunt, wide, and curved laterad along the inner margin and extended laterally to form a blunt salient on the outer edge (Fig. 6.730). Length: 7.9 to 8.6 mm. .....................................Ambrysus (Ambrysus) bifidus La Rivers and Nieser, 1972 (Surinam, Amazonas, Pará, Mato Grosso). - The subgenital plate of the female is not quadrisinuate, or the median sinuosity is not formed by two elevated angles. The genital process of the male is not blunt, wide, and curved laterad along its inner margin (Fig. 6.731). ..................24 24. (23) A distinct median depression is evident on the anterior part of the pronotum. The shape is broadly oval. The excavation at the apex of the female genital operculum is a minute notch (Fig. 6.731). Length: c. 8.5 mm. Maximum width: c. 5.2 mm. ............................................Ambrysus (Ambrysus) acutangulus Montandon, 1897 (Argentina, Rio Grande do Sul). - There is no median depression evident on the anterior part of the pronotum. The body is narrower, with a length of c. 8.2 mm and a width of c. 4.5 mm (Fig. 6.732). .................................................................................................................25

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Fig. 6.731 Ambrysus acutangulus (left to right): antenna, lateral margins of the posteriormost abdominal segments, genital operculum of a female, and a paramere. Based on De Carlo (1950b). 25. (24) The anterolateral angles of the pronotum are acute but rounded at their apices. Fine crenulations are evident on the lateral margins of the pronotum. The head and pronotum are uniformly yellowish. .................................................Ambrysus (Ambrysus) fraternus Montandon, 1897 (Goiás, Mato Grosso). - The angles at the anterolateral corners of the pronotum are obtuse. The lateral margin of the pronotum is not crenulate (Fig. 6.732). ...............................................Ambrysus (Ambrysus) attenuatus Montandon, 1897 (Argentina).

Fig. 6.732 Ambrysus attenuatus: habitus (left), fore-leg in anterior view (center), genital capsule of a male (upper right), and female genitalia (lower right). Based on Lopez Ruf and Hernández (2007).

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Key to the species of adults in the genus Carvalhoiella in South America Information for the key was provided by Nieser et al. (1999). 1. Length: c. 8.5 mm. The dorsal surface of the male is smooth and shiny (Fig. 6.733). The color is an almost uniform brown, but the anterior part of the dorsum has a reddish tinge, and the posterior part is blackish. There is also a faint yellow band across the posterior part of the pronotum and some yellow on the embolium. The female has not been described. ......................................................................Carvalhoiella nitida La Rivers, 1976 (Surinam). - Length: at least 10 mm. The dorsum is covered with punctures and appears dull (Fig. 6.716). ...................................................................................................2

Fig. 6.733 Habitus of a male Carvalhoiella nitida in dorsal view. Based on La Rivers (1976). 2. (1) The length is from 10 to 11 mm. The apical margin of the female operculum is concave (Fig. 6.716). ...................................................Carvalhoiella stysi Nieser, Pelli, and Melo, 1999 (Minas Gerais). - The length is c. 12 mm. The apical margin of the female operculum is straight (Fig. 6.710). The head and pronotum are dark yellow, and the hemielytra are mainly black, except for the outer edges of the embolium, which are bright yellow with dark markings. Only one specimen, a female, has been described. .....................................................................Carvalhoiella beckeri De Carlo, 1963 (Minas Gerais).

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Subfamily Cryphocricinae This subfamily encompasses three genera: Cryphocricos Signoret, 1850, and Procryphocricos Polhemus, 1991, found in South America, and Cataractocoris Usinger, 1941, from Mexico. The South American species are brachypterous, while the hemielytra of Cataractocoris cover the abdomen (Nieser, 1975). Key to the genera of adult Cryphocricinae in South America Information for the key was provided by Polhemus (1991b). 1. In addition to the short setae binding the plastron, there are long, silky setae. The middle and hind femora are not armed with spines or knobs. Each tibia is armed with one irregular subdistal row of spines and one short distal row. The seventh abdominal segment of the male is symmetrical and lacks a spur. The third and fourth antennal segments are subequal in length. Two kinds of ocular setae are present. The abdominal tergites are fused. The penis is only slightly asymmetrical and has a short flagellum (Fig. 6.734). The only known species is about 5.1 mm long. ...........................................................................Procryphocricos Polhemus, 1991 The only known species in this genus is Procryphocricos perlexus Polhemus, 1991, from Venezuela. Only the adult male has been described. - No long, silky setae are present in addition to the short setae of the plastron. The middle and hind femora are armed with spines or knobs. The tibiae are armed with two rows of closely packed spines. The seventh abdominal segment is asymmetrical and bears a spur on the right side. The fourth segment of the antenna is much longer than the third. There are no ocular setae. The abdominal tergites are distinct. The aedeagus is very asymmetrical and bears a long flagellum (Fig. 6.735). .......................................................................Cryphocricos Signoret, 1850..p. 534

Fig. 6.734 Procryphocricos perlexus adult male (left to right): habitus (above) and antenna (below), apex of the abdomen in dorsal and ventral view, and the penis. Based on Polhemus (1991b).

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Fig. 6.735 Cryphocricos daguerrei: habitus of a male (upper left) and a female (upper right), fore-leg of a male (lower left) and a female (lower right center), penis in ventral (lower left center) and lateral view (lower center) and female genitalia (lower right). Based on López Ruf (1991). Key to the species of adult Cryphocricos in South America Information for the key was provided by Usinger (1947), De Carlo (1951b, 1967c), and López Ruf (1991). 1. The lateral margins of the second and third abdominal segments are slightly arcuate (Fig. 6.735). Only brachypterous specimens have been described. Length: c. 9.3 mm. ..................................................................Cryphocricos daguerrei DeCarlo, 1940 (Argentina). - The lateral margins of the second and third abdominal segments follow the general curve of the abdomen (Fig. 6.736). .........................................................2

Fig. 6.736 Habitus of Cryphocricos granulosus. Based on Nieser (1975).

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2. (1) The dorsal surfaces of the body and hemielytra are heavily granulated, and the general color is light to dark brown, sometimes with a reddish tinge (Fig. 6.735). The length apparently ranges between 9.5 and 10.5 mm, but relatively few specimens have been examined. ....................................................................3 - The dorsal surfaces are not heavily granulated (Fig. 6.737). ............................5 3. (2) The hemielytra reach as far posteriad as the middle of the third abdominal segment, and their posterior margins are nearly straight (Fig. 6.736). Length: 9.8 to 10 mm. Color: generally dark with brown legs. ................................................................Cryphocricos granulosus DeCarlo, 1967 (Rio Grande do Sul). - The hemielytra just reach the anterior portion of the second abdominal segment (Fig. 6.738). ............................................................................................4 4. (3) The body is more than half as wide as its total length, the ratio being 33:60. The posterior margins of the hemielytra are curved (Fig. 6.738). There are deep transverse folds on the prosternum. Only brachypterous specimens have been described. Length: c. 10 mm. The color is light brown with a reddish tinge on most of the dorsal and ventral surface. ..........................................................................Cryphocricos rufus DeCarlo, 1940 (Rio de Janeiro). - The body of the male is slightly less than half as wide as its total length, the ratio being 49:100. The posterior margins of the hemielytra are straight (Fig. 6.739). There are no transverse folds on the pleurae and prosternum. Length of male: c. 10 mm. The dorsum is mainly light chestnut brown, but the hemielytra are brownish. Only one male specimen has been described. ......................................................................Cryphocricos montei De Carlo, 1951 (São Paulo).

Fig. 6.737 Cryphocricos vianai (above, left and right): habitus of an adult and a fifth larval instar, and (below, left to right): fore-leg of a male, penis in ventral and lateral view, ventral view of the apex of the abdomen of a male, and fore-leg of a female. Based on López Ruf (1991).

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5. (2) The posterior margins of the hemielytra are straight and rather abruptly rounded laterally (Fig. 6.740). ..............................................................................6 - The posterior margins of the hemielytra are arcuate and broadly rounded laterally (Fig. 6.737). ............................................................................................7

Fig. 6.738 Habitus of a brachypterous Cryphocricos rufus in dorsal view. Based on De Carlo (1941).

Fig. 6.739 Cryphocricos montei male: habitus without appendages (left) and two views of the penis (right). Based on De Carlo (1951b) and López Ruf (1991).

Fig. 6.740 Habitus of a brachypterous Cryphocricos peruvianus in dorsal view. Based on De Carlo (1941).

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6. (5) Length: less than 10 mm. Only brachypterous specimens have been described (Fig. 6.740). ................................................................Cryphocricos peruvianus DeCarlo, 1940 (Peru, Ecuador). - Length: greater than 10 mm for brachypterous and 11 mm for macropterous forms (Fig. 6.712). .....................................................................Cryphocricos barozzii Signoret, 1850 (Colombia, Ecuador, Argentina, Santa Catarina). 7. (5) The lateral margins of the pronotum appear slightly concave in dorsal view, and the posterolateral corners are significantly wider than the anterior part of the pronotum, giving the thorax a wedge-shape (Fig. 6.741). Both brachypterous and macropterous forms are known. ...............................................................Cryphocricos breddini Montandon, 1911 (Peru, Ecuador, Argentina). Syn: Cryphocricus breddini Montandon, 1911 (spelling error). - The lateral margins of the pronotum are convexly curved or straight (Fig. 6.742). Granules are sparce on the pronotum and scutellum, but if their surfaces are rugose, then small granules may cover the hemielytra. ..................................8

Fig. 6.741 Cryphocricos breddini: habitus of an adult (upper left), wing (upper right), fore-leg (lower left), and the female genitalia (lower right). Based on López Ruf (1991). 8. (7) The lateral margins of the pronotum are convex, and the posterior margin is only moderately wider than the anterior part (Fig. 6.742). The surfaces of the body and fore-coxae, trochanters, and femora have a rather sparce covering of small granules, but the middle and hind femora are entirely smooth. Length: c. 10.5 mm. Maximum width: c. 3.8 mm. Color: both the dorsal and ventral surfaces and the legs are predominantly blackish. Only a brachypterous male has been described. .....................................................................Cryphocricos fittkaui De Carlo, 1967 (Peru).

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- The lateral margins of the pronotum are straight, divergent, and lined with about ten coarse teeth on each side (Fig. 6.737). The surfaces of the pronotum and scutellum are rugose, while the hemielytra are covered with small granules. Length: c. 10.5 mm. Maximum width: c. 4.9 mm. .......................................................................Cryphocricos vianai De Carlo, 1951 (Argentina, Rio Grande do Sul, Minas Gerais). Syn: Cryphocricos barozzii Signoret, 1850, pars, specimens from Minas Gerais misidentified by Nieser and Melo (1997), fide Melo and Nieser (2004).

Fig. 6.742 Cryphocricos fittkaui male (left to right): habitus in dorsal view without appendages with an enlarged outline of the lateral pronotal margin to the upper left, the apical segments of the abdomen, and the genital operculum. Based on De Carlo (1968d). Subfamily Limnocorinae In older literature, this subfamily encompassed three nominal genera, two of which, Limnocoris Stål, 1860, and Sattleriella De Carlo, 1966, were known from South America. The consensus today seems to be that all known South American species in this subfamily should be included in one genus: Limnocoris. Key to the species of adult Limnocoris in South America Information for the key was provided by Montandon (1898), De Carlo (1941, 1951c, 1965, 1966b, 1967c), La Rivers (1962b, 1970b, 1974, 1976), Roback and Nieser (1974), Nieser (1975), Nieser et al. (1993), Nieser and Melo (1999b), Nieser and Lopez Ruf (2001), and Pagioro et al. (2007). The species Limnocoris volxemi (Lethierry, 1877), originally described under the name Borborocoris volxemi Lethierry, 1877, is not included in the key. There are no records indicating on or near which continent the type specimens were found, and its natural range remains unknown.

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1. The anterior margin of the head protrudes to form an acute angle in the middle, and its lower margin has a long ventral fold. The posterolateral corners of the pronotum form acute angles. The surface of the pronotum is densely covered by punctures of variable sizes. The posterolateral angles of the connexivum are weakly spinose, more prominently so on the posterior segments. The mesosternal carina is considerably elevated above the mesosternal surface and deeply and widely cleft. The claval suture on the hemielytron is considerably longer than half the length of the scutellum. The membrane is poorly developed. Length: c. 8.8 mm; maximum width: c. 6.9 mm. ...................................................................Limnocoris bouvieri Montandon, 1898 (West Indies, Colombia, Venezuela? Chile?). - The anterior margin of the head is broadly rounded or forms an obtuse angle (Fig. 6.743). The lower margin of the head has no ventral fold or a weak one and is usually vertical. ..........................................................................................2

Fig. 6.743 Limnocoris illiesi, brachypterous specimen (left to right): habitus, apex of the male abdomen in dorsal view, median and dorsal view of the right pseudoparamere and the left pseudoparamere (above to below), and the apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001).

Fig. 6.744 Habitus of Limnocoris bruchi in dorsal view without appendages. Based on De Carlo (1967c).

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2. (1) The body is distinctly egg-shaped with the posterior end narrowed, and the head and thorax broadly rounded anteriorly (Fig. 6.744). The length of the male is approximately 6.4 mm, and its maximum width is c. 4.4 mm. The female is about 6.7 mm long, and its maximum width is c. 4.5 mm. The posterolateral corners of the anterior segments of the connexiva very slightly overlap the segments posterior to them, while those of the three posteriormost segments form distinct spines. There are a few long, fine hairs along the border of the abdomen. The posterior corners are the widest points of the pronotum and rounded, but the widest part of the body is near its midlength, where the embolium extends laterad. The color of the body and legs is brownish yellow with darker membranes of the hemieltyra. ..........................................................................Limnocoris bruchi De Carlo, 1967 (Amazonas, Mato Grosso). - The length is shorter than 6.0 mm or longer than 7.0 mm, or, if not, then one of the three posteriormost segments is not distinctly spinose, or there is a strongly contrasting mottling or other color pattern, or the body shape is oval or elliptical and not distinctly egg-shaped (Fig. 6.743). ...........................................3

Fig. 6.745 Limnocoris pusillus (left to right): habitus of a brachypterous specimen, apex of the male abdomen in dorsal view, mesosternal carina of the fifth larval instar in lateral view, median and dorsal view of the right pseudoparamere (above and below, respectively), right pseudoparamere, and the apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 3. (2) The length is 4.7 to 6.0 mm. At least one laterotergite of the connexiva forms a spine at its posterolateral corner (Fig. 6.743). The general coloration is drab yellow to dark brown without a strongly contrasting pattern. ......................4 - Length: at least 6.5 mm, or, if it is shorter, both of the posteriormost segments of the connexiva are not distinctly spinose (Fig. 6.745). .....................................5

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4. (3) The posterolateral corners of the two posteriormost laterotergites of the connexiva are distinctly spinose. The mesosternal carina is short and appears forked (Fig. 6.743). Length: 4.7 to 5.8 mm. Color: light to medium brown. No macropterous specimens have been found. ...........................................................................Limnocoris illiesi De Carlo, 1967 (Colombia, Amazonas, Pará, Minas Gerais). - Only the posterolateral corners of the fourth laterotergites of the male connexiva bear spines, which are small and fine. In the female, only the fifth laterotergites bear such spines. Length: 5.5 to 6.0 mm. Maximum width: 4.0 to 4.3 mm. Color: dark brown and drab yellow dorsally, producing only a weakly contrasting pattern. The ventral surface is weakly mottled with black and yellow. Only macropterous specimens with fully developed metathoracic wings have been described. ................................................................Limnocoris woytkowskii La Rivers, 1970 (Peru). 5. (3) The lateral margins of the pronotum are broadly rounded and show little trace of a posterolateral corner. The posterolateral corners of the connexival segments form obtuse angles. The lateral margins of the connexiva are finely serrated with 19 to 21 teeth per mm. The genital operculum of the female has the shape of a pentagon with sides of roughly equal length and rounded corners (Fig. 6.746). Length: c. 6.0 to 6.2 mm. Maximum width: c. 3.8 to 3.9 mm. Color: light maroon on the head, pronotum, and ventral surface; dark maroon on the scutellum and hemielytra. ................................Limnocoris gracilis Nieser, Gonzales, and Eichelkraut, 1993 (Colombia). - The lateral margins of the pronotum are usually widest at or near their posterolateral corners, which are angular, rounded near the apex, or slightly truncate (Fig. 6.747). If not, then the length is 6.5 mm or greater. ....................6

Fig. 6.746 Limnocoris gracilis (left to right): anterior part of the habitus in dorsal view, mesosternal keel in lateral view, right lobe of the male fifth tergite, and apex of the female genital operculum. Based on Nieser et al. (1993). 6. (5) The hemielytra of brachypterous specimens do not reach the fourth abdominal tergite, leaving at least some of the third exposed (Fig. 6.747). The length is at least 8 mm. .........................................................................................7

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- The hemielytra of brachypterous specimens reach at least to the fifth abdominal tergite, or only macropterous specimens have been described (Fig. 6.748). ...................................................................................................................8

Fig. 6.747 Limnocoris siolii (left to right): habitus of a brachypterous specimen, apex of the male abdomen in dorsal view, left hemielytron (above) and a ventral view of the mesosternal carina above its lateral view (below), and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001).

Fig. 6.748 Habitus of Limnocoris brauni in dorsal view. Based on De Carlo (1966b). 7. (6) The wings of brachypterous specimens cover part of the third abdominal segment (Fig. 6.747). Length: 8.1 to 8.3 mm. No macropterous specimens have been found. Color: medium brown dorsally with black anterior parts of the connexival segments and some indistinct darker markings; the ventral surface is medium brown to yellowish with yellowish legs. ..........................................................................Limnocoris siolii (De Carlo, 1966) (São Paulo, Paraná). Syn: Sattleriella siolii De Carlo, 1966.

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- The wings of brachypterous specimens do not reach the third abdominal segment. Length: c. 9.5 to 9.8 mm. Width: c. 6.0 to 6.8 mm. The color of brachypterous specimens is reddish-brown or blackish with the anterior parts lighter. Both brachypterous and macropterous specimens are known. ................................................................Limnocoris malkini La Rivers, 1974 pars (Colombia). Syn: Limnocoris pulchellus La Rivers, 1974. See Couplet 46 for the macropterous form.

Fig. 6.749 Limnocoris ovatulus (left to right): habitus of a macropterous female, apex of the male abdomen in dorsal view, the right (above) and left pseudoparamere (below), and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 8. (6) The posterolateral corners of the fourth laterotergites are spinose or obtusely spinose (Fig. 6.749), although the spine may be small; the total length of the insect is clearly greater than 7 mm. ............................................................9 - The posterolateral angles of the fourth laterotergites of the connexivum are not spinose, or if they are, the length does not exceed about 7 mm (Fig. 6.750). ....18 9. (8) The length of the male is c. 8.0 mm, and the maximum width is c. 5.4 mm. The posterolateral corners of the pronotum are broadly rounded, and the hemielytra leave only a small portion of the connexiva uncovered (Fig. 6.748). The embolium does not have a complete transverse suture. The inner margins of the eye are straight. The scutellum is somewhat rugose. The color is almost uniformly brown but with somewhat paler lateral margins of the pronotum and a slightly darker membrane of the hemielytra. The abdomen is dark yellowish and covered with pilosity. The legs are yellowish. Only the male has been described. A remarkably poor description will make this species hard to identify with certainty until a revision is undertaken. ..........................................................................Limnocoris brauni De Carlo, 1966 (Peru).

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- The length is significantly longer or shorter than 8 mm, or, if it is not, the maximum width of the male is greater than 5/8 of the length, or, if the length to maximum width ratio is about 8:5, the color is not relatively uniform, and some markings are evident (Fig. 6.749). .....................................................................10

Fig. 6.750 Limnocoris abbreviatus (above, left to right): habitus of a brachypterous male, apex of the male abdomen in dorsal view, apex of the female abdomen in ventral view, and (below, left to right): mesosternal carina in dorsal and lateral view, apex of the left and the right pseudoparamere. Based on Nieser and Lopez Ruf (2001). 10. (9) The posterolateral corners of the third and fourth segments of the connexivum are spinose, but that of the second is not (Fig. 6.749). The length is usually less than 10 mm. ....................................................................................11 - Either the second and third segments of the connexivum have spinose posterolateral corners, or neither of these corners are spinose (Fig. 6.751). ......12 11. (10) Along the lateral margin of the fourth segment of the connexivum, there are about 20 fine denticles per mm. The posterolateral corners of the third through sixth abdominal segments are obviously spinose (Fig. 6.749), and the connexival segments display obvious dents. The width of the anterior margin of the vertex is 1.1 times the synthlipsis. The claval suture is evident on the hemielytra, which cover all but 15% to 25% of the width of the connexival segments. Length of male: 7.9 to 9.2 mm; female: 8.4 to 9.6 mm. Maximum width of male: 5.40 to 6.18 mm; female: 5.90 to 6.37 mm. Color: almost uniformly light brown with a few dark brown or blackish markings, sometimes including a triangle on the interocular surface of the head, a row of dots along the anterior margin of the pronotum, and costal stripes from the embolium to the membrane. The legs are yellowish. ...................................................................Limnocoris ovatulus Montandon, 1897 (Argentina, Bolivia).

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- Along the lateral margin of the fourth segment of the connexivum, there are about 16 to 18 fine denticles per mm. The width of the anterior margin of the vertex is about 1.25 times the synthlipsis. The mesosternal carina is relatively small (Fig. 6.752). Length of male: c. 9.5 mm; maximum width: c. 6.8 to 6.9 mm. Color: mainly maroon with parts of the hemielytra blackish. The description was based on only one male specimen. ..............................Limnocoris abrasum Nieser, Gonzales, and Eichelkraut, 1993 (Colombia).

Fig. 6.751 The habitus of Limnocoris birabeni in dorsal view. Based on De Carlo (1967c).

Fig. 6.752 Limnocoris abrasum (left to right): anterior part of the body in dorsal view, mesosternal carina in lateral view, right lobe of the fifth abdominal tergite of the male. Based on Nieser et al. (1993).

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12. (10) The posterolateral corners at least of the second through fourth laterotergites of the connexiva are spinose. ........................................................13 - The posterolateral corners of the second and third laterotergites of the connexiva are not spinose, but the corners of the fourth bear obtuse or fine spines (Fig. 6.751). .............................................................................................14 13. (12) The posterolateral corners of the second through fourth laterotergites of the connexiva are spinose. The color is mainly ferrugineous with a yellowish brown head, pronotum, legs, expansions of the embolium, and exposed parts of the connexivum. The lateral borders of the connexiva are serrate, with about 20 teeth per mm along the edge of the fourth laterotergite. The secondary paramere is greatly enlarged at the apex (Fig. 6.753). Length: 10.3 to 11.2 mm. The width of the pronotum is 6.4 to 7.2 mm. .......................................................Limnocoris robustus Roback and Nieser, 1974 (Colombia). - The posterolateral corners of all laterotergites of the connexiva, except for the first, form spines. The color is nearly uniform yellowish without distinctive markings or mottling. The compound eyes are only slightly elevated above the plane of the head, narrow, and strongly divergent. Length: c. 10 mm. Maximum width: c. 7 mm. ...................................................................Limnocoris carcharus La Rivers, 1976 (Venezuela).

Fig. 6.753 Limnocoris robustus male (left to right): metasternal keel with the broken line indicating the base of the fossa, right lobe of the fifth tergite, and the secondary paramere. Based on Roback and Nieser (1974). 14. (12) The length is at least c. 8.3 mm. ...........................................................15 - The length is between c. 7 and 8 mm. .............................................................17

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15. (14) The pronotum is widest across its posterolateral corners, which are scarcely rounded and form acute angles. The cavity in the mesosternal carina is deep (Fig. 6.754). The lateral margins of the connexiva are serrate and bear 19 to 21 tiny denticles per mm. Length: 9.2 to 9.9 mm. Maximum width: 6.5 to 7.0 mm. Head width: 3.13 to 3.25 mm. Synthlipsis: 1.50 to 1.65 mm. Color: mainly maroon with a dark gray band across the posterior part of the pronotum. The prosternum and legs are yellowish. .............................Limnocoris angulatus Nieser, Gonzales, and Eichelkraut, 1993 (Colombia). - The pronotum is widest slightly anterior to the posterolateral corners, which are broadly rounded or appear truncate (Fig. 6.755). .........................................16

Fig. 6.754 Limnocoris angulatus (left to right): anterior part of the body in dorsal view, mesosternal carina in lateral view, right lobe of the fifth abdominal tergite of the male, and the female genital operculum. Based on Nieser et al. (1993).

Fig. 6.755 Limnocoris calii (left to right): anterior part of the body in dorsal view, mesosternal carina in lateral view, right lobe of the fifth abdominal tergite of the male, and the female genital operculum. Based on Nieser et al. (1993).

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Fig. 6.756 Limnocoris trilobatus (left to right): anterior part of the body in dorsal view, mesosternal carina in lateral view, and the posterior margins of the female opercula of two specimens. Based on Nieser et al. (1993).

Fig. 6.757 The habitus of Limnocoris fittkaui in dorsal view. Based on De Carlo (1967c). 16. (15) The synthlipsis is slightly less than half the width of the head. The prosternal carina is triangular with two punctures, and the mesosternal carina extends for little more than half the length of the mesosternum. The genital operculum of the female is roughly pentagonal with rounded corners (Fig. 6.755). Length of the male: 8.5 to 9.0 mm; female: 9.6 to 9.9 mm. Color: maroon on the head, anterior part of the pronotum, connexiva, and legs, dark maroon on the thoracic sternum, and dark to blackish with lighter markings on the rest of the body. ......................................Limnocoris calii Nieser, Gonzales, and Eichelkraut 1993 (Colombia).

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- The synthlipsis of the female is slightly greater than half the width of the head. The mesosternal carina extends for most of the length of the mesosternum. The genital operculum of the female is trilobed along its posterior margin (Fig. 6.756). Length of the female: 10.5 to 11.0 mm. Color: maroon on the head, pronotum, scutellum, and anterior part of the embolium; the eyes and hemielytra are dark, and the connexiva and legs are yellowish red. The ventral surface is dark with yellowish setae. The male has not been described. .............................Limnocoris trilobatus Nieser, Gonzales, and Eichelkraut, 1993 (Colombia). 17. (14) The four dark lines on the interocular area of the head occupy less than ¼ of its width (Fig. 6.751). Length: 6.8 to 7.0 mm. Maximum width: 4.8 to 5.0 mm. The dorsal surface is blackish except for the yellowish head and lateral parts of the body. .......................................................................Limnocoris birabeni De Carlo, 1967 (Amazonas, Pará). The original descriptions of the two species in this couplet appear very similar, suggesting that larger series of specimens should be examined to determine whether the two are really distinct species. - The four dark lines on the interocular area of the head occupy more than ½ of its width (Fig. 6.757). Length: 6.5 to 8.5 mm. Maximum width: 4.8 mm or greater. The dorsal surface is light chestnut brown with yellowish markings on the head and lateral parts of the pronotum, embolium, and connexiva. .........................................................................Limnocoris fittkaui De Carlo, 1967 (Surinam, Amazonas). Specimens from Surinam were assigned to a distinct subspecies, Limnocoris fittkaui surinamensis Nieser, 1975, distinguished by their length of 7.8 to 8.5 mm, as compared with 6.5 to 7.3 mm for Limnocoris fittkaui fittkaui from Amazonas.

Fig. 6.758 Limnocoris burmeisteri: habitus (left), apex of the male abdomen in ventral view (lower center), lobes of the male secondary parameres (upper center), penis and parameres of two specimens (lower right center), the right lobe of the fifth tergites from two males (upper right), posterior margins of two subgenital plates of different females (lower right). Based on De Carlo (1967c) and Nieser (1975), who ascribed some figures to L. bachmanni, now a synonym.

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Fig. 6.759 Limnocoris exogkoma: anterior habitus in dorsal view (left), prosternal carina in lateral view (upper right center), right lobe of the fifth tergite of a male (right), and the genital operculum of a female (lower right center). Based on Manzano et al. (1995).

Fig. 6.760 Limnocoris intermedius: habitus of a brachypterous female (upper left), apex of the male abdomen in dorsal view (upper right), left (lower center) and right pseudoparamere (lower left), and apex of the female abdomen in ventral view (lower right). Based on Nieser and Lopez Ruf (2001).

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18. (8) Posterior to the embolium, the hemielytra curve mesad to expose at least approximately 10% of the maximum abdominal width, measured across the dorsum of macropterous adults and more than 10% of brachypterous specimens (Fig. 6.750). ........................................................................................................19 - Posterior to the embolium, the hemielytra expose no more than 8% of the maximum abdominal width (Fig. 6.758). This couplet is based on the specimens used for the initial descriptions of the species. Because of the known intraspecific variability of the wings of aquatic heteropterans, specimens may vary considerably in this feature. It is therefore recommended to follow both alternatives in this couplet to reduce the possibility of error. .............................32

Fig. 6.761 Limnocoris acutalis (left to right): habitus of a brachypterous male, mesosternal carina in ventral (above) and lateral view (below), left (below) and right pseudoparamere (above), and apex of the male abdomen in dorsal view. Based on Nieser and Lopez Ruf (2001). 19. (18) The prosternal carina is triangular with a process at the apex (Fig. 6.759). The hemielytra lack claval sutures. The connexiva lack spines at the posterolateral corners but have a minute denticulation along the lateral margin, with about 20 to 22 denticles per mm. The maximum width is approximately equal to the greatest width of the pronotum. Length of male: 8.6 to 9.2 mm; female: 9.2 to 9.5 mm. Maximum width of males and females: 6.0 to 6.3 mm. Color: much of the head, pronotum, and legs are light red; the eyes are black, and the central part of the pronotum and much of the rest of the body and hemielytra are dark. .................................Limnocoris exogkoma Manzano, Nieser, and Caicedo, 1995 (Colombia). - The prosternal carina is not triangular with a distinctive process at the apex. If the specimen is between 8.6 and 9.2 mm long, its pronotum at its widest point is narrower than the maximum width of the insect (Fig. 6.750). ...........................20 20. (19) Length: at least 9.2 mm. Width: 6.9 mm or wider. ..............................21 - Length: no more than 9.1 mm. Width: no more than 6.7 mm. ........................22

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21. (20) The anteclypeus extends 0.4 mm anterior to the labium, which is 0.45 mm wide and 0.35 mm in median length (Fig. 6.750). Length: c. 10.3 mm. Width: c. 7.5 mm. Color: dull yellowish without any mottling. ................................................................Limnocoris abbreviatus La Rivers, 1974 (Rio de Janeiro, Santa Catarina, Rio Grande do Sul). Syn: Limnocoris montandoni La Rivers, 1974. - The anteclypeus extends 0.2 mm anterior to the labium or less; the labium is 0.6 mm wide and 0.5 mm in median length (Fig. 6.760). Length: 9.2 to 9.8 mm. No macropterous specimens have been found. Color: medium brown with lighter markings, brown to blackish eyes, yellow legs and ventral surfaces of the head and thorax, and a chestnut brown ventral surface of the abdomen. .............................................Limnocoris intermedius Nieser and Lopez Ruf, 2001 (Paraná). 22. (20) The embolium is triangular with its apex pointed outward and the anterior part of its lateral margins straight, followed by a strong inward curve (Fig. 6.761). Length: c. 7.4 mm. The species is known from only one brachypterous male. .......................................................................Limnocoris acutalis La Rivers, 1974 (Brazil, exact location unknown). - The embolium is not triangular, and the anterior halves of its lateral margins are convex (Fig. 6.762). .....................................................................................23

Fig. 6.762 Limnocoris asper (left to right): habitus of a brachypterous female, apex of the male abdomen in dorsal view, left (above) and right pseudoparamere (below), and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 23. (22) The pronotum of the brachypterous form is widest far anterior of its posterior margin, and its lateral margins are broadly rounded, producing poorly defined posterolateral angles (Fig. 6.762). .........................................................24 - The pronotum of the brachypterous form is widest near the posterior margin, and the posterolateral corners form well-defined acute angles (Fig. 6.763). .....26

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Fig. 6.763 Limnocoris submontandoni: habitus of a macropterous (upper left) and a brachypterous female (upper right), apex of the male abdomen in dorsal view (lower left), apex of the female abdomen in ventral view (lower center), and the right and left pseudoparamere (lower right, above and below, respectively). Based on Nieser and Lopez Ruf (2001). 24. (23) The general coloration is medium brown with the dark brown interocular pattern consisting only of rows of points (Fig. 6.762). Length: 7.2 to 7.6 mm. Maximum width: 5.0 to 5.4 mm. A macropterous form has not been described. .......................................................Limnocoris asper Nieser and Lopez Ruf, 2001 (Minas Gerais). - The general coloration is purplish dark brown with a well-developed interocular pattern (Fig. 6.764). Length: 7.6 to 8.6 mm. Maximum width: 5.6 to 6.4 mm. ..............................................................................................................25 25. (24) The interocular marking is deeply constricted at midlength, giving its anterior part an arrowhead shape (Fig. 6.12). Length: 8.2 to 8.6 mm. Maximum width: 5.9 to 6.4 mm. Color: dark brown with lighter markings and a membrane that is mainly black. Macropterous forms have not been described. .................................................Limnocoris espinolai Nieser and Lopez Ruf, 2001 (Minas Gerais). - The interocular marking is only slightly constricted at midlength (Fig. 6.764). Length: 7.6 to 8.4 mm. Maximum width: 5.7 to 5.9 mm. The ventral surface is mainly pruinose grayish with light to medium brown on parts of the head and pronotum; the legs are also light to medium brown. Only the brachypterous form is known. ...............................................Limnocoris porphyros Nieser and Lopez Ruf, 2001 (Minas Gerais).

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Fig. 6.764 Limnocoris porphyros (left to right): habitus of a brachypterous male, apex of the male abdomen in dorsal view, apex of the female abdomen in ventral view, and (top to bottom): the right pseudoparamere in dorsal and medial view and the left pseudoparamere. Based on Nieser and Lopez Ruf (2001). 26. (23) The length is no more than 8.1 mm. The pronotum reaches a maximum width of about 3.5 mm at its midlength and forms sharp angles at its posterolateral corners (Fig. 6.339). The general color is yellowish brown to medium brown with yellow legs and yellow on lateral parts of the ventral surface. ................................................................................Limnocoris insignis Stål, 1876 (Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina, Rio Grande do Sul). Syn: Limnocoris uhleri Montandon, 1910; Limnocoris admontandoni La Rivers, 1974. - The length is greater than 8.1 mm. ..................................................................27 27. (26) The posterior margin of the embolium is approximately perpendicular to the long axis of the body. The ratio of the vertex width to the synthlipsis is about 1.48:1, and the ratio of the length to the greatest width is 1.28:1. An elongated mesosternal fossa is present (Fig. 6.763). Length: 8.2 to 9.1 mm. Color: the head and pronotum are yellow with small brown dots and a brown posterior margin of the pronotum; the scutellum and hemielytra are medium brown with a darker membrane and sometimes a yellow embolium. The ventral surface is mainly yellow with darker central parts of the abdomen. ...........................................................Limnocoris submontandoni La Rivers, 1974 (São Paulo, Minas Gerais). - The posterior margin of the embolium is not nearly perpendicular to the long axis of the body (Fig. 6.765). .............................................................................28 28. (27) The ratio of the maximum width of the pronotum to the length of the head and pronotum is 2:1 or slightly greater in the brachypterous form (Fig. 6.765). .................................................................................................................29 - The ratio of the maximum width of the pronotum to the length of the head and pronotum of the brachypterous form is less than 2.0:1 (Fig. 6.766). .................30

555

Fig. 6.765 Limnocoris caraceae (left to right): habitus of a brachypterous male, apex of the male abdomen in dorsal view, the right and left pseudoparamere (above and below, respectively), and apex of the female abdomen in ventral view (lower center). Based on Nieser and Lopez Ruf (2001).

Fig. 6.766 Limnocoris nigropunctatus (left to right): habitus of a brachypterous male, left pseudoparamere (lower center), apex of the male abdomen in dorsal view, right pseudoparamere in medial (above) and dorsal view (below), and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 29. (28) There is no distinct anterior tooth on the mesostigmal carina. The pronotum has rounded posterolateral corners. The posterior margin of the female operculum is broadly rounded (Fig. 6.765). Length: c. 8.8 mm. The head near the eyes, marginal parts of the pronotum and scutellum, connexiva, and embolium are yellow. The discs of the pronotum and sometimes also the scutellum are brown. The legs and ventral surface are orange yellow with darker markings on the central parts of the thorax. The eyes are black. .................................................Limnocoris caraceae Nieser and Lopez Ruf, 2001 (Minas Gerais).

556

- There is a distinct anterior tooth on the mesostigmal carina. The posterolateral corners of the pronotum form acute angles. The posterior margin of the female operculum is formed by diagonal sides, which meet along the mid-line at nearly a right angle (Fig. 6.767). Length: 8.2 to 8.8 mm. Color: mainly light brown with small brown spots, larger brown markings, and yellow legs. Brachypterous and macropterous forms have been described. ................................................Limnocoris lanemeloi Nieser and Lopez Ruf, 2001 (Minas Gerais). 30. (28) The posterolateral angle of the embolium is well defined and fairly acute (Fig. 6.766). Length: 8.2 to 8.9 mm. ........................................................Limnocoris nigropunctatus Montandon, 1909 (Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina). Syn: Limnocoris sattleri De Carlo, 1966. - The posterolateral angle of the embolium is broadly rounded and poorly defined (Fig. 6.768). ...........................................................................................31

Fig. 6.767 Limnocoris lanemeloi: habitus of a brachypterous male (upper left), apex of the male abdomen in dorsal view (upper center), apex of the female abdomen in ventral view (upper right), mesosternal ridge in ventral and lateral view (lower left, above and below, respectively), left (lower center) and right pseudoparamere (lower right). Based on Nieser and Lopez Ruf (2001). 31. (30) The right membrane forms a rounded angle; it is about 1/6 as long as a hemielytron (Fig. 6.768). Length of male: 8.1 to 8.3 mm; female: 8.3 to 8.6 mm. Only the brachypterous form is known. The predominant color is orange brown with darker, grayish brown markings, mainly on the median parts of the thorax and abdomen. ..................................................Limnocoris decarloi Nieser and Lopez Ruf, 2001 (Rio de Janeiro, Santa Catarina, Paraná). Syn: Limnocoris sattleri De Carlo, 1966 partim, due to misidentification of specimens from Rio Grande do Sul.

557

- The right membrane of the brachypterous form is broadly rounded; it occupies more than 1/3 of the length of the hemielytron (Fig. 6.769). Length: 7.8 to 8.3 mm. Color: medium brown to brownish yellow, with darker on a brown triangle between the eyes, the central part of the pronotum, and a marking on the scutellum. A macropterous form has not been described. ...................................................................Limnocoris rotundatus De Carlo, 1951 (Paraná, Santa Catarina, Rio Grande do Sul).

Fig. 6.768 Limnocoris decarloi: habitus of a brachypterous female (left), apex of the male abdomen in dorsal view (upper center), apex of the female abdomen in ventral view (upper right), left (lower right) and right pseudoparamere (lower center). Based on Nieser and Lopez Ruf (2001).

Fig. 6.769 Limnocoris rotundatus (left to right): habitus of a brachypterous male, apex of the male abdomen in dorsal view, left and right pseudoparamere (above and below, respectively), and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001).

558

Fig. 6.770 Limnocoris borellii (left to right): habitus of a brachypterous specimen, mesosternal carina of a fifth instar larva in lateral view with its ventral apex to the right (lower left center), apex of the male abdomen in dorsal view, left (below) and right lobe of the eighth abdominal segment (above) together with its respective pseudoparamere, and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 32. (18) Length: 10.8 to 13.3 mm. Maximum width: greater than 8.2 to 9.8 mm. The pronotum has somewhat acute posterolateral angles (Fig. 6.770). Brachypterous and macropterous specimens have been described; the latter were formerly referred to by the synonym, Limnocoris stali. .....................................................................Limnocoris borellii Montandon, 1897 (Central America, Colombia, Venezuela, Peru, Bolivia, Argentina). Syn: Limnocoris stali Montandon, 1897; Limnocoris manco-capaci Poisson, 1954; Limnocoris profundus Stål (nec Say), 1862. - Specimens are usually less than 10 mm long, but if one is longer, then its maximum width is no greater than 7.1 mm (Fig. 6.771). ...................................33 33. (32) Length: 4.7 to 6.5 mm. The posterolateral angles of the two posterior segments of the connexiva are not distinctly spinose. The mesosternal carina lacks a narrow groove on the anterior face of the central body but has a deep, hairy indentation with two teeth anterior to it on the anterior part (Fig. 6.745). ....................................................................Limnocoris pusillus Montandon, 1897 (Colombia, Bolivia, Argentina, Minas Gerais, Rio de Janeiro). Syn: Limnocoris mansosotoi De Carlo, 1951; Limnocoris vianai De Carlo, 1967; Limnocoris illiesi Roback and Nieser, 1974 (nec De Carlo), misidentification of Colombian specimens. - The length is greater than 6.5 mm. ..................................................................34 34. (33) Length: 6.5 to 7.1 mm. ..........................................................................35 - The length is at least 7.3 mm. .........................................................................38 35. (34) The color is mottled shades of dark brown, brown, and yellow. ..........36 - The color is fairly uniform and not mottled. The fossa of the mesosternal carina does not end in a small tubercle. ..............................................................37

559

36. (35) The lateral edges of the anterior part of the connexivum form an almost continuous curve without indentations between the segments. The mesosternal carina is elevated, narrowed in the middle, and with a clearly evident fossa, which proceeds along an oblique laminar structure ending in a small tubercle. The tooth-like process on the right lobe of the fifth tergite is somewhat narrow at the base (Fig. 6.758). Length: 6.2 to 8.0 mm. ..................................................................Limnocoris burmeisteri De Carlo, 1967 (Surinam, Amazonas, Pará). Syn: Limnocoris bachmanni De Carlo, 1967. - The posterolateral corners of the connexivum form moderately large spines and do not form a continuous curve. The tooth-like process on the right lobe of the fifth abdominal segment of the male is fairly short (Fig. 6.772). The hemielytra are uniform in color, except for the lighter emboliar margins, but the anterior parts of the dorsum are mottled and appear generally lighter in color. Length: c. 7 mm. Width: c. 5 mm. ........................................................................Limnocoris menkei La Rivers, 1962 (Venezuela).

Fig. 6.771 Limnocoris minutus (left to right): habitus of a macropterous male, apex of the male abdomen in dorsal view, the lobes on the eighth abdominal segment without the pseudoparameres, and apex of the female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 37. (35) The color is a fairly uniform brownish yellow. The mesosternal carina has a well-developed fossa, which extends for 1/3 or more the length of the carina (Fig. 6.771). Length: 6.7 to 7.1 mm. Maximum width: 4.6 to 4.9 mm. Color: almost uniform brownish yellow but with a few scattered brown spots. ........................................................................Limnocoris minutus De Carlo, 1951 (Argentina, Paraguay, Mato Grosso).

560

- The color of the hemielytra of the brachypterous form is uniformly dark purplish brown, except for the dark central part of the pronotal disc and the strongly contrasting yellowish lateral parts. The mesosternal carina has a small, narrow fossa extending only about 1/6 of the length of the carina (Fig. 6.773). Length: 6.3 to 6.9 mm. The macropterous form has not been described. .....................................................Limnocoris saphis Nieser and Lopez Ruf, 2001 (Minas Gerais). 38. (34) Length: 10.3 to 11.2 mm. The fore-femur of the male has a large, conspicuous black spot, which usually covers most of its ventral surface (Fig. 6.774). ..................................................................Limnocoris brasiliensis De Carlo, 1941 (Minas Gerais, Rio de Janeiro). Syn: Limnocoris bergi De Carlo, 1941. - The length is not greater than 10.1 mm. There is no conspicuous black spot on the ventral surface of the fore-femur (Fig. 6.774). .............................................39

Fig. 6.772 The right lobe of the fifth abdominal segment of a male Limnocoris menkei. Based on La Rivers (1962).

Fig. 6.773 Limnocoris saphis (left to right): habitus of a brachypterous male, apex of the male abdomen in dorsal view, apex of the female abdomen in ventral view, and (top to bottom): the right pseudoparamere in medial and dorsal view and the left pseudoparamere. Based on Nieser and Lopez Ruf (2001). 39. (38) The central part of the mesosternal carina is thin because the central fossa is very wide and deep (Fig. 6.775). Length: 9 to 11 mm. Color: light or medium brown with variable dark mottling; the hemielytra are medium brown with blackish on the corium and membrane. ................................................................Limnocoris pectoralis Montandon, 1897 (Peru, Bolivia, Argentina). Syn: Limnocoris peruvianus Melin, 1930.

561

- The central part of the mesosternal carina is thick because its fossa is shallower. The coloration is uniform except for a darker membrane, or the anterior part of the pronotum has black spots, and the scutellum or elytral margins are partially black, giving the insect a mottled color pattern (Fig. 6.776). Length: 7.2 to 10.1 mm. ........................................................................40 40. (39) The length is shorter than 9 mm, or the ratio of the total length to the maximum width is greater than 1.41:1. ..............................................................41 - The length is greater than 9 mm, or the ratio of the length to the maximum width is less than 1.4:1. If not, the opening to the central fossa of the mesosternal carina is not ovoid with the narrow end almost pointed at the apex (Fig. 6.776). ........................................................................................................44

Fig. 6.774 Limnocoris brasiliensis: habitus of a brachypterous specimen (upper left), left fore-leg of a male (lower left), apex of the male abdomen in dorsal view (upper center), apex of the female abdomen in ventral view (lower center), and right pseudoparamere in inner and dorsal view and left pseudoparamere (right, top to bottom). Based on Nieser and Lopez Ruf (2001). 41. (40) There is no tubercle at either end of the central fossa of the mesosternal carina. .................................................................................................................42 - There is a tubercle at the anterior end of the central fossa of the mesosternal carina. .................................................................................................................43 42. (41) The ratio total length:maximum width is 1.33:1. Length: c. 8.8 mm. The maximum width is c. 6 mm. The mesosternal carina is slightly farther from the surface of the mesosternum at the posterior end than at the anterior end, and its opening becomes slightly wider posteriad. The posterolateral corners of the pronotum form nearly right angles. The scutellum is rugose. The coloration is

562

nearly uniform reddish yellow, but it is slightly paler on the posterior part of the pronotum and on the embolium, and the membrane is black. ................................................................Limnocoris ochraceus Montandon, 1898 (Colombia). - The ratio total length:maximum width is 1.37:1. Length of the female: c. 8.2 mm. The length of the male is reported as somewhat less. The width is c. 6 mm. The opening to the central fossa of the mesosternal carina is ovoid with the narrow end almost pointed at the apex. ...............................................................................Limnocoris rivalis Melin, 1930 (Peru). The original description of this species is exceedingly poor, and a better one is urgently needed.

Fig. 6.775 Limnocoris pectoralis (left to right): habitus of a macropterous female, right (above) and left lobe (below) on the eighth abdominal segment with the pseudoparameres, and apices of the male abdomen in dorsal view and female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001). 43. (41) The posterolateral corners of the pronotum are broadly rounded and not nearly acute. The distal margin of the mesosternal carina is not elevated at either end and runs nearly parallel to the surface of the mesosternum; it has an elongated opening into the central fossa for its entire length, and there is a small tubercle at its truncate anterior end. The embolium on each side is greatly expanded laterad, making the lateral margins of the hemielytra obviously sinuous. Length: c. 8 mm. Maximum width at the embolium: c. 5.7 mm. The color pattern displays strong contrast between the yellow on much of its body, the brown markings on the scutellum and anterior margins of the laterotergites, and the black on its hemielytra, which covers the entire membrane and markings anterior to them. .................................................................Limnocoris bergrothi Montandon, 1898 (Venezuela).

563

- The posterolateral corners of the pronotum are almost acutely pointed but narrowly rounded at the apices of the angles. The opening of the central fossa of the mesosternal carina is rather narrow for its entire length, and it has a tubercle at its anterior end. Length: 7.5 to 8.0 mm. Maximum width: 5.3 to 5.6 mm. Color: mainly pale yellowish with blackish posterior margins of the hemielytra and brownish at the posterior end of the membrane and sometimes on parts of the embolium. .........................................................................Limnocoris pallescens (Stål, 1861) (West Indies, Colombia, Venezuela). Syn: Borborocoris pallescens Stål, 1861. 44. (40) Length: 7.2 to 7.5 mm. Maximum width: 4.9 to 5.0 mm. Ratio of length to width: 1.49:1 to 1.50:1 (Fig. 6.776). The serrated edge of the connexiva has about 20 teeth per mm. Color of the macropterous form: yellowish on most of the head, pronotum, embolium, connexivum, and legs with dark brown or blackish eyes, a brown marking between the compound eyes, and some brown mottling. The scutellum and hemielytra are mainly brown with yellow mottling. The posterior part of the brown marking between the eyes is semicircular, and the anterior part is triangular. A brachypterous form has not been described. ..................................................Limnocoris machrisi Nieser and Lopez Ruf, 2001 (Goiás). - Length: 8.0 to 10.1 mm. The distinctly but finely serrated edge of the connexiva has about 25 teeth per mm, or, if the serrations are coarser, then the ratio of the length to the width is not greater than 1.42:1, or, if not, there are blackish spots on the anterior part of the pronotum, and parts of the scutellum are also black (Fig. 6.777). .................................................................................45

Fig. 6.776 Limnocoris machrisi (above, left to right): habitus of a macropterous male, apex of the male abdomen in dorsal view, apex of the female abdomen in ventral view, and (top to bottom): the right pseudoparamere in medial and dorsal view and the left pseudoparamere, and (lower left center) the mesosternal carina in ventral (above) and lateral view (below). Based on Nieser and Lopez Ruf (2001).

564

45. (44) The dorsal pattern is spotted or mottled. The posterior margin of the pronotum is sinuous. The mesosternal carina is narrow for its entire length. The widest point is at the embolia, and the ratio of the length to the maximum width is c. 1.415:1. Length: c. 8 mm. Maximum width: c. 5.4 mm. Only c. 2 mm of the connexivum is left exposed on each side (Fig. 6.777). Only one specimen has been described, which will make identification tentative until more is learned about the morphological variability of this species. ....................................................................Limnocoris maculatus De Carlo, 1951 (Colombia). - The dorsal pattern is not notably spotted or mottled, or, if it is, the mesosternal carina is wide for part of its length, or the ratio of the length to the maximum width is less than 1.410:1 or more than 1.420:1. ................................................46

Fig. 6.777 Habitus of Limnocoris maculatus. Based on La Rivers (1974). 46 (45) The mesosternal carina of the female is narrow and has a small tubercle at its anterior end. The fourth antenna segment is less than half as long as the third. The ground color is yellowish, but part of the scutellum and spots on the anterior part of the pronotum are black. The genital segments of the female are symmetrical. The length is 8 to 9 mm. Only the female has been described. ......................................................................Limnocoris aymarana Poisson, 1954 (Peru). - The mesosternal carina of the female is wider, or it lacks a tubercle at the anterior end (Fig. 6.778). If not, then the dorsal surface lacks black markings, or the length does not fall in the range of 8 to 9 mm. .............................................47

565

47. (46) The apex of the female subgenital plate is narrowly, evenly rounded. The eye surfaces are in the same plane as the head surface. The lateral margins of the pronotum are smoothly rounded with bluntly curved posterolateral corners. The posterolateral angles of the connexiva form right angles. The connexiva have a ribbed structure. The length is 9.5 to 9.8 mm, and the maximum width is 6.0 to 6.8 mm. The color is almost uniform yellow or somewhat darker without any mottling but with darker wing membranes. The original description is too sketchy to make a positive identification without examining the type specimens. ...............................................................Limnocoris malkini La Rivers, 1974 pars (Colombia). Syn: Limnocoris pulchellus La Rivers, 1974. See Couplet 7 for the brachypterous form. - The length is usually shorter than 9 or longer than 10 mm, but if it is not, then the color pattern shows more contrast, or the ratio of total length:maximum width does not fall in the range between 1.44:1 and 1.58:1, or the connexiva do not form right angles at the posterolateral corners, or the posterolateral corners of the pronotum are more acute (Fig. 6.779). .....................................................48 48. (47) The ratio of the length to the width is 1.43:1 to 1.55:1. The serrations on the connexiva are coarse with teeth numbering 17 to 18 teeth per mm (Fig. 6.779). Length: 8.3 to 9.4 mm. Maximum width: 6.3 to 6.6 mm. ....................49 - The ratio of the length to the width is 1.31:1 to 1.42:1, but it is only rarely more than 1.40:1. The serrations on the connexiva are relatively fine but distinct (Fig. 6.780). Length: 8.2 to 10.1 mm. Maximum width: 5.5 to 6.7 mm. .........50 49. (48) The lateral margins of the labrum are concave (Figs. 6.13, 6.778). ..................................................................Limnocoris aculabrum La Rivers, 1974 (Santa Catarina). - The lateral margins of the labrum are straight (Fig. 6.713). Length: 8.3 to 9.4 mm. ..............................................................Limnocoris maculiceps Montandon, 1897 (Minas Gerais, Santa Catarina, Paraná, Mato Grosso do Sul).

Fig. 6.778 Limnocoris aculabrum (left to right): labrum, mesosternal carina in ventral (above) and lateral view (below), apex of a male abdomen in dorsal view, and apex of a female abdomen in ventral view. Based on Nieser and Lopez Ruf (2001).

566

50. (48) The length of the female is 7.9 to 8.2 mm. The mesosternal carina has a small central part (Fig. 6.779). The male and the macropterous form have not been described. Color: mainly medium brown with darker mottling. The legs and much of the connexivum are yellowish. ...............................................Limnocoris subpauper Nieser and Lopez Ruf, 2001 (São Paulo, Paraná). - The length is at least 8.9 mm. .........................................................................51

Fig. 6.779 Limnocoris subpauper: habitus of a brachypterous female (left) and apex of the female abdomen in ventral view (right). Based on Nieser and Lopez Ruf (2001).

Fig. 6.780 Limnocoris pauper: habitus of a brachypterous female (left), apex of the male abdomen in dorsal view (upper right center), apex of the female abdomen in ventral view (upper right), left pseudoparamere (lower center), and right pseudoparamere in medial and dorsal view (lower right center, above and below, respectively). Based on Nieser and Lopez Ruf (2001).

567

51. (50) The mesosternal carina has a small central part that is about 1/3 of its total length (Fig. 6.780). .....................................................................Limnocoris pauper Montandon, 1897 (São Paulo, Paraná, Santa Catarina, Rio Grande do Sul, “Cumbase,” an unknown locality in Northeastern Brazil, perhaps in Piauí). Syn: Limnocoris plaumanni La Rivers, 1973. - The mesosternal carina of the female has a large central part that is somewhat more than 1/2 of its total length (Fig. 6.781). Length: 8.9 to 9.8 mm. The male and the macropterous form have not been described. .................................................................Limnocoris melloleitaoi De Carlo, 1951 (Paraná).

Fig. 6.781 Limnocoris melloleitaoi: habitus of a brachypterous female (left) and apex of the female abdomen in ventral view (right). Based on Nieser and Lopez Ruf (2001). Subfamily Naucorinae Key to the genera of adults in South America Information for the key was provided by Nieser (1975) and Bachmann and Mazzucconi (1995). 1. The fore-tarsus consists of two segments and bears two claws. The body is rather narrow (Fig. 6.782) with a matt dorsal surface covered by fine punctures. The coloration is dark. ...................................................................Ctenipocoris Montandon, 1897..p. 569 - The fore-tarsus consists of a single segment and bears a single claw (Fig. 6.783). ...................................................................................................................2

568

Fig. 6.782 Habitus of Ctenipocoris brasiliensis in dorsal view. Based on De Carlo (1968d).

Fig. 6.783 Pelocoris magister: habitus in dorsal view (left), fore-leg (upper right center), two views of a male paramere (upper and middle right), and ventral view of the female genital segments (lower right). Based on Lopez Ruf (1994b). 2. (1) The middle femur is expanded in the proximal half. The propleurae are not elevated above the plane of the prosternum. The second valvae of the female are fused into a long and narrow organ bearing short, blunt tubercles along its lateral margins and irregularly placed medial tubercles, all on the apical half (Fig. 6.784). ...................................................................................Placomerus La Rivers, 1956 There is only a single known species in this genus: Placomerus micans La Rivers, 1956, from Paraguay, Bolivia, and São Paulo. It is 6.5 to 7.0 mm long and 3.5 to 4.0 mm wide at its widest point, which is located within the anterior third of its abdomen. The color is mainly reddish brown to blackish on a smooth, polished surface with white punctures. The last three larval instars have been described by López Ruf, M. (1996b).

569

- The middle femur is not expanded in the proximal half (Fig. 6.783). The propleura are elevated above the plane of the prosternum at the posterointernal margins, making the tubular median sclerite posterior to the prosternum appear to be sunken in. The second valvae of the female are fused into a wide, flat organ bearing only microscopic spines. The general habitus of the five larval stages resembles that of the adults (Fig. 6.343). .....................................................................................Pelocoris Stål, 1876..p. 570

Fig. 6.784 Placomerus micans (left to right): sclerite at the base of the adeagus, subgenital plate of the female, right first valva, and fused second valvae in dorsal view. Based on La Rivers (1956). Key to the species of adult Ctenipocoris in South America Information for the key was provided by De Carlo (1940, 1958, 1968d) and López Ruf (2004). The species in this genus were previously considered species of Heleocoris, which seems to be confined to the Old World. López Ruf (2004) confirmed that Ctenipocoris brasiliensis is distinct from the other species but suggested that the other three may not be distinct from one another. However, she did not place them in synonymy. Their cursory descriptions do not provide any differences significant enough to distinguish them with confidence, if such differences exist. A revision of the genus is needed. 1. The extension of the metasternal plate is not elevated above the plane of the plate and does not form a stiff spine. The posterolateral corners of the abdominal segments do not tend to form small, pointed extensions. The connexiva are smooth and lack punctures (Fig. 6.782). Length: c. 8 mm. Maximum width: c. 5 mm. .............................................................Ctenipocoris brasiliensis (De Carlo, 1968) (Pará). Syn: Heleocoris brasiliensis De Carlo, 1968. - The extension of the metasternal plate is elevated above the plane of the plate and forms a stiff spine directed posteriad. The posterolateral corners of some of the abdominal segments tend to form small, pointed extensions (Fig. 6.785). ....2

570

Fig. 6.785 Habitus of Ctenipocoris plaumanni in dorsal view. Based on De Carlo (1968d). 2. The metasternal carina forms a large pointed spine, and the mesosternal carina forms a small tubercle. ..............................................................Ctenipocoris spinipes (Montandon, 1897) (Colombia, Amazonas, Minas Gerais, Rio de Janeiro, Paraná). Syn.: Heleocoris spinipes Montandon, 1897. - The mesosternal carina does not form a small tubercle. ...................................3 3. The connexiva are light brown with a dark marking on each of the first three segments. Length of male: 8.6 to 9.9 mm; female: 9.2 to 9.5 mm. Maximum width: at least approximately 5.3 mm (Fig. 6.5, p.17). ...................................................................Ctenipocoris schadei (De Carlo, 1940) (Paraguay, Argentina, Paraná). Syn: Heleocoris schadei De Carlo, 1940. - The entire ventral surface of the abdomen is dark, except for a pale area on and sometimes adjacent to the genital segments. The widest part of the body is across the posterior half (Fig. 6.785). The Length: c. 8.5 mm. Maximum width: c. 5.0 mm. ..............................................................Ctenipocoris plaumanni (De Carlo, 1968) (Santa Catarina). Syn: Heleocoris plaumanni De Carlo, 1968. Key to the species of adult Pelocoris in South America Information for the key was provided by Nieser (1975, 1994), who noted that a taxonomic revision of the genus is badly needed; Lopez Ruf (1992a, b; 1994a, b), and Lopez Ruf and Nieser (1999). 1. The posterolateral angles of the second through fourth laterotergites are not distinctly spinose. The aedeagus is widened subapically (Fig. 6.786), or it is widest at about mid-length. Length: 8.5 to 9.7 or 16 to 18 mm, according to different authors. ...................................................................................................2

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- At least the posterolateral angles of the fourth laterotergite are spinose (Fig. 6.787). ...................................................................................................................4

Fig. 6.786 Pelocoris poeyi (left to right): right lateral margins of the second through fourth abdominal segments, penis and parameres, and the apex of a female abdomen in dorsal and ventral view. Based on Nieser (1975).

Fig. 6.787 Pelocoris procurrens: habitus (upper left), antenna (lower left center fore-leg (left center), right lateral margins of the second through fourth abdominal segments (upper left center), two views of a male paramere (right center), and ventral view of the female genital segments (right). Based on Nieser (1975) and Lopez Ruf (1994b). 2. Length: 16 to 18 mm. Color: shiny dark brown with small, irregular markings covering the entire dorsum. The penis is widest at about mid-length (Fig. 6.783). ........................................................................Pelocoris magister La Rivers, 1971 (Argentina, Brazil). - Length: 8.5 to 9.7. Color: generally brownish to olive brown with the head and pronotum yellowish brown with dark punctures. The aedeagus is widened subapically (Fig. 6.786). .......................................................................................3 3. The dorsal side of the fore-femur bears a distinct reddish brown patch (Fig. 6.714). ...........................................................................Pelocoris binotulatus (Stål, 1860) (Mexico, Colombia, Argentina, Amazonas, Rio de Janeiro, Minas Gerais). Syn: Naucoris binotulatus Stål, 1860. Both subspecies have been encountered from

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Colombia to Argentina. Pelocoris binotulatus nigriculus Berg, 1879, is still known only from those two countries, while Pelocoris binotulatus binotulatus is presently known from Panama, Paraguay, Argentina, and Brazil. - There is no reddish brown patch on the dorsal side of the fore-femur (Fig. 6.786). Length of male: 8.5 mm; of female: 9.3 mm. ..............................................................Pelocoris poeyi (Guérin-Méneville, 1835) (West Indies, Guyana, Surinam, Pará). 4. Length: 5.4 to 6.4 mm, rarely exceeding 6.5 mm. The dorsal side of the forefemur bears a distinct reddish brown patch. The aedeagus is slender with indistinct chitinous bars laterally (Fig. 6.787). Color: light to medium brown with a yellowish brown head and pronotum bearing darker markings. ..........................................................................Pelocoris procurrens White, 1897 (Paraguay, Argentina, Pará, Amazonas). Syn: Pelocoris minutus Montandon, 1895. - The length exceeds 7 mm. .................................................................................5 5. Length: 11.7 to 13.4 mm. The body shape is nearly oval. The posterolateral angles of the posterior abdominal segments beginning with the third, form short spines. The penis curves more than 90º just beyond the base (Fig. 6.788). Color: brown with darker markings on the scutellum and hemielytra. ....................................................................Pelocoris subflavus Montandon, 1898 (Argentina, Minas Gerais). - The length is shorter than about 10 mm. ...........................................................6

Fig. 6.788 Pelocoris subflavus: habitus in dorsal view (left), fore-leg (upper left center), two views of a male paramere (right center), and ventral view of the female genital segments (right). Based on Lopez Ruf (1994b).

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6. The posterolateral angles of the second laterotergites are acutely spinose. The aedeagus is broad with indistinct chitinous bars laterally. The subgenital plate of the female is narrowed at the apex and not sharply incised (Fig. 6.789). Length: 8.6 to 10.0 mm. Color: brown with the head and pronotum yellowish brown with dark punctures. .....................................................Pelocoris bipunctulus (Herrich-Schäffer, 1853) (Colombia, French Guiana, Surinam, Argentina, Pará, Amazonas, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Rio Grande do Sul). Syn: Pelocoris impicticollis Stål, 1856; Pelocoris lautus Berg, 1879; Pelocoris horvathi Montandon, 1905. - The posterolateral angle of the second laterotergite is bluntly spinose. The aedeagus is slender with distinct chitinous bars laterally. The subgenital plate of the female is not distinctly narrowed at the apex or sharply incised (Fig. 6.790). Length: 7.9 to 8.7 mm. Color: yellowish brown with the head and pronotum shiny and not much lighter than the scutellum. ........................................................................Pelocoris politus Montandon, 1895 (Paraguay, Argentina, Pará, Amazonas, Mato Grosso).

Fig. 6.789 Pelocoris bipunctulus (left to right): right lateral margins of the second through fourth abdominal segments, penis and parameres, and the apex of a female abdomen in dorsal and ventral view. Based on Nieser (1975), who referred to the species as Pelocoris impicticollis.

Fig. 6.790 Pelocoris politus: right lateral margins of the second through fourth abdominal segments (upper left), penis and parameres of two varying specimens (left center to center), posterior margin of the phallotheca at the base of the penis (lower left), and the apex of a female abdomen in dorsal (right center) and ventral view (right). Based on Nieser (1975).

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Potamocoridae Key to the genera in South America Information for the key was provided by De Carlo (1968e), Nieser (1975), and Longo et al. (2005). 1. The hemielytra are entirely chitinized and resemble the elytra of beetles (Fig. 6.791). .............................................................Coleopterocoris Hungerford, 1942.. p.575 - The membrane of the hemielytra is less strongly chitinized than the rest of the structure (Fig. 6.792). ...................................................................Potamocoris Hungerford, 1941..p. 577

Fig. 6.791 Coleopterocoris kleerekoperi: habitus in dorsal view (upper left); head in ventral view (lower left); fore, middle, and hind leg (lower center, left to right); male genital capsule (upper center); apex of the male (upper right) and female abdomen (lower right) in ventral view. Based on Hungerford (1942).

Fig. 6.792 Potamocoris parvus (above, left to right): habitus in dorsal view, apex of the male abdomen in ventral view, male genital capsule, and (below, left to right): head in ventral view, middle and hind leg, and the apical segments of the female abdomen in ventral view. Based on Hungerford (1941).

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Key to the species of Coleopterocoris known from South America Information for the key was provided by Longo et al. (2005). 1. The surfaces of the head and pronotum are covered with many thin setae. The eyes project laterally. The fore-femur is cylindrical and not dilated (Fig. 6.793). .........................................................Coleopterocoris hungerfordi De Carlo, 1968 (São Paulo, Rio de Janeiro, Paraná). - The surfaces of the head and pronotum are smooth and have few thin setae. The eyes do not project laterally (Fig. 6.794). The fore-femur is flattened and dilated. ..................................................................................................................2

Fig. 6.793 Coleopterocoris hungerfordi: habitus without appendages in dorsal view (left) and the apex of a male abdomen in ventral view (right). Based on Longo et al. (2005).

Fig. 6.794 Habitus of Coleopterocoris plaumanni in dorsal view. Based on De Carlo (1968e).

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2. (1) The hemielytra have a symmetrical pattern of black lateral stripes and spots (Fig. 6.794). ...........................................................Coleopterocoris plaumanni De Carlo, 1968 (São Paulo). - The hemielytra lack black spots and stripes (Fig. 6.795). ................................3 3. (2) There are odd pinkish spots between the compound eyes and on the pronotum. The interocular width of the female is about 2.2 times the width of a compound eye. Length: 2.9 to 3.0 mm. ...............................................................Coleopterocoris usingeri De Carlo, 1968 (São Paulo). - There are no odd-looking spots between the compound eyes or on the pronotum. The interocular width of the female is more than 2.7 times the width of an eye (Fig. 6.795). The length is less than 2.9 mm. ......................................4

Fig. 6.795 Coleopterocoris nelsoni (left to right): habitus in dorsal view, apical segments of a male and a female abdomen in ventral view, and the pygofer in posterior view. Based on Longo et al. (2005). 4. (3) The interocular width is 2.7 to 2.8 times the width of a compound eye. Each paramere has a constriction at the apex (Fig. 6.791). The apex of the basal plate is subacutely pointed. Length: 2.5 to 3.0 mm. ....................................................Coleopterocoris kleerekoperi Hungerford, 1942 (Rio Grande do Sul, Mato Grosso) - The interocular width is 3.7 to 4.2 times the width of a compound eye. Each paramere maintains the same width to the apex (Fig. 6.795). The apex of the basal plate is rounded. .............................Coleopterocoris nelsoni Longo, Ribeiro, and Nessimian, 2005 (São Paulo).

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Key to the species of Potamocoris known from South America Information for the key was provided by Hungerford (1941), Polhemus and Polhemus (1983b), and van Doesburg (1984). Until descriptions of the males of two of the three species are available, only female specimens can be distinguished. 1. The interocular space is greater than 1.5 times the width of a compound eye. The posterior margin of the female subgenital plate is sinuate and has a small, narrowly incised notch at the apex. ...................................................................Potamocoris robustus La Rivers, 1969 (Peru, Colombia). - The interocular space is about 1.3 times the width of a compound eye, or less (Fig. 6.792). ..........................................................................................................2 2. (1) The frontal margin of the head is evenly convex and moderately produced anteriad between the compound eyes. The maximum width of the pronotum is about 3½ times its length along the midline. The corium is produced posteriad along its lateral margin to border on the posterolateral margin of the membrane (Fig. 6.344). Length of macropterous specimens: c. 2.75 mm. ...............................................................Potamocoris nieseri van Doesburg, 1984 (Surinam). - The frontal margin of the head is strongly produced anteriad between the compound eyes. The maximum width of the pronotum exceeds 3½ times its length along the midline. The corium is not produced posteriad along its lateral margin and does not border on the posterolateral margin of the membrane (Fig. 6.792). Length: c. 2.8 mm. The color is testaceous with paler markings on the hemielytra; the antennae and legs appear translucent with a greenish tinge. ...................................................................Potamocoris parvus Hungerford, 1941 (Argentina, Paraguay, Peru, southeastern Brazil).

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6.7.3 Infraorder Leptopodomorpha Key to the families in South America Information for the key was provided by Polhemus (1985a). 1. The labium does not reach beyond the apex of the fore-coxa (Fig. 6.9). ...........................................................................................................Leptopodidae Only one species in this family is known to inhabit South America: Saldolepta kistnerorum Schuh and Polhemus, 1980. It has been found in a termite nest in tropical lowland Ecuador and in a seaport during the inspection of plant material from Colombia (Schuh and Slater, 1995) - The labium is long and tapering; it extends posteriad at least as far as the hind coxa (Fig. 6.796). .................................................................................................2 2. (1) The compound eyes cover only a small part of the lateral surface of the head. Ocelli are absent. The scutellum appears as a small wedge. The wings are reduced to small pads apparently without veins, which cover only the anteriormost part of the abdomen. The only known species reaches a length slightly greater than 2 mm. ............................................................................................................Aepophilidae This only known species in this family, Aepophilus bonnairei Signoret, 1879, is known only from the marine intertidal zone in Europe and possibly North Africa. - The compound eyes cover most of the surface of the head. The forewings are macropterous, submacropterous, or beetle-like and cover most of the abdomen (Fig. 6.796). ..........................................................................................................3

Fig. 6.796 Saldula humilis: habitus in dorsal (left) and in ventral view without the legs (lower right) and a hemielytron (upper right). Based on Alvarez and Roldán (1983).

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3. (2) The body is beetle-like and 2 mm or shorter. The compound eyes cover about 1/3 of the lateral surface of the pronotum and extend beyond its collar. ................................................................................................................Omaniidae This family encompasses a few species of the marine intertidal zone in the Indo-Pacific Region. - The length is more than 2.2 mm. The compound eyes reach only as far as the pronotal collar or slightly beyond. Ocelli are present. The hemielytra cover the abdomen, and the scutellum has the shape of an equilateral triangle (Fig. 6.796). ...................................................................................................................Saldidae

Saldidae Key to the genera known from South America Information for the key was provided by Polhemus (1985a, b) and Froeschner (1985). 1. All known South American genera have a long embolar fracture that reaches at least as far as the hind end of the claval suture. The subgenital plate of the female is truncate and usually square, or, if it is produced along the posterior midline, there are five cells in the hemielytral membrane. (Fig. 6.797). ......................................................................Subfamily Chiloxanthinae...............2 - All known South American species have a short embolar fracture that does not reach anteriad more than half way between the fracture on the costal margin and hind end of the claval suture. The subgenital plate of the female is produced posteriad along the midline, or, if not, then the membrane usually consists of only four cells (Fig. 6.798). .................................................................................Subfamily Saldinae...............3

Fig. 6.797 Pentacora sphacelata: hemielytra of two specimens (upper left and center), lateral view of the abdomen (lower left), two aspects of the left male paramere (right, above and below), parandria (lower right center), aedeagus and spermatheca (left of center, left and right, respectively), styloid (center), second gonapophysis (lower left center), and gynatrium (lower right center). Based on Polhemus (1985a).

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2. (1) The hemielytral membrane has five elongate cells of nearly equal size (Fig. 6.797). ...........................................................................................Pentacora Reuter, 1912 No species in this genus have yet been reported from continental South America, but one species, Pentacora sphacelata (Uhler, 1877), inhabits the Galapagos Islands as well as Spain and North and Central America. Its length should fall between about 3.5 to 5.0 mm. Its synonyms include Salda sphacelata Uhler, 1877; Salda rubromaculata Heidemann, 1901; Acanthia rubromaculata Kerkaldy and Torre-Bueno, 1909; Acanthia sphacelata Kerkaldy and Torre-Bueno, 1909; Pentacora rubromaculata Barber, 1925; Pentacora iberica Wagner, 1953. - The hemielytral membrane has only four cells (Fig. 6.799). A postclypeus is lacking. Length of the only species: 2.4 to 2.8 mm. .................................................................Paralosalda Polhemus and Evans, 1969 The only known species in this genus is Paralosalda innova Polhemus and Evans, 1969, which inhabits the rocky intertidal zone along the Pacific coasts of southern Central America and Colombia.

Fig. 6.798 Saldula coxalis: patterns on the hemielytra of two specimens (upper left), ventral view of the fore-femur (lower left), pronotum in dorsal view (center), two aspects of the left paramere of a male (right center), and the parandria (right). Based on Polhemus (1985a). 3. (1) There is no transverse swelling on the head. The length of the only known species is approximately 3.0 mm, and its width is about 1.2 mm. The head is black but covered with short, pale setae. The abdomen is shiny black and distinctly pitted. The metasternum is reddish (Fig. 6.800). Macropterous specimens have never been found. ....................................................................Oiosalda Drake and Hoberlandt, 1952 The only species in this genus, Oiosalda caboti Drake and Hoberlandt, 1952, is known only from Colombia. - The head has a transverse swelling (Fig. 6.801). ..............................................4

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Fig. 6.799 Paralosalda innova: habitus (middle left), spermatheca (upper left), two views of the paramere (lower left and center), parandria (right center), penis in dorsal (upper right) and lateral view showing the filum gonopori (upper middle right), subgenital plate of a female (lower right), and ovipositor (lower right center). Based on Polhemus and Evans (1969).

Fig. 6.800 Habitus of Oiosalda caboti. Based on Drake and Hoberlandt (1952).

Fig. 6.801 Pseudosaldula vulgaris: habitus of a male (upper left) and a female (lower left), male parandria (upper center), head in anterior view (lower center), and parameres (right). Based on Schuh and Polhemus (2009).

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4. (3) The membrane of each hemielytron has five cells (Fig. 6.801). ......................................................................Pseudosaldula Cobben, 1961..p. 583 - The membrane of each hemielytron has four cells (Fig. 6.802). ......................5

Fig. 6.802 Saldula dentulata: a hemielytron with a common color pattern (upper left) and one from a male with an unusual division of one of the cells on the membrane (upper left center), various color patterns on the anterior parts of the hemielytra of six specimens (middle left), two views of the left paramere of a male (upper right center), basal part of the penis filament (lower right center), sclerotized structures in the penis in ventral (lower left) and lateral view (lower left center), parandria (upper right), and the subgenital plate (lower right). Based on Cobben (1960b). 5. (4) The hypocostal ridge is simple; there is no secondary hypocostal ridge. The first and second antennal segments of the male are round in cross section and covered with pubescence of uniformly long setae. The innermost cell of the hemielytral membrane is not produced anteriad more than 1/3 of its length beyond the base of the second cell (Fig. 6.803). ..........................................................................Rupisalda Polhemus, 1985..p. 592 - The hypocostal ridge has a relatively complex structure, and a secondary hypocostal ridge is present. The third and fourth segments of the antenna are no thicker than the distal end of the second. The innermost cell of the hemielytral membrane usually reaches almost to the adjacent cell’s apex (Fig. 6.802). The length of the male is less than 5.5 mm; that of the female is less than 6.0 mm. .............................................................................Saldula van Duzee, 1914..p. 593

583

Fig. 6.803 Rupisalda ventralis: habitus in dorsal view (left) and a hemielytron (right). Based on Champion (1898). Key to the species of Pseudosaldula in South America Information for the key was provided by Signoret (1863), Drake (1949c, 1955), Polhemus (1976b), and Schuh and Polhemus (2009). 1. There is a black marking or lateroventral stripe, and sometimes also a black dorsomedial mark, on the first segment of the antenna (Fig. 6.804). ...................2 - The color of the first antenna segment is uniform, i.e., markings are absent (Fig. 6.805). ..........................................................................................................4

Fig. 6.804 Habitus of Pseudosaldula chilensis. Based on Schuh and Slater (1995).

584

Fig. 6.805 Pseudosaldula andensis male: habitus (left), parandria (right), and parameres (lower right). Based on Schuh and Polhemus (2009). 2. (1) All tibiae lack distinct stripes on the dorsal surface, which may, however, be vaguely darkened. The apices of both male parameres are narrow, subparallel, and rounded (Fig. 6.806). Length of male and female: c. 3.20 to 3.35 mm. ...................................................Pseudosaldula penai Schuh and Polhemus, 2009 (Bolivia, Chile, Argentina). - There is a clearly defined dark stripe on the dorsal surface of the fore and middle tibiae, and sometimes on the hind tibiae as well. The apical portions of the male parameres are narrow, and at least one of them is spindle-shaped or tapers to an acute point at the apex (Fig. 6.807). .................................................3

Fig. 6.806 Pseudosaldula penai male (left to right): habitus, parandria, and parameres. Based on Schuh and Polhemus (2009).

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Fig. 6.807 Pseudosaldula salina: habitus of a male (upper left) and a female (lower left), head in anterior view (lower center), male parandria (upper center), and parameres (right). Based on Schuh and Polhemus (2009).

Fig. 6.808 Pseudosaldula saxicola male: habitus (upper left), head in anterior view (lower left), parandria (center), and parameres (right). Based on Schuh and Polhemus (2009). 3. (2) The coloration is mainly black, the color covering the clavus, corium, and membrane of the hemielytra, but there is a coating of golden setae on the entire dorsum. The golden color shines when the light source is located anterolateral from the insect. There are pruinose areas on the clavus and corium. The legs are almost entirely black with only a pale apical ring on each tibia. Black longitudinal stripes are present on the ventrolateral and dorsomedial surfaces of the first antennal segment (Fig. 6.804). Length of male: c. 3.4 to 3.6 mm; female: c. 4.0 to 4.2 mm. ............................................................Pseudosaldula chilensis (Blanchard, 1852) (Chile, Argentina). Syn: Acanthia chilensis Blanchard, 1852; Acanthia rogeri Kirkaldy, 1899; Acanthia araucanica Kirkaldy, 1899; Pentacora araucanica Drake and Carvalho, 1948; Saldula araucanica Drake and Carvalho, 1948;

586

Saldula rogeri Drake and Hoberlandt, 1951; Pentacora angusta Drake and Carvalho, 1948; Pentacora regilla Drake, 1955; Pseudosaldula rogeri Cobben, 1961; Oreokora regilla Drake, 1962; Oreokora rogeri Drake, 1962; Oreokora angusta Drake, 1962; Oreokora chilensis Drake, 1962. - The setae coating the dorsum are black. The corium has pale markings but never pruinose areas. There is a black longitudinal stripe on the ventrolateral surface of the first antennal segment but no dorsomedial stripe (Fig. 6.807). Length of male: 3.4 to 3.6 mm; female: 3.3 to 3.5 mm. Only macropterous individuals have been found. ..................................................Pseudosaldula salina Schuh and Polhemus, 2009 (Argentina). 4. (1) The surface of the hemielytra is matt except along the overlapping borders of the claval commissure and sometimes on limited areas along the edge of the corium (Fig. 6.808). ..............................................................................................5 - There are shiny surfaces on the embolar area, other parts of the corium, veins, and sometimes areas of the membrane, as well as along the claval commissure (Fig. 6.805). ..........................................................................................................8 5. (4) The dorsal setae are long, simple, erect, dark, and dull, making the insect appear shaggy in lateral view (Fig. 6.809). The hemielytral setae are about twice as long as the diameter of a hind tibia at midlength. Length of male: c. 3.1 to 3.3 mm; female: c. 3.4 to 3.6 mm. ..................................................Pseudosaldula pilosa Schuh and Polhemus, 2009 (Argentina, Chile). - The dorsal setae are short and recumbent, appearing as a short pubescence and never shaggy in lateral view (Fig. 6.808). Some of the setae are shiny and golden in color. The hemielytral setae are only as long as or shorter than the diameter of a hind tibia at midlength. ...................................................................6

Fig. 6.809 Pseudosaldula pilosa: habitus of a male (upper left) and a female (lower left), male parandria (upper center), and parameres (right). Based on Schuh and Polhemus (2009).

587

6. (5) The pronotum is campanulate with slightly to strongly concave margins (Fig. 6.808). The second antenna segment is usually longer than 1.48 times the length of the third. There is an irregular ivory-colored marking on the subapical surface of the exocorium. Length of male: 3.7 to 3.9 mm; female: c. 4.1 to 4.3 mm. ...............................................Pseudosaldula saxicola Schuh and Polhemus, 2009 (Argentina, Peru). - The pronotum is flattened rather than campanulate and has slightly to strongly convex margins (Fig. 6.810). The second antenna segment is usually shorter than 1.38 times the length of the third. There is at most a pale marking on the the exocorium, just proximal to its apex. .............................................................7

Figure 6.810 Pseudosaldula bergi male: habitus (upper left), penis in ventral (upper right) and lateral view (lower right), parandria (lower center), and a paramere (lower left). Based on China (1962), who called his specimen Pseudosaldula cobbeni. 7. (6) Shiny, golden, recumbent setae form a uniform covering on the corium. There are several purplish, pruinose areas on the corium, clavus, and sometimes the membrane. These areas are more conspicuous on dark individuals. The membranes of brachypterous individuals have numerous long setae on the veins, at least on the lateral areas (Fig. 6.801). These setae are less numerous on macropterous specimens. Length of male: c. 3.2 to 3.4 mm; female: c. 3.4 to 3.6 mm. ...............................................Pseudosaldula vulgaris Schuh and Polhemus, 2009 (Bolivia, Peru).

588

- Most setae on the dorsum are black, with only isolated shiny, golden setae on the corium. There are no more than a few, small, inconspicuous purplish areas on the corium, clavus, and membrane (Fig. 6.810). There are only scattered short to medium-length setae on the veins of the membrane. Length: 3.6 to 4.3 mm. ....................................................................Pseudosaldula bergi (Haglund, 1899). (Argentina, Chile). Syn: Salda bergi Haglund, 1899; Saldula sola Drake and Carvalho, 1948; Saldula paralia Torres, 1954; Pseudosaldula cobbeni China, 1962; Oreokora bergi Drake, 1962; Oreokora sola Drake, 1962; Pseudosaldula paralia Polhemus (1976b). 8. (4) The anterior 80% of the exocorium and radial vein are very shiny, while the rest of the corium is dull. Both shiny golden recumbent and erect black setae cover the dorsum, giving the insect a shaggy appearance in lateral view (Fig. 6.811). Both kinds of setae are widely distributed on the surface of the hemielytra. Length of the male: c. 2.65 to 2.8 mm; female: c. 2.75 to 2.9 mm. ........................................Pseudosaldula huamachuco Schuh and Polhemus, 2009 (Peru). - The exocorium is shiny at least on the posterior half; the extent of the shiny area is variable (Fig. 6.805). The average length of males exceeds 2.9 mm, while the average length of females exceeds 3.1 mm. .........................................9

Fig. 6.811 Pseudosaldula huamachuco: habitus of a male (upper left) and a female (lower left), male parandria (center), and parameres (right). Based on Schuh and Polhemus (2009). 9. (8) The smooth and shiny area on the dorsum includes most of the exocorium and at least part of the posterior half of the endocorium (Fig. 6.812). ...............10 - Only the embolar area of the exocorium is smooth and shiny, but if it is shiny only on the posterior half of the embolar area, then the radial vein is also smooth and shiny (Fig. 6.813). .......................................................................................11

589

Fig. 6.812 Pseudosaldula bruesi: habitus of a male (upper left) and a female (lower left), male parandria (upper center), and parameres (right). Based on Schuh and Polhemus (2009). 10. (9) The smooth and shiny area on the dorsum of brachypterous specimens covers the whole exocorium and much of the posterior part of the endocorium, as well as the entire membrane (Fig. 6.805). Only the basal third of the membrane of macropterous specimens is shiny. Length of male: c. 3.50 to 3.65 mm; female: c. 4.15 to 4.30 mm. .................................................................Pseudosaldula andensis (Distant, 1891) (Peru, Ecuador). Syn: Acanthia andensis Distant, 1891; Salda andensis Lethierry and Severin, 1896; Saldula andensis Drake and Hoberlandt, 1951; Pentacora andensis Drake, 1955; Oreokora andensis Drake, 1962. - The smooth, shiny pattern on the exocorium is interrupted by a dull area along the radial vein. Only the basal area and veins on the membranes of brachypterous specimens are smooth and shiny, while the basal third of the membrane of macropterous specimens is shiny (Fig. 6.812). Length of male: c. 3.65 to 3.80 mm; female: c. 3.90 to 4.05 mm. .......................................................................Pseudosaldula bruesi (Drake, 1949) (Peru). Syn: Pentacora bruesi Drake, 1949; Pentacora amazona Drake, 1955; Pentacora bucayana Drake, 1955; Pentacora pillaona Drake, 1955; Oreokora amazona Drake, 1962; Oreokora bucayana Drake, 1962; Oreokora pillaona Drake, 1962; Oreokora pilloana Schuh et al., 1987, with spelling mistake. 11. (9) Only black setae are present on the dorsum; shiny golden setae and pruinose areas are absent. The pronotum is campanulate, and its lateral margins are at least slightly concave (Fig. 6.813). ...........................................................12 - Both black and shiny golden setae are present on the dorsum, which also displays pruinose areas. The pronotum is not obviously campanulate, and its lateral margins may be slightly or strongly convex (Fig. 6.814). ......................13

590

Fig. 6.813 Pseudosaldula yungas: habitus or a male (upper left) and a female (lower left), parandria (upper right), and parameres (lower right). Based on Schuh and Polhemus (2009).

Fig. 6.814 Pseudosaldula aurea: habitus of a male (left), parandria (center), and parameres (right). Based on Schuh and Polhemus (2009).

Fig. 6.815 Pseudosaldula antioquia: habitus of a male (upper left) and a female (lower left), parandria (upper center), and parameres (right). Based on Schuh and Polhemus (2009).

591

Fig. 6.816 Pseudosaldula perula: habitus of a male (upper left) and a female (lower left), parandria (upper center), and parameres (right). Based on Schuh and Polhemus (2009). 12. (11) The setae on the dorsum are long and erect, which is particularly conspicuous in lateral view (Fig. 6.813). The average length of males exceeds 4.3 mm. .................................................Pseudosaldula yungas Schuh and Polhemus, 2009 (Peru, Bolivia). - The setae on the dorsum are erect but only of moderate length, as apparent in lateral view (Fig. 6.815). The mean length of the male is about 2.95 mm. .............................................Pseudosaldula antioquia Schuh and Polhemus, 2009 (Colombia). 13. (11) Only the posterior half of the embolar area and the radial vein are smooth and shiny (Fig. 6.814). The dorsal setae are moderately long and partly erect, giving the insect a shaggy appearance in lateral view. Length of male: c. 3.30 to 3.45 mm; female: 4.15 to 4.30 mm. ...................................................Pseudosaldula aurea Schuh and Polhemus, 2009 (Argentina). - The shiny surface of the embolar area extends for its entire length, but its radial vein is matt. The dorsal setae are of moderate length and suberect, but the insect does not appear very shaggy in lateral view (Fig. 6.816). Length of male: c. 2.80 to 3.00 mm; female: c. 3.55 to 3.75 mm. .......................................................................Pseudosaldula perula (Drake, 1955) (Argentina, Peru). Syn: Pentacora perula Drake, 1955; Oreokora perula Drake, 1962.

592

Key to the species of Rupisalda in South America Information for the key was provided by Drake and Hottes (1951c), Drake (1955), and Polhemus (1985a, b). 1. The pubescence is short, dense, and yellowish brown. It appears to have been clipped. The first segment of the antenna is stout and yellowish, while the second is fuscous but pale at the base, and the third and fourth are dark. The lateral margins of the pronotum are nearly straight with pale marginal stripes visible from both above and below. The male parameres are distinctly sigmoid (Fig. 6.817). The coxae are fuscous brown. The trochanters and bases of the femora are whitish. The apical parts of the femora are fuscous brown, and the tibiae and tarsi are brown. Length: 3.6 to 4.2 mm; width: 1.4 to 1.6 mm. ..........................................................Rupisalda verdica (Drake and Hottes, 1951) (Santa Catarina). Syn: Saldula verdica Drake and Hottes, 1951. - The pubescence is golden or grayish and not very dense. The first antennal segment is either not stouter than the other segments, or it is colored in a shade appearing darker than yellowish. The parameres of known males are not strongly curved (Fig. 6.803). ................................................................................2

Fig. 6.817 The right paramere of a male Rupisalda verdica. Based on Drake and Hottes (1951c). 2. (1) The antennae are brownish fuscous with a short pubescence. The legs are testaceous with a tinge of brown, but the coxae are ferrugineous with whitish bases, and the trochanters and bases of the femora are whitish testaceous. The ventral parts of the acetabula are also whitish. The ventral surface of the forefemur is brown. The ratios of antenna segments I to IV are 22:40:36:40, respectively. The lateral margins of the pronotum are somewhat convex, explanate, and covered with a short gray pubescence. The anterior and posterior ends of the lateral margins are dark, but the explanate portion is whitish yellow in both dorsal and ventral view. Length: 3.35 to 3.80 mm. Maximum width: c. 1.5 mm. ................................................................................Rupisalda zelena (Drake, 1955) (Peru). Syn: Saldula zelena Drake, 1955.

593

- The legs are light brown to pale and lack whitish or whitish testaceous markings on the coxae, trochanters, bases of the femora, or acetabula. ...............3 3. (2) The first segment of the antenna is brownish, and the rest are blackish fuscous. The legs are light brown with the trochanters, bases of the femora, and apical tarsal segments fuscous. The coxae are reddish fuscous. Length: c. 3.5 mm. ............................................................................Rupisalda icolana (Drake, 1955) (Santa Catarina). Syn: Saldula icolana Drake, 1955. - The apical segment of the antenna is lighter in color than blackish fuscous; it has a broad whitish or yellowish ring. The legs, especially the coxae, are relatively pale (Fig. 6.803). ..............................................................................Rupisalda ventralis (Stål, 1860) (Rio de Janeiro, Mato Grosso?). Syn: Salda ventralis Stål, 1860; Acanthia ventralis Stål, 1873; Saldula ventralis Drake and Carvalho, 1948; Saldula brasiliensis Drake and Carvalho, 1948. Key to the species of Saldula in South America Information for the key was provided by Drake and Carvalho (1948), Drake (1955), Polhemus (1968b), and Cobben (1986). The genus is badly in need of a revision encompassing the South American species. The status of many nominal species is doubtful because the only information about them is found in cursory descriptions based on few specimens. 1. The pronotum is pale along its lateral margins (Fig. 6.798). ...........................2 - The color of the pronotum is uniformly dark or black with fuscous brown lateral margins (Fig. 6.818). .................................................................................3 2. (1) The legs are yellowish or leucine with brown markings. The pale stripes along the lateral margins of the pronotum are narrower than the second antennal segment. The lateral margins are not explanate (Fig. 6.798). Length of male: 3.2 to 3.6; female: 3.4 to 4.1 mm. .....................................................................................Saldula coxalis (Stål, 1873) (North and Central America, West Indies, Chile, Argentina, Brazil). Syn: Acanthia coxalis Stål, 1873; Salda argentina Berg, 1979; Acanthia argentina Berg, 1883; Salda coxalis Uhler, 1877; Saldula argentina Drake, 1950 (nec Berg, 1879); Acanthia xanthochila limbosa Barber, 1914. - The legs are mainly testaceous and sometimes marked with brown. The pale markings on the lateral edges of the pronotum do not extend all the way from the anterior to the posterior end. The pronotum narrows considerably toward its anterior end; it is shiny, black, and finely rugose. The lateral margins are deep orange on both the dorsal and ventral sides. The orange marking extends almost to the base and as far anterior as the middle of the callus, occupying about 2/3 of

594

the length of the pronotum. Length of female: c. 4.4 mm. The male has not been described. ...........................................................Saldula doeringi Drake and Carvalho, 1948 (Argentina). 3. (1) The head, pronotum, and scutellum are black and rugulose, sometimes with a slight shine (Fig. 6.802). The length is shorter than c. 3.2 mm. ...............4 - The head, pronotum, and scutellum are smooth and shiny, dull, or covered by pubescence but not rugulose (Fig. 6.818). If calluses make the surface less smooth, then the length is greater than about 4 mm. ............................................5

Fig. 6.818 Habitus of a female Saldula chartoscirtoides in dorsal view. Based on Cobben (1986). 4. (3) The outer margin of the male paramere is curved or hooked without a sign of swelling. A prominent tubercle is evident along the inner surface proximal to the sharp curve. The lateral margins of the pronotum are concave or straight. The head has a dorsomedial sulcus The dorsal pubescence is sparce, recumbent, and golden yellow (Fig. 6.802). Length: 2.7 to 3.1 mm. Maximum width: 1.2 to 1.3 mm. ........................................................................Saldula dentulata (Hodgden, 1949) (Central America, West Indies, Ecuador, Peru, Brazil). Syn: Salda dentulata Hodgden, 1949; Salda sectilis Hodgden, 1949; Saldula sectilis (Hodgden, 1949); Saldula denticulata Drake and Hottes, 1951 (misspelled). - There is a prominent swelling along the outer curvature of the male paramere and only a slight swelling along the inner curvature proximal to the sharp curve. There are two to four light spots on the distal part of the inner corium (Fig. 6.796). Length of male: 2.5 to 2.8 mm; female: 2.7 to 3.1 mm. Maximum width of male: 1.1 to 1.3 mm; female: 1.3 to 1.5 mm. ....................................................................................Saldula humilis (Say, 1832) (North America, West Indies, Surinam, Santa Catarina). Syn: Acanthia humilis Say, 1832; Micranthia humilis (Say, 1832) auctt.

595

5. (3) The lateral margin of the pronotum is concave or straight, and its anterolateral angles are barely wider than the collar (Fig. 6.818). .......................6 - The lateral margins of the pronotum are at least slightly convex, and the pronotum is only moderately narrowed anteriorly (Fig. 6.819). ..........................9 6. (5) The dorsal surfaces of the head and thorax are dull black and covered with a fine silvery or golden pubescence and a few longer black setae on the thorax. The light spots on the head between the ocelli and compound eyes are larger than the ocelli. The ratios of the lengths of antenna segments I to IV are 10:21:16:16. Length: c. 3.2 to 3.8 mm. Maximum width: 1.5 to 1.8 mm. ..........................................................................Saldula peruviana (Cobben, 1986) (Peru). Syn: Micranthia peruviana Cobben, 1986. - The head, pronotum, and scutellum are mainly black and shiny, covered with a sparce, grayish brown or pale pubescence. On the head, there is a yellowish spot between each ocellus and the nearest compound eye (Fig. 6.818). ..............7 7. (6) The hemielytra are dull velvet black with moderately long, golden, semirecumbent pilosity intermixed with some shorter hairs. They are marked with flavous stripes along the outer margins and whitish spots, including an oblong spot in the clavus, four rounded spots and two smaller subapical spots in the inner corium, three spots in each of the three inner cells of the membrane, and two spots in the outermost fourth cell. The ratios of the lengths of antenna segments I through IV are 13:19:17:19, respectively. The fourth antenna segment has a broad white band. The male right paramere is hook-like and somewhat broadened at the apex (Fig. 6.820). Length of male: c. 3.15 mm; female: c. 3.8 mm. Width of male: c. 1.35 mm; female: c. 1.7 mm. The color of the head and thorax is mainly shiny black, less so on the head, which has yellow markings. The pubescence on the head is brownish and gray, and that on the dorsal surface of the thorax is brownish. ..................................................................Saldula scitula Drake and Hottes, 1950 (Santa Catarina). - The pubescence on the hemielytra is long and dark brownish or short and silvery. ..................................................................................................................8 8. (7) The pubescence on the head, pronotum, and scutellum is grayish brown with erect dark setae. The head has a dorsomedial sulcus. The light spots on the head between the ocelli and compound eyes are about the same size as an ocellus. The dorsal surface of the hemielytra is clothed in long, erect, dark brownish setae. Length: c. 3.0 mm. Maximum width: c. 1.25 mm. ..............................................................................Saldula hodgdeni (Drake, 1955) (Bolivia). Syn: Micranthia hodgdeni Drake, 1955. - The pubescence on the head, pronotum, and scutellum is short and pale, admixed with a few longer black setae (Fig. 6.818). The ratios of the lengths of antenna segments I to IV are 10:15:15:17. The dorsal surface of the hemielytra is clothed in a short, silvery pubescence. Length or female: c. 3.7 mm. Maximum width: c. 2.0 mm. Only the female has been described. ..................................................................Saldula chartoscirtoides Cobben, 1986 (Santa Catarina).

596

Fig. 6.819 Saldula galapagosana: habitus in dorsal view (below) and (above, left to right): a hemielytron, penis showing the sclerites, coupling plate, parandria, and two views of the left paramere. Based on Polhemus (1968b) and Froeschner (1985).

Fig. 6.820 The right paramere of a male Saldula scitula. Based on Drake and Hottes (1950). 9. (5) The shape of the female is broadly ovate. The pronotum is mainly black, smooth, and shiny with broadly explanate, fuscous brown lateral margins. The hemielytra are fuscous brown. The ventral surface is brownish black, except for the apex of the posteriormost segment, which is lighter. The legs are brownish. Length: c. 4.35 mm. Maximum width: c. 2.35 mm. The male has not been described. ...............................................................Saldula lynchi Drake and Carvalho, 1948 (Argentina). - The shape is not broadly ovate. The hemielytra have a distinct pattern (Fig. 6.819). .................................................................................................................10

597

10. (9) The length is about 4.35 mm. The shape is oblong ovate. The width of the pronotum is twice its median length because its posterior margin is deeply excavated. The pronotum has expanded and rounded sides, is deeply foveate, and has a moderately raised callus. The head, pronotum, and scutellum are mainly shiny black. The posterior part of the pronotum is finely rugulose. The hemielytra are conspicuously marked. ...........................................................Saldula differata Drake and Carvalho, 1948 (Argentina). - The length is usually no more than 4 mm. ......................................................11 11. (10) The hemielytra lack pruinose areas. The fore-tibia has an uninterrupted dorsal stripe usually extending from the base and a point near the apex (Fig. 6.23). Length: c. 3 to 4 mm. Color: mainly black with testaceous markings and leucine areas. ...........................................................................Saldula pallipes (Fabricius, 1794) (Holarctic Region, Central America, South America). Syn: Cimex pellucidus? Goeze, 1778; Acanthia pallipes Fabricius, 1794; Salda pallipes Fabricius, 1803; Cimex marginalis Turton, 1802; Lygaeus pallipes Latreille, 1804; Lygaeus saltatorius Panzer, 1805; Acanthia striata Billberg, 1820; Acanthia interstitialis Say, 1825; Acanthia dimidiata Curtis, 1835; Salda ocellata A. Costa, 1843; Salda bicolor Fieber, 1859; Salda interstitialis Uhler in Packard, 1871; Acanthia laticollis Reuter, 1875; Salda reperta Uhler, 1877; Salda luctuosa Westhoff, 1880; Acanthia pallipes confluens Reuter, 1891; Salda tropicalis Champion, 1900; Salda saltatoria Champion, 1900 (nec Lennaeus, 1758); Acanthia tropicalis Kirkaldy and Torre Bueno, 1908; Salda inconstans Distant, 1909; Saldula interstitialis Van Duzee, 1914; Saldula laticollis Van Duzee, 1916; Saldula reperta Van Duzee, 1916; Saldula chipetae Drake and Hottes, 1955. - There are pruinose or pubescent areas on the hemielytra. ..............................12 12. (11) The hemielytra, as well as the pronotum and scutellum, are covered by a short, golden pubescence. The pronotum is black and almost completely smooth with a narrowly explanate margin. The pronotum is deeply foveate in the middle and has a wide callus and deeply emarginated posterior margin. The general shape is obovate. The clavus is dark fuscous with a whitish spot near the apex. The corium is dark brown with two or three dark spots. The embolium is nearly hyaline with two or three small brown spots. The femora are pale brown with yellow apices and sometimes small, brown spots. The trochanters are white or yellowish. Length: c. 3.10 mm. Maximum width: c. 2.25 mm. ......................................................Saldula penningtoni Drake and Carvalho, 1948 (Peru, Chile, Argentina, Uruguay, Paraguay, Minas Gerais, Santa Catarina). - There is a bluish white pruinose area on the inner part of the corium and light brown to greenish pruinose areas forming an irregular elongate pattern on the inner part of the corium and on a medial area of the outer part (Fig. 6.819). Length of male: 2.8 to 3.1 mm; female: 3.1 to 3.4 mm. ................................................................Saldula galapagosana (Polhemus, 1968) (Galapagos Islands).

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– Revista de la

Bachmann, A. O., 1965: Sobre la presencia de Ranatridae e Hydrometridae (Hemiptera) en el norte de la Patagonia. – Revista de la Sociedad Entomológica Argentina 27(14):131. Bachmann, A. O., 1966a: Catalogo sistematico y clave para determinacion de las subfamilies, generos y especies de las Gerridae de la Republica Argentina (Insecta, Hemiptera). - Physis 26(71):207-218. Bachmann, A. O., 1966b: Ovametra gualeguay, nueva especie de Gerridae de la Republica Argentina (Hemiptera). - Neotropica 12(39):87-90. Bachmann, A. O., 1966c: Presencia de Sigara (Tropocorixa) hungerfordi Jaczewski en la Republica Argentina (Hemiptera, Corixidae). – Revista de la Sociedad Entomológica Argentina 28(1-4):44. Bachmann, A. O., 1966d: Presencia de Notonecta (Paranecta) polystolisma Fieber en la Republica Argentina (Hemiptera, Notonectidae). – Revista de la Sociedad Entomológica Argentina 28(1-4):60. Bachmann, A. O., 1968 (1967): Las Pleidae de la Republica Argentina (Hemiptera). – Revista de la Sociedad Entomológica Argentina 30(1-4):121-129. Bachmann, A. O., 1971: Catálogo sistemático y clave para determinación de las Notonectidae de la República Argentina (Insecta, Hemiptera). – Physis 30(81):601-617. Bachmann, A. O., 1977a: (1975-76): Nota sobre las Hydrometridae de la Argentina (Heteroptera). – Revista de la Sociedad Entomológica Argentina 35(2-4):143-144. Bachmann, A. O., 1977b: Heteroptera. In: S. H. Hurlbert (Ed.): Aquatic Biota of Austral South America. – San Diege State University, San Diego. pp. 189-212. Bachmann, A. O., 1979a: Notas para una monografia de las Corixidae Argentinas (Insecta, Heteroptera). – Acta Zoologica Lilloana 35:305-350.

602 Bachmann, A. O., 1979b: Heteroptera acuáticos de las Galápagos. - Revista de la Sociedad Entomológica Argentina 37:131-135. Bachmann, A. O., 1979c: Las Pleidae de la Republica Argentina. – Revista de la Sociedad Entomológica Argentina 30:121-129. Bachmann, A. O., 1981: Insecta, Hemiptera, Corixidae. In. R. A. Ringuelet (Ed.), Fauna de Agua Dulce de la República Argentina 35(2):1-270. Bachmann, A. O., 1983 (1982): Notas sobre Corixidae (Heteroptera) (cuarta serie) tres notas distribucionales. – Revista de la Sociedad Entomológica Argentina 41(1-4):197200. Bachmann, A. O., 1985 (1987): Notas sobre Corixidae (Heteroptera) (Quinta Serie). – Revista de la Sociedad Entomológica Argentina 44(1):33-36. Bachmann, A. O., 1997: La familia Pleidae (Heteroptera) en Nicaragua. – Revista Nicaraguense de Entomologia 40:44. Bachmann, A. O., 1998: Heteroptera acuáticos. In: J. J. Morrone and S. Coscarón (Eds.), Biodiversidad de Artrópodos Argentinos. Una perspectiva biotaxonómica. – Ediciones Sur, La Plata. pp. 163-180. Bachmann, A. O., 1999: Catálogo de los tipos de Heteroptera (Insecta) conservados en el Museo Argentino de Ciencias Naturales. – Revista del Museo Argentina de Ciencias Naturales, Nova Serie 1(2):191-230. Bachmann, A. O., and J. L. Farina, 1989 (1986): Nota cientifica: sobre heterópteros acuáticos del área de Mar del Plata (Provincia de Buenos Aires, Argentina). – Revista de la Sociedad Entomológica Argentina 45(1-4):252, 256. Bachmann, A. O., and M. López Ruf, 1994: Los Pleoidea de la Argentina (Insecta, Hemiptera). In: Z. A. de Castellanos (Ed.), Fauna de agua dulce de la República Argentina 35(3):1-32. Bachmann, A. O. and S. A. Mazzucconi, 1995: Insecta Heteroptera (= Hemiptera s. str.). In. E. C. Lopretto and G. Tell (Eds.), Ecosistemas de Aguas Continentales. Editora Sur, La Plata. pp. 1291-1325. Bacon, J. A., 1948: Some new species of Rhagovelia (Hemiptera, Veliidae). – Journal of the Kansas Entomological Society 21(3):71-87. Bacon, J. A., 1956: A taxonomic study of the genus Rhagovelia (Hemiptera, Veliidae) of the Western Hemisphere. – University of Kansas Science Bulletin 38(10):695-913.

603 Berg, C., 1879: Hemiptera Argentina. – Anales de la Sociedad Científica Argentina 8:1316. Berg, C., 1881a: Sinonimia y descripcion de algunos hemípteros de Chile, del Brasil y de Bolivia. – Anales de la Sociedad Científica Argentina 8:19-33, 71-80. Berg, C., 1881b: Hemípteros de Chile, del Brasil y de Bolivia. – Anales de la Sociedad Científica Argentina 12:259-272. Berg, C., 1883a: Addenda et emendanda ad Hemiptera Argentina. – Anales de la Sociedad Científica Argentina 16:105-125. Berg, C., 1883b: Hemiptera Argentina - Addenda. – Anales de la Sociedad Científica Argentina 17:5-25. Berg, C., 1884: Addenda et emendanda ad Hemiptera Argentina. Sociedad Científica Argentina 17(2):97-118.

– Anales de la

Berg, C., 1898: Descriptiones hydrometridarum novarum Republicae Argentinae. – Communicaciones del Museo Nacional de Buenos Aires 1:3-6. Bergroth, E., 1899: A new genus of Corixidae. – Entomological Monthly Magazine 35:282. Boulard, M., and P. Jauffret, 1984: Description d’un Gelastocoris nouveau du Brésil et précisions sur les genitalia mâles et femelles de ce genre d’hemiptères (Gelastocoridae). – Nouvelle Revue d’Entomologie (N.S.) 1(1):7-18. Brailovsky, H., 1989: Un genero y dos especies de hemipteros (Lygaeidae, Bledionotinae, Pamphantini) del Brasil. – Anales del Instituto Biología - Universidad Nacional Autónoma de México, Seria Zoología 59(2):193-202. Breddin, G., 1898: Studia hemipterologica IV. - Jahresbericht und Abhandlungen des Naturwissenschaftlichen Verein in Magdeburg 1896-1898:149-163. Bressa, M. J., and A. G. Papeschi, 2007: New contributions to the study of Corixoidea: cytogenetic characterization of three species of Sigara from Argentina and the plausible mechanisms of karyotype evolution within Nepomorpha. – Revista de la Sociedad Entomológica Argentina 66(3-4):81-89. Brèthes, J., 1904: Insectos de Tucumán. - Anales de Museo Nacional de Buenos Aires 11(3) 4:329-347. Brooks, G. T., 1953: A new subgenus and species of Enithares. – Journal of the Kansas Entomological Society 26(2):74-75.

604 Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954: Classification of Insects. Keys to the living and extinct families of other terrestrial arthropods. – Bulletin of the Museum of Comparative Zoology at Harvard College 108:1-917. Burmeister, H. C. C., 1835: Handbuch der Entomologie, Bd. II. Ordnung Rhynchota. Abt. 1. Hemiptera. - Enslin, Berlin. iv + 400 pp. Buzzetti, F.M., 2006: The genus Potamobates Champion in Ecuador, with description of P. shuar n. sp. (Hemiptera: Gerridae). – Zootaxa 1306: 51-56. Caponigro, M. A., and C. H. Eriksen, 1976: Surface film locomotion by the water strider, Gerris remigis Say. – The American Midland Naturalist 95(2):268-278. Cárcamo, H. A., and J. R. Spence, 1994: Kin discrimination and cannibalism in water striders (Heteroptera: Gerridae): another look. – Oikos 70:412-416. Carpenter, G. H., 1898: Trochopus and Rhagovelia. Magazine, Series 2, 9:109-112.

– Entomological Monthly

Carvalho, J. C. M., 1984: On a new species of intertidal water strider from Brazil (Hemiptera, Gerromorpha, Mesoveliidae). - Amazoniana 8(4):519-523. Carvalho, J. C. M., and R. F. Hussey, 1954: On a collection of Miridae (Hemiptera) from Paraguay, with descriptions of three new species. – Occasional Papers of the Museum of Zoology, University of Michigan, No. 552:1-11. Castanhole, M. M. V., 2009: Spermatogenesis and nucleolar behavior in males of aquatic Heteroptera. – Genetics and Molecular Research 8(3):816. Castanhole, M. M. V., L. L. V. Pereira, H. V. Souza, H. E. M. C. Bicudo, L. L. A. Costa, and M. M. Itoyama, 2008: Heteropicnotic chromatin and nucleolar activity in meiosis and spermiogenesis of Limnogonus aduncus (Heteroptera, Gerridae): a stained nucleolar organizing region that can serve as a model for studying chromosome behavior. – Genetics and Molecular Research 7(4):1398-1407. Champion, G. C., 1897-1901: Biologia Centrali-Americana. Insecta. Rhynchota. Hemiptera-Heteroptera. Volume II. – R. H. Porter, London. 416 pp + 22 pl. Cheng, L., 1997: Disjunct distribution of Halobates in the Pacific Ocean. – Pacific Science 51(2):134-142. China, W. E., 1936: The first genus and species of Helotrephidae (Hemiptera) from the New World. – Annals and Magazine of Natural History, Zoology, Botany, and Geology, Series 10, 10(17):527-538.

605 China, W. E., 1940: New South American Helotrephidae (Hemiptera - Heteroptera). – Annals and Magazine of Natural History (11)5:106-126 + 1 Table. China, W. E., 1962a: South American Peloridiidae (Hemiptera-Homoptera: Coleorrhnycha). – Transactions of the Royal Entomological Society of London 114(5):131-161. China, W. E., 1962b: Hemiptera-Heteroptera collected by the Royal Society Expedition to south Chile 1958-1959. – Annals and Magazine of Natural History, Series 13, 5:705723. Cobben, R. H., 1960a: The Heteroptera of the Netherlands Antilles – I. Forward. Gerridae, Veliidae, Mesoveliidae (water striders). - Studies on the Fauna of Curaçao and other Caribbean Islands 11(50):1-34. Cobben, R. H., 1960b: The Heteroptera of the Netherlands Antilles – III. Saldidae (shore bugs). - Studies on the Fauna of Curaçao and other Caribbean Islands 11(50):4461. Cobben, R. H., 1961: A new genus and four new species of Saldidae (Heteroptera). Entomologische Berichten 21:98-107. Cobben, R. H., 1970: Morphology and taxonomy of intertidal dwarf-bugs (Heteroptera: Omaniidae Fam.-Nov.). – Tijdschrift voor Entomologie 113(2):61-90. Cobben, R. H., 1986: New shorebugs from Solomon Islands, Thailand, India and South America (Heteroptera, Saldidae). – Annales de la Société Entomologique de France (N.S.) 22(2):223-233. Cognato, A. I., 1998: A new Potamobates Champion species (Heteroptera: Gerridae) from Ecuador with new distribution records for P. williamsi Hungerford. – Insecta Mundi 12(1-2):17-22. Consoli, R. A. G. B., M. H. Pereira, A. L. de Melo, and L. H. Pereira, 1989: Belostoma micantulum Stål, 1858 (Hemiptera: Belostomatidae) as a predator of larvae and pupae of Aedes fluviatilis (Diptera: Culicidae) in laboratory conditions. – Memórias do Instituto Oswaldo Cruz 84:577-578. Contartese, A. M. and A. O. Bachmann, 1986 (1985): Distribucion Geografica de las Corixidae Argentinas (Insecta, Heteroptera). – Physis (Buenos Aires), Seccion B:43(105):89-92.

606 Contartese, A. M. and A. O. Bachmann, 1987: Distribucion Geografica de las Belostomatidae Argentinas (Heteroptera). – Physis (Buenos Aires), Seccion B:45(108):25-27. Coscaron, M. C., M. S. Loiacono, and A. O. Bachmann, 2000: Los ejemplares tipo de Leptopodomorpha y Nepomorpha (Heteroptera) depositados en la colección del Museo de la Plata. – Revista del Museo de la Plata, Serie Técnica y Didática (39):39-42. Cranston, P. S., and P. J. Gullan, 2003. Phylogeny of Insects. In: V. H. Resh and R. T. Cardé (Eds.), Encyclopedia of Insects. – Academic Press, Amsterdam. pp. 882-898. Crespo, F. A., 1984: Nuevas citas de Limnogonus (Limnogonus) aduncus y L. (Neogerris) lubricus (Heteroptera: Gerridae). – Physis, Seccion B., 42(103):76. Crespo, F. A., and M. S. Iglesias, 2001 (2000): New records and biological features of Gerridae from Argentina and Uruguay (Insecta, Heteroptera). – Physis, Seccion B 58(134-135):47-48. Cummings, C., 1933: The giant water bugs (Belostomatidae-Hemiptera). University of Kansas Science Bulletin 21(2):197-219.

– The

Deay, H. O., 1930: Six new species of Tenagobia Bergroth (Hemiptera, Corixidae). – Bulletin of the Brooklyn Entomological Society 25(3):171-179 + pl. IX-X. Deay, H. O., 1935: The genus Tenagobia Bergroth (Corixidae, Hemiptera). – The University of Kansas Science Bulletin 22(14):403-477. De Carlo, J. A., 1930: Familia Belostomatidae. Géneros y especies para la Argentina. – Revista de la Sociedad Entomológica Argentina 3(13):101-124 + pl. 5-7. De Carlo, J. A., 1931a: Una nueva especie del género Lethocerus y el alotipo de Lethocerus Del Pontei (Hem. Belostomatidae). – Revista de la Sociedad Entomológica Argentina 3(4):217-218. De Carlo, J. A., 1931b: Géneros y especies nuevas para la Argentina. – Revista de la Sociedad Entomológica Argentina 3(6):329-330. De Carlo, J. A., 1932: Nuevas especies de belostomidos (Hemiptera). – Revista de la Sociedad Entomológica Argentina 5(22):121-126+pl. VI. De Carlo, J. A., 1933: Familia Belostomidae (Hemiptera). Descripción de especies nuevas del Brasil. – Boletim do Museu Nacional do Rio de Janeiro 9:93-98. De Carlo, J. A., 1934: Descripción de especies nuevas de belostómidos (Hemiptera). – Boletim do Museu Nacional de Rio de Janeiro 10:109-111.

607 De Carlo, J. A., 1935a: Familia Belostomatidae. Descripción de una nueva especie y algunas sinonimias. – Revista de la Sociedad Entomológica Argentina 7:203-205 + pl. 16. De Carlo, J. A., 1935b: Catálogo, con la bibliografía más importante de los hemípteros acuáticos y semiacuáticos de Chile. – Revista Chilena de Historia Natural 39:105-111. De Carlo, J. A., 1938a: Los belastómidos Americanos (Hemiptera). – Anales del Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” 39(Entomología 155):189-260. De Carlo, J. A., 1938b: I) Dos nuevas especies del género Abedus Stal. II) Nuevas consideraciones sobre Belostoma costa-limai De Carlo y Lethocerus truncatus Cummings (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 10(1):41-45 + pl. IV-V. De Carlo, J. A., 1939: I) Metamorfosis de Belostoma elegans Mayr. II) Belostoma ellipticum Latreille = Belostoma impavidum Torre Bueno (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 10(2):231-234. De Carlo, J. A., 1940: 1) Descripcion de tres especies nuevas del genero Cryphocricus Signoret. II) Una nueva especie del genero Heleocoris Stal. (Hemiptera – Naucoridae). - Revista de la Sociedad Entomológica Argentina 10(5):426-433. De Carlo, J. A., 1941: I) Descripcion de dos especies nuevas del genero Limnocoris Stal. – II) Nuevas consideraciones sobre Cryphocricus daguerrei y Cryphocricus rufus de Carlo (Hemiptera-Naucoridae). – Revista de la Sociedad Entomológica Argentina 11(1):37-41. De Carlo, J. A., 1942: Descripcion de una especie nueva del genero Belostoma Latr. (Belostomatidae-Hemiptera). – Revista de la Sociedad Entomológica Argentina 11(3):212-213 + pl. 13. De Carlo, J. A., 1946: Los Ranatridae de Sud America (Hemiptera). – Anales del Museo Argentino de Ciencias Naturales «Bernardino Rivadavia», Entomología 42(161):1-38+pl. I-VII. De Carlo, J. A., 1950a: Descripción de especies nuevas de "Ranatridae" y "Belostomatidae" y algunas aclaraciones referentes a otras conocidas (Hemiptera). Revista Brasileira de Biologia 10(4):521-532. De Carlo, J. A., 1950b: Generos y especies de la subfamilia Ambrysinae Usinger (Hemiptera-Naucoridae). – Anales de la Sociedad Cientifica Argentina 150:3-27.

608 De Carlo, J. A., 1951a: Népidos de América (Hemiptera). – Revista del Instituto Nacional de Investigacion de las Ciencias Naturales anexo al Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Ciencias Zoológicas 1(9):385-421. De Carlo, J. A., 1951b: I) Nueva agrupacion en subgenero de las especies del genero Abedus Stal (Hemipt. Belostom.). II) Descripcion de dos especies nuevas del genero Cryphocricos (Hemipt. Naucor.). – Revista de la Sociedad Entomológica Argentina 15(1-3):69-76. De Carlo, J.A., 1951c: Género Limnocoris Stal (Hem. Naucor.). Misiónes de Estudios de Patología Regional Argentina 22:41-51.

– Publicaciones

De Carlo, J. A., 1954a: I. Especies nuevas del género Ranatra (Nepidae). II. Alotipo de Limnocoris bergi y L. brasiliensis (Naucoridae). – Revista de la Sociedad Entomológica Argentina 17(1-2):9-14. De Carlo, J.A., 1954b (1953): Subfamilia Gelastocorinae. Nuevas especies de los géneros Gelastocoris Kirkaldy y Montandonius Melin (Gelastocoridae Hemiptera). – Publicaciones Misiónes de Estudios de Patología Regional Argentina 24(83-84):87-102. De Carlo, J. A., 1956: Neuvas especies de los generos Curicta Stål y Telmatotrephes Stål (Nepidae - Hemiptera). - Boletin del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" 1:1-6 + 5 pl. De Carlo, J. A., 1957 (1956): Especies nuevas de la familia Belostomatidae (Hem.) y algunas consideraciones sobre otras poco conocidas. – Revista de la Sociedad Entomológica Argentina 19(3-4):51-56. De Carlo, J. A., 1958 (1957): Identificación de las especies del género Horvathinia Montandon. Descripción de tres especies nuevas (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 20(3-4):45-52. De Carlo, J. A., 1960a (1959): Descripción de dos especies nuevas del genero Curicta Stal y consideraciones sobre otras poco conocidas. – Actas Trabajos Primer Congreso Sudamericano de Zoologia, Sección IV - Entomologia 3:47-53. De Carlo, J. A., 1960b (1959): Especies nuevas del género Belostoma y consideraciones sobre otras poco conocidas (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 22(1-4):47-59. De Carlo, J.A., 1960c (1959): Generos “Gelastocoris” y “Montandonius” (HemipteraGelastocoridae), Revision de las especies de Guerin y de Carlo consideradas sinonimas. – Acta Zoologica Lilloana 17:53-85.

609 De Carlo, J. A., 1960d: Especies del genero Abedus Stal consideradas erroneamenta sinonimas de otras (Hemiptera-Belostomatidae). - Anales de la Sociedad Cientifica Argentina 175:69-78. De Carlo, J. M., 1962a: Consideraciones sobre la biologia de Lethocerus mazzai de Carlo (Hem. Belostomatidae). - Physis 23(65):143-151. De Carlo, J. A., 1962b: Descripción de una especie nueva del género Lethocerus Mayr. (Hem. Belostomatidae). – Revista de la Sociedad Entomológica Argentina 23(1-4):2123. De Carlo, J. A., 1963a: Aparato genital de macho y hembra de Lethocerus mazzai De Carlo (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 26(1-4):125-128. De Carlo, J. A., 1963b (1961): Una especie nueva del género Ranatra y datos referentes a otras poco conocidas (Hemiptera-Ranatridae). - Revista de la Sociedad Entomológica Argentina 24(1-4):93-96. De Carlo, J. A., 1963c (1961): Una nueva especie del género Ranatra y referencias de Ranatra magna Kuitert (Ranatridae) II. Una nueva especie del género Belostoma Latreille (Belostomatidae). - Revista de la Sociedad Entomológica Argentina 24(14):19-22. De Carlo, J. A., 1963d (1961): Una nuevo género y una nueva especie de la subfamilia Ambrysinae (Hemiptera-Naucoridae). – Revista de la Sociedad Entomológica Argentina 24(1-4):9-11. De Carlo, J. A., 1964a: Los de America (Hemiptera). - Revista Museo Argentino de Ciencias Naturales "Bernardino Rivadavia," Entomología 1:133-216+ 6 pl. De Carlo, J. A., 1964b: Genero Lethocerus Mayr (Hemiptera-Belostomatidae). - Physis 24(68):337-350. De Carlo, J. A., 1965: Segmentos genitales y contenido de la cápsula genital en machos de especies de los géneros Ambrysus, Pelocoris, Limnocoris, Heleocoris y Cryphocricos (Hemiptera-Naucoridae). – Revista de la Sociedad Entomológica Argentina 27(14):101-106. De Carlo, J. A., 1966a: Un nuevo genero, nuevas especies y referencias de otras poco conocidas de la familia Belostomatidae (Hemiptera). – Revista de la Sociedad Entomológica Argentina 28(1-4):97-109.

610 De Carlo, J. A., 1966b: Un nuevo género y nuevas especies de las subfamilias Limnocorinae y Ambrysinae (Hemiptera-Naucoridae). – Revista de la Sociedad Entomológica Argentina 28(1-4):111-117. De Carlo, J. A., 1967a (1966): Nueva especies de los géneros Ranatra y Curicta: referencias de outras poco conocidas (Hemiptera-Ranatridae). – Revista de la Sociedad Entomológica Argentina 29(1-4):30-36. De Carlo, J. A., 1967b: Diferencias entre Ranatridae y Nepidae. Division de la familia Ranatridae en Ranatrinae y Curictinae (Hemiptera). – Revista de la Sociedad Entomológica Argentina 29(1-4):21-29. De Carlo, J. A., 1967c: Una nueva especie del genero Ranatra y nuevas especies de Naucoridae (Hemiptera). - Amazoniana 1(2):189-200. De Carlo, J. A., 1968a: Aparato genital de macho y hembra de Belostoma sp. (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 30(14):45-49. De Carlo, J. A., 1968b: Redescripcion y aparato genital masculino de Austronepa angusta (Hale) (Hemiptera, Ranatridae). - Physis 27(75):343-347. De Carlo, J. A., 1968c: Aparato genital feminino Ranatra (Hemiptera, Ranatridae). Physis 28(76):199-201. De Carlo, J. A., 1968d (1967): Descripcion de especies nuevas de los generos Ambrysus, Cryphocricos y Heleocoris. Allotypus de Cryphocricos daguerrei (Hemiptera – Naucoridae). - Revista de la Sociedad Entomológica Argentina 30(1-4):99-104. De Carlo, J. A., 1968e: Tres especies nuevas del género Coleopterocoris y una especie nueva del género Heleocoris (Hemiptera, Naucoridae). – Physis, Seccion B 28(76):193197. De Carlo, J. M., 1969a: Anatomía microscópica de las bombas alimentarias y salivales de de Lethocerus mazzai. – Revista del Museo Argentino de Ciencias Naturales (Zoología) 10 (12):175-187. De Carlo, J. A., 1969b: Estudio comparativo del aparato genital macho de Belostoma sp. y Abedus sp. (Hemiptera-Belostomatidae). – Revista de la Sociedad Entomológica Argentina 31(1-4):115-119. De Carlo, J. A., 1970: Especies nuevas del género Ranatra Fabricius (Hemiptera, Ranatridae). - Physis 29:311-315.

611 De Carlo, J. A., 1971: Valor sistemático del estudio del aparato genital macho en los hemípteros acuáticos y semiacuáticos. División de la familia Naucoridae en dos familias. Caracteristica externa de la cápsula genital. – Revista de la Sociedad Entomológica Argentina 33(1-4):159-166. De Carlo, J. A., 1973 (1972): Clave de las especies americanas descriptas del genero Ranatra Fabricius (Hemiptera, Ranatridae, Ranatrinae). - Revista de la Sociedad Entomológica Argentina 34(1-2):177-185. De Carlo, J. A., 1978: Aparato genital masculino de Gelastocoris paraguayensis De Carlo y de Nerthra ranina (H. S.). Conclusión. (Hemiptera-Gelastocoridae). – Revista de la Sociedad Entomológica Argentina 37(1-4):113-117. De Carlo, J. A., 1983: Hemipteros acuáticos y semiacuáticos. Estudio en grupos en las partes de igual función de los aparatos genitales masculinos de especies estudiadas. – Revista de la Sociedad Entomológica Argentina 42 (1-4):149-154. De Carlo, J. M., and G. N. Pellerano, 1983: Glandulas odoriferas metasternales de dos especies de Lethocerus (Heteroptera, Belostomatidae). – Comunicaciones del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" Entomologia 1(9):125-132. De Cort-Gommers, M., and N. Nieser, 1966: Records of the Antillean water-striders (Heteroptera). – Studies of the Fauna of Curaçao and Other Caribbean Islands 112:7287. De Coursey, R. M., 1971: Keys to families and subfamilies of the nymphs of North American Hemiptera-Heteroptera. – Proceedings of the Entomological Society of Washington 73(4):413-428. Delétang, L., 1917: Sobre la presencia de Belostoma grandis en Tucuman. – Physis 3:99. De Souza, M. A. A., A. L. de Melo, and G. J. C. Vianna, 2006: Heterópteros aquáticos oriundos do Município de Mariana, MG. – Neotropical Entomology 35(6):803-810. Distant, W.L., 1912: Hemiptera. In: H. Scott (Ed.), Contribution to the knowledge of the fauna of Bromeliaciae. Annals and Magazine of Natural History 8th Series, 10:437438. Doesburg, P. H. van, 1984: A new species of Potamocoris Hungerford, 1941 from Suriname (Heteroptera: Naucoridae). – Zoologische Mededelingen 59(2):19-26. Drake, C. J., 1917: A survey of the North American species of Merragata. – The Ohio Journal of Science 17(4):101-105.

612 Drake, C. J., 1949a: Two new Mesoveliidae, with check list of American species (Hemiptera). - Boletin Entomologica Venezolana 7(3-4):145-147. Drake, C. J., 1949b: Concerning some tropical Rhagovelia (Hemiptera: Veliidae). Boletin Entomologica Venezolana 7(3-4):141-144. Drake, C. J., 1949c: Some American Saldidae (Hemiptera). – Psyche 56(4):187-194. Drake, C. J., 1950: New Neotropical Hydrometridae (Hemiptera). – Acta Entomologica Musei Nationalis Pragae 26(329):1-7. Drake, C. J., 1951a: New water-striders from insular America (Hemiptera: Veliidae). – Proceedings of the Entomological Society of Washington 53(6):338-340. Drake, C. J., 1951b: New water-striders from the Americas (Hemiptera: Veliidae). – Revista de Entomologia 22(6):371-378. Drake, C. J., 1951c: New Neogaean water-striders (Hemiptera-Veliidae). – Proceedings of the Biological Society of Washington 64:75-82. Drake, C. J., 1951d: New Neotropical water-striders (Hemiptera-Veliidae). - Great Basin Naturalist 11(1-2):37-42. Drake, C. J., 1952a: The American species of Cylindrostethus Mayr (Hemiptera, Gerridae). – American Museum Novitates No. 1579:1-3. Drake, C. J., 1952b: Two new Microvelia Westwood (Hemiptera: Veliidae). – Bulletin of the Brooklyn Entomological Society 47(1):13-15. Drake, C. J., 1952c: A new tropical hebrid (Hemiptera). Entomologist 28(4):194.

– The Pan-Pacific

Drake, C. J., 1952d: Concerning American Ochteridae (Hemiptera). – The Florida Entomologist 35(2):72-75. Drake, C. J., 1953a: New Neogaean Rhagovelia (Hemiptera: Veliidae). – Proceedings of the Biological Society of Washington 66:145-152. Drake, C. J., 1953b: Synonymical data and description of a new Hydrometra (Hemiptera-Hydrometridae). – Journal of the Kansas Entomological Society 26(1):4041. Drake, C. J., 1954: Synonymical data: descriptions of new Hydrometridae (Hemiptera). – The Great Basin Naturalist 14(3-4):61-66.

613 Drake, C. J., 1955: New South American Saldidae (Hemiptera). – Journal of the Kansas Entomological Society 28(4):152-158. Drake, C. J., 1956: New Neotropical Hydrometridae (Hemiptera). – Proceedings of the Biological Society of Washington 69:153-156. Drake, C. J., 1957a: New Neotropical Halobatinae (Hemiptera: Gerridae). – Bulletin of the Brooklyn Entomological Society 52(5):128-130. Drake, C. J., 1957b: New Neotropical water-striders (Hemiptera). – Proceedings of the Biological Society of Washington 70:111-118. Drake, C. J., 1957c: A necessary correction in the nomenclature of water-striders (Hemiptera). – Proceedings of the Biological. Society of Washington 70:193-194. Drake, C. J., 1957d: Two new species of rhagovelias (Hemiptera-Veliidae). – Bulletin of the Southern California Academy of Sciences 56(1):47-49. Drake, C. J., 1957e: Notes on the genus Velia (Hemiptera: Veliidae). – Journal of the Kansas Entomological Society 30(4):138-140. Drake, C. J., 1958a: Two new species of water-striders from Peru (Hemiptera: Gerridae). – Bulletin of the Brooklyn Entomological Society 53(5):109-111. Drake, C. J., 1958b: A new Neotropical halobatid (Hemiptera: Gerridae). – Bulletin of the Brooklyn Entomological Society 53:100-102. Drake, C. J., 1958c: New Neotropical Veliidae (Hemiptera). – Proceedings of the Biological Society of Washington 71:133-142. Drake, C. J., 1958d: Two new Neotropical microvelias (Veliidae: Hemiptera). Bulletin of the Brooklyn Entomological Society 53(4):59-61.

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Drake, C. J., 1959: Three new water-striders from Brazil (Hemiptera). – Bulletin of the Southern California Academy of Sciences 58(2):107-111. Drake, C. J., 1961: A new Cylindrostethus from Peru (Hemiptera: Gerridae). Proceedings of the Biological Society of Washington 74:65-66.

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Drake, C. J., 1962: Synonymic data and two new genera of shore-bugs (Hemiptera: Saldidae). – Proceedings of the Biological Society of Washington 75:115-124. Drake, C. J., 1963: A new neotropical water-strider (Hemiptera: Gerridae). – Journal of the Kansas Entomological Society 36(2):93-95.

614 Drake, C. J., and J. M. Capriles, 1952: Water striders from Territorio Amazonas of Venezuela. - Great Basin Naturalist 12:47-54. Drake, C. J., and J. C. M. Carvalho, 1948: Concerning South American Saldidae. – Revista de Entomologia 19:473-479. Drake, C. J., and C. M. Carvalho, 1954: New waterstriders from Brazil (Hemiptera). – Proceedings of the Biological Society of Washington 67:223-226. Drake, C. J., and C. M. Carvalho, 1955: Two new Brazilian rhagovelias (Hemiptera, Veliidae). – Anais da Academia Brasileira de Ciências 27:361-363. Drake, C. J., and H. C. Chapman, 1952: A new species of Micracanthia from Florida (Hemiptera: Saldidae). – The Florida Entomologist 35(4):147-150. Drake, C. J., and H. C. Chapman, 1953a: Distributional data and description of a new hebrid (Hemiptera). – The Great Basin Naturalist 13(1-2):9-11. Drake, C. J., and H. C. Chapman, 1953b: Preliminary report on the Pleidae (Hemiptera) of the Americas. – Proceedings of the Biological Society of Washington 66:53-60. Drake, C. J., and H. C. Chapman, 1954: New American waterstriders (Hemiptera). – The Florida Entomologist 37(3):151-155. Drake, C. J., and H. C. Chapman, 1958a: New Neotropical Hebridae, including a catalogue of the American species (Hemiptera). – Journal of the Washington Academy of Science 48(10):317-326. Drake, C. J., and H. C. Chapman, 1958b: The subfamily Saldoidinae (Hemiptera: Saldidae). – Annals of the Entomological Society of America 51:480-485. Drake, C. J., and R. H. Cobben, 1960: The Heteroptera of the Netherlands Antilles – II. Hebridae. – Studies on the Fauna of Curaçao and other Caribbean Islands 11(51):35-43. Drake, C. J., and J. A. De Carlo, 1953: American species of Ranatra annulipes Stål group (Hemiptera: Ranatridae). – Journal of the Washington Academy of Sciences 43(4):109-117. Drake, C. J., and P. H. van Doesburg Jr., 1966: Water-striders of the American genus Trochopus (Hemiptera: Veliidae). – Studies on the Fauna of Suriname and other Guyanas 8(31):65-76. Drake, C. J., and J. Gomez-Menor, 1954: A new genus of American Ochteridae (Hemiptera). - Eos 30:157-159 + pl. 10.

615 Drake, C. J., and H. M. Harris, 1927: Notes on the genus Rhagovelia, with descriptions of six new species. – Prodeedings of the Biological Society of Washington 40:131-138. Drake, C. J., and H. M. Harris, 1928: Two undescribed water striders from Grenada (Hemiptera). – Florida Entomologist 12:7-8. Drake, C. J., and H. M. Harris, 1930: Notes on some South American Gerridae (Hemiptera). – Annals of the Carnegie Museum 19:235-239. Drake, C. J., and H. M. Harris, 1932a: Some miscellaneous Gerridae in the collection of the Museum of Comparative Zoology (Hemiptera). – Psyche 39(4):107-112. Drake, C. J., and H. M. Harris, 1932b: A survey of the species of Trepobates Uhler (Hemiptera, Gerridae). – Bulletin of the Brooklyn Entomological Society 27:113-122 + pl. 12. Drake, C. J., and H. M. Harris, 1933a: A synopsis of the genus Metrobates Uhler (Hemiptera: Gerridae). – Annals of the Carnegie Museum 21:83-88. Drake, C. J., and H. M. Harris, 1933b: New American Veliidae (Hemiptera). Prodeedings of the Biological Society of Washington 46:45-54.

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Drake, C. J., and H. M. Harris, 1934: The Gerrinae of the Western Hemisphere (Hemiptera). – Annals of the Carnegie Museum 23:179-241. Drake, C. J., and H. M. Harris, 1935a: Notes on some American gerrids (Hemiptera). – Arkiv för Zoologi 28B(2):1-4. Drake, C. J., and H. M. Harris, 1935b: Concerning Neotropical species of Rhagovelia (Veliidae: Hemiptera). – Proceedings of the Biological Society of Washington 48:3338. Drake, C. J., and H. M. Harris, 1936: Notes on American water striders. – Prodeedings of the Biological Society of Washington 49:105-108. Drake, C. J., and H. M. Harris, 1937: Notes on some American Halobatinae (Gerridae, Hemiptera). – Revista de Entomologia 7(4):357-362. Drake, C. J., and H. M. Harris, 1938: «Veliidae» y «Gerridae» sudamericanos descriptos por Carlos Berg. - Notas del Museo de La Plata, Zoologia 3(13):199-204. Drake, C. J., and H. M. Harris, 1940: A new Velia from Peru (Hemiptera, Veliidae). – Pan-Pacific Entomologist 16:31.

616 Drake, C. J., and H. M. Harris, 1941: Concerning some halobatinids from the Western Hemisphere (Hemip, Gerridae). – Iowa State College Journal of Science 15:237-240. Drake, C. J., and H. M. Harris, 1942a: Notas sobre “Rheumatobates”, com descrição uma nova espécie (Hemiptera, Gerridae). - Revista Brasileira de Biologia 2(4):399-402. Drake, C. J., and H. M. Harris, 1942b: A new Brachymetra from Venezuela (HemipteraGerridae). – Boletin de Entomológia Venezolana 1(4):95-96. Drake, C. J., and H. M. Harris, 1943: Notas sobre Hebridae del Hemisferio Occidental (Hemiptera). - Notas del Museo de La Plata, Zoologia 8(64):41-58. Drake, C. J., and H. M. Harris, 1944: A new Rheumatobates from Brazil, with a note on R. imitator (Uhler) (Hemipt. Gerridae). - Revista Entomologica 15:269-272. Drake, C. J., and H. M. Harris, 1945a: Concerning the genus “Metrobates” Uhler (Hemiptera, Gerridae). - Revista Brasileira de Biologia 5(2):179-180. Drake, C. J., and H. M. Harris, 1945b: A new Brachymetra from Venezuela (Hemiptera: Gerridae). - Boletin Entomologica Venezolana 1(4):95-97. Drake, C. J., and H. M. Harris, 1946: A new mesoveliid from Ecuador (Hemiptera, Mesovelidae). – Bulletin of the Brooklyn Entomological Society 41(1):8-9. Drake, C. J., and L. Hoberlandt, 1950: Catalogue of genera and species of Saldidae (Hemiptera). – Acta Entomologica Musei Nationalis Pragae 26(376):1-12. Drake, C. J., and L. Hoberlandt, 1952: A new genus and species of Saldidae from South America (Hemiptera). – Psyche 59(3):85-88. Drake, C. J., and F. C. Hottes, 1950: Three new species of Saldidae (Hemiptera). – Prodeedings of the Biological Society of Washington 63:177-184. Drake, C. J., and F. C. Hottes, 1951a: Stridulatory organs in Saldidae (Hemiptera). Great Basin Naturalist 11(1-2):43-46. Drake, C. J., and F. C. Hottes, 1951b: Notes on the genus Rheumatobates Bergroth (Hemiptera: Heteroptera). – Prodeedings of the Biological Society of Washington 64:147-158. Drake, C. J., and F. C. Hottes, 1951c: Brasilian Saldidae (Hemiptera). – Revista de Entomologia 22(1-3):379-382. Drake, C. J., and F. C. Hottes, 1952: Genus Trepobates Herrich-Schaeffer (Hemiptera; Gerridae). - Great Basin Naturalist 12(1-4):35-38.

617 Drake, C. J., and F. C. Hottes, 1955: Concerning Saldidae (Hemiptera) of the Western Hemisphere. - Boletin Entomologica Venezolana 11(1-2):55-66. Drake, C. J., and R. F. Hussey, 1951: Concerning some American Microvelia (Hemiptera: Veliidae). - Florida Entomologist 34(4):137-145. Drake, C. J., and R. F. Hussey, 1955: Concerning the genus Microvelia Westwood, with descriptions of two new species and a checklist of the American forms (Hemiptera: Veliidae). - Florida Entomologist 38(3):95-115. Drake, C. J., and R. F. Hussey, 1957: Notes on some American Rhagovelia, with descriptions of two new species (Hemiptera: Veliidae). – Occasional Papers of the Museum of Zoology, University of Michigan No. 580:1-6+1 pl. Drake, C. J., and D. R. Lauck, 1959a: The genus Veloidea Gould (Hemiptera: Veliidae). – Proceedings of the Biological Society of Washington 72:161-166. Drake, C. J., and D. R. Lauck, 1959b: Descriptions, synonymy, and check-list of American Hydrometridae (Hemiptera: Heteroptera). – The Great Basin Naturalist 19(23):43-52. Drake, C. J., and A. S. Menke, 1962: Water-striders of the subgenus Stridulivelia from Mexico, Central America, and the West Indies (Hemiptera: Veliidae). – Proceedings of the United States National Museum 113 (3460):413-419. Drake, C. J., and F. Plaumann, 1953: A new waterstrider from Brazil (Hemiptera: Veliidae). – Dusenia 4(5-6):414-416. Drake, C. J., and F. Plaumann, 1955: Some Microvelia from southern Brazil (Hemiptera: Veliidae). – Bulletin, Southern California Academy of Sciences 54(1):22-24. Drake, C. J., and J. A. Roze, 1954: New Venezuelan Gerridae (Hemiptera). Proceedings of the Biological Society of Washington 67:227-230.

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Drake, C. J., and J. A. Roze, 1955: A new species of Veloidea from Venezuela (Hemiptera: Veliidae). – Bulletin of the Brooklyn Entomological Society 50(4):106109. Drake, C. J., and J. A. Roze, 1958: A new Microvelia from Venezuela (Hemiptera: Veliidae). – Bulletin, Southern California Academy of Sciences 57(1):47-48. Dufour, L., 1863: Essai monographique sur les Belostomides. – Annales de la Société Entomologique de France (sér. 4):3:375-400.

618 Egbert, A., 1949: A new Tenagobia from Peru (Hemiptera, Corixidae). – Journal of the Kansas Entomological Society 22(1):35-36. Esaki, T., 1926a: The water-striders of the subfamily Halobatinae in the Hungarian National Museum. – Annales Musei Nationalis Hungarici 23:117-164. Esaki, T., 1926b: On some species of Potamobates (Hemiptera: Gerridae). – Annales Musei Nationalis Hungarici 23:251-257. Esaki, T., 1927: An interesting new genus and species of Hydrometridae (Hem.) from South America. – Entomologist 60:181-184. Esaki, T., and W. China, 1928: A monograph of the Helotrèphidae, subfamily Helotrephinae (Hem. Heteroptera). – Eos 4(2):129-172. Espinola, H. N., 1975: Tenagobia schreiberi sp. n. coletado no Muncipio de Diamantina, Minas Gerais, Brasil (Hemiptera, Corixidae, Micronectinae). – Ciencia e Cultura 27(8):893-900. Estévez, A. L., and M. L. López Ruf, 2006: Subfamily separation in the hemipteran family Gelastocoridae (Heteroptera). – Revista de Biologia Tropical 54(1):1319-1322. Estévez, A. L., and P. J. Pérez Goodwyn, 1999: Morfología externa de los estadíos preimaginales de Lethocerus truncatus Cummings (Heteroptera: Belostomatidae). – Revista Brasileira de Entomologia 43:89-93. Estévez, A. L., and J. T. Polhemus, 2001: The small species of Belostoma (Heteroptera, Belostomatidae). I. Key to species groups and a revision of the denticolle group. – Iheringia, Série Zoologia 91:151-158. Estévez, A. L., and J. T. Polhemus, 2007: The small species of Belostoma (Heteroptera, Belostomatidae). I. Revision of the plebejum group. – Revista de Biologia Tropical 55(1):147-155. Estévez, A. L., and A. C. Armúa de Reyes, 2003: Una nueva especie de Belostoma (Heteroptera: Belostomatidae) de la laguna Iberá (Corrientes, Argentina). – Facena 19:123-127. Estévez, A. L., A. C. Armúa de Reyes, A. O. Bachmann, D. Carpintero, M. L. López Ruf, S. Mazzucconi, P. J. Pérez Goodwyn, and P. Hernández, 2003: Anexo II: Biodiversidad de heterópteros acuáticos y semiacuáticos de los esteros del Iberá. In: B.B. Alverez (Ed.), Fauna del Iberá. – Universidad Nacional Nordeste, Corrientes. pp. 369-375.

619 Estévez, A. L., C. A. de Reyes, and J. A. Schnack, 2006: Successful hatching from eggs carried by females and naturally removed from incumbant males in Belostoma spp. water bugs (Hemiptera: Belostomatidae). – Revista de Biología Tropical 54(2):515-517. Estévez, A. L., and J. A. Schnack, 1977: Una nueva especie del genero Gelastocoris Kirkaldy (Hemiptera, Gelastocoridae). – Neotropica 23 (69):81-83. Estévez, A. L., and J. A. Schnack, 1980a: Las ninfas de Merragata lacunifera (Berg) (Hemiptera, Hebridae). – Neotropica 26(75):29-33. Estévez, A. L., and J. A. Schnack, 1980b: Subfamilia Gelastocorinae Champion. Revision de las especies de America del Sur (Hemiptera, Gelastocoridae). – Revista del Museo de La Plata (n.s.), Obra del Centenario 6:75-89. Estévez, A. L., and J. A. Schnack, 1980c: Descripción de las ninfas de Nerthra ranina (Herrich-Schaffer)(Hemiptera, Gelastocoridae). – Revista de la Sociedad Entomológica Argentina 37(1-4):119-123. Estévez, A. L., and J. A. Schnack, 1980d: Descripción de una nueva especie del genero Nerthra Say (Hemiptera, Gelastocoridae). – Revista de la Sociedad Entomológica Argentina 39(1-2):59-61. Ewald, G., 1963: Vergleichende Studien an Nepiden und Belostomatiden über eine Haftvorrichtung zwischen den Vorderflügeln (Ins., Heteroptera). – Senckenbergiana Biologica 44(5):417-430. Fabricius, J. C., 1803: Systema Rhyngotorum secundum classes, ordines, genera, species adjectis synonymis, locis observationibus, descriptionibus. – Carolus Reichard, Brunswick. vi + 314 pp. Fernández, L. A., and M. López Ruf, 1999: Coleoptera y Heteroptera acuáticos y semiacuáticos de la Isla Martín García (Provincia de Buenos Aires). Physis, Seccion B., 57(132-133):1-4. Fernández, L. A., and M. López Ruf, 2006: Aquatic Coleoptera and Heteroptera inhabiting waterbodies from Berisso, Buenos Aires Province, Argentina. – Revista Biologia Tropical 54(1):139-148. Fieber, F. X., 1851a: Species generis Corisa. Gesellschaft der Wissenschaften. (5)7:213-260.

Abhandlungen der Böhmischen

Fieber, F. X., 1851b: Rhynchotographieen. III. Die Gruppe der Notonectiae. – Gottlieb Haase Söhne, Prague. pp. 43-64.

620 Froeschner, R. C., 1981: Heteroptera or true bugs of Ecuador: a partial catalog. Smithsonian Contributions to Zoology No. 322. iv. + 147 pp. Froeschner, R. C., 1985: Synopsis of the Heteroptera or true bugs of the Galápagos Islands. - Smithsonian Contributions to Zoology No. 407. iii+84 pp. Froeschner, R. C., 1999: True Bugs (Heteroptera) of Panama: A Synoptic Catalog as a Contribution to the Study of Panamanian Biodiversity. – Memoires of the American Entomological Institute, Gainsville, Florida, Vol. 61. pp. i-ii, 1-393. Galactos, K., A. I. Cognato, and F. A. H. Sperling, 2002: Population genetic structure of two water strider species in the Ecuadorian Amazon. – Freshwater Biology 47(3):391399. Galbreath, J. E., 1973: Diapause in Mesovelia mulsanti (Hemiptera: Mesoveliidae). – Journal of the Kansas Entomological Society 46(2):224-233. Galbreath, J.E., 1975: Thoracic polymorphism in Mesovelia mulsanti (Hemiptera: Mesoveliidae). – University of Kansas Science Bulletin 50(10):457-482. Galen, C., and F. F. Herrera, 2006: Fauna cavernicola de Venezuela: una revision. – Boletín de la Sociedad Venezolana de Espeleología 40:39-57. Galileo, M. H. M., 1974: Contribuição ao conhecimento da genitalia dos machos do gênero Horvathinia Montandon, 1911. (Heteroptera, Belostomatidae, Horvathiniinae). – Iheringia, Zoologia 45:26-33. Galliussi, E. A., 1980: Anatomía microscópica de la espermoteca de Tenagobia (Fuscagobia) fuscata (Heteroptera, Corixidae). – Revista de la Sociedad Entomológica Argentina 39(3-4)287-292. Galliussi, E. A., 1989: Desarrollo larval de Tenagobia (Fuscagobia) fuscata (Heteroptera, Corixidae). – Revista de la Sociedad Entomológica Argentina 45(1-4)131144. García, H. A., 2002: Los hemípteros de la película superficial del agua en Colombia. Parte 1. Familia Gerridae. - Academia Colombiana de Ciencias Exactas, Físicas y Naturales Colección Jorge Álvarez Lleras No. 20:i-xiv, 1-239. Gistel, J. N. F. X., 1848: Naturgeschichte des Thierreichs für höhere Schulen. Hoffmann, Stuttgart. 216 pp + 32 pl.

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Gistel, J. N. F. X., 1850: Handbuch der Naturgeschichte aller drei Reiche, für Lehrer und Lernende, für Schule und Haus. – Hoffmann, Stuttgart. 1037 pp.

621 Gittelman, S.H., 1974a: The habitat preference and immature stages of Neoplea striola (Hemiptera: Pleidae). - Journal of the Kansas Entomological Society 47(4):491-503. Gittelman, S.H., 1974b: Descriptions of immature and adult stages of Martarega hondurensis Bore. - Journal of the Kansas Entomological Society 47(2):145-155. Gomes, C. L., 1981: Alguns aspectos morphológicos de Brachymetra albinervus (Amyot & Serville, 1843) (Hemiptera-Gerridae). – Memórias do Instituto Oswaldo Cruz 76(2):169-180. Gorb, S. N., and P. J. Pérez Goodwyn, 2003: Wing-locking mechanisms in aquatic Heteroptera. – Journal of Morphology 157:127-146. Goulart, M., A. Lane de Melo, and M. Callisto, 2002: Qual a relação entre paramêtros ambientais e a diversidade de heterópteros aquáticos em nascentes de altitude? - Bios 10:63-76. Gould, G. E., 1931: The Rhagovelia of the Western Hemispbere, with notes on world distribution (Hemiptera, Veliidae). – The Univeristy of Kansas Science Bulletin 20(1):5-62. Guérin-Menéville, C., 1843: In: Cuvier, Iconographie du Règne Animal VII. (n.v.) Guérin-Menéville, C., 1857: Orthoptera et Hemiptera de l’Île de Cuba 2(7):136-182. – Bertrand, Paris. Guimarães, R. R., H. R. S. Rodrigues, and R. R. Guimarães, 2003: Coincidência no aparecimento de adultos de insetos de duas ordens diferentes em Armação de Búzios, Rio de Janeiro, Brasil. – Boletim S.E.A. 33:274-275. Haddad, C. F. B., and R. P. Bastos, 1997: Predation on the toad Bufo crucifer during reproduction (Anura: Bufonidae). – Amphibia-Reptilia 18(3):295-298. Harris, H. M., and C. J. Drake, 1941: Notes on the family Mesoveliidae (Hemiptera) with descriptions of two new species. – Iowa State College Journal of Science 15:275277. Harris, H. M., and C. J. Drake, 1945: A new Brachymetra from Peru with a list of known species (Hemiptera, Gerridae). – Proceedings of the Entomological Society of Washington 47(7):211-212. Heckman, C. W., 1983: Comparative morphology of arthropod exterior surfaces with the capability of binding a film of air underwater. – Internationale Revue der gesamten Hydrobiologie 68:715-736

622 Heckman, C. W., 1994: New limnological nomenclature to describe ecosystem structure in the tropical wet-and-dry climatic zone. – Archiv für Hydrobiologie 130:385-407. Heckman, C. W., 1998: The Pantanal of Poconé. Biota and ecology in the northern section of the world's largest pristine wetland. – Kluwer, Dordrecht. 662 pp. Herrich-Schäffer, G. A. W., 1845: Die wanzenartige Insekten: getreu nach der Natur abgebildet und beschrieben. Vol. 8. No. 1 – J. R. Lotzbeck, Nüremberg. pp. 1-48 + Figs. 803-804. Herrich-Schäffer, G. A. W., 1848: Die wanzenartige Insekten. Vol. 8. – J. R. Lotzbeck, Nüremberg. 130 pp. Herrich-Schäffer, G. A. W., 1849: Die wanzenartige Insekten: getreu nach der Natur abgebildet und beschrieben. Vol. 9. No. 1. Vol. 8. – J. R. Lotzbeck, Nüremberg. pp. 1-44. Herring, J. L., 1961: The genus Halobates (Hemiptera: Gerridae). – Pacific Insects 3(23): 223-305. Hodgden, B. B., 1949: New Saldidae from the Western Hemisphere (Hemiptera). Journal of the Kansas Entomological Society 22(4):149-165. Horváth, G., 1918: De hydrocorisis nonnullis extraeuropaeis. Nationalis Hungarici 16:140-146.

– Annales Musei

Hungerford, H. B., 1917: The life-history of Mesovelia mulsanti White. – Psyche 24(3):73-84+pl.4. Hungerford, H. B., 1919: Notes on the aquatic Hemiptera. – The Kansas University Science Bulletin 11(5):141-151. Hungerford, H. B., 1922: The life history of the toad bug. – The Kansas University Science Bulletin 14(5):145-171. Hungerford, H. B., 1924: A new Mesovelia with some biological notes regarding it. – Canadian Entomologist 56:142-144. Hungerford, H. B., 1926: Some Notonecta from South America. – Psyche 33(1):1115+pl. 2. Hungerford, H. B., 1927a: A report upon the aquatic and semi-aquatic Hemiptera of the Mulford Biological Expedition to Bolivia, South-America, 1921-22. – Proceedings of the Entomological Society of Washington 29(8):187-190+pl. 10.

623 Hungerford, H. B., 1927b: Trichocorixa and not Corixa for the genus of Corixidae found in America. – Bulletin of the Brooklyn Entomological Society 22:96. Hungerford, H. B., 1927c: A new Ramphocorixa from Haiti (Hemiptera-Corixidae) – American Museum Novitates No. 278:1-2. Hungerford, H. B., 1928a: Notes on the genus Heterocorixa White with the description of some new species (Hemiptera-Corixidae). – Bulletin of the Brooklyn Entomological Society 23(2):99-103 + pl. 3. Hungerford, H. B., 1928b: Some South American Corixidae. – Bulletin of the Brooklyn Entomological Society 23(4):174-180 Hungerford, H. B., 1928c: Two new Notonecta from South America (HemipteraNotonectidae). – Annals of the Entomological Society of America 21(1):119-120. Hungerford, H. B., 1929a: Some new semi-aquatic Hemiptera from South America with a record of stridulatory devices (Veliidae-Velia). Journal of the Kansas Entomological Society 2(3):50-59. Hungerford, H. B., 1929b: A new genus of semi-aquatic Hemiptera. – Bulletin of the Brooklyn Entomological Society 24(5):288-291. Hungerford, H. B., 1929c: A new Velia from Peru (Hemiptera, Veliidae). Entomologische Tidskrift (Stockholm) 50(2):146-147.

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Hungerford, H. B., 1929d: A new Velia from Arizona with notes on other species (Hemiptera-Veliidae). – Annals of the Entomological Society of America 22(4):759761. Hungerford, H. B., 1929e: Two new species of Hemiptera in the collections of the Museum National of Paris. – Bulletin du Muséum, 2nd series 1(3):198-200. Hungerford, H. B., 1930a: Concerning Velia inveruglas Kirkaldy and related forms (Veliidae-Hemiptera). – Annals of the Entomological Society of America 23(1):120125. Hungerford, H. B., 1930b: Three new Velia from South America. - Journal of the Kansas Entomological Society 3(1):23-27. Hungerford, H. B., 1930c: A report on the nomenclature of some neotropical Notonecta with the descriptions of some new species. Bulletin of the Brooklyn Entomological Society 25:138-143.

624 Hungerford, H. B., 1932: A new Potamobates (Gerridae). - Bulletin of the Brooklyn Entomological Society 27:228-230. Hungerford, H. B., 1934 (1933): The genus Notonecta of the world (NotonectidaeHemiptera. - University of Kansas Science Bulletin 21(1):5-195. Hungerford, H. B., 1935: The genus Bacillometra Esaki, including the description of a new species from Peru (Hemiptera, Hydrometridae). – Revista de Entomologia 5(2):117-123. Hungerford, H. B., 1936: A new Potamobates from Peru, S.A. (Hemiptera, Gerridae). Bulletin of the Brooklyn Entomological Society 31:178-180. Hungerford, H. B., 1937: A second new Potamobates from Peru, S.A., with notes on other species (Hemiptera-Gerridae). - Bulletin of the Brooklyn Entomological Society 32:144-146+pl. 2. Hungerford, H. B., 1938a: A third new Potamobates from Peru, S. A. (HemipteraGerridae). – Journal of the Kansas Entomological Society 11(3):85-87. Hungerford, H. B., 1938b: Mesovelia williamsi Hungerford – a note on its distribution. Bulletin of the Brooklyn Entomological Society 33:218. Hungerford, H. B., 1939: Two new Corixidae from Bolivia. – Journal of the Kansas Entomological Society 12(3):97-99. Hungerford, H. B., 1941: A remarkable new naucorid water bug (Hemiptera). – Annals of the Entomological Society of America 34(1):1-4. Hungerford, H. B., 1942: Coleopterocoris, an interesting new genus of the subfamily Potamocorinae (Naucoridae: Heteroptera. – Annals of the Entomological Society of America 35(2):135-139. Hungerford, H. B., 1944: Some Venezuelan aquatic Hemiptera. – Zoologica 29:129. Hungerford, H. B., 1948a: The Corixidae of the Western Hemisphere (Hemiptera). – University of Kansas Science Bulletin 32:4-827. Hungerford, H. B., 1948b: The eggs of Corixidae (Hemiptera). – Journal of the Kansas Entomological Society 21(4):141-142 + Pl. 1. Hungerford, H. B., 1950: Two new generic names. Entomological Society 23(2):73.

– Journal of the Kansas

625 Hungerford, H. B., 1951: A new Metrobates from Brazil, South America (HemipteraGerridae). – Journal of the Kansas Entomological Society 24(2):72-73. Hungerford, H. B., 1953a: Concerning Charmatometra bakeri (Kirkaldy) (Hemiptera: Gerridae). – Entomological News 64:172-175. Hungerford, H. B., 1953b: Concerning Mesovelia douglasensi Hungerford. – Journal of the Kansas Entomological Society 26(1):76-77. Hungerford, H. B., 1954: The genus Rheumatobates Bergroth (Hemiptera-Gerridae). University of Kansas Science Bulletin 36(1):529-588. Hungerford, H. B., 1958: A new genus of Gerridae (Hemiptera) from South America. Florida Entomologist 41:125-128. Hungerford, H. B., and N. E. Evans, 1934: The Hydrometridae of the Hungarian National Museum and other studies in the family. (Hemiptera). – Annales Musei Nationalis Hungarici 28:31-112. Hungerford, H. B., and R. Matsuda, 1957: Descriptions of two new species of the genus Brachymetra (Gerridae, Hemiptera). – Journal of the Kansas Entomological Society 30(1):19-25. Hungerford, H. B., and R. Matsuda, 1958a: Two new genera of Gerridae with the description of a new species. – Journal of the Kansas Entomological Society 31(2):113117. Hungerford, H. B., and R. Matsuda, 1958b: The Tenagogonus-Limnometra complex of the Gerridae. – University of Kansas Science Bulletin 39(9):371-457. Hungerford, H. B., and R. Matsuda, 1958c: Concerning the genus Limnogonus and a new subgenus (Heteroptera: Gerridae). – Journal of the Kansas Entomological Society 32(1):40-41. Hungerford, H. B., and R. Matsuda, 1960: Keys to subfamilies, tribes, genera and subgenera of the Gerridae of the World. – University of Kansas Science Bulletin 41(1):3-23. Hungerford, H. B., and R. Matsuda, 1961: A review of the subgenus Neogerris Matsumura (Hemiptera: Gerridae). – Insecta Matsumurana 24(2):112--114. Hynes, H. B. N., 1948: Notes on the aquatic Hemiptera-Heteroptera of Trinidad and Tobago, B.W.I., with a description of a new species of Martarega B. White (Notonectidae). – Transactions of the Royal Entomological Society of London 99(10):341-360.

626 Iglesias, M. S., and F. A. Crespo, 1999: A new species of the genus Paravelia Breddin 1898 from Argentina (Heteroptera: Veliidae). – Aquatic Insects 21(4):259-265. Iglesias, M. S., M. S. Gaspe, and A. C. Valverde, 2008: A longitudinal study of two species of Belostoma Latreille (Heteroptera: Belostomatidae): allometry and ontogeny. – Neotropical Entomology 37(6):662-667. International Commission on Zoological Nomenclature, 1983: Opinion 1248. Lethocerus Mayr, 1853 (Insecta, Hemiptera): conserved. – Bulletin of Zoological Nomenclature 40:81-82. International Commission on Zoological Nomenclature, 1996: Opinion 1850. Nepa rustica Fabricius, 1781 and Zaitha stollii Amyot & Serville, 1843 (currently Diplonychus rusticus and Belostoma stollii; Insecta, Heteroptera): specific names conserved. – Bulletin of Zoological Nomenclature 53(3):213-214. Ituarte, S., and A. G. Papeschi, 2004: Achiasmatic male meiosis in Tenagobia (Fuscagobia) fuscata (Stål) (Heteroptera, Corixoidea, Micronectidae). – Genetica 122:199-206. Jaczewski, T., 1927a: Argentinian Corixidae collected by Mr. A. C. Jensen-Haarup. – Annales Zoologici Musei Polonici Historiae Naturalis 6(3):251-261 + pl. VII-VIII. Jaczewski, T., 1927b: Corixidae from the State of Paraná. – Annales Zoologici Musei Polonici Historiae Naturalis 6(1):39-59 + pl. II. Jaczewski, T., 1927c: A brief note on the morphology and systematic position of Ectemnostega quadrata (Sign.)(Heteroptera, Corixidae). – Entomologisk Tidskrift 1927:145-149. Jaczewski, T., 1928a: Hydrometridae from the State of Paraná. Zoologici Polonici 7(2-3):81-84 + pl. V.

– Annales Musei

Jaczewski, T., 1928b: Mesoveliidae from the State of Paraná. Zoologici Polonici 7(2-3):75-80 + pl. IV.

– Annales Musei

Jaczewski, T., 1928c: Bemerkungen über die geographische Verbreitung der Corixiden. – Annales Musei Zoologici Polonici 7(1):45-67 + pl. I-III. Jaczewski, T., 1928d: Notonectidae from the State of Paraná. Zoologici Polonici 7(4):121-136 + pl. XIX-XXIII.

– Annales Musei

Jaczewski, T., 1930: Notes on the American species of the genus Mesovelia Muls. (Heteroptera, Mesoveliidae). – Annales Musei Zoologici Polonici 9(21):329-339.

627 Jaczewski, T., 1931a: Notes on Corixidae. VIII – XI. - Annales Musei Zoologici Polonici 9(10):147-154 + Table XIII. Jaczewski, T., 1931b: Die Corixiden (Corixidae, Heteroptera) des Zoologischen Staatsinstituts und Zoologischen Museums in Hamburg. – Mitteilungen des zoologischen Museums Hamburg 44:140-148. Jaczewski, T., 1933a: Notes on some South and Central American Corixidae. – Annales Musei Zoologici Polonici 9(21):149-153 + pl. XLVI. Jaczewski, T., 1933b: Notes on some South and Central American Corixidae. – Annales Musei Zoologici Polonici 9(21):329-339. Jaczewski, T., 1933c: Corixidae collected during the winter voyage, 1931-1932, of the training ship “Dar-Pomorza.”. – Annales Musei Zoologici Polonici 10(1):1-6 + pl 1. Järvinen, O., and K. Vepsäläinen, 1976: Wing dimorphism as an adaptive strategy in water striders (Gerris). – Hereditas 84:61-68. Jordan, K. H. C., 1951: Bestimmungstabellen der Familien von Wanzenlarven. Zoologische Anzeiger 147: 24-31.

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Jurberg, J., and C. L. Gomes, 1983: Estudo morfológico de Limnogonus aduncus Drake & Harris, 1932 e Halobatopsis platensis (Berg, 1879) e suas genitálias externas (Hemiptera-Gerridae). – Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 78(2):199-208. Keffer, S. L., 1997 (1996): Systematics of the New World water scorpion genus Curicta Stål (Heteroptera: Nepidae). – Journal of the New York Entomological Society 104(23):117-215. Keffer, S. L., 1998 (1999): Erratum: The synonymy of C. doesburgi De Carlo with C. montei De Carlo (Heteroptera: Nepidae). – Journal of the New York Entomological Society 106(2-3):114. Keffer, S. L., 1999: Curicta johnpolhemi (Heteroptera: Nepidae): a new waterscorpion species from Brazil. – Journal of the New York Entomological Society 107(2-3):145148. Kenaga, E., 1941: The genus Telmatometra Bergroth (Hemiptera-Gerridae). University of Kansas Science Bulletin 27(9):169-183. Kenaga, E., 1942a: A new genus in the Halobatinae (Gerridae-Hemiptera). – Journal of the Kansas Entomological Society 15:136-141.

628 Kenaga, E., 1942b: Gerrinae in the University of Kansas collections. – University of Kansas Science Bulletin 28:113-143. Kevan, D. K. M., 1948: A note on Mononyx bipunctatus Melin. – Annals and Magazine of Natural History, 11th series, 14(119):812-813. Kirkaldy, G. W., 1899a: On some aquatic Rhynchota from South America in the collections of the Museum of Zoology and Comparative Anatomy of the Royal University of Turin. – Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Università di Torino 14(347):1-2. Kirkaldy, G. W., 1899b: Viaggio del Dr. Enrico Festa nell’Ecuador e regioni vicine. XIX. Aquatic Rhynchota. – Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Università di Torino 14(350):1-9. Kirkaldy, G. W., 1899c: Viaggio del Dr. A. Borelli nella Repubblica Argentina e nel Paraguay. XXIV. Aquatic Rhynchota. – Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Università di Torino 14(351):1-4. Kirkaldy, G. W., 1899d: Viaggio del Dott. A. Borelli nel Chaco boliviano e nella Republica Argentina. XVI. Aquatic Rhynchota. – Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Università di Torino 14(352):1-2. Kirkaldy, G. W., 1899e: Aquatic Rhynchota in the collection of the Royal Museum of Belgium. Notes and descriptions. – Annales del la Société Entomologique de Belgique 43:505-510. Kirkaldy, G. W., 1902: Miscellanea Rhynchotalia – No. 5. – The Entomologist 35:280284, 315-316. Kirkaldy, G. W., 1904: Über Notonectiden (Hemiptera). – Wiener Entomologische Zeitung 23(7):111-135. Kirkaldy, G. W., and J. R. de la Torre Bueno, 1908: A catalogue of American aquatic and semiaquatic Hemiptera. – Proceedings of the Entomological Society of Washington 10(3-4):173-215. Kormilev, N. A., and J. A. De Carlo, 1952: Una especie nueva del genero "Ochterus" Latreille (1807) del Paraguay (Hemiptera. Ochteridae). – Anales de la Sociedad Cientifica Argentina 154:155-159. Kort-Gommers, M. de, and N. Nieser, 1969: Records of Antillean water-striders (Heteroptera). – Studies on the Fauna of Curaçao and other Caribbean Islands 112:7287.

629 Kuitert, L. C., 1942: Gerrinae in the University of Kansas collections. – The University of Kansas Science Bulletin 28:113-143. Kuitert, L. C., 1949a: Some new species of Nepidae (Hemiptera). – Journal of the Kansas Entomological Society 22(2):60-68. Kuitert, L. C., 1949b: Some new Ranatra from the Americas. - Journal of the Kansas Entomological Society 22(1):24-34. La Rivers, I., 1950: A new species of the genus Potamocoris from Honduras (Hemiptera, Naucoridae). – Proceedings of the Entomological Society of Washington 52(6):301-304. La Rivers, I., 1951: Two new ambrysi from South America (Hemiptera, Naucoridae). – Proceedings of the Entomological Society of Washington 53(5):288-293. La Rivers, I., 1952: A new subgenus of Ambrysus from South America (Hemiptera: Naucoridae). – Entomological News 63:33-39. La Rivers, I., 1953: New gelastocorid and naucorid records and miscellaneous notes, with a description of the new species, Ambrysus amargosus (Hemiptera: Naucoridae). – The Wasmann Journal of Biology 11(1):83-96. La Rivers, I., 1954: A new Venezuelan Ambrysus (Hemiptera: Naucoridae). Entomological News 65(8):197-202.

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La Rivers, I., 1956: A new genus and species of naucorid from South America (Hemiptera). – Entomological News 67(9):237-245. La Rivers, I., 1962a: A new Ambrysus from South America (Hemiptera, Naucoridae). – Bulletin of the Southern California Academy of Sciences 61(5):185-187. La Rivers, I., 1962b: A new Limnocoris from Venezuela (Hemiptera: Naucoridae). – Proceedings of the Entomological Society of Washington 64(3):195-196. La Rivers, I., 1963: Two new ambrysi (Hem. - Naucoridae). – Occasional Papers, Biological Society of Nevada, No. 1:1-6. La Rivers, I., 1965: The subgenera of the genus Ambrysus (Hemiptera, Naucoridae). – Occasional Papers, Biological Society of Nevada, No. 4:1-7. La Rivers, I., 1970a: A new Ambrysus from Venezuela (Hemiptera. Naucoridae). – The Great Basin Naturalist 30(1):1-4.

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656 White, F. B., 1879a: Description of new Hemiptera (I). – Journal of the Linnaean Society of London, Zoology 14:482-489. White, F. B., 1879b: List of Hemiptera collected in Amazons by Prof. J. W. Trail, M.A., M.D., in the years 1873-1875, with descriptions of the new species. Part 1. – Transactions of the Entomological Society 1879(4):267-276. Wiley, G. O., 1922: Life history notes on two species of Saldidae (Hemiptera) found in Kansas. – The Kansas University Science Bulletin 14(9):301-311. Wygodzinsky, P., 1948a: Studies on some apterous Aradidae from Brazil (Hemiptera). – Boletim do Museu Nacional, Nova Serie, Zoologia No. 86:1-23+14 pl. Wygodzinsky, P., 1948b: On two genera of “Schizopterinae” (“Cryptostemmatidae”) from the Neotropical region (Hemiptera). – Revista Brasileira de Biologia 8(1):143-155. Wygodzinsky, P. W., and K. Schmidt, 1991: Revision of the New World Enicocephalomorpha (Heteroptera). – Bulletin of the American Museum of Natural History No. 200. 265 pp. Zera, A. J., and K. C. Tiebel, 1991: Photoperiodic induction of wing morphs in the waterstrider Limnoporus canaliculatus (Gerridae: Hemiptera). – Annals of the Entomological Society of America 84(5):509-516.

Index akanthinomeros, Neosigara 353, 354, 355 albinerva, Brachymetra 38, 85, 135 albinerva, Brachymetra albinerva 135 albinervis, Brachymetra 135 albinervus, Helobates 135 albonotata, Microvelia 248 albotrimaculata, Paravelia 223, 225 albotrimaculata, Velia 225 alia, Stridulivelia 213, 214 alia, Velia 214 amazona, Oreokora 589 amazona, Pentacora 589 amazonensis, Gelastocoris 441, 442 amazonensis, Rhagovelia 203 amazonicus, Tropicoparapamphantus 18 amazonum, Belostoma 429 amblydonti, Metrobates 79, 103 Ambrysinae 514, 518 Ambrysus 12, 518, 519, 520-522, 523, 524-531 Ambrysus 522 ameghinoi, Ranatra 380 americana, Nerthra 448 americana, Notonecta 496, 500 americanus, Ochterus 461 amici, Belostoma 429, 430, 434 amnigenopsis, Buenoa 483, 484 amnigenus, Anisops 483 amnigenus, Buenoa 483 amoena, Mesovelia 275, 279 amoena, Paravelia 223 amoena, Velia 223 Amorgius 397, 398, 400 Amphischizops 357, 358, 370 amplicollis, Mononyx 453, 454 amplicollis, Nerthra 452, 453 anchicaya, Potamobates 129, 130

abbreviatus, Limnocoris 544, 552 Abedus 395 abrasum, Limnocoris 545 abrupta, Rhagovelia 180, 181 absidata, Buenoa 472, 473 absona, Neoplea 510, 511 absona, Plea 511 Acanthia 580, 585, 589, 593, 594, 597 acapulcana, Hebrus 271 accedens, Rhagovelia 185 accola, Brachymetra 131 aconita, Tenagobia 302, 303 aculabrum, Limnocoris 24, 565 acuta, Telmatometra 104, 105 acutalis, Limnocoris 551, 552 acutangulus, Ambrysus 530, 531 acutangulus, Ochterus 459, 460 Acyttarsus 520 adamsoni, Tachygerris 85, 138 adamsoni, Tenagogonus 138 adelomorpha, Ranatra 391, 392 admontandoni, Limnocoris 554 aduncus, Limnogonus 38, 75, 147 adusta, Zaitha 434 advena, Euvelia 209, 210 Aedes 54 aegypti, Aedes 54 aemulana, Microvelia 253 aeneifrons, Ochterus 458, 462 aeneifrons, Ochterus aeneifrons 462 aeneifrons, Pelogonus 462 Aenictopecheidae 15 Aenictovelia 211 Aepophilidae 54, 578 Aepophilus 578 agenor, Hydrometra 261 agra, Rhagovelia 162, 184 Aieyrodoidea 11 aiuruoca, Rhagovelia 160, 187 657

658

ancona, Microvelia 245 andensis, Acanthia 589 andensis, Oreokora 589 andensis, Pentacora 589 andensis, Pseudosaldula 584, 589 andensis, Salda 589 andensis, Saldula 589 andersoni, Notonecta unifasciata 495, 497 andinus, Gelastocoris 436, 440 andromeda, Gerris 148 anduzei, Brachymetra 132 anduzei, Heterocorixa 310, 315 angulata, Notonecta unifasciata 495, 497 angulatus, Gelastocoris 437, 438, 439, 440 angulatus, Limnocoris 547 angulatus, Montandonius 439 angusta, Oreokora 586 angusta, Pentacora 586 angustipes, Rhagovelia 178 angustum, Belostoma 408, 409 Anisopinae 464 Anisops 470, 472, 473, 476, 482484 annulipes, Amorgius 398, 400 annulipes, Belostoma 398, 400, 402 annulipes, Lethocerus 48, 400 annulipes, Lethocerus 398 annulipes, Ranatra 388 annulipes, Ranatra 378 anta, Paravelia 232 anta, Stridulivelia 216 antigone, Anisops 475 antigone, Buenoa 474, 475 antigone, Buenoa antigone 474, 475 antiguensis, Arctocorisa 336 antioquia, Pseudosaldula 590, 591 antioquiae, Rhagovelia 181, 183 anura, Zaitha 413, 422 anurum, Belostoma 420 anurus, Diplonychus 420

anurus, Zaitha 420, 423 apache, Belostoma 435 Aphelocheiridae 291 Aphelosigara 337 Aphidoidea 11 apureensis, Gelastocoris 445 Aquarius 88, 89 Aradidae 20 Aradomorpha 18, 20 araucanica, Acanthia 585 araucanica, Pentacora 585 araucanica, Saldula 585 arca, Microvelia 239 Arctocorisa 336, 338, 341 Arctocorixa 326, 338, 339, 341, 343, 344, 346, 349 arcuatus, Rhagovelia 161, 162 argemii, Horvathinia 403 argentata, Microvelia 246, 252 argentina, Acanthia 593 argentina, Hydrometra 265 argentina, Neoplea 513, 514 argentina, Plea 513 argentina, Salda 593 argentina, Saldula 593 argentiniensis, Sigara 345, 346 argentiniensis, Sigara 338, 346 aristera, Neosigara 351, 354 ashlocki, Microvelia 158, 241 asiatica, Zaitha 422 asiaticum, Belostoma 422 asiaticum, Belostoma 413, 420 asper, Limnocoris 552, 553 astralis, Stridulivelia 216 astralis, Velia 216 ater, Mononyx 454 ater, Nerthra 453, 454 atra, Paravelia 225, 226 atra, Velia 225 atrekes, Eurygerris 149, 150 attenuata, Ranatra 378 attenuatus, Ambrysus 531 aurea, Pseudosaldula 590, 591

659

aurivilliana, Zaitha 410 aurivillianum, Belostoma 410 australis, Velia 237 ayacuchana, Microvelia 244 ayacucho, Stridulivelia 215 aymarana, Limnocoris 564 azulita, Rhagovelia 194, 195 bachmanni, Belostoma 426 bachmanni, Curicta 365 bachmanni, Limnocoris 549, 559 Bacillometra 255, 256 Bacillometra 266, 267 Bacillometroides 255, 257, 266, 267 baconi, Rhagovelia 198 bahiensis, Sigara 344 bakeri, Brachymetra 85 bakeri, Charmatometra 77, 85 balia, Tenagobia 295 Baliagobia 295, 296 banksii, Ochterus 463 barberi, Ochterus 287, 463 barozzii, Cryphocricos 516, 537 barozzii, Cryphocricos 538 basalis, Paravelia 221, 222 basalis, Velia 221 bassleri, Cylindrostethus 120 beckeri, Carvalhoiella, 515, 532 beckeri, Curicta 363 beebei, Trichocorixa 326, 326 beieri, Eurygerris 149, 150 beieri, Gerris 149 bella, Ovatametra 109 Belostoma 39, 45,46, 48, 52-54, 63, 66, 285, 395, 404-434, 435 Belostoma 398, 400, 402, 407-409, 413, 414, 419, 420, 423, 429-431, 434, 435 Belostomatidae 14, 15, 29, 38, 39, 42, 47, 48, 53, 55, 56, 60, 62, 64, 284, 393 Belostomatinae 393 bentoi, Martarega 491, 493

berezowskii, Potamometra 74 bergi, Ambrysus 528 bergi, Belostoma 412, 413 bergi, Belostoma 413 bergi, Limnocoris 560 bergi, Oreokora 588 bergi, Pseudosaldula 587, 588 bergi, Salda 588 bergi, Zaitha 413 bergrothi, Limnocoris 562 bergrothi, Rheumatobates 99 biae, Paravelia 220, 229 bicavum, Belostoma 426, 431 bicirca, Notonecta 465, 497 bicircoidea, Notonecta 465, 497 bicolor, Paravelia 221 bicolor, Salda 597 bidentatus, Ochterus 462, 463 bidentatus, Potamobates 122 bifasciata, Notonecta 496, 497, 503 bifasciata, Notonecta 502 bifidus, Ambrysus 530 bifoveolata, Zaitha 409 bifoveolatum, Belostoma 39, 43, 409 bila, Mesovelia 280 bilobata, Curicta 359, 360 bilobatus, Cylindrostethus 118, 119 binotulatus, Naucoris 571 binotulatus, Pelocoris 517, 571, 572 binotulatus, Pelocoris binotulatus 572 bipunctata, Notonecta polystolisma var. 503 bipunctatus, Mononyx 456 bipunctulus, Pelocoris 290, 573 birabeni, Limnocoris 545, 549 bisignata, Mesovelia 279 bisignata, Mesovelia mulsanti 279 bisignata, Rhagovelia 177 blackburni, Arctocorixa 326 blackburni, Corixa 326

660

blackburni, Trichocorixa 326 bocaina, Rhagovelia 186, 207 boliviana, Paravelia 224 boliviana, Rhagovelia 208 bolivianus, Gelastocoris 445, 446 boliviensis, Heterocorixa 319 boliviensis, Sigara 343 boliviensis, Trepobatoides 80 bollei, Horvathinia 403 bonaerensis, Curicta 284, 361 bonäerensis, Helotenthes 361 bonaerensis, Nepa 361 bonariensis, Rheumatobates 94, 95 bonnairei, Aepophilus 578 boops, Zaitha 413, 420, 422 Borborocoris 538, 563 bordoni, Belostoma 415, 416 borealis, Microvelia 241 borealis, Nerthra 456 borellii, Curicta 283, 365 borellii, Curicta 362 borellii, Limnocoris 558 borellii, Neoplea 509, 510 boscii, Belostoma 420, 423 bosqi, Belostoma 411, 413 bosqi, Belostoma 423 bourquini, Ambrysus 529 bouvieri, Limnocoris 539 brachialis, Paravelia 237 brachialis, Platyvelia 157, 237 brachialis, Velia 237 Brachymetra 38, 50, 84, 85, 131, 132-134, 135 Brachymetra 85, 134, 135 brachypala, Arctocorixa chrostowskii var. 349 brachypala, Sigara 349 brasilana, Hydrometra 261 brasiliensis, Curicta 365 brasiliensis, Belostoma 414 brasiliensis, Ctenipocoris 568, 569 brasiliensis, Guzmania 50, 236 brasiliensis, Heleocoris 569

brasiliensis, Heterocorixa 312 brasiliensis, Limnocoris 560, 561 brasiliensis, Martarega 492 brasiliensis, Paravelia 221 brasiliensis, Ranatra 370, 390, 391 brasiliensis, Saldula 593 brasiliensis, Telmatotrephes 369 brauni, Limnocoris 542, 543 braziliensis, Enithares 466, 503 braziliensis, Heterocorixa 44, 311, 312 braziliensis, Microvelia 245 braziliensis, Sigara 348 breddini, Cryphocricos 537 breddini, Cryphocricus 537 brevicauda, Ranatra 373, 374 bridarolli, Montandonius 440 bridarollii, Gelastocoris 438, 440 bruchi, Lethocerus 399, 400 bruchi, Limnocoris 539, 540 bruesi, Pentacora 589 bruesi, Pseudosaldula 589 brunnea, Velia 224 brunneus, Cryptobatoides 81, 82 brunneus, Ochterus 461 bucayana, Oreokora, 589 bucayana, Pentacora, 589 Buenoa 30, 40, 464, 466, 467-474, 475, 476-488 Buenoa 282,284 buenoi, Nerthra 450 bullialata, Paravelia 234, 235 burmeisteri, Limnocoris 549, 559 caboti, Oiosalda 580, 581 cachoeirinhensis, Belostoma 419 calcaris, Rhagovelia 207 cali, Rhagovelia 164, 182 calii, Limnocoris 547, 548 calilegua, Ambrysus 527 Callicorixa 351 callida, Rhagovelia 172, 173 calopa, Rhagovelia 171, 172 camposi, Amorgius 397

661

camposi, Hebrus 271 camposi, Lethocerus 397 camposi, Ranatra 389 campula, Tenagobia 308 candidulum, Belostoma 430, 431 capillata, Paravelia 228 capillata, Velia 227, 228 capitata, Microvelia 248 caraceae, Limnocoris 555 caraiba, Gerris 261 caraiba, Hydrometra 69, 261 caraiba, Limnobates 261 caraiba, Mesovelia mulsanti 279 carajaensis, Belostoma 431 carapachay, Tenagobia 306, 308 carbonaria, Zaitha 407, 420 carcharus, Limnocoris 546 carinata, Buenoa antigone 475 carinata, Curicta 363, 364 carinata, Curicta 365 carniventris, Eurygerris 150 carniventris, Gerris 150 carvalhoi, Potamobates 124, 125 carvalhoi, Ranatra 381 carvalhoi, Rheumatobates 97 carvalhoi, Telmatotrephes 368, 369 Carvalhoiella 515, 519, 532 castanea, Rhagovelia 204 castanea, Tenagobia 296 castilloi, Horvathinia 403 Cataractocoris, 518 533 cauca, Rhagovelia 188, 190, 192 cavernula, Microvelia 50, 248 celeris, Limnogonus 139, 142, 143 celeris, Neogerris 141, 142, 143 celeris, Neogerris 141-143 celocis, Tachygerris 138 celocis, Tachygonus 138 celocis, Tenagogonus 138 Centrocorisa 321, 323, 336, 337 Ceratocombidae 19 Cercopoidea 11 chagasi, Ranatra 390

championiana, Hydrometra 261 chapadiensis, Heterocorixa 318, 319 chapadiensis, Heterocorixa Hesperia var. 318 Charmatometra 77, 85 Charmatometrinae 77, 131 Charmometra 85 chartoscirtoides, Saldula 594, 595 Cheirochelinae 514 Chepuvelia 71, 72 chichinal, Sigara argentiniensis 338 chilena, Microvelia 253 chilensis, Acanthia 585 chilensis, Aquarius 88, 89 chilensis, Gerris 88 chilensis, Hydrometra 265 chilensis, Limnometra 265 chilensis, Limnotrechus 88 chilensis, Oreokora 586 chilensis, Pseudosaldula 583, 585 Chiloxanthinae 579 chinai, Martarega 489 chipetae, Saldula 597 chrostowskii, Arctocorixa 349 chrostowskii, Sigara 349, 350 chrysocastanis, Halobatopsis 112 Cicadelloidea 11 Cicadoidea 11 Cimex 597 Cimicomorpha 21, 22 cinctipes, Stridulivelia 70, 211, 212 cinctipes, Velia 211, 212 citata, Rhagovelia 189 citrinus, Poicilocapsus 22 clanis, Rheumatobates 94 cobbeni, Pseudosaldula 587, 588 Coccoidea 11 cochisiana, Notonecta unifasciata 495, 497 cognata, Paravelia 227 cognata, Velia 227 cognata, Velia capillata var. 227

662

Coleoptera, 22, 57 Coleopterocoris 574-576 colombiana, Notonecta 495, 496 colombianus, Rhagovelia 161, 163 colombicus, Ambrysus 523 columbiae, Belostoma 421 columbiensis, Neosigara 351, 352, 355, 356 columbiensis, Paravelia 224, 225 columbiensis, Velia 224 comata, Hydrometra 258 comitialis, Trepobates 111 commitialis, Trepobates 111 communis, Buenoa 478, 479 compressicollis, Ranatra 357, 358, 370 conata, Paravelia 226 conata, Velia 226 concava, Euvelia 155, 209 concinnus, Hebrus 272, 274 concinnus, Naeogeus 274 confluens, Acanthia pallipes 597 confusa, Notonecta 496, 497 confusa, Paravelia 222, 227 confusa, Rhagovelia 171, 173 confusa, Velia 227 consolidus, Hebrus 272 consolidus, Naeogeus 272 constricta, Tenagobia 28, 303, 304 constricta, Tenagobia 304 Corisa 326, 336, 338, 341 Corisae 336 Corixa 326, 330, 338, 341, 351 Corixidae 15, 27, 29, 30, 42, 46, 282, 292 Corixinae 292, 293, 320 Corixoidea 14, 42, 52, 58 correntina, Paravelia 222, 231 costaiana, Microvelia 253 costalimai, Belostoma 405, 407 costalimai, Ranatra 385 costalimai, Rhagovelia 195, 199 costaricana, Tenagobia 300

coxalis, Acanthia 593 coxalis, Salda 593 coxalis, Saldula 580, 593 crassifemur Rheumatobates 98, 100, 101 crassifemur, Rheumatobates crassifemur 98, 100 crassipes, Anisops 470 crassipes, Buenoa 470 creaseri, Notonecta mexicana 493 crenulatus, Ambrysus 526 cruzi, Ranatra 383 Cryphocricinae 514, 533 Cryphocricos 29, 516, 533-538 Cryphocricus 537 Cryptobatoides 81, 82 Cryptovelia 276, 277 Ctenipocoris 515, 516, 567, 568, 569, 570 cubae, Corisa 336 cubae, Corisae 336 cucullatus, Gelastocoris 442 cucunubensis, Buenoa, 40, 481 culebrana, Rhagovelia 169 cummingsi, Belostoma 48, 419 cunucunumana, Oiovelia 151, 218, 219 cupariana, Paravelia 236 Curicta 283, 284, 356, 357, 359, 360, 364, 365, 366, 367 Curicta 361-365, 367 curracis, Metrobates 102, 103 curracis, Rheumatobates 93 curtafemorata, Ranatra 383 curtum, Belostoma 398 curtus, Lethocerus 398 Cylindrostethinae 76, 117 Cylindrostethus 38, 76, 84, 118-121 Cylindrostethus 121 czakii, Arctocorixa 343 czakii, Sigara 343 daguerrei, Cryphocricos 534 dallasi, Belostoma 423

663

dallasi, Belostoma 420 darpomorza, Trichocorixa mendozana 329 darwini, Ectemnostega 330 Darwinivelia 277, 278, 281 decarloi, Curicta 364, 365 decarloi, Gelastocoris 443, 444 decarloi, Limnocoris 556, 557 decarloi, Temnocephala 48 delectus, Halobatopsis 113 delpontei, Lethocerus 394, 401, 402 delpontei, Lethocerus 398 deminuta, Rhagovelia 173 denseconscripta, Arctocorisa 341 denseconscripta, Arctocorixa 341 denseconscripta, Corixa 341 denseconscripta, Sigara 339, 341 denseconscriptoidea, Arctocorixa 344 denseconscriptoidea, Sigara 344 dentata, Zaitha 417 dentatum, Belostoma 43, 417, 418 denticolle, Belostoma 424, 428 denticulata, Saldula 594 dentiferus, Tachygerris 137 dentipes, Buenoa 484 dentulata, Salda 594 dentulata, Saldula 582, 594 depressus, Platygerris 83 Diaprepocorinae 292 differata, Saldula 597 difficilis, Zaitha 434 diffidentis, Husseyella 254 dilatata, Curicta 364 dilatata, Paravelia 233, 234 dilatata, Zaitha 447 dilatatum, Belostoma 409 dilatus, Lethocerus 401 dimidiata Acanthia, 597 Diplonychus 420 Dipsocoridae 19 Dipsocoromorpha 19 discala, Euvelia 209, 210

discretum, Belostoma 411, 412 distanti, Microvelia 245 disturbata, Notonecta 499 dita, Arctocorixa 349 dita, Sigara 347 349 doellojuradoi, Horvathinia 403 doeringi, Saldula 594 doesburgi, Belostoma 415 doesburgi, Buenoa 478, 481 doesburgi, Curicta 367 doesburgi, Ranatra 379 380 dolichodentata, Ranatra 382 douglasensis, Mesovelia 279 drakei, Belostoma 434 drakei, Rheumatobates 94, 95 dufouri, Belostoma 408 duidaensis, Enithares 503, 504 duidana, Microvelia 246 duolineatus, Tenagogonus 138 duplicatus, Gelastocoris 440 dureti, Curicta 365 Ectemnostega 320, 322, 330 Ectemnostegella 294, 321, 322, 330, 331-336 ecuadorensis, Mononyx amplicollis var. 454 ecuadorensis, Nerthra 453, 454 ecuadorensis, Telmatotrephes 369 ecuadoriensis, Ranatra 370, 377, 378 ecuadoris, Hebrus 275 egbertae, Sigara 349 egregia, Platyvelia 238 egregia, Velia 238 elegans, Belostoma 43, 408, 409 elegans, Rhagovelia 26, 195, 199 elegans, Zaitha 409 elimatus, Hebrus 268, 272 ellipticum, Belostoma 420, 421 elongatum, Belostoma 422, 423 Emydocoris 20 engaeus, Hebrus 274 Enicocephalidae 15, 16

664

Enicocephalomorpha 15 Enicocephalus 16 Enithares 466, 503, 505 Enitharoides 503 Eobates 84, 85, 86 Eotreches 78 Eotrechinae 77, 78 ephydros, Rhagovelia 159, 161 ephydros, Trochopus 161 equatoria, Rhagovelia 203, 208 erythropus, Cylindrostethus 119 erythropus, Hydrometra 119 esakii, Rheumatobates crassifemur 100, 101 espinolai, Limnocoris 24, 553 estevezae, Belostoma 432, 433 Euhemiptera 11 eumorpha, Zaitha 418 Eurygerris 16, 87, 88, 148, 149, 150 Eurygerris 148 Euvelia 155, 209, 210 evidis, Rhagovelia 176 exalla, Hydrometra 262 excavata, Buenoa 485 excella, Hydrometra 262 exogkoma, Limnocoris 550, 551 falloui, Curicta 361 falloui, Nepoidea 361 Fasciagobia 298 fasciata, Buenoa 287, 288 fasciata, Notonecta polystolisma var. 503 fasciata, Tenagobia 297, 298 fazi, Arctocorixa 341 fazi, Notonecta 499, 500 femoralis, Anisops 475 femoralis, Buenoa 473, 475 femoralis, Curicta 365 femoralis, Rhagovelia 197, 198 femoridens, Sigara 342 festae, Rhagovelia 167 fittkaui, Ambrysus 522

fittkaui, Belostoma 405, 406 fittkaui, Buenoa 477 fittkaui, Cryphocricos 537, 538 fittkaui, Limnocoris 548, 549 fittkaui, Limnocoris fittkaui 549 flaviclavus, Ochterus 460 flavidus, Rheumatobates minutus 93 flavolineatus, Eurygerris 87, 148 flavolineatus, Gerris 148 flavomarginata, Paravelia 225, 226 flavomarginata, Velia 225 flavus, Galgulus 440, 445 flavus, Gelastocoris 444, 445 flavus, Gelastocoris 439 flokata, Ranatra 374, 375 foersteri, Ochterus 461 fontanalis, Rhagovelia 170, 171 forciceps, Arctocorisa 338 forciceps, Arctocorixa 338 forciceps, Corisa 338 forciceps, Corixa 338 forciceps, Sigara 338, 339 forciceps, Sigara 340 forsteri, Horvathinia 403 fossatus, Ambrysus 526 fosteri, Darwinivelia 277, 281 foveata, Merragata 269 foveata, Paravelia 230, 231 foveolata, Zaitha 409 foveolatum, Belostoma 414, 415 foveolatum, Belostoma 419 fragosus, Neotrephes 508 franciscanus, Gerris 146, 148 franciscanus, Limnogonus 51, 145, 146, 148 fraternus, Ambrysus 531 fruhstorferi, Hydrometra 260, 261 fuallagana, Bacillometra 267 fuallagana, Bacillometroides 266, 267 fuanucana, Hydrometra 261 fucatus, Ambrysus 527, 528 fugientis, Metrobates 102

665

Fulgoroidea 11 furva, Brachymetra 50, 134 fusca, Ovatametra 110 fusca, Telmatometra 107 Fuscagobia 304, 305 fuscata, Sigara 305 fuscata, Tenagobia 42, 45, 305 fuscinervis, Brachymetra 148 fuscinervis, Eurygerris 16, 148, 149 fuscinervis, Gerris 148 fuscipennis, Anisops 484 fuscipennis, Buenoa 482, 484 fuscipes, Mononyx 454 fuscipes, Nerthra 449, 450 fuscus, Gelastocoris 442 gaigei, Rhagovelia 195, 196 galapagosana, Saldula 596, 597 Galgulus 440, 443, 445 gaucha, Nerthra 447, 452 gauchita, Neoplea 512, 513 Gelastocoridae 25, 27, 30, 52, 62, 286, 435 Gelastocorinae 435, 437 Gelastocoris 32, 64, 284, 435, 436, 437, 438-446 Gelastocoris 436, 439, 440, 442, 444, 445 gemignanii, Ambrysus 526 genikos, Metrobates plaumanni 104 genticus, Limnogonus 142 genticus, Neogerris 142 genupes, Heterocorixa 315, 316 Geocorisa 13, 14 Gerridae 30, 36, 38, 60, 64, 69, 72 Gerrinae 72, 77, 135 Gerris 88, 89, 90, 148 Gerris 88, 134, 146-149, 261 Gerroidea 14 Gerromorpha 14, 17, 23, 25, 27, 29, 34-36, 38, 41, 47, 49, 51, 56, 68 gestroi, Belostoma 411, 413 gigantea, Veloidea 155, 211 glauca, Notonecta 494

globoidea, Neoplea 513 gloriosus, Hebrus 273 gonostyla, Martarega 490 gorgona, Rhagovelia 196, 197 gracilis, Buenoa 477, 478 gracilis, Limnocoris 541 grande, Belostoma 398 grandicollis, Telmatotrephes 358, 369 grandicollum, Belostoma 407 grandis, Belostoma 398 grandis, Curicta 362, 363 grandis, Iliastus 398 grandis, Lethocerus 396, 398 grandis, Lethocerus 398 grandis, Nepa 398 granulosa, Curicta 364, 365 granulosus, Cryphocricos 534, 535 Griffini, Corixa 351 griffini, Neosigara 350, 351 griffini, Sigara 351 gualeguay, Ovatametra 109, 110 Guazuplea 510 guerini, Gerris 146, 148 guerini, Limnogonus 146, 148 guianae, Belostoma 423 guianana, Hydrometra 263 guianana, Rhagovelia 206, 208 guttata, Notonecta polystolisma var. 503 Guzmania 50, 235 halirrhyta, Mesovelia 278, 279 Halobates 38, 49, 51, 60, 75, 114116, 117 Halobates 112, 115, 121 Halobatinae 60, 76, 114 Halobatopsis 38, 82, 83, 112, 113 halophila, Husseyella 154, 254 hamatus, Rheumatobates 98 hambletoni, Mesovelia 278, 279 hambletoni, Microvelia 252 hambletoni, Rhagovelia 165 hammoniorum, Peloridium 13

666

harrisi, Belostoma 417, 419 Hebridae 37, 70, 267 hebroides, Merragata 267, 269 Hebrus 70, 267, 268, 269, 270, 271, 272, 273, 274, 275 Hebrus 271 helenae, Paravelia 233, 236 helenae Velia, 236 Heleocoris 515, 569 Heleocoris 569, 570 Helobates 135 Helotenthes 361 Helotrephidae 41, 42, 55, 290, 504 Hemiptera 11, 12, 53, 61 henryi, Rhagovelia 197 hesione, Neogerris 88, 140, 142 hesperia Heterocorixa 311, 312 Hesperia, Heterocorixa 318 Heterocleptinae 255 Heterocorixa 44, 293, 310-320 Heterocorixa 312, 318 Heterocorixinae 292, 293, 310 Heteroptera 11-15, 22, 23, 27, 30, 31, 33-35, 38, 41, 43-49, 55-65, 67, 68, 551 heydeni, Ranatra 359, 381, 382 hinei, Microvelia 251 hintoni, Heterocorixa 312, 313 hintoni, Paratrephes, 504, 505 hirsutum, Belostoma 406, 407 hodgdeni, Micranthia 595 hodgdeni, Saldula 595 Homoptera 11-13 horvathi, Ambrysus 523 horvathi, Belostoma 430 horvathi, Belostoma 430 horvathi, Pelocoris 573 horvathi, Potamobates 123 horvathi, Ranatra 385 Horvathinia 393, 394, 403 Horvathinia 403 Horvathiniinae 394 hosfordae, Sigara 349

hosfordi, Arctocorixa 349 howardii, Notonecta 500 huallagana, Hydrometra 261 huamachuco, Pseudosaldula 588 huila, Rhagovelia 194 huincamoreni, Notonecta 502 humboldti, Rhagovelia 187, 188 humilis, Acanthia 594 humilis, Micranthia 594 humilis, Saldula 278, 594 hungerfordi, Coleopterocoris 575 hungerfordi, Cylindrostethus 118, 121 hungerfordi, Gelastocoris 438, 442 hungerfordi, Hebrus 274 hungerfordi, Heterocorixa 317 hungerfordi, Martarega 389, 390 hungerfordi, Microvelia 245 hungerfordi, Notonecta 498 hungerfordi, Ochterus 459 hungerfordi, Paravelia 227, 232 hungerfordi, Ranatra 379 hungerfordi, Rhagovelia 180 hungerfordi, Sigara 347, 348 hungerfordi, Tenagobia 304, 306 hungerfordi, Velia 232 Husseyella 154, 158, 254 husseyi, Belostoma 435 husseyi, Hydrometra 265 hyalina, Hydrometra 146 hyalinus, Gerris 146 hyalinus, Limnogonus 87, 145, 146 Hydrocorisa 13, 14 Hydroessa 241 Hydrometra 69, 255, 256, 257-260, 261, 262-265 Hydrometra 119, 146, 261, 262, 265, 266 Hydrometridae 37, 69, 255 Hydrometrinae 255 Hymenobator 99 Hymenoptera 57 Hynesia 92

667

Hypsipteryigdae 19 iberica, Pentacora 580 icolana, Rupisalda 593 icolana, Saldula 593 ida, Buenoa 468, 475 ignotus, Limnogonus 144 Iliastus 398 illiesi, Limnocoris 539, 541 illiesi, Limnocoris 558 imitator, Hymenobator 99 imitator, Rheumatobates 73, 99 imitatrix, Rhagovelia 168, 169 immaculata, Notonecta polystolisma var. 503 impavidum, Belostoma 420 impensa, Rhagovelia 187, 189, 192 impicticollis, Pelocoris 573 inannana, Microvelia 250 inca, Notonecta 501 incasiana, Gelastocoris 440 incerta, Microvelia pulchella 241 incerta, Tenagobia 309 Incertagobia 307-309 incisa, Brachymetra albinerva 135 incompta, Buenoa 486 inconstans, Salda 597 indentata, Telmatometra 105, 106 indica, Notonecta 500 indica, Notonecta 496 innova, Paralosalda 580, 581 insignis, Limnocoris 288, 554 insignis, Microvelia 245 insularis, Rhagovelia 26, 199 intermedia, Curicta 366, 367 intermedia, Curicta 365 intermedia, Nepoidea 367 intermedius, Limnocoris 550, 552 interstitialis, Acanthia 597 interstitialis, Salda 597 interstitialis, Saldula 597 inveruglas, Paravelia 221, 223 inveruglas, Velia 223 ioana, Microvelia 247

irrorata, Corisa 336 isabelae, Microvelia 242, 243 itaguaiensis, Itagunannus 19 Itagunannus 19 itatiaiana, Rhagovelia 184, 186 itatiayana, Paravelia 232 jaczewskii, Heterocorixa 313, 314 jaczewskii, Neotrephes 506, 507 jamesi, Ectemnostegella 332 janeira, Rhagovelia 174 jensenhaarupi, Sigara 346 jensen-haarupi, Sigara 346 jimenezasuai, Lethocerus 396, 399 johnpolhemi, Curicta 361, 362 jubata, Rhagovelia 205 juruana, Paravelia 235, 236 kahli, Eurygerris 16, 148 kahli, Gerris 148 kahli, Paravelia 230 kahli, Velia 230 kalidasa, Eotreches 78 Kallistometra 111 kirkaldyana, Hydrometra 265 kistnerorum, Saldolepta 21, 578 klagei,Rheumatobates 23, 100 kleerekoperi, Coleopterocoris 574, 576 kleopatra, Brachymetra 133, 134 kleopatra, Brachymetra 134 kleopatra, Gerris 134 klugi, Notonecta, 493 koina, Buenoa 468, 469 kolla, Ambrysus 521, 527 kollari, Centrocorisa 323, 336 kollari, Corisa 336 kollarii, Centrocorisa 336 konta, Buenoa 467, 477 kontos, Neogerris 140, 141 Laccocorinae 515 lacunana, Microvelia 158 lacunana, Xiphovelia 158 lacunifera, Merragata 268, 269 lacuniferus, Lipogomphus 268

668

laetus, Metrobates 102 lamprus, Ambrysus 521, 523 lanei, Ranatra 387, 388 lanei, Temnocephala 48 lanemeloi, Limnocoris 556 lanemeloi, Neotrephes 288, 508 largus, Lethocerus 398 lariversi, Belostoma 424, 425 lata, Brachymetra 133, 135 lata, Euvelia 210 lata, Nerthra 454 laticollis, Acanthia 597 laticollis, Saldula 597 latioculata, Tenagobia 303 latus, Mononyx 454 latus, Neotrephes 505 lautus, Pelocoris 290, 573 lenti, Curicta 359, 367 lenti, Horvathinia 403 lenti, Ranatra 370, 392 Leptopodidae 43, 54, 578 Leptopodoidea 14 Leptopodomorpha 13-15, 21, 22, 25, 27, 31, 34, 43, 47, 49, 54, 56, 60, 68, 578 Lethocerinae 53, 393 Lethocerus 33, 48, 53, 285, 393, 394, 395, 396, 397, 398, 399-401 Lethocerus 285, 398, 400, 401 leucothea, Microvelia 239, 240 limaiana, Microvelia 246 limbata, Zaitha 434 limbosa, Acanthia xanthochila 593 limnaeus, Hebrus 274 Limnobates 261 Limnocorinae 516, 538 Limnocoris 24, 30, 48, 64, 288, 517, 538, 539-567 Limnocoris 543, 552, 554, 556, 558-560, 565, 567 Limnogonus 58, 51, 75, 87, 88, 144, 145, 146, 147 Limnogonus 139-143, 146, 147

Limnometra 137, 138, 265 Limnotrechus 88 lindbergi, Merragata 269 linearis, Cylindrostethus 118, 119 linearis, Cylindrostethus 121 lingula, Neoplea 511, 512 Lipogomphus 267, 268 longimanus, Curicta 359, 365 longipes, Microvelia 243 longipes, Rhagovelia 168 longirostrum, Belostoma 407 longisetosus, Rheumatobates 97 longixiphus, Heterocorixa 316 loprettoai, Belostoma 427, 428 lotus, Limnogonus 142 lotus, Neogerris 142, 143 lubricus, Limnogonus 141, 143 lubricus, Neogerris 140, 141, 143 lucida, Rhagovelia 185 luctuosa, Salda 597 lujanana, Microvelia 249 lundbladi, Belostoma 408 lundbladi, Ectemnostegella 331 lundbladi, Heterocorixa 311, 313 lundbladi, Tenagobia 307 Lygaeus 597 lynchi, Saldula 596 macarena, Rhagovelia 190, 191 machrisi, Belostoma 430 machrisi, Buenoa 467, 468 machrisi, Limnocoris 563 machrisi, Ranatra 373, 374 macrocephala, Notonecta 493 macrophthalma, Ranatra 370, 381, 382 macrotrichia, Buenoa 471 Macroveliidae 71 macta, Rhagovelia 186 maculata, Arctocorixa 341 maculata, Corisa 341 maculata, Corixa 341 maculatus, Limnocoris 564

669

maculiceps, Limnocoris 48, 517, 565 maculosa, Neoplea 510, 512 maculosa, Zaitha 434 magister, Pelocoris 568, 571 magna, Ranatra 379, 381 magnus, Neogerris 139 magnus, Neogerris celeris 139 major, Gelastocoris 436, 440 major, Hebrus 70, 271 major, Zaitha mayri forma 418 makrocheira, Orocorixa 324, 325, 353, 354 malkini, Belostoma 416, 418 malkini, Limnocoris 543, 565 malkini, Rhagovelia 191 mallochi, Buenoa 482 manausana, Paravelia 229, 236 manco-capaci, Limnocoris 558 mangrovensis, Rheumatobates 92, 93 manni, Ochterus 463 mansosotoi, Gelastocoris 439 mansosotoi, Limnocoris 558 mansosotoi, Montandonius 439 manzanoae, Potamobates 124 manzanoi, Rhagovelia 182, 183 marginalis, Cimex 597 marginata, Microvelia 249 marginata, Microvelia 241 marginatus, Gerris 89, 90 marginatus, Gerris 146, 148 marginatus, Limnogonus 146 margineguttata, Zaitha 407 mariae, Corixa 326 mariae, Trichocorixa 326 marinus, Trachopus, 161 marinus, Trochopus 161 maritima, Paravelia 237 maritima, Platyvelia 237 maritimus, Trochopus, 161 marmorata, Tenagobia 299, 300

Martarega 30, 42, 286, 488, 489493 martinezi, Belostoma 450 martinezi, Gelastocoris 442 martini, Belostoma 43, 409, 410 Matinus 447 maximus, Lethocerus 397, 398 mayri, Belostoma 400 mayri, Belostoma annulipes var. 400 mayri, Zaitha 418 mayri, Zaitha mayri forma 418 mazzai, Lethocerus 401, 402 mcateei, Martarega 465, 492 mediana, Ranatra 391 medium, Belostoma 398 medius, Lethocerus 53, 398, 399 Megochterus 457 melini, Mononyx 452, 453 melini, Tenagobia 292, 306 Melloiella 520, 522, 523 melloleitaoi, Lethocerus 402 mello-leitãoi, Lethocerus 402 melloleitaoi, Limnocoris 567 membranacea, Martarega 286, 488 mendozana, Microvelia 251 mendozana, Trichocorixa 328, 329 mendozana, Trichocorixa mendozana 328, 329 menkei, Limnocoris 559, 560 mensor, Hydrometra 265 mera, Brachymetra 132 merga, Rhagovelia 153, 199 meridionalis, Mesovelia mulsanti 279 Merragata 267, 268, 269 Merragata 269 Mesovelia 35, 36, 71, 275, 278-280 Mesovelia 279 Mesoveliidae 37, 68, 71, 275 Mesoveloidea 275, 276, 280, 281 metator, Hydrometra 258, 259

670

Metrobates 29, 30, 79, 101, 102, 103 mexicana, Notonecta 493, 494 mexicana, Notonecta mexicana 493 mexicana, Tenagobia 309 meyeri, Horvathinia 403 micans, Halobates 51, 115, 117 micans, Placomerus 568, 569 micantula, Zaitha 434, 435 micantulum, Belostoma 43, 52, 285, 435 Micranthia 594, 595 Micronectinae 30, 292, 294 Microvelia 50, 152, 158, 239, 240, 241-246, 247, 248, 249, 250, 251, 252, 253 Microvelia 158, 241, 243, 245, 248, 249, 251 Microveliinae 15, 152, 239 milicorum, Trichocorixa 321, 328 mimula, Microvelia 251 minima, Microvelia 251 minima, Ovatametra 108 minimus, Rheumatobates 95 minor, Belostoma 404 minuscula, Zaitha 434 minusculum, Belostoma 432, 434 minuta, Heterocorixa 317 minuta, Notonecta 498 minuta, Tenagobia 300 minutus, Limnocoris 559 minutus, Neotrephes 506 minutus, Pelocoris 572 minutus, Rheumatobates 93 minutus, Rheumatobates minutus 93 mixta, Ranatra 370, 376, 377 moderata, Ranatra 380 modesta, Microvelia 243 modesta, Rhagovelia 177, 178 Mononyx 447, 451-454, 456 monrosi, Gelastocoris 446 montana, Ectemnostegella 335 montandoni, Limnocoris 552

montandoni, Nerthra 451 Montandonius 437, 439, 440 montei, Cryphocricos 535, 536 montei, Curicta 359, 367 montei, Ranatra 48, 371 mulfordi, Bacillometra 266 mulfordi, Bacillometroides 266 mulfordi, Hydrometra 266 mulsanti, Mesovelia 35, 36, 279 murilloi, Neosigara 352, 353, 356 mutabilis, Buenoa 480 myersi, Microvelia 251 Naeogeus, 272, 274 naias, Anisops 484 naias, Buenoa 484 nama, Velia 213 Naucoridae 30, 39, 42, 53, 290, 514 Naucorinae 516, 567 Naucoris 448, 456, 571 Naucoroidea 14 53 nebulosus, Galgulus 445 nebulosus, Gelastocoris 444 neivai, Ranatra 376 nelsoni, Coleopterocoris 576 Neogerris 87, 88, 139-143 Neogerris 141-143 Neoplea 289, 509-514 Neosigara 324, 325, 350-356 Neotrephes 288, 504-598 Nepa 356 Nepa 361, 398 nepaeformis, Naucoris 456 nepaeformis, Nerthra 455, 456 Nepidae 29, 39, 42, 53, 283, 356 Nepoidea 14, 52, 53 Nepoidea 361, 363, 365, 367 Nepomorpha 14, 17, 24-27, 29, 31, 33, 34, 38, 43, 47, 49, 52, 53, 55, 68, 282 Nerthra 25, 30, 61, 287, 435, 437, 447-453, 454, 455, 456 Nerthra 453-454 nessimiani, Belostoma 434, 435

671

nexa, Paravelia 230 nexa, Velia 230 nicaeum, Belostoma 425 nieseri, Potamocoris 291, 577 nigra, Heterocorixa 318 nigra, Notonecta 494 nigriculus, Pelocoris binotulatus 517, 571 nigripennis, Centrocorisa 336, 337 nigripennis, Corisa 336 nigripennis, Sigara 336 nigropunctatus, Limnocoris 555, 556 nitida, Buenoa 475, 476 nitida, Carvalhoiella 532, 533 Notonecta 465, 466, 493, 494-503 Notonecta 472, 493, 496-498, 502, 503 Notonectidae 29, 30, 33, 41, 42, 288, 464 Notonectinae 464 Notonectoidea 14, 52, 53 noualhieri, Belostoma 404, 408 novana, Microvelia 244, 254 novana, Rhagovelia 167, 168 Nychina 466 obesa, Ovatametra 108, 109 obscura, Ranatra 377, 378 obscuratus, Ambrysus 529 occulcata, Rhagovelia 170 ocellata, Salda 596 ochraceus, Ambrysus 528 ochraceus, Limnocoris 562 ochroischion, Rhagovelia 167, 171 ochrothoe, Notonecta 496 ochrothoe, Notonecta shooteri var. 496 Ochteridae 17, 27, 52, 286, 457 Ochterus 287, 457, 458-463 Ochterus 457 oculata, Buenoa 485, 486 oculatus, Gelastocoris 32, 440, 443 Ocyochterus 457

Oiosalda 580, 581 Oiovelia 151, 156, 157, 217, 218, 219 olallai, Hydrometra 259, 260 oliveiracesari, Ranatra 370, 384, 387 ollalai, Heterocorixa wrighti 314 Omaniidae 54, 578 opaca, Limnometra 138 opacus, Tachygerris 138, 139 opacus, Tenagogonus 138 oporapa, Rhagovelia 193 orbiculatum, Belostoma 426, 427 oreia, Buenoa 468, 469 Oreokora 586, 588, 589, 591 orinocensis, Trichocorixa 328 orinocoensis, Trichocorixa 327, 328 ornata, Rhagovelia 209 ornatus, Rheumatobates 95, 96 ornitheia, Ranatra 370, 391, 392 Orocorixa 324, 325, 353, 354 osborni, Potamobates 123 osborniana, Paravelia 224 osborniana, Velia 224 Ovatametra 69, 81, 107, 108, 109, 110 ovatulus, Limnocoris 543, 544 oxyura, Zaitha 429 oxyurum, Belostoma 43, 45, 46, 53, 419, 429 oxyurum, Belostoma 430 pacayana, Rhagovelia 179, 180 pacifica, Martarega 488, 489 palea, Rhagovelia 204, 205 pallens, Anisops 473 pallens, Buenoa 470, 473 pallescens, Borborocoris 563 pallescens, Limnocoris 563 pallidum, Belostoma, 407 pallipes, Acanthia 597 pallipes, Anisops 472 pallipes, Buenoa 472

672

pallipes, Lygaeus 597 pallipes, Notonecta 472, 500 pallipes, Salda 597 pallipes, Saldula 43, 597 palmaris, Cylindrostethus 38, 76, 121 panamensis, Microvelia 242 panamensis, Telmatometra 104, 105 panamensis, Trepobates 112 paolettii, Paravelia 234 paraensis, Lethocerus 398 paraguayensis, Curicta 362 paraguayensis, Gelastocoris 445 paralia, Pseudosaldula 588 paralia, Saldula 566 Paralosalda 580, 581 paramo, Neosigara 325, 352 parana, Microvelia 427 paranensis, Buenoa 464, 480 Paraplea 289, 509 Paratrephes 504, 505 Paravelia 30, 50, 64, 156, 157, 219, 220-229, 230, 231-237 Paravelia 221, 237, 238 parilis, Paravelia 230 parilis, Velia 230 partridgei, Ambrysus 525 parva, Telmatometra 106, 107 parvoculum, Belostoma 431 parvula, Nerthra 452, 453, 454 parvula, Ovatametra 108, 109 parvula, Ranatra 385, 386 parvulus, Hebrus 272, 273 parvum, Belostoma 433, 434 parvus, Ochterus 460 parvus, Potamocoris 574, 577 paulana, Rhagovelia 175 paulus, Hebrus 273 pauper, Limnocoris 566, 567 paxilla, Paravelia 156, 227 paxilla, Velia 277 pectoralis, Limnocoris 560, 562

peculiaris, Rheumatobates 92 pelleranoi, Curicta 360, 361 pelleranoi, Horvathinia 403 pellucidus, Cimex 597 Pelocoris 290, 517, 568, 569, 571573 Pelocoris 290, 572, 573 pelocoroides, Horvathinia 393, 403 Pelogonus 457, 458 Peloridium 13 Peltopterus 447 penai, Pseudosaldula 584 penningtoni, Saldula 597 Pentacora 579, 580 Pentacora 580, 585, 586, 589, 591 Pentatomomorpha 18, 20 perbosci, Ochterus 458 perbosci, Pelogonus 458 perfidiosa, Rhagovelia 200, 202 perija, Rhagovelia 191, 192 perlexus, Procryphocricos 533 perseus, Gerris 148 peruana, Ectemnostegella 333, 334 peruana, Tenagobia 296, 300 peruensis, Gelastocoris andinus 436, 440 perula, Oreokora 591 perula, Pentacora 591 perula, Pseudosaldula 591 peruviana, Curicta 360, 362 peruviana, Mesoveloidea 280 peruviana, Micranthia 595 peruviana, Mononyx 451 peruviana, Nerthra 451 peruviana, Notonecta 502 peruviana, Saldula 595 peruvianum, Belostoma 418 peruvianus, Ambrysus 526 peruvianus, Cryphocricos 536, 537 peruvianus, Limnocoris 560 peruvianus, Potamobates 84, 126 peruviensis, Microvelia 247 Phintius 447

673

Picrops 519, 522 pillaona, Oreokora 589 pillaona, Pentacora 589 pilloana, Oreokora 589 pilosa, Pseudosaldula 586 pilosafrons, Ectemnostegella 334, 335 pilosidorsus, Hebrus 269, 270 placita, Hydrometra 264 Placomerus 568, 569 planum, Belostoma 414 planus, Ambrysus 524 planus, Ambrysus planus 524 platensis, Halobates 112 platensis, Halobatopsis 38, 112, 113 platensis, Paravelia 231 platensis, Sigara 338, 340, 344 platensis, Velia 231 platycnemis, Anisops 482 platycnemis, Buenoa 481, 482 Platygerris 83 Platyvelia 157, 158, 237, 238 plaumanni, Coleopterocoris 575, 576 plaumanni, Ctenipocoris 570 plaumanni, Hebrus 270, 271 plaumanni, Heleocoris 570 plaumanni, Limnocoris 567 plaumanni, Metrobates 103 plaumanni, Metrobates plaumanni 103 plaumanni, Neotrephes 507, 508 plaumanni, Rhagovelia 202 plax, Ambrysus planus 524 Plea 289, 511, 512, 513 plebeja, Zaitha 434 plebejum, Belostoma 43, 433, 434 Pleidae 41, 42. 55, 286, 290, 509 plumbea, Rhagovelia 161, 162 plumbea, Trochopus 161 plumbeus, Taochopus 161 plumbeus, Trochopus 161

podargus, Cylindrostethus 121 poeyi, Pelocoris 570, 571 Poicilocapsus 22 polhemi, Darwinivelia 281 politus, Pelocoris 572, 573 polystolisma, Notonecta 501, 503 porphyros, Limnocoris 48, 553, 554 porteri, Belostoma 395, 417 potama, Microvelia 254 Potamobates 38, 84, 121, 122-130 Potamocoridae 291, 573 Potamocoris 291, 573, 574, 577 Potamometra 73, 74 praecipua, Nerthra 61, 437, 448 priscus, Hebrus 273 problematicus, Gelastocoris 442 probolicornis, Rheumatobates 91, 95 Procryphocricos 553 procurrens, Pelocoris 571, 572 profugus, Limnogonus 144, 146, 147 profundus, Limnocoris 558 prostatus, Rheumatobates 96 pseudoguianae, Belostoma 421 pseudoromani, Tenagobia 301, 302, 304 Pseudosaldula 581, 582, 583-591 Pseudosaldula 586, 587, 588 Psylloidea 11 Ptilomerinae 73 pudoris, Microvelia 249 puella, Paraplea 289, 509 pulchella, Hydroessa 241 pulchella, Microvelia 241 pulchella, Microvelia pulchella 241 Pulchella, Velia (Microvelia) 241 pulchellus, Limnocoris 543, 565 pulchra, Notonecta 497, 499 pulchra, Tenagobia 295, 298 pusillus, Limnocoris 540, 558 pygmaea, Corisa 326 pygmaea, Trichocorixa 326

674

pygmeum, Belostoma 405, 425 quadrata, Corixa 330 quadrata, Ectemnostega 322, 330 quadrilineata, Limnometra 137 quadrilineatus, Tachygerris 137, 138 quadrilineatus, Tenagogonus 137 quadrimaculatus, Galgulus 443 quadrimaculatus, Gelastocoris 441, 443 quadrimaculatus, Gelastocoris 445 quadrispina, Hydrometra 257, 265 quadrispinosa, Stridulivelia 214, 217 quadrispinosa, Velia 217 quechua, Ectemnostegella 334, 335, 336 quieta, Merragata 269 quieta, Microvelia 252, 253 quinquedentata, Nerthra 454, 455 rabida, Ranatra 384, 385 Ramphocorixa 324 Ranatra 30, 41, 42, 48, 357, 358, 359, 370-377, 378, 379-392 Ranatra 378, 380, 381, 389, 391 Ranatridae 283 Ranatrinae, 30 ranina, Nerthra 25, 287, 452, 453 raptoria, Nerthra 449, 450 raspa, Stridulivelia 217 raspa, Velia 217 recens, Paravelia 50, 235, 236 recifana, Microvelia 247 recurvus, Limnogonus 146, 147 regilla, Oreokora 286 regilla, Pentacora 586 regulus, Cylindrostethus 121 regulus, Halobates 121 rehi, Sigara 338, 339 reinhardti, Heterocorixa 318 relicta, Rhagovelia 207, 208 reperta, Salda 597 reperta, Saldula 597

reticulata, Corisa 326 reticulata, Trichocorixa 326, 327 retusa, Telmatometra 105, 106 retusum, Belostoma 427, 428 Rhagadotarsinae, 72, 90 Rhagovelia, 26, 36, 38, 50, 151, 152, 153, 159, 160, 161, 162-209 Rhagovelia 26, 166, 171, 173, 180, 195, 199, 200, 202 Rhagoveliinae 151, 159 Rhamphocorixa 324 Rheumatobates 23, 73, 79, 90, 9199, 100, 101 Rheumatobates 93, 94 rhomboides, Weberiella 394 Rhynchota 11 Rhyngota 11 ribeiroi, Belostoma 407, 408 riggii, Curicta 365 rioana, Rhagovelia 168 ripicolum, Belostoma 409 rivalis, Limnocoris 562 rivicola, Oiovelia 157, 218 rivulosa, Rhagovelia 179 roberti, Sigara 346, 347 robusta, Microvelia 241 robusta, Ranatra 372 robusta, Rhagovelia 207 robustus, Halobates 116, 117 robustus, Limnocoris 546 robustus, Potamocoris 577 rogeri Acanthia 585 rogeri, Oreokora 586 rogeri, Pentacora 586 rogeri, Pseudosaldula 586 rogeri, Saldula 586 roldani, Rhagovelia 199, 200 romani, Tenagobia 302, 303 Romanogobia 300, 301, 303, 304 rostra, Buenoa 471, 472, 481 rotundanotata, Paravelia 228 rotundanotata, Velia 228 rotundatus, Limnocoris 557

675

rotundocephala, Ramphocorixa 324 rozeboomi, Telmatometroides 80, 81 rozeboomi, Telmatometra 80 rubra, Rhagovelia 201, 202 rubromaculata, Acanthia 580 rubromaculata, Pentacora 580 rubromaculata, Salda 580 rubyae, Sigara 339, 341 rubyi, Arctocorixa 339, 341 rubyi, Sigara 339 rudis, Mononyx fuscipes var. 454 rudis, Nerthra, 453, 454 ruficeps, Belostoma 400 rufus, Cryphocricos 535, 536 rugosa, Naucoris 448 rugosa, Nerthra 447, 448 Rupisalda 582, 583, 592, 583 sabrina, Rhagovelia 177, 178 Salda 580, 588, 589, 593, 594, 597 Saldidae 13, 14, 43, 54, 579 Saldinae 579 Saldolepta 21, 578 Saldula 43, 578, 580, 582, 593, 594, 595, 596, 597 Saldula 585, 586, 588, 589, 592594, 597 salina, Pseudosaldula 585, 586 salina, Rhagovelia 160, 161 salinus, Trochopus 161 saltatoria, Salda 597 saltatorius, Lygaeus 597 salutis, Buenoa 476, 482 sanctulum, Belostoma 427, 429 sanctulum, Belostoma 430 sanmartini, Curicta 365 santiagiensis, Arctocorixa 349 santiagiensis, Sigara 348, 349 saphis, Limnocoris 48, 560 sarmientoi, Ranatra 380 sarpta, Microvelia 247 sattleri, Belostoma 416 sattleri, Limnocoris 556

sattleri, Ranatra 370, 382, 383 Sattleriella 538, 542 saxicola, Pseudosaldula 585, 587 sayagoi, Belostoma 406, 407 sbolos, Rhagovelia 164, 177 schadei, Arctocorixa rubyi var. 341 schadei, Ctenipocoris 17, 570 schadei, Heleocoris 570 schadei, Sigara 341, 342 schadei, Tenagobia 304, 306 Schadeogobia 306 Schizoporidae 19 Schizopteridae 19 schoutendeni, Curicta 365 schreiberi, Tenagobia 297, 298 schroederi, Rheumatobates crassifemur 100, 101 schubarti, Horvathinia 403 schuhi, Enicocephalus 16 schuhi, Veliometra 255 scitula, Rhagovelia 205 scitula, Saldula 595, 596 scolius, Ambrysus 12, 524 sculpticollis, Telmatotrephes 368 scutellaris, Notonecta variabilis var. 500 Scylaecus 447 sectilis, Salda 594 sectilis, Saldula 594 seducta, Sigara 307 segrega, Ranatra 370, 374 selecta, Tenagobia 301, 304 sellaris, Corixa 326 sellaris, Trichocorixa verticalis var. 326 sellata, Notonecta 502, 503 semipicta, Neoplea 512 semipicta, Plea 512 sericeus, Halobates 51, 75, 116, 117 serrana, Buenoa 467, 470 serrata, Tenagobia 308 sessoris, Merragata 268 shawi, Brachymetra 134

676

shooteri, Notonecta 496 shuar, Potamobates 127, 128 Sigara 321, 323, 337, 338-350 Sigara 305, 307-309, 336, 338-340, 344, 346, 349, 351 signata, Sigara 307 signata, Tenagobia 307 Signoreti, Belostoma 398, 400 signoreti, Ranatra 370, 385, 386 Signoretiella 492 similis, Heterocorixa 316, 317 simulans, Sigara 307 sinuata, Rhagovelia 50, 201, 202 siolii, Ambrysus 529 siolii, Limnocoris 542 siolii, Ranatra 388, 390 siolii, Sattleriella 542 siviae, Mononyx melini forma 454, 453 sjostedti, Ranatra 387 slossoni, Merragata 269 sobrina, Sigara socialis var. 307 sobrinus, Halobates 115, 116, 117 socialis, Sigara 307, 308, 309 socialis, Tenagobia 282, 308 sola, Oreokora 588 sola, Saldula 588 spatulata, Notonecta polystolisma var. 503 sphacelata, Acanthia 580 sphacelata, Pentacora 579, 580 sphacelata, Salda 580 spiculus, Hebrus 272, 273 spiculus, Potamobates 126 spinifera, Paravelia 220, 221 spinifera, Tenagobia 306, 307 spinipes, Ctenipocoris 515, 570 spinipes, Heleocoris 570 spiniventris, Halobatopsis 82, 113 spinosa, Rhagovelia 169 spinosus, Rheumatobates 96, 98 spinulatus, Tachygerris 135, 136 splendens, Halobates 114, 115, 117

splendidulus, Ochterus 457 splendidulus, Pelogonus 457 splendoris, Paravelia 228 splendoris, Velia 228 spumicola, Oiovelia 218 stagnalis, Velia 237 stali, Ambrysus 518, 529 stali, Gelastocoris 445 stali, Limnocoris 558 Steinovelia 154, 158, 238 stellata, Microvelia 240 stellata, Microvelia 245 Stemmocrypidae 19 stenoptera, Paravelia 30, 219, 220 sterea, Neosigara 352, 353, 355 Sternorrhyncha 11 stolli, Belostoma 407 stolli, Zaitha 407 stollii, Belostoma 413, 414 stollii, Zaitha, 414, 420 striata. Acanthia 597 stridulata, Ectemnostegella 294, 335 stridulata, Stridulivelia 212, 215 stridulata, Velia 215 Stridulivelia, 30, 70, 153, 156, 211, 212-217 strigosa, Stridulivelia 215, 216 striola, Neoplea 289 striola, Plea 289 stygius, Cylindrostethus 120 stysi, Carvalhoiella 519, 532 subflavus, Pelocoris 572 subinermis, Ranatra 375 submontandoni, Limnocoris 48, 553, 554 subpauper, Limnocoris 566 subspinosa, Zaitha 420 sumaco, Potamobates 38, 122, 130 summersi, Microvelia 250 surinamensis, Heterocorixa 320 surinamensis, Limnocoris fittkaui 549

677

surinamensis, Ochterus aeneifrons 458, 462 surinamensis, Ranatra 381, 389 surinamensis, Tachygerris 136 suspecta, Curicta 363 Syncollis 520, 524, 525, 529 sztolemani, Hydrometra 264 Tachygerris 85, 86, 135, 136-139 Tachygonus 138 tantilla, Rhagovelia 166 Taochopus 161 tarsalis, Buenoa 474, 475 taylori, Kallistometra 111 taylori, Trepobates 111 Telmatometra 81, 104-107 Telmatometra 80, 104, 105 Telmatometroides 80, 81 Telmatotrephes 356, 358, 368, 369 Telmatotrephes 369 Temnocephala 48 Temnocephalidae 48 Tenagobia 28, 30, 42, 45, 282, 292, 294-309 Tenagobia 300, 304, 306, 308 Tenagogonus 86, 137, 138 tenebrosa, Nerthra 456 tenebrosus, Ochterus 462 tenuipes, Rhagovelia 170, 171, 173 tenuistyla, Neoplea 511, 513 termasensis, Sigara 345, 346 Termitaphididae 18, 20 terrestris, Cryptovelia 276, 277 terrestris, Mononyx 458 terrestris, Nerthra 455, 456 tersa, Stridulivelia 153, 213 tersa, Velia 213 testacea, Tenagobia 299 testaceopallidum, Belostoma 48, 406, 407 teutonius, Ambrysus 521, 525 thaumana, Ovatametra 69, 107 thaumana, Rhagovelia 176 thomasi, Hydrometra 264

thomasi, Potamobates 125 tibialis, Buenoa 487 tibialis, Curicta 357, 363 tibialis, Nepoidea 363 tijuca, Rhagovelia 197, 198 timida, Microvelia 152, 254 titschacki, Mononyx 456 torquata, Rhagovelia 180, 182 townsendi, Arctocorixa chrostowskii var. 349 townsendi, Sigara 349 Trachopus 161 traili, Rhagovelia 200, 202 transversa, Stridulivelia 213, 214 transversa, Velia 214 transversus, Neotrephes 506, 507 travassosi, Ranatra 382 trepida, Rhagovelia 185, 186 trepidus, Trepobates 74, 111 Trepobates 74, 82, 111, 112 Trepobates 111 Trepobatinae 73, 101 Trepobatoides 80 triangula, Rhagovelia 183, 184 triangularis, Buenoa 479 trianguloides, Rhagovelia 181, 184 triangulum, Belostoma 431, 432 Trichocorixa, 320, 321, 325-329 Trichocorixa 326, 327 tricuspis, Ambrysus 523 tridentatus, Potamobates 129, 130 trilobatus, Limnocoris 548, 549 trimaculata, Corixa 341 trimaculata, Sigara 340, 341 trinidalis, Rhagovelia 199 trinitatis, Hynesia 92 trinitatis, Rheumatobates 91, 92 trista, Rhagovelia 181, 182 Trochopus 159, 161 tropicalis, Acanthia 597 tropicalis, Salda 297 Tropicoparapamphantus 18 Tropocorixa 337, 349

678

truncata, Tenagobia 298 truncaticollis, Ambrysus 522, 523 truncatus, Lethocerus 285, 400 truxali, Belostoma 407 truxali, Buenoa 486, 487 tuberculifrons, Ranatra 371, 375 tucma, Sigara 337, 338 tumidacephala, Ectemnostegella 332, 333 turmalis, Rhagovelia 174, 175 uhleri, Limnocoris 554 ujhelyii, Telmatometra 105 unca, Brachymetra 131, 132 uncinata, Rhagovelia 198, 199 undulata, Notonecta 496, 500 unguis, Buenoa 482, 484 unicornis, Nerthra 450, 451 unidentata, Ranatra 41, 370, 375 unidentatus, Potamobates 124, 125 unifasciata, Notonecta 495, 496, 497 unifasciata, Notonecta unifasciata 495, 496 uruguayensis, Martarega 492, 493 uruguayensis, Signoretiella 492 usingeri, Ambrysus 520, 522 usingeri, Chepuvelia 71, 72 usingeri, Coleopterocoris 576 usingeri, Emydocoris 20 usingeri, Neotrephes 506 vandamepompanoni, Gelastocoris 284, 443 variabilis, Notonecta 500, 502 variabilis, Potamobates 127, 128 variegatus, Gelastocoris oculatus 443 variegatus, Neotrephes 504, 507 Velia 152 Velia 211-217, 221, 223-232, 236238, 241 Veliidae 15, 29, 30, 36-38, 47, 50, 60, 69, 151, 152 Veliinae 152, 209

Veliometra 255 velocis, Rhagovelia 166 Veloidea 155, 211 venezolana, Curicta 464, 465 venezolana, Veloidea 211 venezuelae, Belostoma 414, 422 venezuelana, Heterocorixa hesperia 311, 312 venezuelana, Rhagovelia 206 ventralis, Acanthia 593 ventralis, Bacillometra 256 ventralis, Rupisalda, 583, 593 ventralis, Salda 593 ventralis, Saldula 593 venturii, Ectemnostegella 322, 332 venustatis, Microvelia 250 verdica, Platyvelia 238 verdica, Rupisalda 592 verdica, Saldula 592 verdica, Velia 238 vereertbruggheni, Notonecta 498, 500 versuta, Rhagovelia 164, 166 verticalis, Corisa 326 verticalis, Corixa 326 verticalis, Trichocorixa 326 verticalis, Trichocorixa verticalis 326 vianai, Cryphocricos 535, 538 vianai, Gelastocoris 445 vianai, Limnocoris 558 vicinus, Gelastocoris 444, 446 victor, Ochterus 457 victor, Ocyochterus 457 victor, Pelogonus 457 vigilis, Metrobates 102, 103 vinnula, Paravelia 238 vinnula, Steinovelia 238 vinnula, Velia 238 virescens, Notonecta 465, 497 virescens, Notonecta 500 virgata, Paravelia 238 virgata, Steinovelia 154, 238

679

virgata, Velia 238 viridifrons, Ochterus 459 viriosa, Rhagovelia 172 visendus, Limnogonus 140, 143 visendus, Neogerris 140, 141, 143 vittata, Brachymetra 85 vittatus, Eobates 85, 86 vivata, Rhagovelia 180, 181 vivatus, Potamobates 124 volxemi, Borborocoris 238 volxemi, Curicta 359, 365, 366 volxemi, Nepoidea 365 vonprahli, 177 vonprahli, Rhagovelia 165, 166 vulgaris, Pseudosaldula 581, 587 vuriloche, Sigara 345, 346 vuriloche, Sigara argentiniensis 346 wagneri, Ranatra 372, 373 wallengreni, Corixa 326 wallengreni, Trichocorixa 326 weberi, Ranatra 375 Weberiella 394 westermanni, Heterocorixa 293, 313 whitei, Rhagovelia 38, 204, 205 whitei, Telmatometra 81, 104 willei, Paravelia 223 willei, Velia 223 williamsi Corixa, 351 williamsi, Heterocorixa 310, 311 williamsi, Hydrometra 263 williamsi, Martarega 491 williamsi, Mesoveloidea 276, 281 williamsi, Nerthra 448, 449 williamsi, Paravelia 229

williamsi, Potamobates 38, 129, 130 williamsi, Ranatra 391 williamsi, Rhagovelia 203 williamsi, Velia 229 willneri, Montandonius 439 willneri, Gelastocoris 436, 439 woytkowskii, Bacillometra 267 woytkowskii, Bacillometroides 257, 267 woytkowskii, Ectemnostegella 331, 333 woytkowskii, Heterocorixa 315, 318 woytkowskii, Limnocoris 541 woytkowskyi, Potamobates 125, 126 wrighti, Heterocorixa 314 wrighti, Heterocorixa wrighti 314 wrighti, Rheumatobates 94 Wuellersdorffi, Halobates 115 xanthochila, Acanthia 593 Xiphovelia 158 yacuivana, Rhagovelia 166 yala, Sigara 323, 343 yanomamo, Rhagovelia 189, 190 yungas, Pseudosaldula 590, 591 Zaitha 407, 409, 410, 413-415, 417, 418, 420, 422, 423, 429, 434, 435 zela, Rhagovelia 179 zelena, Rupisalda 592 zelena, Saldula 592 zelotypus, Zaitha 435 zeteki, Hydrometra 260 zeteki, Mesovelia 71, 280 zeteki, Ranatra 385, 386 zeteki, Rhagovelia 201