This book is an anniversary present from his colleagues from 20 countries and recognition of Michail Josifov’s remarkable contribution to the investigation of Heteroptera. The 32 articles, written by 49 authors, present new data on the origin, zoogeography and species composition of heteropteran fauna from different parts of all continents. Numerous nomenclature and taxonomic decisions are suggested. Various problems of phylogeny, biology, karyosystematics and phylogeorgaphy of different Heteroptera species are treated. This book includes the description of 2 genera and 19 species, 1 genus and 14 species of which are named after Michail Josifov. The book also contains short biographical notes, a complete bibliography of Michail Josifov’s scientific articles, and a list of all taxa described by him or named after him. The articles in this book are not only a wonderful present, they are another step forward in the study of the splendid world of true bugs. They, we hope, will be useful to everybody devoted to this cause.
Advances in Heteroptera Research
In the end of 2007, the prominent Bulgarian entomologist, acknowledged specialist in taxonomy and biology of Heteroptera, our colleague and friend, Michail Josifov, celebrated his 80th birthday. An author of 126 scientific works and more than 50 popular articles and books, he has described 9 new genera and 144 new species of true bugs. In recognition of his contribution to Bulgarian science, he has been awarded the Order of Saints Cyril and Methodius.
Advances in Heteroptera Research Festschrift in Honor of 80th Anniversary of Michail Josifov Editors Snejana Grozeva & Nikolay Simov
Michail Josifov – 80th anniversary 1
Advances in Heteroptera Research Festschrift in Honor of 80 th Anniversary of Michail Josifov
2 P. Beron
Michail Josifov – 80th anniversary 3
ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honor of 80 th Anniversary of Michail Josifov Edited by S. Grozeva & N. Simov
Pensoft Publishers National Museum of Natural History – Sofia Institute of Zoology – Sofia Sofia–Moscow 2008
4 P. Beron
Advances in Heteroptera Research Festschrift in Honor of 80 th Anniversary of Michail Josifov Editors: S. Grozeva & N. Simov
First published 2008 ISBN 978-954-642-311-5 (HB) ISBN 978-954-642-421-1 (e-book) Pensoft Series Faunistica No 82 ISSN 1312-0174
Photo on the cover: Lygaeus creticus LUCAS, 1854 by N. Simov
© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.
Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359-2-870-42-82
[email protected] www.pensoft.net
Printed in Bulgaria, June 2008
Contents 5
Contents Preface .............................................................................................................. 13 P. Beron Michail Josifov - 80th anniversary .................................................................... 15 N. Simov Michail Josifov – Bibliography, described and dedicated taxa ....................... 27 B. Aukema Psallus (Apocremnus) montanus JOSIFOV, 1973, in The Netherlands (Heteroptera, Miridae) .................................................................................... 49 P.-p. Chen & N. Nieser The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) ................................................................ 55 J. Damgaard MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) ............................................................... 65 R.K. Duwal & T. Yasunaga A new species of the pilophorine plant bug genus Pilophorus HAHN from Nepal (Heteroptera, Miridae, Phylinae) ......................................................... 79 D. Forero, H.R. Gil-Santana & P.H. van Doesburg Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) ............................................................................ 85 D.A. Gapon A revision of Leprosoma BAERENSPRUNG, 1859 (Heteroptera: Pentatomidae) ................................................................................................ 105 A. Gogala Survival of the endemic Hemiptera species in Slovenia during the Holocene ........................................................................................................ 121
6 Advances in Heteroptera Research
V.B. Golub A new species of lacebug fromTajikistan, Catoplatus josifovi nov. sp. (Heteroptera, Tingidae) ................................................................................ 129 J. Gorczyca A new species of Euchilofulvius POPPIUS, 1909 from Malaysia (Heteroptera: Miridae: Cylapinae) ................................................................ 133 M. Goula, M. Costas, S. Pagola-Carte, M. Baena, T. López, A. Vázquez, F. Gessé, J. Ribes & E. Ribes On some threatened Heteroptera from the Iberian fauna ........................... 139 J. Grazia & C.F. Schwertner Review of Parachinavia ROCHE (Hemiptera, Pentatomidae, Pentatominae) ................................................................................................ 159 S. Grozeva & N. Simov Cytotaxonomy of two Cremnocephalus species (Heteroptera, Miridae) ....... 171 H. Günther A new tingid species from Southern Spain: Dictyonota michaili nov. sp. (Heteroptera: Tingidae) ................................................................................ 181 I.M. Kerzhner A new species of Pilophorus from the Far East (Heteroptera, Miridae) ....... 185 P. Kment & Z. Jindra Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia ......................................................................................... 189 F.V. Konstantinov Review of Omocoris LINDBERG, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus WAGNER 1961 (Heteroptera: Miridae: Phylinae) ......................................................................................... 215 R.E. Linnavuori & E. Heiss A new subspecies of Lygaeidae from northern Iran: Raglius alboacuminatus josifovi nov. ssp. (Heteroptera, Lygaeidae) ........................... 231 P. Moulet Alary polymorphism and new localities in palaearctic Oncocephalus KLUG, 1830 (Heteroptera, Reduviidae, Stenopodainae) ............................... 235
Contents 7
N. Nieser, P.-p. Chen & P. Leksawasdi The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification and five new records for Thailand ........... 241 J.T. Polhemus & D.A. Polhemus A new genus of Microveliinae from the Austral Islands, French Polynesia (Heteroptera, Veliidae) .................................................................. 293 Lj. Proti Contribution to the knowledge of the Isometopinae (Heteroptera, Miridae) of the Balkan Peninsula.................................................................. 303 W. Rabitsch The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria................................................................... 309 D. Rédei First record of Pinochius CARAYON, 1949 from the Oriental Region, with description of a new species from Vietnam (Heteroptera: Schizopteridae)....................................................................... 327 M.D. Schwartz, C.W. Schaefer & J.D. Lattin The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World: Chorosoma josifovi nov. sp. .............. 339 N. Simov A new Loricula species from Bulgaria (Heteroptera, Microphysidae) ......... 349 P. Štys & P. Baa Xenicocephalus - an enigmatic genus of American Enicocephalidae (Heteroptera): a new male-based species from Suriname ............................ 357 M.H. Sweet The Recognition of “Botocudo” ornatulus (BERGROTH, 1895) (Hemiptera, Prosorrhyncha, Rhyparochromidae, Antillocorini) of Australia, the Designation of the Lectotype, and a Redescription of the Type Specimen, with a Discussion of the Nomenclature of Botocudo KIRKALDY.......................................................................................... 377 N.N. Vinokurov Calacanthia josifovi nov. sp., a new species of shore bugs (Heteroptera, Saldidae) from Xizang, China ....................................................................... 385
8 Advances in Heteroptera Research
C. Weirauch Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar ..................................................... 391 T. Yasunaga & R.K. Duwal New species of the mirine plant bug genus Castanopsides YASUNAGA and its assumed sister genus Mahania POPPIUS from Nepal, with a new synonymy of the genus Liocapsus POPPIUS (Heteroptera, Miridae, Mirinae) ................................................................... 403 Appendix: List of new taxa described in this volume........................................................419
Contents 9
Съдържание Предговор .............................................................................................................................. 14 П. БЕРОН Михаил Йосифов на 80 години ....................................................................................... 17 Н. СИМОВ Михаил Йосифов – Библиография, описани таксони и посветени на негово име таксони ........................................................................................................... 43 B. AUKEMA Psallus (Apocremnus) montanus JOSIFOV, 1973 в Холандия (Heteroptera, Miridae) ......................................................................................................... 52 P.-P. CHEN & N. NIESER Семейство Corixidae (s. str.) в Tайланд и описание на нов вид от род Sigara (Heteroptera, Nepomorpha).................................................................................. 62 J. DAMGAARD Многообразие по митохондриална ДНК и филогеография на пет палеарктични вида от сем. Gerridae (Hemiptera-Heteroptera) .......................... 75 R.K. DUWAL & T. YASUNAGA Нов вид хетероптера от род Pilophorus HAHN от Непал (Heteroptera, Miridae, Phylinae) ................................................................................................................ 84 D. FORERO, H.R. GIL-SANTANA & P. VAN DOESBURG Преописание на неотропичния род Aristathlus (Heteroptera, Reduviidae, Harpactorinae) .............................................................................................100 Д.A. ГАПОН Ревизия на род Leprosoma BAERENSPRUNG, 1859 (Heteroptera, Pentatomidae) ....119 A. GOGALA Оцеляването на ендемични видове от разред Hemiptera в Словения през Холоцена......................................................................................................................127
10 Advances in Heteroptera Research
В. ГОЛУБ Нов вид в сем. Tingidae от Таджикистан, Catoplatus josifovi nov. sp. (Heteroptera, Tingidae) .....................................................................................................132 J. GORCZYCA Нов вид в род Euchilofulvius POPPIUS, 1909 от Малайзия (Heteroptera: Miridae: Cylapinae) ............................................................................................................136 M. GOULA, M. COSTAS, S. PAGOLA-CARTE, M. BAENA, T. LÓPEZ, A. VÁZQUEZ, F. GESSÉ, J. RIBES & E. RIBES За някои застрашени видове от разред Heteroptera в иберийската фауна ..........................................................................................................154 J. GRAZIA & C.F. SCHWERTNER Ревизия на род Parachinavia ROCHE (Hemiptera, Pentatomidae, Pentatominae) ......................................................................................................................168 С. ГРОЗЕВА & Н. СИМОВ Цитотаксономия на два вида от род Cremnocephalus (Heteroptera: Miridae) ....................................................................................................... 177 H. GÜNTHER Нов вид от семейство Тingidае от Южна Испания: Dictyonota michaili nov. sp. (Heteroptera: Tingidae)......................................................................184 И.M. KEРЖНЕР Нов вид от род Pilophorus от Далечния Изток (Heteroptera, Miridae) ...... 187 P. KMENT & Z. JINDRA Ревизия на семейство Gelastocoridae (Heteroptera: Nepomorpha) от Юго-Източна Азия...........................................................................................................209 Ф.В. KOНСТАНТИНОВ Ревизия на род Omocoris LINDBERG, 1930 и описание на нов род за Eurycolpus dimorphus WAGNER, 1961 (Heteroptera: Miridae: Phylinae) ..............229 R.E. LINNAVUORI & E. HEISS Нов подвид от семейство Lygaeidae от Северен Иран: Raglius alboacuminatus josifovi nov. ssp. (Heteroptera, Lygaeidae) .....................................234
Contents 11
P. MOULET Крилен полиморфизъм и нови находища на палеарктичните видове от род Oncocephalus KLUG, 1830 (Heteroptera, Reduviidae, Stenopodainae) ...239 N. NIESER, P.-P. CHEN & P. LEKSAWASDI Семейство Notonectidae (Heteroptera, Nepomorpha) в Тайланд и прилежащите райони, с ключове за определяне, и пет нови съобщения от Тайланд ..........................................................................................................................288 J.T. POLHEMUS & D. A. POLHEMUS Нов род в подсемейство Microveliinae от островите Тубуай, Френска Полинезия (Heteroptera, Veliidae) .................................................................................301 LJ. PROTIĆ Принос към изучаването на подсемейство Isometopinae (Heteroptera, Miridae) на Балканския полуостров ..........................................................................306 W. RABITSCH Времената се менят: движещите сили на съвременните изменения на хетероптерната фауна на Австрия ........................................................................321 D. RÉDEI Първо съобщение за род Pinochius Carayon, 1949 в Ориенталската област, с описание на нов вид от Виетнам (Heteroptera: Schizopteridae) ....336 M.D. SCHWARTZ, C.W. SCHAEFER & J.D. LATTIN Първи вид от род Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) за Новия свят: Chorosoma josifovi nov. sp. ................................347 Н. СИМОВ Нов вид Loricula от България (Heteroptera: Microphysidae) .............................356 P. ŠTYS & P. BAŇAŘ Xenicocephalus – един загадъчен род aмерикански Enicocephalidae (Heteroptera): нов вид, описан по мъжки индивиди от Суринам ...................376 M. SWEET Признаване на вида “Botocudo” ornatulus (BERGROTH, 1895) (Hemiptera, Prosorrhyncha, Rhyparochromidae, Antillocorini) от Австралия, означаване на лектотип, и преописание на типовия екземпляр, обсъждане на номенклатурата на род Botocudo KIRKALDY ................................381
12 Advances in Heteroptera Research
Н.Н. ВИНОКУРОВ Calacanthia josifovi nov. sp., нов вид от сем. Saldidae (Heteroptera, Saldidae) от Тибет, Китай ...........................................................................................389 C. WEIRAUCH Mangabea barbiger, нов вид от трибус Collartidini (Heteroptera: Reduviidae: Emesinae) от Мадагаскар .......................................................................401 T. YASUNAGA & R.K. DUWAL Нови видове мириди от род Castanopsides YASUNAGA и приетия за сестрински нему род Mahania POPPIUS от Непал, както и нов синоним на род Liocapsus POPPIUS (Heteroptera, Miridae, Mirinae) .....................................416 Приложение: Cписък на описаните в този том таксони .........................................................419
Michail Josifov – 80th anniversary 13
Preface The winter of 1943/1944. The culmination of World War II. Sofia has been bombed several times. After another hard bombing, a youngster, still a schoolboy, is walking in the streets of the almost ruined city pulling his sledge behind him. He’s not hurrying to reach the evacuation point. Instead, he’s urgently trying to locate something among the burning ruins. Horrified, he sees that the building he is looking for is also hit. Suddenly, a man walks out of it and asks him in German what he wants. Embarrassed, the youngster hands out a parcel receipt message and to his greatest amazement the man enters the building which is still burning. Shortly after, he comes back and handles a parcel to the boy. That’s how the schoolboy Michail Josifov receives his first book on Heteroptera, Stichel’s “Illustrierte Bestimmungstabellen der Deutschen Wanzen” 1. With the same ardent desire to study and the same dedication to science, he continues his researches and helps others to discover the extremely interesting world of insects, especially of Heteroptera. We, as well as the authors of this 420-page Festschrift and all the colleagues and friends who have contributed to its creation, are very happy to greet Dr. Josifov with his 80th birthday. This book, for sure, would not have been written and published without the 49 authors; without the help and precious comments of the reviewers of the articles included: B. Aukema, P. Baňař, W. Cai, A. Carapezza, G. Cassis, J. Damgaard, D. Forero, E. Guilbert, A. Gogala, V. Golub, M. Goßner, E. Heiss, I. Kerzhner, P. Kment, E. Kondorosy, F. Konstantinov, V. Kuznetsova, R. Linnavuori, A. Matocq, S. Nokkala, J. Péricart, J. Polhemus, Lj. Protić, D. Rédei, D. Rider, C. Schaefer, N. Tatarnik, J. DavidováVilímová, M. Wall, T. Weir, C. Weirauch, T. Yasunaga, H. Zettel; as well as the gracious support of Dr. Lyubomir Penev and PENSOFT Publishers, the assistance of the colleagues from the National Museum of Natural History and the Institute of Zoology, and the complementary financial support of the National Science Fund. We express our deepest gratitude to all of them, and both on ours and on their behalf we wish Dr. Josifov a lot of health, happiness, good luck and new scientific discoveries from the world of Heteroptera in Bulgaria and around the world. Ad multos annos! The editors
1
This is a true story and M. Josifov tells it every now and then as an example of the bookstore owner’s remarkable accuracy
14 P. Beron
Предговор Зимата на 1943/44 година. Втората световна война е в разгара си. София е бомбардирана многократно. След поредната тежка бомбардировка, един младеж, все още ученик, ходи със своята шейна из пустите улици на полуразрушения град. Не бърза да избяга и да се отправи към мястото за евакуация, а настойчиво търси нещо сред горящите развалини. С ужас забелязва, че и търсената от него сграда е поразена. Ненадейно от нея излиза човек и го пита на немски какво иска. Притеснен младежът подава известие за получаване на пратка и много изненадан наблюдава как човекът влиза във все още не загасената сграда. След малко се появява с един пакет и му го подава. Така ученикът Михаил Йосифов получава първата си книга за хетероптери: Stichel’s “Illustrierte Bestimmungstabellen der Deutschen Wanzen”. 1 Със същия младежки плам за знания и отдаденост на науката той продължава да изследва, и помага на други да открият безкрайно интересния свят на насекомите, и в частност хетероптерите. За нас, заедно с авторите на 420-те страници на този юбилеен том и с всички помогнали при създаването му приятели и колеги, е голямо щастие, че можем да го поздравим с неговия 80ти рожден ден. Тази книга, разбира се, не би била факт без ръкописите на 49-тe автори; без помощта и полезните коментари на колегите, които рецензираха статиите, включени в тома: B. Aukema, P. Baňař, W. Cai, A. Carapezza, G. Cassis, J. Damgaard, D. Forero, E. Guilbert, A. Gogala, V. Golub, M. Goßner, E. Heiss, I. Kerzhner, P. Kment, E. Kondorosy, F. Konstantinov, V. Kuznetsova, R. Linnavuori, A. Matocq, S. Nokkala, J. Péricart, J. Polhemus, Lj. Protić, D. Rédei, D. Rider, C. Schaefer, N. Tatarnik, J. DavidováVilímová, M. Wall, T. Weir, C. Weirauch, T. Yasunaga, H. Zettel; както и без безрезервното спомоществувателство от д-р Любомир Пенев и издателство ПЕНСОФТ, подкрепата на ръководството и колегите от Националния природонаучен музей и Института по зоология, както и частичната финансова подкрепа на Фонд „Научни изследвания“. На всички тях благодарим, и от тяхно и наше име желаем на д-р Йосифов много здраве, щастие, късмет и нови открития при изучаването на хетероптерите на България и света. За много години! От редакторите
1
Историята е истинска и много рядко е разказвана от М. Йосифов като пример за акуратността на собственика на книжарницата.
Michail Josifov – 80th anniversary 15
Michail Josifov - 80th anniversary P. Beron I haven’t even realized that a whole decade has somehow slipped by since I last wrote a piece about the most prominent contemporary Bulgarian entomologist, in Historia Naturalis Bulgarica, N 8. It’s a pleasure to see that our scientist, colleague and friend Michail Josifov is still full of energy and keeps working. His 80th anniversary gives us a good reason to express our respect to the person whom we’d still ask for advice or with whom we’d still have a friendly chat. Let us remember of the special evenings at academic Ivan Buresch’s house. Let us remember the words that he, the patriarch of Bulgarian zoology, has said: “True, among all the young keen zoologists around, Josifov is the most distinguished one”. Those evenings, we were drinking every single word of his stories about the “good old chieftains” of our science. Michail Vladimirov Josifov was born in Sofia on 14 November 1927. His interest in insects - first in butterflies and later in true bugs - dates back in his school years. He graduated Biology at the Sofia University in 1950. He obtained a PhD in 1964. In 1981 he became Doctor of Science, and in 1983 - senior researcher. A constantly ascending path towards the peaks of the world entomology. From all the publications of Josifov’s, which have long ago outnumbered one hundred, only a couple have not been dedicated to his favourite true bugs (Heteroptera). Yet in the distant 1950, G. Dechev and he contributed to knowing more about the winter life in the Rila lakes. And the overview on Bulgarian bats that he published in 1958 together with V. Hanák is still one of the fundamental works of Bulgarian chiropterologists. 1954 was a year of several rather important events in M. Josifov’s life. He miraculously survived after finding himself trapped under an avalanche in Rila mountain on December the 9th. And on that same day was he officially hired at the Institute of Zoology. In the same year, the first of his numerous works dedicated to Heteroptera was published. Nowadays he is a world-wide acknowledged authority on everything concerning the thousands of Palearctic taxa of this large and significant insects group. His concepts are based on vast knowledge as well as on a collection, perfectly maintained and complied for years. It’s rather doubtful that he himself would be able to list all the countries in which he has described new taxa or where he has contributed to the thorough studying of the local fauna. But we should definitely mention North Korea to which he dedicated a dozen of field trips and many years of study. I was lucky to join one of these trips and I got convinced that on field M. Josifov was just the same as he was in the lab – dedicated, systematic and composed. The other trip we have ever undertaken together was to the
16 P. Beron
Mozambique savannas – one of his few adventures in the tropics. And in the end, let’s not forget his expeditions in Central Asia and Vietnam. Actually, my reason to visit him in 1958 and introduce myself as a student interested in bats, was exactly his research on bats. The gentleman who met me was tall and wearing eye-glasses and his manners seemed to be so European, that they were in big contrast with the general post-war environment of the Institute of Zoology. Later on, I realized that Josifov preferred the silence of study and the diligent and conscious labour to any group meetings and pseudo-public activity that were so popular at that time. He was never tempted to get involved in politics or in the superstitious academic vanity. Being a prominent and respected zoologist, however, he could not avoid taking part in scientific councils, nor he could escape all the administrative work. Until 1992, he was Head of the Department of Taxonomy, Faunology and Zoogeography at the Institute of Zoology. In 1986 M. Josifov was awarded the order of St. Cyrill & Methodius for his contribution to Bulgarian science. In 1964, Juli Stefanov, untimely died paleontologist, took the initiative to provide an adequate translation of the International Code of Zoological Nomenclature in Bulgarian. The first to be involved in this project from all Bulgarian zoologists was Josifov, who had already been recognized as the best Bulgarian expert in this field. Even today anyone who feels uncertain about something in this specific area would address him, and the Faculty of Biology in Sofia University has invited him for lecturer on Taxonomy, Faunistics and Zoological Nomenclature. Besides the dozens of precisely written and elaborately illustrated taxonomic works, Josifov has also published interesting analyses on the origins of Balkan and European Heteroptera. He is one of the authors of the Catalogue of the Heteroptera of the Palaearctic Region. And we should not neglect all his remarkable books and articles addressed to the general public. The increased diopter of his eye-glasses seems to be the biggest reward he has received for his hard work. Yet, there is another reward, too. Josifov has gained the most valuable thing a scientist could hope for – the respect and recognition of his fellow colleagues, and their warm feelings, as shared in this book and expressed on the day of his 80th Anniversary. He has just entered the ninth decade of his life – I wonder how many other valuable works we’d have by 14 November 2017! We’ll still need taxonomists at that time, for sure... May he be healthy and happy for many more years!
Michail Josifov – 80th anniversary 17
Михаил Йосифов на 80 години П Берон Някак неусетно се изтъркаля едно десетилетие, откакто писах за най-видния съвременен български ентомолог в кн. 8 на Historia naturalis bulgarica. С удоволствие виждаме, че нашият учен, колега и приятел Михаил Йосифов е все така жизнен и продължава да работи. За нас настоящата 80 годишнина е повод да изкажем уважението си към човека, при когото и сега се отбиваме за съвет и добронамерен разговор. Да си припомним празничните вечери у академик Иван Буреш и думите на патриарха на българската зоология: „Да, измежду новите зоолози с мерак, най-изтъкнат е Йосифов“. Вечерите, когато ние попивахме с интерес разказите за „вехтите войводи“ в нашата наука. Михаил Владимиров Йосифов е роден в София на 14 ноември 1927 г. Интерес към насекомите проявява още от ученическите години, първо към пеперудите и после към хетероптерите. Завършил е биология в Софийския университет през 1950 г., получил е к.б.н. през 1964, доктор на биологическите науки (1981), старши научен сътрудник I ст. (1983). Един път към върховете на световната ентомология. Само две от публикациите на Йосифов, които отдавна са надхвърлили стотицата, не са посветени на любимите му дървеници (хетероптери). Още в далечната 1950 г. заедно с Г. Дечев той допринася за опознаването на зимния живот в рилските езера. А прегледът на българските прилепи, публикуван през 1958 заедно с V. Hanáк, и сега е сред основните трудове на българските хироптеролози. През 1954 г. се случват някои съдбоносни за М. Йосифов неща. На 9 декември оцелява по чудо, затрупан от лавина на Рила, и в този ден го назначават на работа в Зоологическия институт. През същата година излиза първата от многобройните му работи върху разред Heteroptera и сега той е световно признат авторитет по хилядите палеарктични представители на тази голяма и практически важна група насекоми. Преценките му се основават както на обширни познания, така и на събраната през десетилетията огромна и образцово поддържана колекция. Надали той сам може да изброи страните, откъдето е описал нови таксони и е допринесъл за изучаването на фауната им. Не можем обаче да пропуснем Северна Корея, на която са посветени десетина пътувания и многогодишни изследвания. Имах възможността да бъда на едно от тези пътувания и се убедих, че и на терена Йосифов е такъв, какъвто е и в лабораторията – прецизен, системен и спокоен. Другото ни съвместно пътуване беше в саваните на Мозамбик – една от малкото му авантюри в тропиците. Да не забравяме и експедициите му в Средна Азия и Виетнам.
18 P. Beron
Всъщност, прилепарският му труд беше и поводът да му се представя през 1958 г. като студент, който се интересува от прилепите. Посрещна ме висок човек с очила с подчертано европейски обноски, който силно контрастираше на общия следвоенен фон на Зоологическия институт. После установих, че пред модните тогава събрания и псевдообществена дейност Йосифов предпочита тишината на кабинета и упорития, честен труд. Никога не остави да го забъркат в политиката или във висшата академична суета. Като изявен и авторитетен зоолог обаче той не можеше да отбегне нито участие в научни съвети, нито административната работа. До 1992 г. той беше ръководител на секция Таксономия, фаунистика и зоогеография в Института по зоология. През 1986 г. М. Йосифов е удостоен с орден „Св. Св. Кирил и Методий“ II-ра степен за заслуги в развитието на българската наука. През 1964 г. рано загиналият палеонтолог Юли Стефанов поде инициативата да се направи качествен български превод на Международния кодекс на зоологичната номенклатура. От зоолозите пръв беше привлечен да сътрудничи на това дело Йосифов, който вече се беше утвърдил като най-добрият специалист в тази област в България. И днес всеки, който среща затруднения в тази специфична материя, се обръща към него, а ръководството на Биологическия факултет на Софийския университет го покани да чете курса по Таксономия, фаунистика и зоологическа номенклатура. Освен десетките прецизни, прекрасно илюстрирани с отлично изработени рисунки, таксономични работи, Йосифов публикува и интересни анализи за произхода на балканската и европейската хетероптерна фауна. Той е и един от авторите на Каталога на палеарктичните хетероптери. Не можем да отминем и прекрасните му научнопопулярни книги и статии. Увеличените диоптри са главното, което получи в награда за този труд. Има обаче и друга награда. М. Йосифов получи и получава най-ценното, на което един учен може да се надява – уважението и признанието от страна на колегите си, а и техните топли чувства, изразени на вълнуващия празник за 80 годишнината му, и в този том. Е, тече деветото му десетилетие – колко ли ценни трудове ще видим на 14 ноември 2017 годинa! И тогава ще са нужни таксономи... Да ни е жив и здрав още много години!
Michail Josifov – 80th anniversary 19
Fig. 1: M. Josifov and acad. I. Buresch in Rila Mts., Bulgaria. Фиг. 1: М. Йосифов заедно с академик И. Буреш в Рила палнина.
Fig. 2: M. Josifov with E. Wagner and H. Eckerlein at Munich Railway Station, 1962. Фиг. 2: М. Йосифов, изпращан на гара Мюнхен от видните европейски хетероптеролози E. Wagner и H. Eckerlein, 1962 г.
20 P. Beron
Fig. 3: M. Josifov in Norway, 1962. Фиг. 3: М. Йосифов в Норвегия през 1962.
Fig. 4: M. Josifov in Tajikistan, above Kondara, 2200 m with Prangos pabularia, a plant several Heteroptera species are trophically associated with. As a result of Dr. Josifov’s 5 expeditions in Middle Asia, a great amount of collected material, and his long-term fruitful collaboration with the hemipterologists from the former Soviet Union I.M Kerzhner, N.N. Muminov etc, a considerable number of new Heteroptera taxa have been described, 1970. Фиг. 4: М. Йосифов в Таджикистан, над Кондара, 2200 м, с растението Prangos pabularia, с което трофично са свързани няколко вида хетероптери. В резултат на петте му посещения в Средна Азия, огромното количество събран материал, и дългогодишното ползотворно сътрудничество с колегите хемиптеролози от бившия Съветски съюз И.М.Кержнер, Н.Н. Муминов и др., са описани от много нови таксони хетероптери, 1970 г.
Michail Josifov – 80th anniversary 21
Fig. 5: M. Josifov collecting true bugs on pistachio above Arslanbob in Kyrgyzstan, 1975. Фиг. 5: М. Йосифов събира хетероптери от храст на шам-фъстък в Киргизия над Арсламбоб, 1972 г.
Fig. 6: M. Josifov in Vietnam – seaside in the vicinity of Saigon (now Ho Chi Minh City), 1975. Фиг. 6: М. Йосифов във Виетнам Крайбрежието в околностите на Сайгон (сега Хо Ши Мин), 1975 г.
22 P. Beron
Fig. 7: M. Josifov in Vietnam – Botanical garden of Saigon (now Ho Chi Minh City), 1975. Фиг. 7: М. Йосифов във Виетнам - Ботаническата градина на Сайгон (сега Хо Ши Мин), 1975 г.
Fig. 8: Members of the expedition of the National Museum of Natural History in Mosambique, 1983: señor Langa, V. Ivanov, P. Beron, M. Josifov, V. Beshkov (from left to right). Фиг. 8: Част от членовете на експедицията на Националния природонаучен музей в Мозамбик, 1983. От ляво на дясно: сеньор Langa, В. Иванов, П. Берон, М. Йосифов, Вл. Бешков.
Michail Josifov – 80th anniversary 23
Fig. 9: M. Josifov in North Korea on an ancient dolmen in the region of Taesˇong, 30 km to the north of P’yˇongyang. In 1970-1990, M. Josifov has visited this closed and isolated country 10 times. Perhaps he is one of the few foreigners who have spent so much time in North Korea visiting different regions. Paradoxically, during the same period M. Josifov was not allowed to visit a neighboring with Bulgaria country due to political reasons. Фиг. 9: М. Йосифов в Северна Корея, седнал върху древен долмен в района на Тесонг, 30 km северно от Пхенян. В периода 1970 – 1990 М. Йосифов посещава 10 пъти тази затворена и изолирана от света страна. Може би е един от малкото чужденци, прекарали толкова време в нея, и то в различни райони. Парадоксалното е, че по политически причини М. Йосифов не получава разрешение да посети съседна на България страна в този период.
Fig. 10: M. Josifov above the glacier where A. Piccard’s high altitude balloon has landed. This is one of his expeditions to the Austrian Alps during his teamwork with E. Heiss. Фиг. 10: М. Йосифов по време на съвместната работа с E. Heiss в Австрийските Алпи на 3000 m, върху ледникът, на който се е спуснал балонът на A. Piccard.
24 P. Beron
Fig. 11: Heteropterologists at a SIEEC symposium in Gödöllö, Hungary, 6th September 1991. From left to right, sitting: M. Goula, J. Pericart, M. Angeles Vasquez, standing: A. Carapezza, N. Nieser, J. Lis, F. Faraci, I.M. Kerzhner, M. Josifov, C. Schaefer and standing behind: J. Gorczyca, E. Heiss, B. Aukema. Фиг. 11: Хетероптеролозите присъствали на XIII Международен симпозиум по ентомофауната на Средна Европа в Гьодьольо, Унгария, 6 септември 1991. От ляво на дясно седнали: M. Goula, J. Pericart, M. Angeles Vasquez, прави отпред: A. Carapezza, N. Nieser, J. Lis, F. Faraci, I.M. Kerzhner, М. Йосифов, C. Schaefer и прави заден ред: J. Gorczyca, E. Heiss, B. Aukema.
Fig. 12: M. Josifov during the preparation of third volume of the Catalogue of the Heteroptera of the Palaearctic Region, Institute of Zoology, Sofia, 1992. Фиг. 12: М Йосифов по време на подготовката на том 3 от Каталога на палеарктичните хетероптери в Института по зоология, София, 1992.
Michail Josifov – 80th anniversary 25
Fig. 13: M. Josifov in the Entomological collection of Institute of Zoology, Sofia, 2005. Фиг. 13: М. Йосифов по време на работа в депото на Ентомологичната колекция на Института по зоология, София, 2005.
26 P. Beron
S. Grozeva &Michail N. Simov (Eds) 2008 Josifov – Bibliography, described and dedicated taxa 27 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 27-47. © Pensoft Publishers Sofia–Moscow
Michail Josifov – Bibliography, described and dedicated taxa* N. Simov National Museum of Natural History, Tzar Osvoboditel 1, 1000 Sofia, Bulgaria. E-mail:
[email protected]
ABSTRACT The paper summarizes the scientific opus of Michail Josifov (126 publications) and lists of described (9 genera and subgenera and 144 species) and dedicated taxa (3 genera and 31 species). Over 50 popular articles are not included in this paper. Keywords: Heteroptera, bibliography, Josifov, Miridae
INTRODUCTION For generations of Bulgarian zoologists and entomologists, Michail Josifov has been a teacher and an example of devotion to science, perfect work, knowledge, ground-breaking ideas and unresting enthusiasm about knowledge and new discoveries. His always helping hand, his advice, and the friendly conversations with Prof. Josifov remain priceless for anyone who knows him. The present work is an attempt at summarizing M. Josifov’s scientific career by including a bibliographic list of his scientific works from 1950 until 2007, a list of the described by him 144 species and 9 genera of bugs, as well as listing all the taxa (3 genera and 31 species) named in his honor. Excluded were the more than 50 popular articles and books. * I dedicate this work to my teacher, friend, colleague and famous scientist M. Josifov in honor of his 80th anniversary and in recognition of his remarkable contribution to the research of Heteroptera. I want to thank him for all his encouragement, good discussions and truly friendship.
28 N. Simov
LIST OF SCIENTIFIC PUBLICATIONS OF MICHAIL JOSIFOV (1)
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ЙОСИФОВ М. & Г. ДЕЧЕВ [ Josifov M. & G. Detschev] (1950). Хидробиологични изследвания на Рилските езера през зимата. [Hydrobiologischen Untersuchungen der Gebirgsseen im Rila-Geb. im Winter.] — Годишник на Софийския университет, Природо-математически факултет [Annuaire de l’Université de Sofia. Faculté de Sciences], 46 (3): 197-199. (In Bulgarian) ЙОСИФОВ М. [ Josifov M.] (1954). Нови и редки за българската фауна хемиптери Hemiptera-Heteroptera. [Neue und seltene für die bulgarische Fauna HemipteraHeteroptera.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 3: 237-242. (In Bulgarian, German summary) Josifov M. (1955). Hemipterologische Mitteilung. — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 4/5: 461-462. ЙОСИФОВ М. [ Josifov M.] (1957a). Ceresa bubalus F. (Hem. Hom. Membracidae) – един нов неприятел на овощните дървета и люцерната у нас [Ceresa bubalus F. (Hem. Hom. Membracidae) – ein neuer Schädling auf Ostbäumen und der Luzerne bei uns.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 6: 569-573. (In Bulgarian, German summary) ЙОСИФОВ М. [ Josifov M.] (1957b). Хемиптерологично съобщение. II. [Hemipterologische Mitteilung II.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 6: 577-579. (In Bulgarian, German summary) Josifov M. (1957c). Forma macroptera von Diomphalus hispidulus Fieb. (Hem. Het.). — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 6: 581-583. Josifov M. (1957d). Forma macroptera von Halosalda lateralis Fall. (Heteroptera, Saldidae). — Acta Entomologica Musei Nationalis Pragae, 31 (470): 69-71. Josifov M. (1957e). Über Eurydema spectabilis Horv. (Heteroptera, Pentatomidae). — Acta Entomologica Musei Nationalis Pragae, 31 (474): 103-108. ЙОСИФОВ М. [ Josifov M.] (1958a). Хемиптерологично съобщение. III. [Hemipterologische Mitteilung III.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 7: 343-347. (In Bulgarian, German summary) Josifov M. (1958b). Eine neue Aradus- Art (Hem. Het.) aus Bulgarien. — Acta Entomologica Musei Nationalis Pragae, 32: 261-263. Josifov M. (1958c). Über Odontotarsus confraginosus Hoberlandt, 1956 (Heteroptera, Pentatomidae). — Acta Entomologica Musei Nationalis Pragae, 32 (502): 265-268. Josifov M. (1958d). Eine neue Gampsocoris- Art aus Bulgarien: G. lilianae n. sp. (Hem. Het. Berytidae). — Acta Entomologica Musei Nationalis Pragae, 32 (503): 269-270. Josifov M. (1958e). Zwei neue Dicyphus (Brachyceraea)- Arten aus Bulgarien (Het.). — Acta Entomologica Musei Nationalis Pragae, 32 (504): 271-274.
Michail Josifov – Bibliography, described and dedicated taxa 29
(14) Josifov M. (1958f ). Heterocordylus cytisi n. sp. – eine neue Miriden-Art aus Bulgarien (Hem. Heteroptera). — Fragmenta Balcanica, 2 (11): 83-89. (15) Hanák V. & M. Josifov (1959). Zur Verbreitung der Fledermäuse Bulgariens. — Säugetierkundliche Mitteilungen, 7 (4): 145-151. (16) Josifov M. (1959a). Eine neue Dichrooscytus- Art aus Bulgarien (Hem. Het. Miridae). — Entomologische Berichten, 19: 63-64. (17) Josifov M. (1959b). Gampsocoris eckerleini n. sp. – eine neue Heteropterenart aus Griechenland (Hem. Het. Neididae). — Fragmenta Balcanica, 2 (18): 155-158. (18) ЙОСИФОВ М. [ Josifov M.] (1959c). Хемиптерологично съобщение. IV. [Hemipterologische Mitteilung IV.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 8: 201-203. (In Bulgarian, German summary) (19) ЙОСИФОВ М. [ Josifov M.] (1959d). Hemiptera – Heteroptera от остров Тасос. [Hemiptera-Heteroptera von der Insel Thasos.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 8: 265-268. (In Bulgarian, German summary) (20) Josifov M. (1959e). Über zwei Miriden-Arten aus dem östlichen Mittelmeerraum. — Acta Entomologica Musei Nationalis Pragae, 33 (554): 359-362. (21) ЙОСИФОВ М. [ Josifov M.] (1960). Видов състав и разпространение на насекомите от разред Heteroptera в България. Част I. [Artenzusammensetzung und Verbreitung der Insekten von der Ordnung Heteroptera in Bulgarien, Teil I.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 9: 107-177. (In Bulgarian, German summary) (22) Josifov M. (1961a). Macrolophus nubilis geranii n. subsp. (Hem. Het. Miridae). — Comptes rendus de l’Académie bulgare des Sciences, 14 (1): 87-89. (23) ЙОСИФОВ М. [ Josifov M.] (1961b). Халобионтните и халофилните хетероптери по Черноморското крaйбрежие. [Die halobionten und halophilen Heteropteren an der Schwarzmeerküste.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 10: 5-34. (In Bulgarian, German summary) (24) Josifov M. (1961c). Eurydema eckerleini n. sp. – eine neue Pentatomidenart von der Insel Kreta — Comptes rendus de l’Académie bulgare des Sciences, 14 (4): 397-400. (25) Josifov M. (1961d). Was ist Megalonotus chiragra var. emarginatus (Rey) 1888 (Heteroptera, Lygaeidae). — Acta Entomologica Musei Nationalis Pragae, 34 (582): 117-119. (26) Josifov M. (1961e). Über drei in Bulgarien anzutreffende Miridenarten (Hemiptera, Heteroptera). — Fragmenta Balcanica, 4 (3): 21-26. (27) Seidenstücker G. & M. Josifov (1961). Auchenodes joakimoffi n. sp., eine neue ostmediterrane Oxycareninae (Heteroptera, Lygaeidae). — Acta Entomologica Musei Nationalis Pragae, 34 (574): 27-32. (28) ЙОСИФОВ М. [ Josifov M.] (1962a). Количествени и качествени проучвания на ентомофауната на люцернови ниви в Софийско с оглед насекомите от разреда Heteroptera. [Quantitative und qualitative Untersuchungen über die Entomofauna der Luzernefelder im Hindblick auf den Insekten von der Ordnung Heteroptera.]
30 N. Simov
(29) (30)
(31) (32) (33)
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(35) (36) (37) (38) (39) (40) (41) (42) (43)
(44)
— Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 11: 117-137. (In Bulgarian, German summary) Josifov M. (1962b). Eine neue Miridenart von der Insel Kreta (Hemiptera, Heteroptera). — Annales historico-naturalis Musei Naturalis Hungarici, 54: 377-379. ЙОСИФОВ М. [ Josifov M.] (1963a). Полукрили насекоми (Heteroptera) от околностите на Петрич (Югозападна България). [Heteropteren aus der Umgebung von Petric (SW Bulgarien).] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 13: 93-131. (In Bulgarian, German summary) Josifov M. (1963b). Eine neue Rhyparochromus- Art von der Insel Kreta (Hem. Het. Lygaeidae). — Acta Entomologica Musei Nationalis Pragae, 35: 123-126. Seidenstücker G. & M. Josifov (1963). Die Odontotarsus oculatus-Gruppe (Hemiptera, Heteroptera, Pentatomidae). — Reichenbachia, 2 (46): 67-75. ЙОСИФОВ М. [ Josifov M.] (1964a). Heteroptera от Тракийската низина. [Heteroptera aus der Thrakischen Tiefebene.] — Фауна на Тракия, Част I [Die Fauna Thrakiens, Band I]: 207-245. (In Bulgarian, German summary) ЙОСИФОВ М. [ Josifov M.] (1964b). Видов състав и разпространение на насекомите от разред Heteroptera в България. Част II. [Artbestand und Verbreitung der Insekten von der Ordnung Heteroptera in Bulgarien, Teil II.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 16: 83-149. (In Bulgarian, German summary) Josifov M. (1964c). Cryptostemma remanei n. sp., eine neue Heteropteren-Art aus Bulgarien. — Mitteilungen der Deutschen Entomologischen Gesellschaft, 23 (5/6): 93-94. Josifov M. (1964d). Über die Gattung Orthonotus Stephens, 1829 (Heteroptera, Miridae). — Reichenbachia, 4 (18): 151-156. Josifov M. (1965a). Eine neue Paredrocoris- Art aus Bulgarien (Heteroptera, Miridae). — Reichenbachia, 5 (16): 157-159. Josifov M. (1965b). Zur Systematik der Gattung Melanocoryphus Stål (Hem. Het. Lygaeidae). — Acta Entomologica Musei Nationalis Pragae, 36: 311-334. Josifov M. (1965c). Eine neue Melanocoryphus- Art aus China (Heteroptera, Lygaeidae). — Acta Entomologica Musei Nationalis Pragae, 36: 383-384. Josifov M. (1965d). Zwei neue Emblethis- Arten (Heteroptera, Lygaeidae). — Reichenbachia, 5 (23): 203-206. Josifov M. (1965e). Zur Systematik der Gattung Gampsocoris Fuss, 1852 (Heteroptera, Berytidae). — Reichenbachia, 5 (33): 285-289. Josifov M. (1966). Zur Systematik der Gattung Maccevethus Dallas, 1852 (Hemiptera, Heteroptera, Rhopalidae). — Reichenbachia, 8 (7): 55-60. Kerzhner I.M. & M. Josifov (1966). Beschreibung neuer Arten von Landwanzen (Heteroptera) aus der Mongolischen Volksrepublik und Bemerkungen über Phytocoris turkestanicus Pop. — Bulletin de l’Académie polonaise des Sciences, 14 (9): 627-634. Josifov M. (1967a). Eine neue Acalypta- Art aus Anatolien (Heteroptera, Tingidae). — Reichenbachia, 8 (26): 211-213.
Michail Josifov – Bibliography, described and dedicated taxa 31
(45) Josifov M. (1967b). Zur Systematik der Gattung Cryptostemma H.-S. (Heteroptera). — Annales zoologici, 25 (2): 215-225. (46) Josifov M. & I.M. Kerzhner (1967). Heteroptera von der Mongolischen Volksrepublik. — Fragmenta faunistica, 14 (1): 1-15. (47) Josifov M. (1968a). Eine Heteropterenausbeute von der Insel Kreta. — Annales zoologici, 25 (11): 453-457. (48) Josifov M. (1968b). Die Paläarktischen Arten der Gattung Phymata Latreille, 1802 (Hem. Het. Reduviidae). — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 26: 29-32. (49) Josifov M. (1968c). Ferganocoris kerzhneri n. gen. n. sp. – eine neue asiatische Phyline (Heteroptera, Miridae). — Reichenbachia, 10 (3): 13-17. (50) Josifov M. (1968d). Eine neue Chorosoma- Art aus Bulgarien (Heteroptera, Rhopalidae). — Reichenbachia, 10 (33): 255-158. (51) ЙОСИФОВ М. [ Josifov M.] (1969a). Видов състав и разпространение на насекомите от разред Heteroptera в България. Част III. [Artenzusammensetzung und Verbreitung der Insekten von der Ordnung Heteroptera in Bulgarien, Teil III.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 29: 29-82. (In Bulgarian, German summary) (52) Josifov M. (1969b). Einige neue Miriden aus Bulgarien (Hemiptera, Heteroptera). — Reichenbachia, 12 (4): 29-36. (53) Josifov M. (1969c). Eine neue Psallus- Art aus Bulgarien (Heteroptera, Miridae). — Reichenbachia, 12 (8): 69-71. (54) Josifov M. (1969d). Neue asiatische Derephysia- Arten (Heteroptera, Tingidae). — Acta Entomologica Musei Nationalis Pragae, 38: 59-66. (55) Josifov M. (1969e). Über die Taxonomie von Ventocoris (Selenodera) falcatus (Cyrillus, 1787). — Acta Entomologica Musei Nationalis Pragae, 38: 67-72. (56) Josifov M. (1970). Ergebnise der Albanien-Expedition 1961 des Deutschen Entomologischen Institutes. 82. Beitrag. Heteroptera. — Beiträge zur Entomologie, 20 (7/8): 825-956. (57) Josifov M. (1971). Die Gattung Elasmucha Stål, 1864, im östlichen Mittelmeerraum (Heteroptera, Acanthosomatidae). — Reichenbachia, 13 (25): 239-243. (58) Josifov M. & I.M. Kerzhner (1972). Heteroptera aus Korea. I. Teil (Ochteridae, Gerridae, Saldidae, Nabidae, Anthocoridae, Miridae, Tingidae und Reduviidae). — Annales zoologici, 29 (6): 147-179. (59) Josifov M. (1973a). Eine neue Pilophorus- Art aus Portugal (Heteroptera, Miridae). — Reichenbachia, 14 (19): 151-152. (60) Josifov M. (1973b). Beitrag zur Taxonomie der Gattung Psallus Fieb., 1858 (Hemiptera, Heteroptera, Miridae). — Reichenbachia, 14 (31): 245-248. (61) ЙОСИФОВ М. [ Josifov M.] 1974a. Полутвърдокрилите насекоми (Heteroptera) от Българското Черноморско крайбрежие. [Die Heteropteren der bulgarischen Schwarzmeerküste.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 39: 5-25. (In Bulgarian, German summary)
32 N. Simov
(62) Josifov M. (1974b). Eine neue Phylinae aus Bulgarien (Heteroptera, Miridae) — Comptes rendus de l’Académie bulgare des Sciences, 27 (4): 541-542. (63) ЙОСИФОВ М. [ Josifov M.] (1974c). Хемиптерологично съобщение. V. [Hemipterologische Mitteilung. V.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 40: 217-221. (In Bulgarian, German summary) (64) Josifov M. (1974d). Neue südpaläarktische Miriden (Hemiptera, Heteroptera). — Reichenbachia, 15 (8): 61-68. (65) Josifov M. (1974e). Neue südpaläarktische Phytocoris- Arten (Heteroptera, Miridae). — Reichenbachia, 15 (11): 77-84. (66) Josifov M. (1974f ). Eine neue Psallus- Art aus Bulgarien und eine neue Orthotylus- Art aus Kirgisien (Heteroptera, Miridae). — Reichenbachia, 15 (13): 89-92. (67) ЙОСИФОВ М. [ Josifov M.] (1974g). Дендробионтните полукрили насекоми (Heteroptera) на Стара планина. [Dendrobionte Heteropteren im Balkangebirge.] — Известия на Зоологическия институт с музей [Bulletin de l’Institut de Zoologie et Musée], 41: 95-105. (In Bulgarian, German summary) (68) Josifov M. (1974h). Beitrag zur Systematik der paläarktischen Dichrooscytus- Arten (Heteroptera, Miridae). — Reichenbachia, 15 (20): 149-173. (69) Josifov M. & I.M. Kerzhner (1974). Zur Systematik der ostpaläarktischen Mezira-Arten (Heteroptera, Aradidae). — Reichenbachia, 15 (7): 49-59. (70) Josifov M. (1975). Eine neue Metatropis- Art aus Korea (Heteroptera, Berytidae). — Reichenbachia, 15 (34): 285-286. (71) ЙОСИФОВ М. [ Josifov M.] (1976a). Видообразуване сред хетероптерите в Средиземноморието като резултат от постглациална дизюнкция на ареалите им. [Artbildung bei den Heteropteren im Mittelmeerraum als Folge der postglazialen Disjunktion ihrer Areale.] — Acta zoologica bulgarica, 4: 11-19. (In Bulgarian, German summary) (72) Josifov M. (1976b). Eine neue Lygaeidae-Art (Heteroptera) aus der Balkanhalbinsel. — Acta zoologica bulgarica, 4: 65-66. (73) Josifov M. (1976c). Drei neue Orthotylus-Arten aus Korea (Heteroptera, Miridae). — Reichenbachia, 16 (11): 143-146. (74) Josifov M. (1977). Zur Systematik der Gattung Harpocera Curtis, 1838 (Heteroptera, Miridae). — Acta zoologica bulgarica, 6: 50-53. (75) Josifov M. (1978a). Neue Miridenarten aus Nord-Korea (Heteroptera). — Acta Entomologica Musei Nationalis Pragae, 39: 280-287 (1977). (76) ЙОСИФОВ М. [ Josifov M.] (1978b). Дендробионтни и дендрофилни полутвърдокрили насекоми (Heteroptera) по дъба в България. [Dendrobionte und dendrophile Halbflugler (Heteroptera) an der Eiche in Bulgarien.] — Acta zoologica bulgarica, 9: 3-12. (In Bulgarian, German summary) (77) Josifov M. & I.M. Kerzhner (1978). Heteroptera aus Korea. II. Teil (Aradidae, Berytidae, Coreidae, Urostylidae, Acanthosomatidae, Scutelleridae, Pentatomidae, Cydnidae, Plataspidae). — Fragmenta faunistica, 23 (9): 137-195. (78) Josifov M. (1979). Kerzhneriola gen. n., eine neue Phylinen- Gattung aus Asien (Heteroptera, Miridae). — Reichenbachia, 17 (25): 215-218.
Michail Josifov – Bibliography, described and dedicated taxa 33
(79) Josifov, M. (1980). Über die Verbreitung mancher Heteropterenarten auf der Balkanhalbinsel. — Acta zoologica bulgarica, 15: 20-23. (80) ЙОСИФОВ М. [ Josifov M.] (1981a). Heteroptera, Pentatomoidea. — Фауна на България [Fauna bulgarica] 12. БАН, София: 1-205. (In Bulgarian) (81) Josifov M. (1981b). Die verkannte Dichrooscytus valesianus Fieber, 1861 und der übersehene Dichrooscytus gustavi sp. n. aus Mitteleuropa (Heteroptera, Miridae). — Reichenbachia, 19 (7): 43-45. (82) ЙОСИФОВ М. [ Josifov M.] (1982a). Фаунистични комплекси сухоземни животни. [Faunistische Komplexe der Landtiere.] — В: ГЪЛЪБОВ Ж. (Ред.): География на България. Физическа география. Природни условия и ресурси. БАН, София. [In: Galabov Zh. (Ed.): Geography of Bulgaria. Physical Geography. Natural conditions and resources, Bulgarian Academy of Sciences, Sofia]: 461-466. (In Bulgarian) (83) Josifov M. (1982b). Eine neue Uhlerites- Art aus Korea (Heteroptera, Tingidae). — Reichenbachia, 20 (11): 98-99. (84) Josifov M. (1983a). Beitrag zur Taxonomie der ostpaläarktischen DeraeocorisArten (Heteroptera, Miridae). — Reichenbachia, 21 (12): 75-86. (85) Josifov M. (1983b). Über die Verbraitung mancher Heteropterenarten auf der Balkanhalbinsel. II. — Acta zoologica bulgarica, 22: 3-8. (86) Josifov M. (1983c). Neue Psallus- Arten aus Nord-Korea (KDVR) (Heteroptera, Miridae). — Reichenbachia, 21 (35): 197-211. (87) Josifov M. (1984a). Zur Taxonomie von Aelia klugi Hahn, 1831 und Aelia sibirica Reuter, 1884 (Heteroptera, Pentatomidae). — Acta zoologica bulgarica, 25: 45-49. (88) Josifov M. (1984b). Trophische Verbindungen und Verbraitung der Heteropterenarten. — In: Kaszab Z. (Ed.): Verhandlungen des zehnten Internationalen Symposiums uber Entomofaunistik Mitteleuropas, 15-20 August 1983, Budapest, Muzsák Közmüvelödési Kiadó, Budapest: 99-101. (89) Josifov M. & I.M. Kerzhner (1984). Zur Systematik der Gattung Dryophilocoris Reuter, 1875 (Heteroptera, Miridae). — Reichenbachia, 22 (310): 215-226. (90) Josifov M. & U. Göllner-Scheiding (1984). Bibliographie der faunistischen Literatur über die Heteropteren der Balkanhalbinsel (Insecta). —Faunistische Abhandlungen Staatliches Museum füer Tierkunde in Dresden, 12 (2): 25-37. (91) Josifov M. (1985). Lygocoris (Arbolygus) kerzhneri sp. n. – eine neue ostpaläarktische Miridenart (Heteroptera). — Reichenbachia, 23 (16): 91-93. (92) Josifov M. (1986a). Über die Verbreitung mancher Heteropterenarten auf der Balkanhalbinsel und der Insel Zypern. — Acta zoologica bulgarica, 31: 42-48. (93) Josifov M. (1986b). The zoogeographical character of Balkan Heteroptera. — In: Drosopoulos S. (Ed.): Proceedings of the 2nd International Congress Concerning the Rhynchota Fauna of Balkan and Adjacent Regions. 18-22 August, Mikrolimni-Prespa, Greece: 6-7. (94) Josifov M. (1986c). Eine neue Aethus-Art von der Balkanhalbinsel (Heteroptera, Cydnidae). — Reichenbachia, 23 (22): 127-129.
34 N. Simov
(95) Josifov M. (1986d). Verzaichnis der von der Balkanhalbinsel bekannten Heteropterenarten (Insecta, Heteroptera). — Faunistische Abhandlungen Staatliches Museum füer Tierkunde in Dresden, 14 (6): 61-93. (96) Josifov M. (1987a). Einige neue Miriden aus Nordkorea (KVDR) (Heteroptera). — Reichenbachia, 24 (15): 115-122. (97) Josifov M. (1987b). Über die Synonimie mancher Lygaeiden (Insecta, Heterroptera). — Acta zoologica bulgarica, 33: 3-9. (98) Josifov M. & J.M. Štusák (1987). A new genus and two new species of Afrotropical Metacanthinae (Heteroptera, Berytidae). — Acta Entomologica Bohemoslovaca, 84: 287-294. (99) Josifov M. (1988). Über den zoogeographischen Charakter der Süderupäischen Insektenfauna unter besonderer Berücksichtigung der Heteropteren. — Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck, 75: 177-184. (100) Josifov M. (1989). Beitrag zur Taxonomie der europäischen Pilophorus- Arten (Insecta, Heteroptera: Miridae). — Reichenbachia, 27 (2): 5-12. (101) Josifov M. & E. Heiss (1989). Eine neue Acrosternum- Art von der Insel Kreta (Heteroptera aus Kreta VII). — Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck, 76: 141-145. (102) Heiss E. & M. Josifov (1990). Vergleichende Untersuchung über Arten-spectrum, Zoogeographie und Ökologie der Heteropteren-Fauna in Hochgebirden Österreichs und Bulgariens. — Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck, 77: 123-161. (103) ЙОСИФОВ М. [ Josifov M.] (1990a). Върху появата на неарктичния вид Corytucha ciliata (Say, 1832) (Heteroptera, Tingidae) в България. [On the appearance of the nearctic spacies Corytucha ciliata (Say, 1832) (Heteroptera, Tingidae) in Bulgaria] — Acta zoologica bulgarica, 39: 53-55. (In Bulgarian, English summary) (104) Josifov M. (1990b). Über die Verbraitung mancher Heteropterenarten auf der Balkanhalbinsel. III. — Acta zoologica bulgarica, 40: 3-11. (105) Josifov M. (1990c). Eine neue Miridenart aus Bulgarien (Insecta: Heteroptera). — Reichenbachia, 27 (16): 95-96. (106) ЙОСИФОВ М. [ Josifov M.] (1990d). Полутвърдокрилите насекоми (Insecta, Heteroptera) в пояса на бука, смърча и субалпийския пояс на Витоша. [Heteroptera (Insecta, Heteroptera) in the belts of the beech, spruce and the subalpine belt of Vitoša.] — Фауна на Югозападна България [Fauna of Southwestern Bulgaria], 3: 88-101. (In Bulgarian, English summary) (107) Park Hee Cheon & M. Josifov (1991). Contribution to a Knowledge of the Heteroptera of North Korea. — Insecta Koreana, 8: 91-103. (108) Josifov M. (1992a). Zur taxonomie der paläarktischen Macrolophus- Arten (Insecta, Heteroptera: Miridae). — Reichenbachia, 29 (1): 1-4. (109) Josifov M. (1992b). Eine neue Lygus- Art aus Tadshikistan (Insecta, Heteroptera: Miridae). — Reichenbachia, 29 (2); 5-7. (110) Josifov M. (1992c). Contribution to a Knowledge of the family Miridae of North Korea (Heteroptera). — Insecta Koreana, 9: 115-128.
Michail Josifov – Bibliography, described and dedicated taxa 35
(111) Josifov M. (1992d). Neue Miriden aus Korea (Insecta, Heteroptera). — Reichenbachia, 29 (20): 105–118. (112) Josifov M. (1992e). Steht die Phänologie der Miridenarten (Heteroptera) in gewissem Zusammenhang mit ihren zoogographischen Charakteristiken? — In: Zombori L. & Peregovits L. (Eds): 1992. Proceeding of the Fourtth European Congress of Entomology and the XIII. Internationale Symposium für die Entomofaunistik Mitteleuropas, Gödölo, Hungary, 1991, Hungarian Natural History Museum, Budapest: 634-636. (113) Josifov M. (1993a). Beitrag zur Verbreitung mancher Heteropterenarten auf der Balkanhalbinsel samt Bemerkungen über die Synonymie mancher Miridenarten (Insecta, Heteroptera). — Acta zoologica bulgarica, 46: 50-57. (114) Josifov M. (1993b). Eine neue mediterrane Isometopus- Art (Insecta: Heteroptera: Miridae: Isometopinae). — Reichenbachia, 30 (3): 7-8. (115) Josifov M. (1993c). Ein kleiner Beitrag zur Systematik der Miriden (Insecta: Heteroptera). — Reichenbachia, 30 (4): 9-15. (116) Josifov M. (1997). Heteroptera. — In: Sakalian V. (Ed.): Endemic and relict insects in the Pirin National Park, Bulgaria. Pensoft Publishers, Sofia-Moscow: 31-42. (117) Georgiev V., Beshovski V., Russev B., Kumanski K., Josifov M. & V. Sakalian (1998). Insects of Bulgaria, Part 1: Odonata, Ephemeroptera, Plecoptera, Homoptera (Auchenorrhyncha), Heteroptera, Coleoptera. — In: Meine C. (Ed.): Bulgarian’s Biological Diversity: Conservation Status and Needs Assessment 1, Washington D. C.: 163-209. (118) Josifov M. (1999). Heteropterous insects in the Sandanski-Petrich Kettle, Southwestern Bulgaria. — Historia naturalis ÿÿlgarica, 10: 35-66. (119) Kerzhner, I.M. & M. Josifov (1999). Family Miridae Hahn, 1833. — In: Aukema, B. & C. Rieger (Eds): Catalogue of the Heteroptera of the Palaearctic Region, The Netherlands Entomological Society, Amsterdam, Vol. 3: 1-576. (120) Hubenov. Z., Beshšvski V., Josifov M., Popov A., Kumanski K., Sakalian V., Abadjiev S., Vidinova Y. & T. Lyubomirov (2000). Entomofaunistic diversity of the Rila National Park. — In: Sakalian M. (Ed.): Biological diversity of the Rila National Park, USAID, Sofia: 285-331. (121) Hubenov Z., Beshšvski V., Josifov M., Popov A., Kumanski K., Sakalian V., Abadjiev S., Vidinova Y. & T. Lyubomirov (2000). Entomofaunistic diversity of the Central Balkan National Park. — In: Sakalian M. (Ed.): Biological diversity of the Central Balkan National Park, USAID, Sofia: 319-362. (122) Josifov M. & N.Simov (2004). Heteroptera in Bulgarian side of Eastern Rhodopes (Insecta). — In: Beron P.& A. Popov (Eds): Biodiversity of Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece).Pensoft & Nat. Mus. Natur. Hist., Sofia: 331-369. (123) Simov N. & M. Josifov (2004). Contribution to the Bulgarian fauna of Heteroptera. — Historia naturalis bulgarica, 16: 89-94.
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(124) Josifov M. & N. Simov (2006). Endemism among the Heteroptera on the Balkan Peninsula. — In: Rabitsch W. (Ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia, 19: 879-898. (125) Grozeva S., Simov N. & M. Josifov (2007). Karyotaxonomy of some European Macrolophus species (Heteroptera: Miridae). — Mainzer Naturwissenschaftliches Archiv, 31: 81-87. (126) Josifov M. & N. Simov (2008). Contribution to the taxonomy of Dicyphus constrictus (Boheman, 1852) (Heteroptera: Miridae) — Historia naturalis bulgarica, 19 (in press).
NEW TAXA DESCRIBED BY MICHAIL JOSIFOV Taxa are ordered by alphabet. After the year of publication the number of the paper in the list is shown. Genera and subgenera Family Berytidae Gampsoacantha Josifov & Stusak, 1987 – 1987: 287 (98). (Typus generis: Gampsoacantha beroni Josifov & Stusak, 1987). Lindbergius Josifov, 1965 – 1965e: 287. (41). (Typus subgeneris: Gampsocoris pallidipes Lindberg, 1958). [Syn. Henry 1997: 53, = Gampsocoris s.str.] Family Lygaeidae Horvathiolus Josifov, 1965 – 1965b: 316 (38). (Typus generis: Melanocoryphus superbus Pollich, 1781). Family Miridae Baginocoris Josifov, 1992 – 1992d: 112 (111). (Typus generis: Baginocoris alienae Josifov, 1992). Ferganocoris Josifov, 1968 – 1968c: 13 (49). (Typus generis: Ferganocoris kerzhneri Josifov, 1968). [Syn. КЕРЖНЕР [Kerzhner] 1988b: 75, = Orthonotus Stephens, 1829]. Kerzhneriola Josifov, 1979 – 1979: 215 (78). (Typus generis: Kerzhneriola asiatica Josifov, 1979). [Syn. КЕРЖНЕР [Kerzhner] 1988a: 856]. Loristes Josifov & Kerzhner, 1972 – 1972: 162 (58). (Typus generis: Adelphocoris decoratus Reuter, 1908). Putshkoviattus Josifov, 1993 – 1993c: 9 (115). (Typus generis: Putshkoviattus muminovi Josifov, 1993). [Syn. Kerzhner 1997: 246, = Glaucopterum Wagner, 1963]. Tricholygus Josifov, 1992 – 1992d: 108 (111). (Typus subgeneris: Lygocoris (Tricholygus) niger Josifov, 1992). [Nomem novum: Kerzhner & Schuh 1995: 2, = Josifovolygus Kerzhner & Schuh, 1995].
Michail Josifov – Bibliography, described and dedicated taxa 37
Species and subspecies Family Acanthosomatidae Elasmucha eckerleini Josifov, 1971 – 1971: 240 (57). (Iran). Elasmucha eckerleini cypria Josifov, 1971 – 1971: 240 (57). (Cyprus). Family Aradidae Aradus bureschi Josifov, 1958 – 1958b: 261 (10). (Bulgaria). [Syn. Tamanini 1972: 26, = Aradus krueperi Reuter, 1884] Mezira hoberlandti Josifov & Kerzhner, 1974 – 1974: 52 (69). (Russia FE, Korea). [nomen novum: Mezira ludviki Josifov & Kerzhner, 1978 ( Josifov & Kerzhner 1978: 139)]. Mezira subsetosa Josifov & Kerzhner, 1974 – 1974: 49 (69). (Russia FE). Mezira subtilis Josifov & Kerzhner, 1974 – 1974: 51 (69). (Russia FE). Family Berytidae Gampsoacantha beroni Josifov & Stusak, 1987 – 1987: 288 (98). (Mozambique). Gampsoacantha pumilio Josifov & Stusak, 1987 – 1987: 291 (98). (Nigeria, Chad, Ivory Cost). Gampsocoris eckerleini Josifov, 1959 – 1959b: 155 (17). (Greece & Crete). [downgraded by Péricart 1984, = Gampsocoris culicinus eckerleini Josifov] Gampsocoris lilianae Josifov, 1958 – 1958d: 269 (12). (Bulgaria). Gampsocoris minutus Josifov, 1965 – 1965e: 288 (41). (Iran: Schachrud). Metatropis tesongsanicus Josifov, 1975 – 1975: 285 (70). (Korea). Family Cydnidae Aethus balcanicus Josifov, 1986 – 1986c: 125 (94). (Geece & Crete, Macedonia, Bulgaria). [in genus Byrsinus Fieber, 1860 (Lis 1999)] Aethus riedeli Josifov & Kerzhner, 1978 – 1978: 186 (77). (Korea). [Syn. Lis 1999: 194, = Byrsinus varians (Fabricius, 1803)]. Chilocoris nigricans Josifov & Kerzhner, 1978 – 1978: 190 (77). (Russia FE, Korea). Family Dipsocoridae Cryptostemma carpaticum Josifov, 1967 – 1967b: 222. (45). (Poland). Cryptostemma gracile Josifov, 1967 – 1967b: 222 (45). (Tadzhikistan). Cryptostemma remanei Josifov, 1964 – 1964c: 93 (35). (Bulgaria). Cryptostemma roubali Josifov, 1967 – 1967b: 220 (45). (S France). Family Lygaeidae Auchenodes joakimoffi Seidenstücker & Josifov, 1961 – 1961: 27 (27). (Turkey: Asian part, Bulgaria). [Syn. Péricart 1994: 97- 98, = Auchenodes costalis (Lethierry, 1877)].
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Cymus elegans Josifov & Kerzhner, 1978 – 1978: 141 (77). (Korea). Cymus koreanus Josifov & Kerzhner, 1978 – 1978: 148 (77). (Korea). Emblethis persicus Josifov, 1965 – 1965d: 206 (40). (Iran: Isfagan). Emblethis robustus Josifov, 1965 – 1965d: 203 (40). (Greece: Crete, Cyprus). Geocoris megacephalus creticus Josifov, 1968 – 1968a: 454 (47). (Greece: Crete). [Syn. Josifov 1987b: 4, = Geocoris pubescens ( Jakovlev, 1871)]. Geocoris variabilis minusculus Kerzhner & Josifov, 1966 – 1966: 631 (43). (Mongolia). Horvathiolus kiritshenkoi Josifov, 1965 – 1965b: 322 (38). (Armenia, SW Iran, Turkey). Ischnocoris bureschi Josifov, 1976 – 1976b: 65 (72). (Bulgaria, Greece, Albania). Melanocoryphus kerzhneri Josifov, 1965 – 1965c: 383 (39). (China). Plinthisus (Dasythisus) bembidioides Josifov & Kerzhner, 1978 – 1978: 148 (77). (Korea). Rhyparochromus (Neoxanthochilus) creticus Josifov, 1963 – 1963b: 123 (31). (Greece & Crete). [Syn. Péricart 1998, = Xanthochilus minusculus Reuter, 1885]. Trichodrymus pallipes Josifov & Kerzhner, 1978 – 1978: 150 (77). (Korea). Family Miridae Adelphocoris ponghvariensis Josifov, 1978 – 1978a: 282. (75). (Korea). Alloeotomus linnavuorii Josifov & Kerzhner, 1972 – 1972: 153 (58). (Korea). [Syn. КЕРЖНЕР [Kerzhner] 1978: 37, = Alloeotomus simplus (Uhler, 1896)]. Baginocoris alienae Josifov, 1992 – 1992d: 113. (111). (Korea). Blepharidopterus mesasiaticus Josifov, 1993 – 1993c: 10 (115). (Kirgizia). Cinnamus inexpectatus Josifov, 1978 – 1978a: 279 (75). (Korea). [in genus Acrorrhinium Noualhier, 1895 (Kerzhner & Josifov 1999)]. Compsidolon (Apsinthophylus) balcanicum Josifov, 1993 – 1993c: 12 (115). (Bulgaria). [Syn. Konstantinov 2006: 499, = Compsidolon (Apsinthophylus) pumilum ( Jakovlev, 1876)]. Daraeocoris (Knightocapsus) ulmi Josifov, 1983 – 1983a: 83 (84). (Korea). Daraeocoris (s.str.) castaneae Josifov, 1983 – 1983a: 79 (84). (Korea). Daraeocoris (s.str.) kerzhneri Josifov, 1983 – 1983a: 77 (84). (Russia FE, Japan). Daraeocoris (s.str.) pallidicornis Josifov, 1983 – 1983a: 77 (84). (Russia FE, Korea). Daraeocoris (s.str.) salicis Josifov, 1983 – 1983a: 81 (84). (Russia FE, Korea). Deraeocoris (Camptobrochis) onphoriensis Josifov, 1992 – 1992d: 105 (111). (Korea). [Syn. Lee, Miyamoto & Kerzhner 1994: 4, = Deraeocoris pulchellus (Reuter, 1906)]. Deraeocoris (s.str.) ventralis megophthalmus Josifov & Kerzhner, 1972 – 1972: 155 (58). (Russia FE, Korea). Dichrooscytus algiricus bureschi Josifov, 1974 – 1974h: 164 (68). (Bulgaria). [upgraded by Günther 1990, = Dichrooscytus bureschi Josifov] Dichrooscytus altaicus Josifov, 1974 – 1974h: 164 (68). (Russia: Altai). Dichrooscytus asanovae Josifov, 1974 – 1974h: 158 (68). (Kirgizia). Dichrooscytus bulgaricus Josifov, 1959 – 1959a: 63 (16). (Bulgaria). [Syn. Josifov, 1981b: 43, = Dichrooscytus valesianus Fieber, 1861] Dichrooscytus gustavi Josifov, 1981 – 1981b: 44 (81). (Germany).
Michail Josifov – Bibliography, described and dedicated taxa 39
Dichrooscytus kerzhneri Josifov, 1974 – 1974h: 163 (68). (Kirgizia). Dichrooscytus kiritshenkoi Josifov, 1974 – 1974h: 159 (68). (Kazakhstan). Dichrooscytus persicus Josifov, 1974 – 1974d: 66 (64). (Iran). Dichrooscytus putshkovi Josifov, 1974 – 1974h: 153. (68). (Russia: Dagestan). Dichrooscytus seidenstueckeri Josifov, 1974 – 1974h: 151 (68). (Turkey: Asian part). Dicyphus (Brachyceroea) digitalidis Josifov, 1958 – 1958e: 271 (13). (Bulgaria). Dicyphus (Brachyceroea) martinoi Josifov, 1958 – 1958e: 273 (13). (Bulgaria). [in subgenus Mesodicyphus Wagner, 1951 (Wagner 1967)]. Dryophilocoris (s.str.) jenjouristi Josifov & Kerzhner, 1984 – 1984: 218 (89). (Russia FE, Korea). Dryophilocoris (s.str.) kanyukovae Josifov & Kerzhner, 1984 – 1984: 220 (89). (Russia FE, Korea). Dryophilocoris pallidulus Josifov & Kerzhner, 1972 – 1972: 169 (58). (Korea). Ferganocoris kerzhneri Josifov, 1968 – 1968c: 14 (49). (Kirgizia) [in genus Orthonotus Stephens, 1829 (КЕРЖНЕР [Kerzhner] 1988b: 75)]. Halticus apterus koreanus Josifov, 1987 – 1987a: 115 (96). (Korea). Halticus comitans Josifov & Kerzhner, 1972 – 1972: 167 (58). (Russia FE, Korea, China). Harpocera choii Josifov, 1977 – 1977: 50 (74). (Korea). Harpocera koreana Josifov, 1977 – 1977: 51 (74). (Korea). Heterochlorillus nathaliae Josifov 1974 – 1974d: 63. (64). (Tadzhikistan). Heterocordylus cytisi Josifov, 1958 – 1958f: 84 (14). (Bulgaria). Isometopus longirostris Josifov, 1993 – 1993b: 7 (114). (Bulgaria). Kerzhneriola asiatica hissarensis Josifov, 1979 – 1979: 216 (78). (Kazakhstan, Kirgizia, Tadzhikistan, Armenia, Azerbaijan, Russia: Dagestan). [in Atomoscelis Reuter, 1875 (КЕРЖНЕР [Kerzhner] 1988a)]. Kerzhneriola asiatica Josifov, 1979 – 1979: 216 (78). (Korea). [in genus Atomoscelis Reuter, 1875 (КЕРЖНЕР [Kerzhner] 1988a)]. Lygocoris (Arbolygus) kerzhneri Josifov, 1985 – 1985: 91 (91). (Korea, Russia FE). [in genus Castanopsides Yasunaga, 1992 (Yasunaga 1998)]. Lygocoris (Neolygus) mjohjangsanicus Josifov, 1992 – 1992d: 106 (111). (Korea). Lygocoris (Neolygus) sylvaticus Josifov, 1992 – 1992d: 106 (111). (Korea). Lygocoris (Tricholygus) niger Josifov, 1992 – 1992d: 109 (111). (Korea). [in genus Josifovolygus Kerzhner & Schuh, 1995 (Kerzhner & Schuh 1995)]. Lygus dracunculi Josifov, 1992 – 1992b: 5 (109). (Tadzhikistan). [Syn. Aglyamzyanov 2005: 76, = Lygus poluensis Wagner, 1967]. Macrolophus insignis Josifov, 1968 – 1968a: 453 (47). (Bulgaria, Greece: Crete). [Syn Josifov 1992a: 2, = Macrolophus pygmaeus (Rambur, 1839)]. Macrolophus nubilis (sic!) geranii Josifov, 1961 – 1961a: 87 (22). (Bulgaria). [upgraded by Grozeva et al. 2007, = Macrolophus geranii Josifov]. Macrotylus soosi Josifov, 1962 – 1962b: 377 (29). (Greece & Crete). Mecomma ambulans montanus Josifov, 1969 – 1969b: 35 (52). (Bulgaria).
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Orthonotus kiritschenkoi Josifov, 1964 – 1964d: 153 (36). (Azerbaijan). Orthonotus pseudoponticus Josifov, 1964 – 1964d: 154 (36). (Bulgaria). Orthotylus (Melanotrichus) choii Josifov, 1976 – 1976c: 144 (73). (Korea). Orthotylus (Melanotrichus) namphoensis Josifov, 1976 – 1976c: 144 (73). (Korea). [Syn. КЕРЖНЕР [Kerzhner] 1988b: 834, = Orthotylus parvulus Reuter, 1879]. Orthotylus (Orthotylus) bureschi Josifov, 1969 – 1969b: 32 (52). (Bulgaria) [in subgenus Parapachylops Ehanno & Matocq, 1990 (Kerzhner & Josifov, 1999: 269)]. Orthotylus (Orthotylus) tenellus meridionalis Josifov, 2006 – 2006: 881 (124). (Bulgaria). Orthotylus (s.str.) kogurjonicus Josifov, 1992 – 1992d: 111 (111). (Korea). Orthotylus (s.str.) kymgangsanicus Josifov, 1987 – 1987a: 121 (96). (Korea). [Syn. Lee, Miyamoto & Kerzhner 1994: 13, = Orthotylus salicis Jakovlev, 1893]. Orthotylus (s.str.) sophorae Josifov, 1976 – 1976c: 145 (73). (Korea). Orthotylus junipericola balcanicus Josifov, 1974 – 1974d: 66 (64). (Bulgaria). [in subgenus Parapachylops Ehanno & Matocq, 1990 (Carapezza 1997)]. Orthotylus putshkovi Josifov, 1974 – 1974f: 91 (66). (Kirgizia). [in subgenus Parapachylops Ehanno & Matocq, 1990 (Carapezza 1997)]. Paredrocoris seidenstueckeri Josifov, 1965 – 1965a: 157 (37). (Bulgaria). Phylus (Teratoscopus) coryloides Josifov & Kerzhner, 1972 – 1972: 173 (58). (Russia FE, China, Korea). Phytocoris (Compsocerocoris) strymonensis Josifov, 1990 – 1990c: 95. (105). (Bulgaria). Phytocoris (Ktenocoris) muminovi Josifov, 1974 – 1974e: 80 (65). (Tadzhikistan). Phytocoris (Phytocoris) thrax Josifov, 1969 – 1969b: 31 (52). (Bulgaria). Phytocoris (Ribautomiris) dichrooscytoides Josifov, 1974 – 1974e: 77 (65). (Libya). Phytocoris (Ribautomiris) pseudoscytulus Josifov, 1974 – 1974e: 79 (65). (Algeria). Pilophorus alni Josifov, 1987 – 1987a: 117 (96). (Korea). [Syn. Kerzhner 1993: 100, = Pilophorus erraticus Linnavuori, 1962)]. Pilophorus choii Josifov, 1978 – 1978a: 285 (75). (Korea). Pilophorus coiffaiti Josifov, 1973 – 1973a: 151 (59). (Portugal). [Syn. Wagner 1975: 284, = Pilophorus angustulus Reuter, 1888]. Pilophorus dianae Josifov, 1989 – 1989: 9 (100). (Bulgaria). Pilophorus koreanus Josifov, 1978 – 1978a: 283 (75). (Korea). Pilophorus pseudoperplexus Josifov, 1987 – 1987a: 118 (96). (Korea). Pilophorus simulans Josifov, 1989 – 1989: 8 (100). (Russia, Croatia, Bulgaria, Romania, France). Placochilus seladonicus mediterraneus Josifov, 1969 – 1969b: 35 (52). (Bulgaria). Psallus (Apocremnus) ater Josifov, 1983 – 1983c: 198 (86). (Korea). Psallus (Apocremnus) atratus Josifov, 1983 – 1983c: 197 (86). (Korea). Psallus (Apocremnus) betuleti montanus Josifov, 1973 – 1973b: 246 (60). (Bulgaria). [upgraded by Rieger & Rabitsch 2006, = Psallus montanus Josifov, 1973] Psallus (Apocremnus) niger Josifov, 1992 – 1992d: 113 (111). (Korea). [nomen novum: Kerzhner & Schuh 1995: 4, = Psallus (Apocremnus) michaili Kerzhner & Schuh, 1995]. Psallus (Mesopsallus) muminovi Josifov, 1973 – 1973b: 245 (60). (Tadzhikistan). [in genus Lepidargyrus Muminov, 1962 (Drapolyuk 1993)].
Michail Josifov – Bibliography, described and dedicated taxa 41
Psallus (Mesopsallus) samdzijonicus Josifov, 1983 – 1983c: 200 (86). (Korea). Psallus (Phylidea) castaneae Josifov, 1983 – 1983c: 202 (86). (Korea). Psallus (Phylidea) kerzhneri Josifov, 1992 – 1992d: 115 (111). (Korea). Psallus (Phylidea) ulmi Kerzhner & Josifov, 1966 – 1966: 627 (43). (Russia ES, Mongolia, China). Psallus (Pityopsallus) lapponicus kimi Josifov, 1983 – 1983c: 210 (86). (Korea). [upgraded by Vinokurov 1998: 285, = Psallus kimi Josifov, 1983]. Psallus (Psallus) balcanicus Josifov, 1969 – 1969c: 29 (53). (Bulgaria) [Syn. Kerzhner & Josifov 1999: 416, = Psallus lucanicus Wagner, 1968]. Psallus (Psallus) helenae Josifov, 1969 – 1969c: 69 (53). (Bulgaria). Psallus (s.str.) amoenus Josifov, 1983 – 1983c: 208 (86). (Korea). Psallus (s.str.) bagjonicus Josifov, 1983 – 1983c: 205 (86). (Korea). Psallus (s.str.) faniae Josifov, 1974 – 1974d: 61 (64). (Bulgaria). Psallus (s.str.) koreanus Josifov, 1983 – 1983c: 209 (86). (Korea). Psallus (s.str.) milenae Josifov, 1974 – 1974f: 89. (66). (Bulgaria). Psallus (s.str.) sanguinolentus Josifov, 1983 – 1983c: 209 (86). (Korea). [nomen novum: Psallus sanguinarius Josifov, 1999 (Kerzhner & Josifov, 1999: 417)]. Psallus (s.str.) tesongsanicus Josifov, 1983 – 1983c: 207 (86). (Korea). Psallus pseudoquercus Josifov, 1974 – 1974b: 541 (62). (Bulgaria). Putshkoviattus muminovi Josifov, 1993 – 1993c: 9 (115). (Tadzhikistan). [in genus Glaucopterum Wagner, 1963 (Kerzhner 1997)]. Reuteria castaneae Josifov, 1987 – 1987a: 115 (96). (Korea). Rubrocuneocoris quercicola Josifov, 1987 – 1987a: 122 (96). (Korea). Tytthus coreanus Josifov & Kerzhner, 1972 – 1972: 171 (58). (Korea). Family Pentatomidae Acrosetnum malickyi Josifov & Heiss, 1989 – 1989: 141 (101). (Greece: Crete). Arma koreana Josifov & Kerzhner, 1978 – 1978: 181 (77). (Korea, China). Codophila (Antheminia) mongolica Kerzhner & Josifov, 1966 – 1966: 633 (43). (Mongolia). Eurydema eckerleini Josifov, 1961 – 1961c: 397 (24). (Greece & Crete). Homalogonia grisea Josifov & Kerzhner, 1978 – 1978: 175 (77). (Korea, China). Rhacognathus lamellifer Josifov & Kerzhner, 1978 – 1978: 184 (77). (Russia FE, NE China, Korea). Family Rhopalidae Chorosoma gracile Josifov, 1968 – 1968d: 255 (50). (Bulgaria). Family Tingidae Acalypta anatolica Josifov, 1967 – 1967a: 211 (44). (Turkey) [Syn. Péricart 1982: 352, = Acalypta uniseriata (Puton, 1879)]. Biskria nigricosta Kerzhner & Josifov, 1966 – 1966: 629 (43). (Mongolia, Russia WS). [in genus Dictyonota Curtis, 1827 (ГОЛУБ [Golub] 1975)].
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Biskria pulchricornis Kerzhner & Josifov, 1966 – 1966: 630 (43). (Mongolia, Russia ES and FE). [in genus Dictyonota Curtis, 1827 (ГОЛУБ [Golub] 1975)]. Derephysia bucharensis Josifov, 1969 – 1969d: 62 (54). (Uzbekistan). Derephysia gracilicornis Josifov, 1969 – 1969d: 65 (54). (Armenia, Azerbaijan). Derephysia kiritshenkoi Josifov, 1969 – 1969d: 59 (54). (Armenia, Tadzhikistan, NO Iran) [downgraded by Péricart 1978: 92, = Derephysia (s.str.) rectinervis kiritshenkoi Josifov]. Derephysia minuta Josifov, 1969 – 1969d: 63 (54). (Tadzhikistan). Uhlerites gracilis Josifov, 1982 – 1982b: 97 (83). (Korea). Table 1. Summary of new taxa described by Michail Josifov. Berytidae Lygaeidae Miridae
Family
Genera and subgenera 2 1 6
Acanthosomatidae Aradidae Berytidae Cydnidae Dipsocoridae Lygaeidae Miridae Pentatomidae Rhopalidae Tingidae
Species and subspecies 2 4 6 3 4 13 97 6 1 8
MICHAIL JOSIFOV DEDICATED TAXA Taxa are ordered by chronology and then by alphabet. a) Genera [1]. Michailocoris Štys, 1985 (Heteroptera, Miridae) [2]. Josifovolygus Kerzhner & Schuh, 1995 (Heteroptera, Miridae) [3]. Josifovius Konstantinov, 2008 (Heteroptera, Miridae) b) Species and subspecies [4]. Streptanus josifovi Dlabola, 1957 (Cicadomorpha: Cicadellidae) [ = Streptanus albanicus (Horváth, 1916)] [5]. Orthotylus josifovi Wagner, 1959 (Heteroptera, Miridae)
Michail Josifov – Bibliography, described and dedicated taxa 43
[6]. Acetropis (Acetropis) josifovi Wagner, 1967 (Heteroptera, Miridae) [=Acetropis sinuata Wagner, 1951] [7]. Adelphocoris josifovi Wagner, 1968 (Heteroptera, Miridae) [8]. Biskria josifovi Seidenstücker, 1968 (Heteroptera, Tingidae) [= Dictyonota opaca (Linnavuori, 1965)] [9]. Raphidia (Bureschiella) josifovi Popov, 1974 (Raphidioptera: Raphidiidae) [= Mongolraphidia (Usbekoraphidia) josifovi (Popov, 1974)] [10]. Agriphila beieri josifovi Ganev, 1975 (Lepidoptera: Crambidae) [11]. Leptus josifovi Beron, 1975 (Acari: Erythraeidae) [12]. Parapiesma josifovi Péricart, 1977 (Heteroptera, Piesmatidae) [13]. Bubastia (Bubastia) josifovi Dlabola 1980 (Fulgoromorpha: Issidae) [14]. Michailocoris josifovi Štys, 1985 (Heteroptera, Miridae) [15]. Deraeocoris josifovi Kerzhner, 1988 (Heteroptera, Miridae) [16]. Oxyethira josifovi Kumanski, 1990 (Trichoptera: Hydroptilidae) [17]. Luperomorpha josifovi Gruev, 1994 (Coleoptera: Chrysomelidae) [18]. Dicyphus (Dicyphus) josifovi Rieger, 1995 (Heteroptera, Miridae) [19]. Psallus (Apocremnus) michaili Kerzhner & Schuh, 1995 (Heteroptera, Miridae) [20]. Dichrooscytus josifovi Kerzhner, 1997 (Heteroptera, Miridae) [21]. Calacanthia josifovi Vinokurov, 2008 (Heteroptera, Saldidae) [22]. Castanopsides michaili Yasunaga & Duwal, 2008 (Heteroptera, Miridae) [23]. Catoplatus josifovi Golub, 2008 (Heteroptera, Tingidae) [24]. Chorosoma josifovi Schwartz, Schaefer & De Lattin, 2008 (Heteroptera, Rhopalidae) [25]. Dictyonota michaili Günther, 2008 (Heteroptera, Tingidae) [26]. Euchilofulvius josifovi Gorczyca, 2008 (Heteroptera, Miridae) [27]. Loricula (Myrmedobia) josifovi Simov, 2008 (Heteroptera, Microphysidae) [28]. Pilophorus josifovianus Duwal & Yasunaga, 2008 (Heteroptera, Miridae) [29]. Pilophorus josifovi Kerzhner, 2008 (Heteroptera, Miridae) [30]. Pinochius josifovi Rédei, 2008 (Heteroptera, Schizopteridae) [31]. Raglius alboacuminatus josifovi Linnavuori & Heiss, 2008 (Heteroptera, Lygaeidae) [32]. Sigara (Pseudovermicorixa) josifovi Chen & Nieser, 2008 (Heteroptera, Corixidae) [33]. Tubuaivelia michaili Polhemus & Polhemus, 2008 (Heteroptera, Veliidae) [34]. Xenicocephalus josifovi Štys & Baa, 2008 (Heteroptera, Enicocephalidae)
РЕЗЮМЕ За поколения български зоолози и ентомолози Михаил Йосифов е учител и пример за подражание с отдадеността си на науката, перфектната работа, огромната си ерудираност, новаторските си идеи и нестихващия ентусиазъм за нови знания и от-
44 N. Simov
крития. За тези, които го познават и които са имали възможността да работят с него, винаги подадената ръка за помощ, съветите, напътствията и приятелските разговори са безценни. Настоящата работа е опит за обобщение на творческия път на Михаил Йосифов до момента и включва библиографска справка за научните му трудове за периода 1950-2007 г., списък на описаните от него 9 рода и 144 вида хетероптери, както и наименованите на негово име таксони (3 рода и 31 вида). Не са включени неговите над 50 научнопулярни статии и книги.
REFERENCES List of the cited in the text publications (except these from the list of M. Josifov’ publications), as well the papers of dedicated taxa (mentioned by the number of the taxon in the list) ГОЛУБ В.Б. [Golub V.B.] (1975). Обзор клопов-кружевиц рода Dictyonota Curtis (Heteroptera, Tingidae) фауны СССР и Монголии. [Review of the lacebugs of the genus Dictyonota Curtis (Heteroptera, Tingidae) of the fauna of the USSR and Mongolia.] — Насекомые Монголии [Insects of Mongolia], 3: 55-78. (In Russian) КЕРЖНЕР И.М. [Kerzhner I.M.] (1978). Полужесткокрылые (Heteroptera) Сахалина и Курильских островов. [Heteroptera of Saghalien and Kurile Islands]. — Труды Биолого-почвенного института Далневосточного научнаго центра АН СССР. Новая серия. [Proceedings of the Institute of biology and soil sciences, Far East Branch of the Acad. Sci. USSR, Vladivostok, new ser.], 50 (153): 31-57. (In Russian) КЕРЖНЕР И.М. [Kerzhner I.M.] (1988a). Инфраотряд Cimicomorpha. 21. Сем. Miridae (Capsidae) – Слепняки [Infraorder Cimicomorpha. 21. Family Miridae (Capsidae)], 778-857. — In: ЛЕР П.А. [Ler, P. A.] (Ed.): Определитель насекомых Дальнего Востока СССР. Т. II. Равнокрылые и полужесткокрылые. [Keys to the identification of insects of the Soviet Far East. Vol. 2: Homoptera and Heteroptera.] Наука, Ленинград [Nauka, Leningrad]: 1-972. (In Russian) [15] КЕРЖНЕР И.М. [Kerzhner I.M.] (1988b). Новые и малоизвестные полужесткокрылые насекомые (Heteroptera) с Дальнего Востока СССР. [New and little known heteropteran insects (Heteroptera) from the Far East of the USSR]. ДВНЦ АН СССР, Владивосток [Far East Branch of the Acad. Sci. USSR, Vladivostok], 1987: 1-88. (In Russian) Aglyamzyanov R.S. (2005). Lygus poluensis Wagner and its synonyms (Heteroptera: Miridae). Zoosystematica Rossica, 14(1): 76 [11] Beron P. (1975). Erythraeidae (Acariformes) larvaires de Bulgarie. – Acta zoologica bulgarica, 1: 45-75. Carapezza A. (1997). Heteroptera of Tunisia. — Il Naturalista Siciliano, 21 (suppl. A): 1-331. [32] Chen P-p. & N. Nieser (2008). The Corixidae (s. str.) of Thailand with description of a new species of Sigara (Heteroptera, Nepomorpha). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 55-63.
Michail Josifov – Bibliography, described and dedicated taxa 45
[4] Dlabola J. 1957. The problem of the genus Delphacodes and Calligypona, three new species and other Czechoslowakian faunistics (Hom. Auchenorrh.). — Acta Musei Nationalis Pragae, 31 (476): 113-110 [13] Dlabola J. 1980. Tribus-Einteilung, neue Gattungen und Arten der Subf. Issinae in der eremischen Zone (Homoptera, Auchenorrhyncha). — Acta Musei Nationalis Pragae, 36 B (4): 173-247. Drapolyuk I.S. (1993). Review of the capsid bugs of the genus Lepidargyrus (Heteroptera: Miridae). — Zoosystematica Rossica, 2: 107-119. [28] Duwal R.K. & T. Yasunaga (2008). A new species of the pilophorine plant bug genus Pilophorus Hahn from Nepal (Heteroptera, Miridae, Phylinae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 79-84. [10] Ganev J. (1985). Agriphila beieri josifovi n. ssp. aus der Türkei. – Sonderabdruck aus dem Nachrichtenblatt der Bayerischen Entomologen, 34 (4): 139-141. [23] Golub V. (2008). A new species of lacebug from Tajikistan, Catoplatus josifovi nov. sp. (Heteroptera, Tingidae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 129-132. [26] Gorczyca J. (2008). A new species of Euchilofulvius Poppius, 1909 from Malaysia (Heteroptera: Miridae: Cylapinae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 133-137. [17] Gruev B. (1994). New Distributional Data about Some Leafbeetles in the Korean Peninsula and Description of Four New Species (Coleoptera, Chrysomelidae) — Insecta Koreana, 11: 75-84. Günther H. (1990). Contribution to the Heteroptera fauna of the Balkan Peninsula. — Scopolia, (suppl. 1): 39-52. [25] Günther H. (2008) A new tingid species from Southern Spain: Dictyonota michaili nov. sp. (Heteroptera: Tingidae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 181-184. Henry T.J. (1997). Cladistic analysis and revision of the stilt bug genera of the world (Heteroptera: Berytidae). — Contributions of the American Entomological Institute, 30(1): 1-100. Kerzhner, I.M. (1993). Notes on synonymy and nomenclature of Palearctic Heteroptera. — Zoosystematica Rossica, 2: 97-105. Kerzhner, I.M. (1997). Notes on taxonomy and nomenclature of Palearctic Miridae (Heteroptera). — Zoosystematica Rossica, 5 (2): 245-248. [20] Kerzhner I.M. (1997). Type specimens of some Palaearctic Miridae in the Zoological Museum, Helsinki (Heteroptera). — Zoosystematica Rossica, 6: 115-121. [29] Kerzhner I.M. (2008). A new species of Pilophorus from the Far East (Heteroptera, Miridae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, SofiaMoscow: 185-188. [2, 19] Kerzhner I.M. & Schuh R.T. (1995). Homonymy, synonymy, and new combinations in the Miridae (Heteroptera). — American Museum Novitates, 3137: 1-11.
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Konstantinov F.V. (2006). Two new species of Phylini (Heteroptera, Miridae, Phylinae) from Middle Asia and Caucasus with notes on Compsidolon pumilum ( Jakovlev, 1876) — In: Rabitsch W. (Ed): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia, 19: 493-502. [3] Konstantinov F. (2008). Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus Wagner 1961 (Heteroptera: Miridae: Phylinae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, SofiaMoscow: 215-230. [16] Kumanski K. (1990). Studies on the fauna of Trichoptera (Insecta) of Korea I. Superfamily Rhyacophiloidea. — Historia naturalis bulgarica, 2: 36-60. Lee C.E., Miyamoto S. & I.M. Kerzhner (1994). Additions and corrections to the list of Korean Heteroptera. — Nature and Life (Korea), 24: 1-34. [31] Linnavuori R.E. & E. Heiss (2008). A new subspecies of Lygaeidae from northern Iran: Raglius alboacuminatus josifovi nov. ssp. (Heteroptera, Lygaeidae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 231-234. Lis J. (1999). Burrower bugs of the Old World – a catalogue (Hemiptera: Heteroptera: Cydnidae). — Genus, 10(2): 165-249. [12] Péricart J. (1977). Un Hémiptère Piesmatidae nouveau de Corée et un Hémiptère Berytidae nouveau d’Afrique du Nord. — Bulletin de la Société Entomologique de France, 82: 33-38. Péricart J. (1978). Révision systématique des Tingidae Ouest-paléarctiques. 3. Subdivision du genre Derephysia et revue critique des espéces (Hemiptera). — Annales de la Société Entomologique de France (N.S.), 14 (1): 87-94. Péricart J. (1982). Révision systématique des Tingidae Ouest-paléarctiques. 9. Compléments et corrections. — Annales de la Société Entomologique de France (N.S.), 14 (4): 349-372. Péricart J. (1984). Hémiptéres Berytidae euro-méditerranéens. — Faune de France 70: i-v, 1-172. Péricart J. (1994). Lygaeidae paléarctiques: synonymies, combinations nouvelles et description de deux Geocoris nouveaux (Hemiptera). Bulletin de la Société entomologique de France, 99 (1): 93-105. Péricart J. (1998). Hémiptéres Lygaeidae euro-méditerranéens. Vol.3. — Faune de France 84C: i-vi, 1-487. [33] Polhemus J.T. & D.A. Polhemus (2008) A new genus of Microveliinae from the Austral Islands, French Polynesia (Heteroptera, Veliidae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 293-302. [9] Popov A. (1974). Eine neue Raphidia aus Tadshikistan (Raphidioptera). — Nouvelle Revue d’Entomologie, 4 (2): 153-158. [30] Rédei D. (2008). First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species from Vietnam (Heteroptera: Schizopteridae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 327-337. [18] Rieger C. (1995). Zwei neue Miriden von de Insel Kreta (Heteroptera). — Entomologische Berichten, Amsterdam 55: 79-82.
Michail Josifov – Bibliography, described and dedicated taxa 47
Rieger C. & W. Rabitsch (2006). Taxonomy and distribution of Psallus betuleti (Fallén) and P. montanus Josifov stat. nov. (Heteroptera, Miridae). — Tijdschrift voor Entomologie, 149: 161-166. [24] Schwartz M.D., C.W. Schaefer & J.D. De Lattin (2008). The first species of Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World: Chorosoma josifovi sp. n. — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 339-348. [8] Seidenstücker G. (1968). Biskria josifovi n.sp. (Heteroptera, Tingidae). — Reichenbachia, 10 (36): 267-268. [27] Simov N. (2008). A new Loricula species from Bulgaria (Heteroptera: Microphysidae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 349-356. [1, 14] Štys P. (1985). A new genus of Palearctic Bryocorinae related to Afrotropical Rhodocoris (Heteroptera, Miridae). — Acta Entomologica Bohemoslovaca, 82: 407-425. [34] Štys P. & P. Baa (2008). Xenicocephalus – an enigmatic genus of American Enicocephalidae (Heteroptera): a new male-based species from Suriname. — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 357-376. Tamanini L. (1972). Osservazioni sugli Aradus krueperi Rt., bureschi Jos., ribauti Wgn. e Iongirostris Gyll. — Bolletino della Società Entomologica Italiana, 104 (1-3): 23-27. Vinokurov N.N. (1998). Asian plant bugs of the subgenus Pityopsallus E. Wagn., genus Psallus Fieb. (Heteroptera: Miridae). — Zoosystematica Rossica, 7: 285-296. [21] Vinokurov N.N. (2008). Calacanthia josifovi nov. sp., a new species of shore bugs (Heteroptera, Saldidae) from Tibet. — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 385-389. [5] Wagner E. (1959). Orthotylus josifovi nov. spec. (Hem. Het. Miridae). — Deutsche Entomologische Zeitschrift (N.F.), 6: 145-147. [6] Wagner E. (1967). Über Acetropis Fieb. (Heteroptera, Miridae). — Reichenbachia, 8: 205209. Wagner E. (1967). Zur Systematik der Gattung Dicyphus Fieber, 1856 (Heteroptera, Miridae).— Reichenbachia, 8: 111-121. [7] Wagner E. (1968). Die Adelphocoris vandalicus Gruppe (Hemiptera, Heteroptera, Miridae). — Reichenbachia, 10 (14): 119-125. Wagner E. (1975). Über die Synonymie einiger Miriden-Arten (Hem. Het.).— Reichenbachia, 15: 283-284. Yasunaga T. (1998). Revision of the mirine genus Castanopsides Yasunaga from the eastern Asia (Heteroptera: Miridae). — Entomologica Scandinavica, 29: 99-119. [22] Yasunaga T. & R.K. Duwal (2008) New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania Poppius from Nepal, with a new synonymy of the genus Liocapsus Poppius (Heteroptera, Miridae, Mirinae). — In: Grozeva S. & N. Simov (Eds): Advances in Heteroptera Research. Festschrift in honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow: 403-417.
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S. Grozeva & N. Simov1973, (Eds) 2008 Psallus (Apocremnus) montanus Josifov, in Th e Netherlands (Heteroptera, Miridae) 49 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 49-54. © Pensoft Publishers Sofia–Moscow
Psallus (Apocremnus) montanus JOSIFOV, 1973, in The Netherlands (Heteroptera, Miridae) B. Aukema Zoölogisch Museum, Afdeling Entomologie, Plantage Middenlaan 64, 1018 DH Amsterdam, Nederland. E-mail:
[email protected]
ABSTRACT Dutch material identified as Psallus betuleti is revised. Both P. betuleti and P. montanus occur in The Netherlands, the latter being the more common of the two. Distribution and phenology of both species are illustrated. Keywords: Psallus montanus, Psallus betuleti, The Netherlands, distribution, phenology
INTRODUCTION Psallus (Apocremnus) betuleti montanus was described by Josifov (1973) as a subspecies of P. (Apocremnus) betuleti (Fallén, 1826) from material from Bulgaria collected on Vitosha Mt. at 1400 m on European white birch (Betula pendula Roth = B. verrucosa Ehrh.). At first, it was considered an endemic Bulgarian subspecies, restricted to mountain areas (Heiss & Josifov 1990), but Bacchi & Rizzotti Vlach (1994) changed the picture by reporting it from Italy (Venetia, Lombardy) and Austria (North Tyrol). Subsequently, Rieger & Rabitsch (2006) raised Psallus montanus to species rank based on constant and distinct differences in male and female genitalia.
The distribution of P. montanus is only partly known, because P. betuleti usually was identified without dissecting the genitalia. Rieger & Rabitsch (2006) consider it a Palaearctic species, introduced into eastern North America. They list the species from Austria, Bulgaria, Czech Republic, Germany, Italy, Russia (Caucasus) and North America (Canada and USA). Aukema et al. (2007) give Belgian records and Nau (2007) confirms its presence in Great Britain. The identity of P. betuleti from other countries listed by Kerzhner &
50 B. Aukema
Josifov (1999): Byelorussia, Denmark, Kazakhstan, Finland, France, Hungary, Ireland, Latvia, Luxembourg, Norway, Poland, Romania, Slovakia, Slovenia, Spain, Sweden, Switzerland, Ukraine, Armenia, China, Georgia, Korea and Asian Russia, remains to be checked. A problem may arise if P. montanus also occurs in Sweden, because P. betuleti was described from that country and the type material could not be located (Kerzhner & Josifov 1999).
RESULTS AND DISCUSSION Dutch collection material identified as P. betuleti was checked, it was verified that both species are present in The Netherlands and P. montanus is the more common species. It is represented by 349 specimens collected over the period 1916-2007 in 59 10x10 kilometre squares (Fig. 1), from all provinces except Zeeland in the southwest. P. betuleti is less common, with 76 specimens collected over the period 1918-2007 in 34 10x10 kilometre squares (Fig. 2). So far, no records are known from Groningen in the north, the new polders (Flevoland) in the centre, and Zeeland in the southwest. Both species occur on the West Frisian Islands: P. montanus was collected on Terschelling and Vlieland, and P. betuleti on Texel and Terschelling. Despite the observed differences, the overall distribution pattern of the two species is quite similar and frequently they are found at the same locality on the same day. The observed coincidence of the species, and the greater abundance of P. montanus, also seem to be a general Central European pattern (Rieger & Rabitsch 2006).
Fig. 1: Distribution of Psallus montanus in The Fig. 2: Distribution of Psallus betuleti in The Netherlands Netherlands.
Psallus (Apocremnus) montanus Josifov, 1973, in The Netherlands (Heteroptera, Miridae) 51
There is no evidence that both species occur on the same tree, but from experience with other Psallus species-pairs this seems not unlikely. As far as food plants are cited, both species are nearly always found on Betula sp. This applies for the records in The Netherlands as well as other European countries (Rieger & Rabitsch 2006). Records from other tree species, e.g. Quercus sp., might be accidental or for reasons other than larval development (e.g. resting, adult feeding on aphids etc.). Both species fly frequently and are often captured in window traps and light traps, indicating a high dispersal power. The phenology of both species is also quite similar and confirms findings in other European countries (Rieger & Rabitsch 2006). The first adults of P. montanus appeared in the first decade (10-day period) of May, most were found in June and the last were found at the end of July (Fig. 3). In P. betuleti a similar pattern was found but over a shorter period, from the second decade of May to the second decade of July (Fig. 4), however this apparent difference may be due to the smaller sample of the latter species. The colour pattern of the males of Dutch material shows more variation as suggested by Rieger & Rabitsch (2006): two male P. betuleti and a male P. montanus had a colour pattern “typical” of females of the latter.
Number of records
50 45
females
40
males
35 30 25 20 15 10 5 0 5-1
5-2
5-3
6-1
6-2
6-3
7-1
7-2
7-3
Decade
Number of records
Fig. 3: Phenology of Psallus montanus, by 10-day periods. 15 females
10
males
5 0 5-1 5-2 5-3 6-1 6-2 6-3 7-1 7-2 7-3 Decade
Fig. 4: Phenology of Psallus betuleti.
52 B. Aukema
ACKNOWLEDGEMENTS Thanks are due to Willem Hogenes, Zoölogisch Museum, Amsterdam, Rob de Vries, Naturalis, Leiden, Antoon Loomans, Plantenziektenkundige Dienst, Wageningen and Yde Jongema, Universiteit Wageningen for giving access to the collections and to Kees den Bieman, Ulvenhout, Bas Drost, Wadenoijen, Dik Hermes, Geldrop, Sjoerd Tiemersma, Wezep and Philip Zeinstra, Ter Idzard for the use of their material. Bernard Nau, Dunstable, corrected the English text.
РЕЗЮМЕ Mатериал от Холандия, определен като Psallus betuleti, е ревизиран. Установено е, че видовете P. betuleti и P. montanus се срещат в Холандия, като от двата последният е по-обичаен. Разпространението и фенологията на двата вида са илюстрирани. REFERENCES Aukema B., J. Bruers & G. Viskens (2007). Nieuwe en zeldzame Belgische wantsen II (Hemiptera: Heteroptera). — Bulletin van de Koninklijke Belgische Vereniging voor Entomologie, 143: 83-91. Bacchi I. & M. Rizzotti Vlach (1994). Quattro specie del genre Psallus Fieber, 1858 nuove per la fauna italiana (Heteroptera, Miridae). — Atti XVII Congresso Nazionale Italiano di Entomologia, Udine, 13-18 giugno 1994: 191-194. Heiss E. & M. Josifov (1990). Vergleichende Untersuchung über Artenspektrum, Zoogeographie und Ökologie der Heteroptera-Fauna in Hochgebirgen Österreichs und Bulgariens. — Berichte des Naturwissenschaftlich-Medizinischen Vereins Innsbruck, 77: 123-161. Josifov M. (1973). Beitrag zur Taxonomie der Gattung Psallus Fieb., 1858 (Hemiptera, Heteroptera, Miridae). — Reichenbachia, 14: 245-248. Kerzhner I.M. & M. Josifov (1999). Cimicomorpha II. Miridae. In: Aukema B. & Chr. Rieger (Eds): Catalogue of the Heteroptera of the Palaearctic Region 3 — Netherlands Entomological Society, Amsterdam. Nau B.S. (2007). From the regions. Bedfordshire. — Het News, 9: 14. Rieger Chr. & W. Rabitsch (2006). Taxonomy and distribution of Psallus betuleti (Fallén) and P. montanus Josifov stat. nov. (Heteroptera, Miridae). — Tijdschrift voor Entomologie, 149: 161-166.
Psallus (Apocremnus) montanus Josifov, 1973, in The Netherlands (Heteroptera, Miridae) 53
APPENDIX Material examined The following abbreviations for collectors and collections are used: BA: B. Aukema, Renkum; DH: D.J. Hermes, Geldrop; LUWC: Wageningen Universiteit en Research Centrum, Wageningen; PDWC: Plantenziektenkundige Dienst, Wageningen; PZ: Philip Zeinstra, Ter Idzard; RMNH: Nationaal Natuurhistorisch Museum Naturalis, Leiden; ZMAN: Zoölogisch Museum, Amsterdam. Unless indicated otherwise, the specimens are in the collection of the collector. Psallus montanus THE NETHERLANDS. Friesland. Hemrik, Wijnjeterper Schar, 1.vi.1999, 1♀ from Betula, PZ; Terschelling, Hoorn, 23.vi.2007, 2♀♀ from Betula and 27.vi.2007, 1♀ from Betula, BA; Terschelling, Formerum-Noord, 27.vi.2007, 1♂, 4♀♀ from Betula, BA; Vlieland, 28.vi.2007, 10♀, BA. Groningen. Sellingen, Ter Borg, 19.vi.1968, 1♀, Ph. Pronk (RMNH); Wollinghuizen, 11.vi.1986, 1♂ from Quercus, K. den Bieman (ZMAN). Drenthe. Dwingeloo, 12.vi.1971, 1♂, J. Teunissen (ZMAN); Kraloër Heide, 12.vi.1971, 1♀, C. de Jong (RMNH); Hooghalen, Kamp Westerbork, vi.1944, 1♂, 1♀, E.A.M. Speijer (RMNH); Wijster, 29.vi.1942, 2♀♀, W. Beijerinck (RMNH). Overijssel. Diepenveen, 22-23.vi.1967, 2♂♂ and 11-12.vii.1967, 1♂ at light, J.J. Meurer (ZMAN); Holten, 4.vi.1960, 1♀, S. van Heijnsbergen (ZMAN); Ootmarsum, Springendal, 18.vi.1970, 1♂, 1♀ and 2.vii.1970, 2♀♀, BA (ZMAN); Vasse, dal van de Mosbeek, 14.vi.2003, 1♀, B. Drost. Flevoland. Biddinghuizen, 21.vi.1981, 1♀, BA. Gelderland. Bennekom, 30.v.1952, 1♀, R.H. Cobben (LUWC); Deelerwoud, 9.vii.1977, 1♂ and 10.vii.1977, 1♀, C. Dolleman (ZMAN); Doornspijk, 8.vi.1930, 1♀, A. Reclaire (RMNH); Ede, 24.vi.1958, 1♀, J.J. Meurer (ZMAN); Eerbeek, Coldenhove, 20.vi.1944, 1♂, 1♀, J. van Hespen (RMNH); Eerbeek, 6.vii.1944, 2 ♀♀, C. de Jong (RMNH); Epe, Eperholt, 5.vii.1980, 1♀, BA; Heveadorp, Duno, 4.vi.2007, 9♂♂, 2♀♀ from Betula, BA; Oldebroek, 12.vi.1970, 1♀, BA (ZMAN); Overasselt, Uiversnest, 29.v.1988, 1♀, DH; Rheden, Posbank, 27.vi.1987, 1♀, BA; Wageningen, 20.vi.1949, 2♀♀, R.H. Cobben (LUWC); ibid., De Eng, 16.v-11.vi.2007, 15♂♂, 9♀♀ in window traps, BA; ibid., Oranje Nassau’s Oord, 20.vi.1949, 1♀, R.H. Cobben (LUWC); ibid., 10.vi.2006, 2♂♂, 19.vi.2006, 1♂, 2.vii.2006, 1♂, 2♀♀, 3.vii.2006, 1♀ and 9.vii.2006, 1♀ at light, BA; ibid., Wageningse Berg, 4.vi.1953, 1♀, R.H. Cobben (LUWC). Utrecht. Baarn, 25.vi.1927, 1♂, A. Reclaire (ZMAN); ibid., 13.vii.1929, 1♀, A. Reclaire (RMNH); Driebergen, 28.v.1946, 1♀ and 14.vi.1942, 3♀♀, W.H. Gravestein (ZMAN); Leusden, 1.vi.1952, 2♀♀, S. van Heijnsbergen (ZMAN). Noord-Holland. Aerdenhout, vi.1950, 1♀, S. van Heijnsbergen (ZMAN); Amsterdam, 4.vii.1977, 1♀, W.H. Gravestein (ZMAN); ibid., Amsterdamse Bos, 14.vi.1951, 2♂♂, 1♀, P.A.A. Loof and 24.vi.1973, 2♀♀, BA (ZMAN) and 2♀♀, ibid. (BA); Amsterdam, Vondelpark, 29.vi.1944, 3♀♀, W.H. Gravestein (ZMAN); Bergen, 20.vi.1942, 1♀, W.H. Gravestein (ZMAN); Bergen, Verbrande Pan, 21.vi.1954, 3♀♀, J.H. Woudstra (ZMAN); Heemskerk, 18.vi.1952, 1♀ and 19.vi.1958, 1♀, J.H. Woudstra (ZMAN); Heemstede, 16.vii.1955, 1♀, 3-4.vii.1956, 1♂ at light, 15-16.vii.1957, 1♀ at light, 17-18.vi.1957, 2♂♂, 1♀ at light, 18-19.vi.1957, 2♂♂ at light, 23-24. vii.1957, 1♀ at light, 30.vi.1957, 2♀♀, 6.vii.1957, 2♂♂, 1♀ and 6-7.vii.1957, 3♂♂, 1♀ at light, J.J. Meurer (ZMAN); Hilversum, 6.vii.1921, 1♂, 2.vii.1927, 2♂♂, 14.vii.1929, 1♂, 20.vi.1937, 1♀, 2.vi.1943, 2♀♀ and 4.vi.1943, 5♀♀, A. Reclaire (ZMAN); ibid., 7.vi.1947, 1♀, 9.vi.1947, 1♀ and 10.vi.1947, 1♀, A. Reclaire (RMNH); Kortenhoef, 30.v.1964, 1♀, S. van Heijnsbergen (ZMAN); Laren, 21.vi.1942, 3♀♀, W.H. Gravestein (ZMAN); Oud-Naarden, 5.vi.1982, 1♀, S. van Heijnsbergen (ZMAN); Wijk aan Zee, 8.vi.1952, 1♂ and 10.vi.1952, 1♀, J.H. Woudstra (ZMAN). Zuid-Holland. De Zilk, 20.vi.1957, 1♂ and 26.vi.1957, 3♀♀, J.J. Meurer (ZMAN); Wassenaar, Bierlap, 21.vi.1935, 1♀, A.M. Scholte (RMNH); Wassenaar, Meijendel, 17.vi.1923, 1♀, 19.vi.1924, 1♀ and 23.vi.1925, 1♂, H.C. Blöte (RMNH); ibid., 17.vi.1923, 1♂, J. van der
54 B. Aukema
Vecht, RMNH. Noord-Brabant. Eindhoven, TUE-terrein, 7.vi.1989, 1♀ from Betula, DH; Galder, 14.vi.1951, 8♂♂, 6♀♀, R.H. Cobben (LUWC); Geldrop, Gijzenrooise Zegge, 27.v.1989, 1♀, DH; ibid., Rulse Dijk, 17.vii.1996, 2♂♂, 5♀♀, DH; ibid., 19.vi.1998, 2♀♀, DH; ibid., Strabrechtse Heide, Scheidingsven, 13.vi.1991, 1♂, DH (ZMAN); Heeze, Groote Heide, 11.vi.2006, 1♀, DH; Lage Mierde, De Utrecht, 10.vi.2006, 1♀, B. Drost; Maarheeze, De Pan, 11.vi.1994, 1♀, DH; Middelbeers, 10.vi.2006, 1♂, Sj. Tiemersma; Nuenen, Eeneind, 3.vii.1998, 1♀, DH; Oisterwijk, 28.v.1949, 1♀, P.J. Brakman (RMNH); Riethoven, Keersopper Beemden, 1.vi.1999, 2♀♀, DH; Strijbeek, 17.vi.1967, 1♀, J.H. Woudstra (ZMAN); Tilburg, Kaaistoep, 30.v.2007, 18♂♂, 72♀♀ from Betula, BA; Zundert, 14.vi.1968, 1♂, J.H. Woudstra (ZMAN). Limburg. Arcen, 18.vi.1966, 1♀, W.H. Gravestein (ZMAN); Arcen, De Hamert, 18.vi.1968, 1♂, 2♀♀, C. de Jong (RMNH); Bunde, 22.v.1952, 1♀, Excursie Museum Leiden (RMNH); Epen, 31.v.1949, 1♀, P.J. Brakman (RMNH); Epen, 21.vi.1946, 1♀, W.H. Gravestein (ZMAN); ibid, 1♀ (PDWC); Heythuizen, 11.vi.1944, 1♀, R.H. Cobben (LUWC); Horn, 5.vi.1945, 1♀, R.H. Cobben (LUWC); Lomm, Ravenvennen, 6.vi.1982, 1♀, BA; Lomm, 9.vi.1982, 2♀♀, BA; Lottum, 16.vi.1982, 1♀, BA; Rijckholt, Zure Dries, 8.vi.2007, 1♂, 2♀♀ from Betula, BA; Schaesberg, 22.vi.1987, 1♀, G.R. Langohr (RMNH); Valkenburg, vi.1923, 1♂, 3♀♀, H.C.L. van Eldik (RMNH); Venlo, vi, 1♂, R.A.H. van den Brandt (ZMAN); ibid., 9.vi.1988, 1♀, E.A.M. Speijer (RMNH); Vijlen, 31.v.1949, 1♀, P.J. Brakman (RMNH); Vijlen, Kerperbosch, 9.vi.2007, 1♂, 1♀ from Betula, BA; Vlodrop-Station, 25.vi.1987, 1♀, G.R. Langohr (RMNH); Well, 29.vi.1987, 2♀♀, G.R. Langohr (RMNH); Wieler, 1.vii.1974, 1♂, BA. Psallus betuleti THE NETHERLANDS. Friesland. Hemrik, Wijnjeterper Schar, 1.vi.1999, 3♂♂ from Betula, PZ; Nijeberkoop, Boschhoeve, 25.v.2000, 1♂ from Betula, PZ; Nijeholtpade, 30.v.1999, 1♂ from Betula, PZ; Terschelling, Hoorn, 23.vi.2007, 1♂ from Betula, BA. Drenthe. Diever, Berkenheuvel, 27.vi.1942, 1♀, W. Beijerinck (RMNH); Mantinge, Mantingerbos, 15.vi.1980, 1♀, BA; Ruinen, Moddergat, 7.vii.1984, 1♀, BA; Wijster, IJsbaan, 18.vi.1977, 1♂, 1♀, BA. Overijssel. Ommen, vi.1918, 1♂, D. MacGillavry (ZMAN). Gelderland. Doornspijk, 8.vi.1930, 2♂♂, A. Reclaire (ZMAN); Leuvenum, 19.vi.1938, 1♀, A. Reclaire (ZMAN); Lunteren, 24.v.1990, 1♂, K. den Bieman; Nunspeet, De Vennen, 16.vi.1985, 1♂, P.H. van Doesburg (RMNH); Voorthuizen, 1.viii.1962, 1♀, J.H. Woudstra (ZMAN); Wageningen, De Eng, 16.v.2007, 2♀♀ in windowtrap, BA. Utrecht. Bilthoven, 6.vi.1971, 1♀and 21.vi.1975, 1♂, C. de Jong (RMNH); Driebergen, 3.vi.1946, 3♂♂, W.H. Gravestein (ZMAN). Noord-Holland. Amsterdam, Vondelpark, 29.vi.1944, 3♂♂ from Betula, W.H. Gravestein (ZMAN); Ibid., Amsterdamse Bos, 14.vi.1951, 1♀, P.A.A. Loof (ZMAN); ibid., 24.vi.1973, 2♂♂, BA and 1♀, BA (ZMAN); Heemstede, 15-16. vii.1956, 1♂ and 3-4.vii. 1957, 1♂ at light, and 6.vii.1957, 1♂, J.J. Meurer (ZMAN); Hilversum, 1.vii.1922, 1♂, 12.vi.1931, 1♀, 26.vi.1937, 1♂ and 21.vi.1941, 1♂, A.Reclaire (ZMAN); ibid., 9.vi.1947, 1♀, A. Reclaire (RMNH); Kortenhoef, 30.v.1964, 1♂, S. van Heijnsbergen (ZMAN); Laren, 21.vi.1942, 2♂♂, W. H. Gravestein (ZMAN); Oud-Naarden, 5.vi.1982, 1♂, S. van Heijnsbergen (ZMAN); Texel, De Koog, ‘t Mientje, 13.vii.1996, 1♂, DH; Westzaan, Guisveld, 29.vi.1987, 1♂, R. de Vos (ZMAN); Wijk aan Zee, 10.vi.1952, 1♂, J.H. Woudstra (ZMAN). Zuid-Holland. Wassenaar, Kijfhoek, 19.vi.1935, 1♂, A.M. Scholte (RMNH); Ibid., Meijendel, 22.vi.1925, 1♂, H.C. Blöte (RMNH). Noord-Brabant. Eindhoven, TUE-terrein, 7.vi.1989, 1♂, DH; Maarheeze, De Pan, 11.vi.1994, 1♂, DH; Tilburg, Kaaistoep, 30.v.2007, 2♂♂, 2♀♀ from Betula, BA.. Limburg. Bemelerberg, 22.v.1952, 1♂, Museum Leiden (RMNH); Lomm, 5.vi.1982, 1♂, BA; Lomm, Ravenvennen, 6.vi.1982, 1♀, BA; Lottum, 14.vi.1982, 1♀, BA; St Odiliënberg, 20.v.1948, 1♀ from Quercus, R.H. Cobben (LUWC); Vijlen, 30.v.1949, 1♂, P.J. Brakman (RMNH).
S. Grozeva & N. Simov 2008 The Corixidae (s. str.) of Thailand, with description of a new(Eds) species of Sigara (Heteroptera, Nepomorpha) 55 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 55-63. © Pensoft Publishers Sofia–Moscow
The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) P.-p. Chen & N. Nieser ABSTRACT Three new records and a new species, Sigara (Pseudovermicorixa) josifovi, of Corixidae from Thailand, with a key to Thai species. Keywords: Corixidae, key, new species, Sigara, Thailand.
INTRODUCTION Although the Corixidae s. str. (sensu Nieser, 2002) with about 450 species worldwide (Chen et al. 2005) form the largest family of Nepomorpha, they are poorly represented in tropical areas. The only area between 23.5° North and South latitude which is comparatively rich in species and specimens, is the higher elevations of the Andean Region of South America (Hungerford 1948; Nieser & Padilla Gil 1992). The reason for this scarcity is probably that nearly all non ephemeral waters in warmer areas are infected with fish, many species of which prey on Corixidae. Only a few specimens of Corixidae were collected during our trips to Thailand. As precise literature references for Thailand are lacking we report here on our own material and include in the key one additional species which, judging from its distribution, may occur in Thailand. The measurements of the new species are based on five specimens of each sex, presented as the range with the mean values in italics and the value of the holotype added between {} or as mean values only.
56 P.-p. Chen & N. Nieser
Depositories nctn Nieser collection, Tiel, The Netherlands. rmnh collection of the Nationaal Natuurhistorisch Museum Naturalis, Leiden, The Netherlands. Specimens studied are in nctn, unless otherwise indicated.
SYSTEMATIC PART Key to species of Corixidae possibly occurring in Thailand (Primarily applicable to males) 1. -.
2.
-. 3.
Pronotum and hemielytra concolorous. Body shape broad, length 6.5-8.0 mm, head width 2.3-2.9 mm, Fig. 1......................... Agraptacorixa hyalinipennis (Fabricius) Pronotum and hemielytra brown with yellowish transverse bands which are interrupted on hemielytra. Body shape more elongate, length 4.5-7.0 mm, head width 1.5-2.2 mm, Fig. 2 (genus Sigara).........................................................................2 Pronotum with 8-11, often anastomotic, transverse yellowish lines; male vertex in dorsal view anteriorly pointed (Fig. 3); right paramere (Fig. 5) apically slightly swollen ................................................. Sigara (Pseudovermicorixa) josifovi nov. sp. Pronotum with 6-8 (occasionally 9) usually regular transverse yellowish lines; male vertex in dorsal view anteriorly rounded (Fig. 2) ..................................................3 Right paramere distally slender (Fig. 13)............................................................... ...................................................... Sigara (Pseudovermicorixa) septemlineata Paiva
Fig. 1: Agraptacorxa hyalinipennis (Fabricius), Fig. 2: Sigara (Tropocorixa) paivai Lundblad, male, length 7.7 mm (from Chen et al. 2005). male, length 5.0 mm (from Chen et al. 2005).
The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) 57
-. 4. -.
Right paramere distally wide or widened (Figs 14, 15) .......................................4 Right paramere with a median bulge (Fig. 14) ...................................................... .................................................................. Sigara (Tropocorixa) paivai Lundblad Right paramere distally widened “birdhead-like” (Fig. 15) .................................... .................................................................... Sigara (Tropocorixa) distorta Distant Agraptacorixa hyalinipennis (Fabricius) Fig. 1.
Sigara hyalinipennis Fabricius, 1803: 105. Agraptacorixa hyalinipennis; Hutchinson 1940: 411-412. Agraptacorixa hyalinipennis; Jansson 1995: 36.
Material examined. Thailand: Chiang Mai Prov., Muang District, at foot of Doi Suthep, drying pond in Agro-forestry Research Station, Faculty of Agriculture, Chiang Mai University, 14.XII.2004, leg. P. Chen & N. Nieser, CN04100, 1♀. First record for Thailand. Diagnosis. A medium sized, short and broad species, length 6.5-8.5, width of head 2.7-3.1 mm (Fig. 1). Pronotum and hemielytra concolorous, pale. Within the continental SE Asian fauna this species can be recognized at once by these characteristics. Distribution. From India to S. China including Taiwan and Japan (Okinawa) and through SE Asia to Indonesia (Moluccas) and the Philippines (Mindanao) ( Jansson 1995; Nieser & Chen 1999).
Sigara (Pseudovermicorixa) josifovi nov. sp. Figs 3-5, 7-12.
Type material. Holotype male (RMNH) THAILAND, Chiang Mai Prov., Chom Thong Distr., Baan Luang Subdist., Doi Inthanon Nat. Park, Khun Huay Haeney, 18°34’23.5”N, 98°31’04.9”E, 1567m a.s.l., 15 Jan.2005. Permanent pond along road, in decidious forest, heavily shaded, no aquatic vegetation, pH 7.0, leg. P.-p. Chen & N. Nieser, CN0508. Paratypes, 5♂, 9♀, same data as holotype (NCTN, 1♀ RMNH); Chiang Mai Prov., Chom Thong Distr., Baan Luang Subdistr., Doi Inthanon Nat. Park, Huay Ton Phueng, 18°39’11.7”N, 98°27’29.7”E, 1357m a.s.l., 15 Jan.2005. Narrow muddy stream in mixed pine forest, virtually stagnant, shallow pools with bottom of thin black mud connected by short stretches with low velocity of current; water colourless, clear, pH 6.5, temp. 17°C, leg. P.-p. Chen & N. Nieser, CN0507, 2♂, 3♀; Chiang Mai Prov., Fang District, Doi Ang Khang, Agricultural Horticulture Station, pond, 30 Oct. 2004. Stream pond at station office, stagnant water with trees and low grasses, leg. P.-p. Chen, N. Nieser & R. Thapa, CN0455, 1♂ 1♀. Dimensions. Length ♂ 5.5-5.6-5.8 {5.6}, ♀ 5.5-5.6-5.7; width of head ♂ 1.801.81-1.82 {1.80}, ♀ 1.80-1.91-1.97.
58 P.-p. Chen & N. Nieser
5 3 6
4
7
Fig. 3-5: Sigara (Pseudovermicorixa) josifovi nov. sp. 3 head an pronotum of holotype male, dorsal view; 4 head of holotype anterior view; 5 right paramere of paratype. Fig. 6: Sigara (Pseudovermicorixa) kempi Hutchinson, right paramere (after Hutchinson, 1940, not on scale). Fig. 7: Sigara (Pseudovermicorxa) josifovi nov. sp., paratype, left paramere.
8 10
9
11
Fig. 8-11: Sigara (Pseudovermicorixa) josifovi nov. sp., paratype; 8 abdominal tergite VI; 9 abdominal tergite VII, 10 strigil; 11 hind femur, ventral view.
The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) 59
Colour. General facies dark brown to blackish. Pronotum (Fig. 3) with 8-11 transverse yellowish bands often some anastomozing, especially in anterior half of pronotum. Exposed part of propleura in dorsal view blackish. Clavus with fairly regular transverse yellowish pattern especially in anterior half; corium with vermiculate broken yellowish pattern, yellowish marks about as broad as alternating dark brown to blackish marks. Corium and membrane separated by yellowish line. Pruinose area of embolium black. Head and legs yellowish, eyes light castaneous. Thoracic venter yellowish with medially variable dark brown to blackish markings. Venter of abdomen in males greyish brown; in females light brown. Structural characteristics. Lateral angles of pronotum rounded leaving a distinct dorsal part of the propleura uncovered. Lateral lobe of prothorax slightly longer than its basal width (2.8/2.50) its lateral margins slightly tapering to a rounded apex. Metaxyphus shorter than its basal width (2.6/3.4), triangular, with broadly rounded apex. Postnodal pruinose area of embolium slightly shorter than claw of middle leg (1.08/1.11), no apparent difference between sexes. Middle leg: femur : tibia : tarsus : claw 2.19: 1.06: 0.73: 1.12, claws in males slightly longer than in females (1.15 and 1.09 respectively). Hind leg: femur : tibia : tarsi : tarsII 1.14: 1.12: 1.40: 0.59. Hind femur in ventral view (Fig. 11) border between pubescent and glabrous area well distally of middle of femur; spines on glabrous area: three spines along anterior margin, two to three spines posterodistally, about five in posterior half and one in anterior half of glabrous surface. Male. Median length of head subequal to median length of pronotum (0.87/0.87, Fig. 3). Vertex in dorsal view acutely produced anteriad (Fig. 3); facial impression large, in frontal view occupying nearly the entire area of frons dorsally of rostrum (Fig. 4). Synthlipsis subequal to width of an eye (measured along posterior margin of eye, 1.04/1.03). Fore leg (Fig. 12): pala elongate with about 27 pegs, 30-34 bristles in upper, 23-26 bristles in lower palmar row. Abdominal tergite VI (Fig. 8) with medium sized strigil consisting of 5 combs of teeth (Fig. 10); tergite VII (Fig. 9) with submedian lobe short and broad with about 30 caudal bristles. Right paramere (Fig. 5) slender with a slight swelling in apical third; left paramere (Fig. 7) rather short and broad. Female. Median length of head about two thirds median length of pronotum (0.63/0.95). Vertex in dorsal view rounded anteriorly; frons without facial impression. Synthlipsis on average slightly larger than width of eye (1.10/1.07). Comparative notes. The pointed vertex in dorsal view reminds at first sight of Sigara (Vermicorixa) scripta (Rambur) which, however occurs in the Mediterranean reaching not farther east than Iraq ( Jansson 1986). S. scripta differs among others in having a infuscated second tarsal segment of the hind leg, males lack a strigil and have a small tubercle on the proximal dorsal angle of the pala. The general description of Sigara kempi Hutchinson, 1940 suggests that this species is quite similar to S. josifovi, however, S. kempi is slightly larger, length of male about 6.5, of female about 6.8, the male has a rounded vertex in dorsal view and the right paramere is different distinctively (Fig. 6). Jaczewski (1962b) suggested that S. kempi might belong to the subgenus Pseudovermicorxa Jaczewski 1962. Jansson (1995) placed it, without comments, in the subgenus Vermicorixa Walton, 1940. In view of its close relationship to S. (P.) josifovi we follow
60 P.-p. Chen & N. Nieser
Jaczewski and place it in Pseudovermicorixa. In Thailand occurs another species of the subgenus Pseudovermicorixa, S. septemlineata Paiva, which is smaller; males have a rounded vertex in dorsal view, a smaller strigil and a different right paramere (Fig. 13). Finally Sigara (Pseudovermicorixa) matsumurai Jaczewski (1968) (= Sigara parvula Matsumura 1905 nec Champion 1901) has also a similar right paramere but the male has a rounded vertex in dorsal view and lacks a strigil. Etymology. With great pleasure we dedicate this species to our, always helpful, colleague Michail Josifov, for his great contribution to the knowledge of Hemiptera. Distribution. So far only known from the type area, Chiang Mai Province in N. Thailand.
Sigara (Pseudovermicorixa) septemlineata (Paiva) Fig. 14. Corixa septemlineata Paiva, 1918: 30. Corixa (Vermicorixa) septemlineata; Hutchinson 1940: 430-432. Sigara (Pseudovermicorixa) septemlineata; Jaczewski 1962b: 183. Sigara (Pseudovermicorixa) septemlineata; Jansson 1995: 49.
Material studied. THAILAND: Chiang Rai Prov. Mae Fah Luang District, hilltribe village 15 km N of Maesalong village, 29.X.2004, small muddy rain puddle at side of road, leg. P. Chen, N. Nieser & R. Thapa, CN0449, 17♂, 29♀. First record for Thailand. Diagnosis. Length 4.3-4.8, width of head 1.52-1.72 mm. Pronotum with seven to eight (occasionally nine) transverse yellow lines. Male, facial fovea large but shallow, strigil small with three to five combs of pegs; apical part of right paramere parallel-sided with a small apical hook (Fig. 13). Distribution. Burma, N. Thailand, N. Vietnam, throughout China including Taiwan and Korea to far eastern Russia and Japan.
Sigara (Tropocorixa) distorta (Distant) Figs 15, 16. Corixa distorta Distant, 1911: 343-344. Corixa (Tropocorixa) distorta; Hutchinson 1940: 440-445. Sigara (Tropocorixa) distorta; Jaczewski 1962a: 27.
Diagnosis. Length 5.0-5.5, width of head 1.7 mm. Pronotum with seven regular transverse yellowish lines, distinctly narrower than the intervening brown spaces. Male. Strigil rather small with six combs of pegs; right paramere apically widened (Fig. 15); left paramere with a cleft and lobe ventrally (Fig. 16), this latter characteristic distinguishes this species from all other Sigara.
The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) 61
Distribution. From Pakistan, Nepal and India through N. Vietnam to SW China including Taiwan and Japan (Okinawa) ( Jansson 1995). Not recorded from Thailand but may occur in the North. Sigara (Tropocorixa) paivai Lundblad Sigara paivai Lundblad, 1928: 241. Sigara connexa Lundblad, 1933: 83-85. Sigara paivai; Jansson 1995: 54-55.
Material studied. THAILAND: Nakhon Ratchasima Prov., Bua Yai District, 116 km S of Khon Kaen City, pond beside road No2, shallow, 26.XII.1994, leg. P. Chen, T. Jamganya & Jum, 2♂, 2♀. Diagnosis. Length 4.9-5.5, width of head 1.53-1.78 mm. Pronotum with six (rarely seven) regular transverse yellow lines. Male, strigil elongate but narrow, consisting of one
12 15 13 16 14 Fig. 12: Sigara (Pseudovermicorixa) josifovi nov. sp., paratype, anterior leg. Figs 13-15: Sigara spp., right paramere; 13 S. (Pseudovermicorxa) septemlineata Lundblad; 14 S. (Tropocorixa) paivai Lundblad; 15 S. (Tropocorixa) distorta Distant. Fig. 16: S. (Tropocorixa) distorta Distant, left paramere.
62 P.-p. Chen & N. Nieser
comb of pegs; right paramere (Fig. 14) with a ventral bulge halfway, this latter characteristic separates this species from other Sigara in SE Asia. Distribution. NE India, Burma, Thailand, Indonesia (Sumatra) and Korea (Hutchinson 1940; Jansson 1995).
ACKNOWLEDGEMENTS We would like to thank the Uyttenboogaart-Eliasen Stichting for the promotion of entomological research, Amsterdam, The Netherlands, for two grants to the authors supporting the collecting trip to Thailand in 2004/2005. We are indebted to the following persons from universities and the research institute for their warm hospitality and the great help during our field work in Thailand: C. Kasemset, J. Kaewthong, Family P. Leksawasdi, M. Isenstadt, W. Sommit (Chiang Mai), S. Jinahyon, W. Sirichana, R. Thapa (Chiang Rai), T. Jamganya (Khon Kaen University), S. Wongsiri (Bangkok).
РЕЗЮМЕ Описан е нов за науката вид хетероптера Sigara (Pseudovermicorixa) josifovi от Тайланд. Съобщени са три нови за фауната на страната представители от същото семейство. Даден е ключ за тайланските видове от семейство Corixidae.
REFERENCES Champion G.C. (1901). Insecta Rhynchota (Hemiptera-Heteroptera) 2. — In: Godman, F.D. & O. Salvin (Eds.): Biologia Centrali Americana, London: I-xvi, 345-416. Chen P.-p, N. Nieser & H. Zettel (2005). The aquatic and semiaquatic bugs (Heteroptera: Nepomorpha & Gerromorpha) of Malesia. — Fauna Malesiana Handbooks 5: i-x, 1-546. Distant L., (1911). The Fauna of British India including Ceylon and Burma. Rhynchota 5. — Taylor & Francis, London: i-xii + 1-362. Fabricius J.C. (1803). Systema Rhyngotorum. — Reichard, Burnsvigae: i-iv, 1-314. Hungerford, H.B., 1948. The Corixidae of the Western Hemisphere (Hemiptera). — The University of Kansas Science Bulletin, 32: 1-827. Hutchinson G.E., (1940). A revision of the Corixidae of India and adjacent regions. — Transactions of the Connecticut Academy of Arts and Sciences , 33: 339-476, pls 1-36. Jaczewski T. (1962a). Notes on some Corixidae (Heteroptera) from Viêt-Nam. — Bulletin de l’Académie Polonaise des Sciences, 10: 23-28. Jaczewski T. (1962b). Pseudovermicorixa subg. n. in the genus Sigara Fabr. (Heteroptera, Corixidae).— Bulletin de l’Académie Polonaise des Sciences, 10: 181-184. Jaczewski T. (1968). Notes concerning the taxonomy and nomenclature of some Corixidae (Heteroptera). — Bulletin de l’Académie Polonaise des Sciences, 16: 299-301.
The Corixidae (s. str.) of Thailand, with description of a new species of Sigara (Heteroptera, Nepomorpha) 63
Jansson A. (1986). The Corixidae (Heteroptera) of Europe and some adjacent regions. — Acta Entomologica Fennica, 47: 1-94. Jansson A. (1995). Family Corixidae Leach, 1815 - water boatmen. — In: Aukema, B. & C. Rieger (Eds.). Catalogue of the Heteroptera of the Palaearctic Region 1. The Netherlands Entomological Society: 26-56. Lundblad O. (1928). Beitrag zur Kenntnis der Corixiden. — Entomologisk TidskriftI, 48: 219-243. Lundblad O. (1933). Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von Sumatra, Java und Bali. — Archiv für Hydrobiologie, 12, Supplement Tropische Binnengewässer 4: 1-195, 263-498. Matsumura S. (1905). Die Wasser-Hemipteren Japans. — Journal of the Sapporo Agricultural College, 2: 53-66. Nieser N. (2002). Guide to aquatic Heteroptera of Singapore and Peninsular Malaysia IV. Corixioidea. — The Raffles Bulletin of Zoology 50: 263-274. Nieser N. & P.-p. Chen (1999). Sixteen new species of Nepomorpha (Heteroptera) mainly from Sulawesi (Indonesia). — Tijdschrift voor Entomologie, 142: 77-123. Nieser N. & D.N. Padilla Gil (1992). Three new species of Corixidae from Colombia (Heteroptera). — Entomologische Berichten Amsterdam, 52: 38-46. Paiva C.A. (1918). Aquatic Rhynchota from the Southern Shan States. — Records of the Indian Museum, 14: 21-32. Walton G.A. (1940). Walton’s classification of the family. — In: Hutchinson G.E., (1940). A revision of the Corixidae of India and adjacent regions. — Transactions of the Connecticut Academy of Arts and Sciences, 33: 339-476.
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S. Grozeva N. Simov (Eds) 2008 MtDNA diversity and phylogeography of five & Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 65 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 65-78. © Pensoft Publishers Sofia–Moscow
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) * J. Damgaard Laboratory of Molecular Systematics, Botanical Garden and Museum, The Natural History Museum of Denmark, Sølvgade 83, Opg. S., DK-1307 Copenhagen K, Denmark. E-mail
[email protected]
ABSTRACT Genetic diversity and phylogeography was examined in a survey of 769 bp of COI mtDNA from multiple representatives of five Palaearctic water strider species with different habitat preferences, phenologies and frequencies of various wing morphs: Aquarius najas (De Geer, 1773), A. paludum (Fabricius, 1794); Gerris lacustris (Linnaeus, 1758), G. costae (HerrichSchaeffer, 1850) and G. thoracicus Schummel, 1832. Comparisons of uncorrected p sequence divergences with standard deviations showed significant differences between the five species with Aquarius najas being the most diverse species (>2%), followed by A. paludum and Gerris lacustris with intermediate diversity (~1%), and with G. costae and G. thoracicus showing the lowest sequence diversity (<0.5%). As expected there was a strong negative correlation between the frequencies of macropterous individuals and the amount of sequence divergences. When sequence divergences were displayed as a distance tree it was shown that Aquarius najas showed most spatial variation hereby indicating a high degree of geographical isolation especially around the Mediterranean Sea; Gerris lacustris had deep sequence divergences in Turkey and Armenia, and Aquarius paludum, Gerris costae and G. thoracicus all have shallow divergences and no geographical structure. The results are used to discuss molecular markers, species taxonomy, Pleistocene refugia and postglacial dispersal routes.
* This paper is dedicated to Dr. Michail Josifov for the occasion of his 80th anniversary and recognition of his remarkable contribution to the investigations of Miridae and the Palaearctic fauna of true bugs.
66 J. Damgaard
INTRODUCTION In the northern Hemisphere the Pleistocene glaciations were arguably the most significant historical events to have occurred during the evolutionary lifespan of most extant species (Hewitt 1996, 2001). Freshwater habitats in particular were altered on an unprecedented scale by climatic oscillations leading to destruction of old systems and widespread creation of new lakes and rivers, and limnic organisms were strongly affected because of the island-like nature of their habitats. The combination of phylogenetics and population genetics with biogeography, commonly termed phylogeography, has revolutionized our abilities to investigate the impact of such historical impacts on the population structures of extant species (Avise 2000). So far, most studies of freshwater organisms have addressed economically important fish, and have let to some generalizations the impacts of historical events on the genetic composition and structure of modern populations (Bernatchez & Wilson 1998). However, fishes are powerful dispersers and are highly susceptible to human mediated activities such as pollution, stocking and releases, and may therefore not be representative of freshwater life at large. Water striders (Hemiptera-Heteroptera: Gerridae) are semi-aquatic bugs that offer great potentials for comparative phylogeographic studies for several reasons. Firstly, water striders are relatively large insects that are present on the water surface for their entire life, except for overwintering on land and occasional dispersal flights, thus making them easy to observe, collect and cultivate. Secondly, water striders are not directly dependent of the water quality (oxygen content, temperature, submerged vegetation, substrate etc.) and therefore less affected by human mediated impacts, such as pollution, eutrophication, and habitat changes as compared to other limnic organisms, and may even benefit from creation of new lakes, canals and other water bodies. Four studies have recently addressed the genetic diversity and phylogeography of five Palaearctic water strider species groups: The Aquarius paludum group (Damgaard & Zettel 2003); the A. najas group (Damgaard 2005); the Gerris costae and G. thoracicus groups Damgaard 2006); and the G. lacustris group (Damgaard 2008) (Fig. 1). The five nominal species of these species groups are well known and widely distributed species in Europe and neighboring regions, but also species that differ in habitat preferences, phenologies, and life history adaptations (Vepsäläinen 1973; 1974; Savage 1989). Of greatest interest here is that the five species differ significantly in the frequencies of different wing morphs, meaning that populations or generations of the same species have different dispersal abilities as a result of differences in wing length and/or development of flight musculature (Andersen 1982). While the adaptive significance of wing polymorphism in water striders has been given considerable attention (Ahlroth et al. 1999; Andersen 1973, 1993, 2000), little empirical evidence is known about the influence of dispersal on the overall genetic structure of water strider species. Zera (1981) found significant spatial variation and abundant fixation or near fixation of allele frequencies in the nearly wingless Aquarius remigis (Say, 1832), but
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 67
A. philippinensis A. distanti A. antigone A. fabricii
A. paludum group
A. adelaidis A. lili
A. p. amamiensis A. conformis A. nebularis A. elongatus A. cinereus A. najas A. ventralis
A. najas A. conformis group group
A. p. paludum
G. swakopensis gr. G. costae gr.
G. argenticollis gr. G. latiabdominis gr.
Gerris
G. lacustris gr. G. thoracicus gr. G. marginatus gr.
G. odontogaster gr. G. nepalensis gr. G. gillettei gr. Macrogerris Gerriselloides L. dissortis L. notabilis L. genitalis L. rufoscutellatus
L. rufoscuteatus gr.
“A. remigis gr.”
L. canaliculatus gr. L. esakii gr. G. gigas “A. chilensis gr.”
Fig. 1: Phylogenetic relationships among species, species groups and subgenera of Limnoporus, Aquarius and Gerris based on Damgaard & Cognato (2005).
68 J. Damgaard
geographically homogenous allele frequencies in the wing polymorphic Limnoporus canaliculatus (Say, 1832), both species being widespread in the U.S.A. The present study surveys the genetic diversity of the five Palaearctic species in order to examine the genetic consequences of different dispersal abilities, and searches for congruent phylogeographic patterning, which may indicate common Pleistocene refugia and postglacial dispersal routes.
MATERIALS AND METHODS Presentation of taxa Aquarius paludum (Fabricius, 1794) is one of the most widely distributed water strider species, and is recorded from throughout the Palaearctic Region and extends into the Oriental Region (Aukema & Rieger 1995). A subspecies, A. p. amamiensis (Miyamoto, 1958) is recorded on Okinawa and the Ryukyu Islands south of Japan (Andersen 1990). Aquarius najas (De Geer, 1773) is recorded from throughout Europe and expands into North Africa (Morocco, Tunisia) (Andersen 1990). In the western Mediterranea A. najas is sympatric with A. cinereus (Puton, 1869) (Zimmermann & Scholl 1993), and in southeastern Europe and the Middle East A. najas is replaced by A. ventralis (Fieber, 1861), even though the exact geographical border between the two species is still uncertain (Andersen 1990). Gerris thoracicus Schummel, 1832 is distributed throughout Europe and Central Asia, and expands into the Oriental Region (Aukema & Rieger 1996). G. costae (Herrich-Schäffer, 1850) is distributed in the western part of the Palaearctic region, but is absent from Scandinavia, even though it lives in Germany and on the British Isles (Aukema & Rieger 1995). Andersen (1995) listed the three subspecies costae (Herrich-Schäffer, 1850) from Central Europe, poissoni Wagner & Zimmermann, 1955 from Western Europe and fieberi Stichel, 1938 from southeastern Europe, but Damgaard (2006) questioned the taxonomic validity of these subspecies and down-ranked G. sahlbergi Distant, 1879 from Central Asia to be yet another subspecies of G. costae. Finally, G. lacustris (Linnaeus, 1758) is widespread in the Palaearctic Region from the Atlantic Coast to Korea and China (Aukema & Rieger 1995). Aquarius paludum lives on larger stagnant water bodies, especially oligotrophic lakes, but may also be encountered on canals and slow flowing rivers (Savage 1989; Nieser & Wasscher 1996; Damgaard & Andersen 1996). A. paludum has a macropterous overwintering generation which gives rise to a direct breeding and brachypterous second generation. This generation completes its development during summer, and their eggs develop into macropterous individuals that enter diapause and overwinter. A. najas is one of the few rheophilic water striders in Europe, and are found on larger streams and smaller rivers, but may even be encountered on large, oligotrophic lakes (Brinkhurst 1966; Damgaard & Andersen 1996). It is univoltine throughout most
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 69
of its distribution, but presence of a small number of early instar nymphs late in the season may indicate a partial second generation (Murray & Giller 1990). A. najas is predominantly apterous in northern latitudes (Vepsäläinen 1974; Damgaard & Andersen 1996), but the frequency of macropterous individuals increase with decreasing latitudes (Poisson 1957). Gerris thoracicus is common in small and weedy cattle ponds, ditches, and other artificial habitats, and is often encountered in brackish habitats (Vepsäläinen 1973; Savage 1989). G. thoracicus is predominantly univoltine and macropterous, but brachypterous specimens may indicate a partial second generation (Vepsäläinen 1974). G. costae is found in lentic habitats, often in moorlands and mountain regions, and is predominantly univoltine and macropterous, but brachypterous individuals are sometimes encountered (Southwood & Leston 1959). Finally, G. lacustris lives in both lentic habitats and slowly flowing streams and rivers; is univoltine in northern Europe and at higher altitudes but bivoltine in central- and southern Europe, and is permanently wing dimorphic. Andersen (1993: Table 1) listed the distribution, habitat preferences, voltinism, and wing morph frequencies of most species of Limnoporus, Aquarius and Gerris, and the information of the five relevant species is summarized in Table 1. Table 1. Species
Habitat
Voltinism
Wingmorph
A. najas
Lotic
Univoltine
Mainly brachypterous
A. paludum
Lentic
Bivoltine
Seasonal dimorphic
G. thoracicus
Lentic
Partly bivoltine
Mainly macropterous
G. costae
Lentic
Partly bivoltine
Mainly macropterous
G. lacustris
Lentic
Partly bivoltine
Permanent dimorphic
Individuals sampled Damgaard & Zettel (2003) included 29 specimens of Aquarius p. paludum and one specimen of A. p. amamiensis. Damgaard (2005) included 65 specimens of A. najas; Damgaard (2006) included 20 specimens each of G. thoracicus and G. costae (incl. G. c. sahlbergi); and Damgaard (2008) included 34 specimens of G. lacustris. These publications should be consulted for information about collecting events, DNAextraction numbers and Haplotype numbers. The published taxon samples represent large parts of the species distribution ranges, but new material is here included that further expand the geographical sampling of each species: G. costae and G. lacustris from the British Isles; G. costae from Sicily; Aquarius najas from Sardinia and A. paludum from Northern Europe, the Middle East, Japan, Taiwan and Laos. More detailed information will appear in a forthcoming publication, but can until then be obtained by contacting the author.
70 J. Damgaard
Characters In the studies of the Aquarius najas-, Gerris costae-, G. thoracicus- and G. lacustris groups (Damgaard 2005, 2006, 2008) amplified and sequenced 772-777 bp of COI by using the primers C1-J-2183 ( Jerry) – TL2-N-3014 (Pat) from Simon et al. (1994), while Damgaard & Zettel (2003) used just 425 bp of the 5´end of COI for the Aquarius paludum group using Jerry and C1-N-2669 from Damgaard et al. (2000a). All new DNA extractions and the original extractions from Damgaard & Zettel (2003) were tested with Jerry – Pat in order to match the sequence data from the other studies. Details about DNA extraction, PCR amplification and cycle sequencing are explained in detail in Damgaard et al. (2000b).
Genetic analyses The sequences were assembled in Sequencher 4.7 (Gene Codes Corporation Inc., Ann Arbor, Michigan) and aligned according to their amino acid sequences. The assembled consensus files were exported as a NEXUS file for later analyses. Uncorrected p genetic distances with standard deviations were calculated in MEGA2 (Kumar et al. 2001) using default options including 500 bootstrap replications and with missing data represented by questionmarks (“?”). The data was then filtered for identical and redundant haplotypes were in MacClade (Maddison & Maddison 1992) and haplotypes were numbered according to previous studies or, in the case of new haplotypes, numbered according to the specimens DNA extraction number(s). In order to make an uncorrected distance tree, the data was analyzed in PAUP* (Swofford 2004) using the neighbor-joining algorithm and uncorrected (p) sequence divergences.
RESULTS Genetic analyses The new specimen of Gerris costae (1799) from the British Isles belongs to haplotype III of this species, which was previously recorded from Schwitzerland, while the new specimen of G. thoracicus (1710) from Sicily belongs to haplotype XVIII of this species, which was previously recorded from Crete (Damgaard 2006). The new specimens of Aquarius najas from Sardinia (1569, 1570) have two new haplotypes compared to Damgaard (2005), and so has the new specimen of G. lacustris (1793) from the British Isles when compared to Damgaard (2008). It is more complicated for Aquarius paludum because the addition of data to already published sequences (Damgaard & Zettel 2003) has shown that specimen 520 from Bulgaria is different from other specimens having haplotype IV and
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 71
3.0 % 2.5 2.0 1.5 1.0 0.5 0.0 A. najas
A. paludum
G. costae
G. thoracicus
G. lacustris
Fig. 2: Uncorrected (p) genetic distances of 769 bp of COI from 62 specimens of A. najas, 45 specimens of A. paludum, 21 specimens of G. costae, 21 specimens of G. thoracicus and 35 specimens of G. lacustris.
therefore represents a unique haplotype. Of the newly sequenced material, specimen 1624 from Japan belongs to haplotype II; 1616 from Iran and 1618 + 1619 from Iraq belong to haplotype IV; 1620 from Denmark and 1622 from Sweden belong to haplotype V; 1612 and 1625 from the Netherlands belong to haplotype VIII and 1615 from Poland belongs to haplotype IX. Specimens 678, 695 and1614 (all Denmark), 1611 (Laos), 1608 (Taiwan), and 1617 (Germany) all have unique haplotypes. The average uncorrected (p) genetic distances including standard deviations are shown in Fig. 2. Aquarius najas has by far the largest genetic divergence reaching an average of 2.2% ± 0.4, followed by Gerris lacustris (1.1% ± 0.3), Aquarius paludum (0.9% ± 0.3), G. thoracicus (0.4% ± 0.2) and G. costae (0.3% ± 0.2). The differences between the most (Aquarius najas), the intermediate (A. paludum, Gerris lacustris), and the least divergent (G. costae, G. thoracicus) species are thus significant. The genetic differentiation within the five species is fully correlated with differences in their wing length frequencies in that the highest diversity is found in the species with the highest frequency of specimens unable to disperse by flight (Aquarius najas) followed by species that are either permanently (Gerris lacustris) or seasonally (Aquarius paludum) wing dimorphic, meaning that at least some individuals can migrate, and with the least diversity found in species that are almost entirely macropterous (Gerris costae and G. thoracicus), meaning that most individuals can migrate. When filtered for identical and redundant haplotypes and displayed as a neighborjoining tree (Fig. 3), it is shown that the genetic divergences are well reflected in the geographical distributions of haplotypes. For Aquarius najas there are very deep divergences between specimens sampled on the Mediterranean islands, especially Crete (Haplotypes XXXII-XXXIV) and the Tyrrhenian Islands (Corsica + Sardinia) (Haplotypes XXX, XXXI, XXVIII, XXXIX and the two new haplotypes from specimens 1569 and 1570), but also considerable differences between populations from Balkan (Haplotypes XXV-XXVII), North Africa (Haplotypes XXII-XXIV), and the Iberian Peninsula including the Balearic Islands (Mallorca) (Haplotypes XV-XXI). The largest cluster of haplotypes includes from northern and Central Europe (Haplotypes I-XIII) but also Haplotype VII, which is found from Scandinavia to northern Italy, and Haplotype XIV from Greece (Nomos Drama Rhodope Mts.) (see Damgaard 2005 for more details). Gerris lacustris has a relatively deep
72 J. Damgaard
Aquarius najas
Aquarius paludum
Gerris costae
Gerris thoracicus
Gerris lacustris
A.najas.VII A.najas.XII A.najas.V A.najas.XI North and A.najas.VI A.najas.VIII Central A.najas.IV A.najas.IX Europe A.najas.I A.najas.II A.najas.III A.najas.X A.najas.XIV A.najas.XIII A.najas.XV A.najas.XVI Iberian A.najas.XVII A.najas.XVIII peninsula A.najas.XIX A.najas.XX A.najas.XXI A.najas.XXV A.najas.XXVI Balkan A.najas.XXVII A.najas.XXIII North A.najas.XXIV A.najas.XXIIAfrica A.najas.XXVIII A.najas.1570* Tyrrhenian A.najas.XXXIX A.najas.XXX Islands A.najas.XXXI A.najas.1569* A.najas.XXXII A.najas.XXXIII Crete A.najas.XXXIV A.paludum.520* A.paludum.1617* A.paludum.VI A.paludum.695* A.paludum.V A.paludum.VII A.paludum.IV A.paludum.678* A.paludum.1608* A.paludum.1611* A.paludum.IX A.paludum.VIII A.paludum.I A.paludum.III A.paludum.1614* A.paludum.II G.costae.V G.costae.III.1799 G.costae.IV G.costae.VI G.costae.I G.costae.II G.costae.XI G.costae.XIII G.costae.VIII G.costae.XII G.costae.VII G.costae.IX G.costae.X G.thoracicus.XVIII.1710 G.thoracicus.XXIII G.thoracicus.XIV G.thoracicus.XVII G.thoracicus.XXII G.thoracicus.XVI G.thoracicus.XX G.thoracicus.XV G.thoracicus.XXI G.thoracicus.XIX G.lacustris.IV G.lacustris.1793* G.lacustris.V Turkey, Europe G.lacustris.II G.lacustris.IX G.lacustris.III G.lacustris.VIII Turkey, Armenia G.lacustris.VI Italy G.lacustris.I Gigantometra gigas
0.001 substitutions/site
Fig. 3: Neighbor joining tree of unique haplotypes. The information of individual haplotypes is available from the publication of Damgaard & Zettel (2003) and Damgaard (2005; 2006; 2008). Haplotypes that have come from the present study are named by their DNA extraction number, and new and unique haplotypes are marked with an asterisk (*).
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 73
divergence between populations from Italy (Haplotype I), Armenia and Turkey (Haplotypes III, VI and VIII) and Turkey and Central and northern Europe (Haplotypes II, IV, V, IX and the new specimen 1793). Gerris costae, G. thoracicus and Aquarius paludum all have relatively shallow divergences and widely distributed haplotypes, especially Aquarius paludum where haplotypes such as V is shared between Sweden and Thailand and VIII found from Central Europe to Japan (see also Damgaard & Zettel 2003).
DISCUSSION COI as a molecular marker Mitochondrial DNA is a popular tool for surveys of genetic differentiation in animals, and the advantages and disadvantages of using mitochondrial markers such as COI has been addressed extensively by Simon et al. (1994), Caterino et al. (2000), and Funk & Omland (2003). Because of the high substitution rate, high copy number and lack of recombination, COI has been implemented as a universal “DNA barcoding” sequence for identification (Hebert et al. 2003) and even description of species (Tautz et al. 2003). However, since mitochondrial DNA is not linked to sexual reproduction, it is considered more susceptible to factors leading to differences in gene and species histories, and both introgression (Abe et al. 2005) and incomplete linage sorting (Damgaard 2005, 2008) have been claimed relevant for the lack of reciprocal monophyly among closely related water strider species. So far, no shared haplotypes are found between different species, meaning that “DNA barcoding” of water striders is possible as long as it is done via pattern recognition and not via phylogenetic analyses (e.g. Petersen et al. 2007).
Historical biogeography and phylogeography Damgaard & Cognato (2005) found that Limnoporus, Aquarius and Gerris, as well as most species groups contained therein most likely had originated in the Palaearctic Region and then dispersed to other zoogeographic regions. However, the Aquarius paludum group, with geographically highly disjunct taxa in the Palaearctic, Oriental, Afrotropical and Australian regions, may have obtained their present distribution by means of vicariance, which place their basal divergences in the Mesozoic (Damgaard & Zettel 2003). Both Aquarius paludum and Gerris lacustris span most of the Palaearctic Region, while G. costae and G. thoracicus extend deep into Asia. By comparison, Aquarius najas is restricted to the western Palaearctic Region, but is still widely distributed in Europe and North Africa and with populations on most of the larger islands in the Mediterranean Sea (Andersen 1990). The Pleistocene glaciations were undoubtedly the single most important factor in shaping their present genetic diversity in these species, especially by long time isolation in ice-free refugia. Such refugia may be identified by the presence
74 J. Damgaard
of many distinct lineages, while more recently colonized regions will be dominated by a single genetic lineage (Bernatchez & Wilson 1998). For Aquarius paludum, Gerris costae and G. thoracicus there is no geographical substructure giving any information about the localization of such a refugium, while G. lacustris has a deep divergence between populations from Turkey and Armenia, thus indicating the eastern Mediterranea and Caucasus as a refugium. Aquarius najas is much more geographically subdivided, and there is evidence of multiple isolated populations around the Mediterranean Sea, including the Iberian Peninsula, North Africa, Balkan, and some of the Mediterranean islands, including Crete, Corsica and Sardinia. The deep divergences between these areas could result from an accelerated substitution rate in mtDNA in this species, but is more likely to be due to an older diversification that probably predates the onset of the Pleistocene. It is not unlikely that A. najas populations on some of these islands are very old and relictual, and may even qualify as distinct species as already been suggested by Zimmermann & Scholl (1993) and Damgaard (2005). According to Andersen (1998) a fossil, A. lunpolaensis (Lin) from Miocene Tibet (5-20 mya) is superficially very similar to A. najas, indicating that the A. najas group was distributed far more to the east than seen today. The nesting of a Mallorcan specimen (Haplotype XVIII) within the cluster from the Iberian Peninsula shows that the Balearic Islands were probably colonized after the separation from the Tyrrhenian Islands approximately 20 mya (De Jong 1998). It was mentioned earlier that populations from both Greece and Italy are nested within the cluster of more northern sequences, thus indicating that one or both of these regions was the source of the lineage(s) that colonized northern and Central Europe after the end of the glaciation periods. In order to get a better localization of such refugia and get a clearer picture of the postglacial dispersal routes, more material of all species should be included from areas that are still not sampled. Of particular interest is Aquarius najas from southern Italy including Sicily and from the Balkans, which may also show hybridization zones between different lineages and therefore potential reproductive barriers between different cryptic species. For other species, especially Gerris thoracicus from northern India, North Africa, the Canary Islands and Madeira, and G. lacustris from East Asia and North Africa are desirable in order to inspect whether one or more of these regions played a role as refugia during the glaciation periods.
CONCLUSIONS The correlation between dispersal abilities and genetic differentiation is best explained by a high frequency of macropterous individuals ensures an effective mixture of populations, while a low frequency of macropterous individuals leads to genetic isolation and differentiation of populations as a result of mutation, natural selection, and genetic drift (Avise et al. 1987). The differences in dispersal abilities among the five selected water
MtDNA diversity and phylogeography of five Palaearctic water striders (Hemiptera-Heteroptera: Gerridae) 75
strider species are also shown in the phylogeographical patterning of different mitochondrial lineages between species, with a clear geographical subdivision in Aquarius najas; less so in Gerris lacustris, and more or less panmixing in Aquarius paludum, Gerris costae and G. thoracicus. There is no doubt that the allocation of resources from the development of dispersal abilities and to the development of eggs (the “flight-oogenesis syndrome” of Johnson 1969) gives a selective advantage within populations, but also the risk of extinction if the quality of water body changes dramatically (Andersen 1993, 2000). Although still poorly understood, these differences in dispersal abilities are probably determined by a combination of genetic and environmental factors, mainly changes in day-length and temperature (Andersen 1973; Ahlroth et al. 1999). For Aquarius najas a few macropterous individuals have been found in northern Europe (Vepsäläinen 1974; Huldén 1979; Damgaard & Andersen 1996) and it has been demonstrated that development under high temperature and short day-length can induce the macropterous morph in otherwise micropterous Finnish populations (Ahlroth et al. 1999). Even though average temperature in northern Europe has changed periodically since the end of the Pleistocene glaciations, the average day-length has not, and it is therefore still an open question how A. najas managed to colonize vast areas of Northern Europe, when the predominantly macropterous Gerris costae could not.
ACKNOWLEDGEMENTS The material upon which this study is based was generously donated by the following colleagues and friends: Nils Møller Andersen, Filippo Maria Buzetti, Anthony I. Cognato, Victor Diaz, Hans Dreisig, Koen de Gelas, Eric Guilbert, Martin Bay Hebsgaard, Mogens Holmen, Reza Hossseini, Javier Juste, Mark Kalashan, Ole Karsholt, KlausDieter Klass, Finn E. Klausen, Petr Kment, Philippe Magnien, Ole Martin, Robert Merritt, Verner Michelsen, Bo Vest Pedersen, Børge Pedersen, Jan Pedersen, Xana Sá Pinto, Lisbeth S. Poulsen, Richard Preziosi, Wolfgang Rabitsch, Ignacio Ribera, Birgit C. Schlick-Steiner, Ole Seberg, Thomas J. Simonsen, Nikolai Simov, Florian M. Steiner, Jurga Turcinaviciene, Jitka Villimova and Herbert Zettel. The study was supported by grants from the Carlsberg Foundation (grant no. 0254/60), the VILLUM KANN RASMUSSEN FOUNDATION, and the Danish Natural Science Research Council.
РЕЗЮМЕ Проведено е изследване на 769 bp COI mtDNA от многобройни представители на пет палеарктични вида от сем Gerridae с различни хабитатни предпочитания, фенология и вариабилност на формата на крилата. Изследвани са генетичното разнообразие и филогеографията на петте вида: Aquarius najas (DE GEER, 1773), A. paludum (FABRICIUS, 1794); Gerris lacustris (LINNAEUS, 1758), G. costae (HERRICH-SCHAEFFER, 1850) and G. thoracicus
76 J. Damgaard
SCHUMMEL, 1832. Сравнението на некоригираните p на дивергенциите на секвенциите със стандартното отклонение показа статистически значими различия между петте вида - Aquarius najas показа най-голямо отклонение (>2%), следван от A. paludum и G. lacustris с междинно отклонение (~1%), и G. costae и G. thoracicus с най-малки отклонения (<0.5%). Както можеше да се очаква, най-силна отрицателна корелация имаше между честотите на пълнокрилите индивиди и степента на дивергенция на секвенциите. Когато дивергенциите бяха представени като дендрограма стана ясно, че Aquarius najas има най-голяма пространствена изменчивост – показател за висока степен на географска изолация, особено в района на Средиземно море; Gerris lacustris показва значителна изменчивост на секвенциите в Турция и Армения, а Aquarius paludum, Gerris costae и G. thoracicus имат незначителна дивергенция и липса на географска структура. Резултатите за използвани за обсъждане на молекулярните маркери, таксономията на видовете, плейстоценските рефугиуми и пътищата на постглациално разселване.
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Avise J.C., J. Arnold, R. Ball, E. Bermingham & T. Lamb (1987) Intraspecific phylogeography: the mitochondrial DNA bridge between population genetics and systematics. — Annual Review of Ecology and Systematics, 18: 489-522. Avise J.C. (2000) Phylogeography: The history and formation of species. — Cambridge, MA: Harvard University Press: 1-477. Bernatchez L. & C.C. Wilson (1998) Comparative phylogeography of Nearctic and Palearctic fishes. — Molecular Ecology, 7: 431-452. Brinkhurst R.O. (1966) Population dynamics of the large pond skater A. najas DeGeer (Hemiptera-Heteroptera). — Journal of Animal Ecology, 35: 13-25. Caterino M.S., S. Cho & F.A.H. Sperling (2000) The current state of insect molecular systematics: a thriving Tower of Babel. — Annual Review of Entomology, 45: 1-54. Damgaard J. (2005) Genetic diversity, taxonomy and phylogeography of the western Palaearctic water strider Aquarius najas (De Geer) (Heteroptera: Gerridae). — Insect Systematics and Evolution, 36: 395-406. Damgaard J. (2006) Phylogeny and mtDNA phylogeography of two widespread European pond skater species (Hemiptera-Heteroptera: Gerridae: Gerris Fabricius). — Insect Systematics and Evolution, 37: 335-350. Damgaard J. (2008) MtDNA diversity and species phylogeny of western Palaearctic members of the Gerris lacustris group (Hemiptera-Heteroptera: Gerridae) with implications for “DNA barcoding” of water striders. — Insect Systematics and Evolution, 39: 7-12. Damgaard J. & N.M. Andersen (1996) Distribution, phenology, and conservation status of the larger water striders in Denmark. — Entomologiske Meddelelser, 64: 289-306 Damgaard J., N.M.Andersen, & Sperling F.A.H. (2000a) Phylogeny of the water strider genus Aquarius Schellenberg (Heteroptera: Gerridae) based on nuclear and mitochondrial DNA sequences and morphology. — Insect Systematics and Evolution, 31: 71-90. Damgaard J., N.M. Andersen, L. Cheng & F.A.H. Sperling (2000b) Phylogeny of sea skaters, Halobates Eschscholtz (Hemiptera, Gerridae), based on mtDNA sequence and morphology. — Zoological Journal of the Linnaean Society, 130: 511-526. Damgaard J. & A.I. Cognato (2005) Phylogeny and reclassification of species groups in Aquarius Schellenberg, Limnoporus Stål and Gerris Fabricius (Insecta: HemipteraHeteroptera, Gerridae). — Systematic Entomology, 31: 93-112. Damgaard J. & H. Zettel (2003) Genetic diversity, species phylogeny and historical biogeography of the Aquarius paludum group (Heteroptera: Gerridae). — Insect Systematics and Evolution, 34: 313-328. De Jong H. (1998) In search of historical biogeographic patterns in the western Mediterranean terrestrial fauna. — Biological Journal of the Linnean Society, 65: 99-164. Funk D.J. & K.E. Omland 2003 Species level paraphyly and polyphyly: Frequency, causes, and consequences, with insights from animal mitochondrial DNA. — Annual Review of Ecology, Evolution and Systematics, 34: 397-423. Hebert P.D.N., A. Cywinska, S.L. Ball & J.R. DeWaard (2003) Biological identifications through DNA barcodes. — Proceedings of the Royal Society, Biological Sciences, Series B, 270: 313-321. Huldén L. (1979) Gerris najas (De Geer 1773) (Heteroptera, Gerridae) populationsdynamik och utbredningshistorie i Finland. — M.sc. thesis, Helsingfors Universitetets Zoologiska Institution.
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Hewitt G.M. (1996) Some genetic consequences of ice ages, and their role in divergence and speciation. — Biological Journal of the Linnaean Society 58: 247-276. Hewitt G.M. (2001) The genetic legacy of the Quaternary ice ages. — Nature, 405: 907-913. Johnson C.G. (1969) Migration and Dispersal in Insects by Flight. Methuen, London, 763 pp. Kumar S., K. Tamura, I.B. Jakobsen & M. Nei (2001) MEGA2: Molecular Evolutionary Genetics Analysis software. — Bioinformatics, 17 (12): 1244-1245. Maddison W.P. & D.R. Maddison (1992) MacClade: Analysis of Phylogeny and Character Evolution, version 3.05. — Sunderland, Massachusetts. Murray A.M. & P.S. Giller (1990) The life-history of Aquarius najas De Geer (Hemiptera: Gerridae) in southern Ireland. — Entomologist, 109: 53-64. Nieser N. & M. Wasscher (1986) The status of the larger waterstriders in the Netherlands (Heteroptera: Gerridae). — Entomologisches Berichtungen, Amsterdam, 46: 68-76. Petersen F.T., J. Damgaard & R. Meier (2007) DNA taxonomy: How many DNA sequences are needed for solving a taxonomic problem? The case of two parapatric species of louse flies (Diptera: Hippoboscidae: Ornithomyia Latreille, 1802). — Arthropod Systematics & Phylogeny, 65(2): 111-117. Poisson P. (1957) Hémiptéres aquatique. Faune de France, 61. — Fédération Française de Sociétés Naturelles. Paris : 1-263. Savage A.A. (1989) Adults of the British aquatic Hemiptera Heteroptera. A key with ecological notes. — Scientific Publications from the Freshwater Biological Association, 50: 1-173. Simon C., F. Frati, A. Beckenbach, B. Crespi, H. Liu & P. Flook (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. — Annals of the Entomological Society of America, 87: 651-701. Southwood T.R.E. & D. Leston (1959) Land and water bugs from the British Isles. — London & New York: 1-436. Swofford D.L. (2004). PAUP*: Phylogenetic analysis using parsimony (and other methods). — Sinauer Associates, Inc., Sunderland, Massachusetts. Tautz D., P. Arctander, A. Minelli, R.H. Thomas & A.P. Vogler (2003) A plea for DNA taxonomy. — Trends in Ecology and Evolution, 18: 70-74. Vepsäläinen K. (1973) The distribution and habitats of Gerris Fabr. species (Heteroptera, Gerridae) in Finland. — Annales Zoologici Fennici, 10: 419-444. Vepsäläinen, K. (1974) The life cycles and wing lengths of Finnish Gerris Fabr. Species (Heteroptera, Gerridae) in Finland. — Annales Zoologici Fennici, 141: 1-73. Zimmermann M. & A. Scholl (1993) Specific status of Aquarius cinereus (Puton) and A. najas (De Geer) (Hemiptera: Gerridae) and the extent of hybridization in the Mediterranean region. — Entomologica Scandinavica, 24: 197-210. Zera A. (1981) Genetic structure of two species of waterstriders (Gerridae: Hemiptera) with differing degrees of winglessness. — Evolution, 35(2): 218-225.
S. Grozeva & N. Simov (Eds) 2008 A new species of the pilophorine plant bug genus Pilophorus Hahn from Nepal 79 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 79-84. © Pensoft Publishers Sofia–Moscow
A new species of the pilophorine plant bug genus Pilophorus HAHN from Nepal (Heteroptera, Miridae, Phylinae) R.K. Duwal1 & T. Yasunaga2 1
Laboratory of Insect Biosystematics, Division of Entomology, College of Agriculture and Life Sciences, Seoul National University, San 56-1, Silim9-dong, Gwanak-gu, Seoul 151-742, Korea. E-mail:
[email protected] 2 Research Associate, American Museum of Natural History, Central Park W at 79th Street, New York, NY 10024, c/o Nameshi 2-33-2, Nagasaki, 852-8061, Japan. E-mail:
[email protected]
ABSTRACT A new species of the ant-mimetic pilophorine plant bug genus Pilophorus Hahn, P. josifovianus, is described from Kathmandu Valley, Nepal. A finding of this unique species represents the first record of this genus as well as the tribe Pilophorini from Nepal. Keywords: Miridae, Nepal, new species, Pilophorini, Pilophorus, Phylinae.
INTRODUCTION A new species of the ant-mimetic plant bug genus Pilophorus Hahn, belonging to the tribe Pilophorini of the subfamily Phylinae, is herein described from Nepal. The finding of this species represents the first record for the genus as well as the tribe from Nepal. The biological information, that has not been available for any known relatives with the strongly and mesially constricted pronota, is briefly reported, and clear photographic images of the live individuals are also provided. All measurements are given in millimeters. Digital images of live individuals were made by TY, using Canon EOS Kiss Digital camera body + Canon-Olympus mount
80 R.K. Duwal & T. Yasunaga
adapter + Olympus Macrophoto System (Auto Extension Tube with 38 mm macrolens and T10 Ringflash). All type specimens designated in this study were collected by the authors as part of a cooperative program between JICA Senior Volunteer and Natural History Museum, Tribhuvan University, Kathmandu, and will be deposited in the collections of American Museum of Natural History, New York (AMNH); Natural History Museum, Tribhuvan University, Kathmandu (NMTU); Seoul National University, Korea (SNU) and TY personal collection, Nagasaki (YCN).
DESCRIPTION OF A NEW SPECIES Genus Pilophorus HAHN, 1826
Discussion. This ant-mimetic genus currently comprises approximately a hundred species worldwide (Schuh 1995). Most members are known from the Old World tropics and subtropics where many undescribed species appear to remain. The present discovery of a new species in Nepal represents the first record of the genus from this small but species-rich Himalayan country. Within described species of Pilophorus, some members (P. maculatus (Schuh), P. myrmecoides (Carvalho), P. palawana (Schuh) etc.) were previously placed in Bilirania proposed by Carvalho (1956), a synonym of Pilophorus, on the basis of the unique, strongly and mesially constricted pronotum, which results in the more conspicuous ant-mimetic form. The new species found in Nepal evidently is a member of the group in Pilophorus. These conspicuous ant-mimetic species with the modified pronota are usually rare and available from very limited number of material (Schuh 1984, 1989, 1991). During our investigation, we collected the species described below by a light trap, and by sweeping shrubs and herbs. When collected by sweeping plants, all individuals were found with ants (Fig. 5).
Pilophorus josifovianus DUWAL & YASUNAGA nov. sp. (Figs 1-9)
Diagnosis. Recognized by the large size, generally chestnut brown, elongate, ant-mimetic body, distinct necklike constriction of the pronotum, widely castaneous mesocoxa, and structure of the male genitalia. Description. Body generally chestnut brown, antlike; dorsum rather dull, with densely distributed, golden, woolly setae on posterior lobe of pronotum, scutellum, clavus and corium, and partly with silvery scale-like setae on hemelytron. Head blackish brown, shining, with densely distributed, dark, erect setae on vertex and frons; head in front
A new species of the pilophorine plant bug genus Pilophorus Hahn from Nepal 81
below compound eyes more or less castaneous. Antenna chestnut brown; ventral part of segment I and basal 1/3-1/2 of segment II somewhat paler; segment II incrassate, a little flattened, almost equal in length to metafemur; basal half of segment III creamy yellow. Labium shiny brown, slender, sharp, reaching intermediate part between pro- and mesocoxae. Pronotum with a distinct, necklike constriction; anterior lobe dark brown, shining, with a circular depression medially; scutellum with silvery pubescence along lateral margins. Hemelytron castaneous, widely provided with soft, golden vestiture and partly with silvery scale-like setae (Fig. 1), strongly declivous at cuneal fracture especially in ♂; corium darkened posteriorly, with bands of silvery setae; clavus pruinosed or minutely shagreened posteriorly; cuneus shiny dark brown; membrane dark smoky brown. Leg chestnut brown; basal half of procoxa and more than apical 2/3 of metacoxa creamy yellow; all trochanters yellowish brown; apical half of each tibia more or less pale; tarsomere III dark brown. Abdomen shiny dark brown. 2
1
5
3
4
Figs 1-3: Pilophorus josifovianus, ♂. Fig. 4: Ditto, ♀. Fig. 5: Comparison with an ant that cooccurred with P. josifovi.
82 R.K. Duwal & T. Yasunaga
Male genitalia (Figs 6-9): Right paramere small, straight (Fig. 6); left paramere with elongate hypophysis (Fig. 7). Phallotheca broad (Fig. 8). Vesica simply J-shaped, not broadened or narrowed, with a median spine and series of spine-like processes near secondary gonopore (Fig. 9). Measurements (♂/ ♀): Body length 3.16-3.76/ 3.52-3.89; head width across eyes 0.82-0.84/ 0.81-0.86; vertex width 0.32-0.36/ 0.34-0.41; length of antennal segments I- IV: 0.28-0.34, 1.50- 1.66, 0.48-0.52, 0.55-0.60/ 0.30-0.32, 1.44-1.47, 0.43-0.45, 0.480.50; total labial length 1.29/ 1.35; mesal pronotal length 0.99-1.00/ 1.00-1.05; basal pronotal width 1.05-1.18/ 0.93-1.01; width across hemelytron 1.26-1.42/ 1.27-1.34; and length of metafemur, tibia and tarsus: 1.44-1.50, 2.26-2.50, 0.37-0.40/ 1.44-1.49, 2.25-2.40, 0.36-0.41. Holotype ♂: NEPAL, Kathmandu, Samakhusi (Gongabu), light trap, 15. vi. 2005, T. Yasunaga (AMNH). Paratypes: NEPAL. Bhaktapur: Dadhikot, 22. vii. 2006, 1♂, R. Duwal (SNU); same locality and collector, 2. ix. 2006, 1♂ & 8. v. 2007, 1♂ (NMTU, SNU); same locality, 15. viii. 2007, T. Yasunaga (YCN). Kathmandu: same data as for holotype, 1♀ (YCN); same data except for date, 27. vi., 30. vi. & 23. vii. 2005, 1♂2♀ (YCN).
6 7
9
8
Figs 6-9: Male genitalia of Pilophorus josifovianus. Fig. 6: Right paramere. Fig. 7: Left paramere. Fig. 8: Phallotheca. Fig. 9: Vesica.
A new species of the pilophorine plant bug genus Pilophorus Hahn from Nepal 83
Etymology. Named for Dr. Michail Josifov, in celebration of his 80 years birthday; a noun in genitive case. Distribution. Nepal (Kathmandu Valley, between 1,200 m and 1,500 m alt.). Discussion. This new species is similar in general appearance to P. myrmecoides (Carvalho) known from the Philippines. The former can be distinguished from the latter by the larger size, longer antennal segment II, widely chestnut brown mesocoxa, longer hypophysis of the left paramere, broader phallotheca and narrower apex of the vesica. The vesical structure of P. josifovianus resembles that of P. palawana (Schuh) (Palawan, Philippines), but P. josifovianus has the narrower anterior lobe of the pronotum, widely darkened mesocoxa and broadened phallotheca. As mentioned in generic discussion, the present new species was found to have co-occurred with many workers of an unidentified ant in Dadhikot, Bhaktapur (Fig. 10). Overall appearance and size of them are very similar to one another (Fig. 5). The host plant association of this mirid remains unknown. A male was observed to have sucked on plant sap (Figs 2, 3) although the species is supposed to be predominantly predaceous. Collection records suggest P. josifovianus has two generations per year.
Fig. 10: Shrubs along a stream in Dadhikot, Bhaktapur, Nepal, habitat of Pilophorus josifovianus. Several individuals of this new species were collected together with many ants by sweeping the shrubs.
84 R.K. Duwal & T. Yasunaga
ACKNOWLEDGEMENTS Special thanks are due to Dr. R. T. Schuh (AMNH) for his generous support. We are indebted to Dr. K. Shrestha and Assoc. Prof. P. K. Shrestha (NMTU) for encouraging our research activities in Nepal, and to Dr. S. H. Lee (SNU) and Dr. S. Miyamoto (Fukuoka, Japan) for continuing guidance. We are grateful to Drs. S. Grozeva and N. Simov (Sofia, Bulgaria) who kindly invited us to this commemorative volume for Dr. Josifov. Thanks are extended to an anonymous reviewer for helpful comments on the manuscript. This research was partly supported by JICA ( Japan International Cooperation Agency) Senior Volunteer Cooperative Program and Planetary Biodiversity Inventory (PBI) Project organized by Dr. Schuh.
РЕЗЮМЕ Описана е нова за науката мравкоподобна хетероптера от род Pilophorus HAHN, P. josifovianus от долината на Катманду в Непал. Това е първото съобщение за този род и за представители на трибус Pilophorini от Непал.
REFERENCES Carvalho J.C.M. (1956). On a new genus of mirid from the Biliran Island (Hemiptera). — Anais da Academia Brasileira de Ciencias, 28: 215-216. Schuh R.T. (1984). Revision of the Phylinae (Heteroptera, Miridae) of the Indo- Pacific. — Bulletin of the American Museum of Natural History, 177: 1-476. Schuh R.T. (1989). Old World Pilophorini: Descriptions of nine new species with additional synonymic and taxonomic changes (Heteroptera: Miridae: Phylinae). — American Museum Novitates, 2945: 1-16. Schuh R.T. (1991). Phylogenetic, host and biogeographic analyses of the Pilophorini (Heteroptera: Miridae: Phylinae). — Cladistics, 7: 157-189. Schuh R.T. (1995). Plant bugs of the world (Insecta: Heteroptera: Miridae). Systematic catalog, distributions, host list and bibliography. — The New York Entomological Society: xii + 1-1329.
S. Grozeva & genus N. Simov (Eds)(Heteroptera, 2008 Redescription of the Neotropical Aristathlus Reduviidae, Harpactorinae) 85 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 85-103. © Pensoft Publishers Sofia–Moscow
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) * D. Forero1, H.R. Gil-Santana2 & P.H. van Doesburg3 1
Division of Invertebrate Zoology (Entomology), American Museum of Natural History, New York, New York 10024–5192; and Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853–2601, USA. E-mail:
[email protected] 2 Departamento de Entomologia, Instituto Oswaldo Cruz, Avenida Brasil 4365, Rio de Janeiro, 21045-900, Brazil. E-mail:
[email protected] 3 Nationaal Natuurhistorisch Museum, Postbus 9517, 2300 RA Leiden, The Netherlands. E-mail:
[email protected]
ABSTRACT The Neotropical genus Aristathlus Bergroth 1913, is redescribed. Digital dorsal habitus photographs for A. imperatorius Bergroth and A. regalis Bergroth, the two included species, are provided. Selected morphological structures are documented with scanning electron micrographs. Male genitalia are documented for the first time with digital photomicrographs and line drawings. New distributional records in South America are given for species of Aristathlus. Keywords: Harpactorini, Hemiptera, male genitalia, Neotropical region, taxonomy.
INTRODUCTION Reduviidae is the second largest family of Heteroptera with more than 6000 species described (Maldonado 1990). Despite not having an agreement about the suprageneric classification of Reduviidae (e.g., Putshkov & Putshkov 1985; Maldonado 1990), *
This paper is dedicated to Michail Josifov on the occasion of his 80th birthday.
86 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
it is evident that the tribe Harpactorini (Harpactorinae) is the most diversified group (Davis 1969; Maldonado 1990). Members of Harpactorinae are commonly associated with plant crops and some have been studied as biological control agents of crop pests (e.g., Grundy & Maelzer 2000, 2003; Grundy et al. 2000; Jahnke et al. 2003; Nishi et al. 2004; Chandral et al 2005; Grundy 2007; Ishikawa et al. 2007). In the Neotropical Region there are 51 genera of Harpactorini (Maldonado 1990, 1992; Carvalho et al. 2001; Bérenger 2003, 2007). The only and outdated key to the genera of the Neotropics is that of Stål (1872). Despite their importance as crop pest predators, species identification of Harpactorini is sometimes challenging. After Stål’s key (1872), many genera have been described as new, for instance Atopozelus (Elkins 1954a), Carmenula (Maldonado 1992), Coilopus (Elkins 1969), Ecelenodalus (Elkins & Wygodzinsky 1957), Harpactorella (Wygodzinsky 1947), Iquitozelus (Bérenger 2003), Marjoriana (Bérenger 2007), Mucrolicter (Elkins 1962), Notocyrtoides (Carvalho et al. 2001), and Orbella (Maldonado 1987). Only a fraction of the Harpactorini genera have been revised or redescribed in tropical America, for instance, Atopozelus Elkins (Hart 1972b), Atrachelus Amyot & Serville (Elkins 1954b), Castolus Stål (Maldonado 1976b), Diarthrotarsus Bergroth (Wygodzinsky 1948), Doldina Stål (Hussey & Elkins 1955), Erbessus Stål (Wygodzinsky 1947), Harpactor Laporte (Wygodzinsky 1947), Heza Amyot & Serville (Maldonado 1976a, 1983), Ischnoclopius Stål (Hart 1975), Nesocastolus (Maldonado 1993), Notocyrtus Burmeister (Carvalho & Costa 1992, 1993), Sosius Champion (Maldonado & Carpintero 1993), and Sava Amyot & Serville (Coscarón et al. 1999). Hart (1986, 1987) treated the speciose genus Zelus Fabricius for North America and the West Indies, but most of the species distributed in South America remain unpublished (Hart 1972a). Bergroth (1913) described the genus Aristathlus to include two new species, A. imperatorius and A. regalis (Putshkov & Putshkov 1985; Maldonado 1990). Beyond catalog entries, these species have not been mentioned again in the literature, with the exception of Gil-Santana (2007), who recorded Aristathlus from Brazil and documented the pronotal color variation in A. imperatorius. The aim of this paper is to redescribe the genus, add new distributional information, and describe and illustrate male genitalic characters. This information will facilitate the identification of Aristathlus.
MATERIAL AND METHODS Specimens studied are deposited in the following institutions (acronyms follow Evenhuis 2007): American Museum of Natural History, New York (AMNH), Instituto de Ciencias Naturales, Universidad Nacional, Bogotá, Colombia (ICN), Universidade Federal do Rio de Janeiro, Brazil (MNRJ), National Museum of Natural History, Leiden, The Netherlands (RMNH), United States National Museum of Natural History, Washington D.C. (USNM).
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 87
Observations were made with a Nikon SMZ1500 stereoscope, and drawings with a camera lucida attached to it. Measurements were made with a micrometer eyepiece. Scanning electron micrographs (SEM) were taken with uncoated specimens in a Zeiss environmental SEM, EVO 60EP. Digital dorsal habitus images and male genitalia images were taken using a Microptics-USA photomicrographic system, with Infinity K2 lens and CF-1 and CF-4 objectives. All measurements are in millimeters unless otherwise stated. Dissections of the male genitalia were accomplished by removing the pygophore from the abdomen with a pair of forceps, and clearing it in a warm 10% KOH solution. The pygophore was then rinsed in water and dehydrated in 70% ethanol. Structures were then dissected, studied, and drawn in glycerin. The endosoma of the phallus was everted by gently pulling the dorsal endosomal processes with a fine forceps. Locality data of the specimens were georeferenced with GEOLocate (Rios & Bart 2005) and the aid of gazetteers and regional maps of South America. Decimal degree coordinates for these localities were then processed in DIVA-GIS (Hijmans et al. 2007) with a digital elevation model of South America to produce a distribution map of the species. Terminology: Morphological terms mostly follow Davis (1969) and Lent & Wygodzinsky (1979). Reduviidae have four labial segments as observed in other Heteroptera, although the first segment is reduced in most subfamilies (C. Weirauch, pers. com.). The labial segments are numbered in the descriptive parts of the text from second to fourth, corresponding to the visible segments (see e.g., Weirauch & Forero 2007). Terms of pretarsal structures follow Weirauch (2005). Davis (1966) and Carrera & Osuna (1996) are followed for male genitalic terms, and Scudder (1959) for female genitalic terminology. Abbreviations: 1gcx, first gonacoxa; 1gpo, first gonapophysis; bp, basal plates; bstr, base of struts; end, sclerotized processes of the apex of the endosoma; ep, lateral endosomal processes; gpl, gonoplac; mp, median apical process of the dorsal phallothecal sclerite; ptg 8, paratergite 8; pht, dorsal phallothecal sclerite; sp 8, spiracle 8; st 6-7, sternites 6 and 7; syn 9/10, syntergite 9/10.
Aristathlus BERGROTH, 1913 Bergroth 1913: 240 [new genus]; Wygodzinsky 1949: 36 [catalog]; Putshkov & Putshkov 1985: 26 [catalog]; Maldonado 1990: 166 [catalog].
Type species: A. imperatorius Bergroth, 1913 (by original designation). Diagnosis: Recognized by its dark coloration with U-shaped yellow marking on the pronotum; body elongate, much longer than wide, total length 5.6 – 5.9 times longer as humeral width; third (second visible) labial segment the longest, fourth (third visible) the shortest; anterior lobe of pronotum with two rounded lobes on disc, elevated in males; posterior lobe of pronotum without spines; mesepisternum without tubercle on anterior margin; fore and middle femora stout, hind femur less stout; scapus the longest antennal
88 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
segment, pedicel and basiflagellomere together subequal to scapus; pygophore with single median posterior process and elongate, slender parameres; waxy areas on the pleura. Bergroth (1913) noted that Aristathlus is similar to Zelus, but did not mention specifically which characters separate the two genera. Representatives of both genera have elongate bodies, with the head longer than wide, and without spines next to the antenniferous tubercles. A number of genera in the Neotropical Region superficially share the condition of elongate body and head. Apart from Zelus, those genera include Atopozelus, Ischnoclopius, Iquitozelus, and Heza. The structure of the fore and middle femora separate Aristathlus form Zelus and Atopozelus, because they are robust in Aristathlus and long and delicate in Zelus and Atopozelus (Hart 1972a, b, 1986, 1987). Furthermore, Atopozelus does not have parameres (Hart 1972b), which are present in Aristathlus. Aristathlus can be distinguished from Ischnoclopius by the elongate abdomen with parallel margins, and by having the fore femur slightly longer than the middle femur. In Ischnoclopius the abdomen is widened apically and the fore femur is much longer than the middle femur (Hart 1975). Aristathlus is distinguished from Iquitozelus mainly by the parallel abdomen, which in the latter is lobate at the level of the abdominal segments six and seven (Bérenger 2003). Species of Heza have a spatulate abdomen, with the connexivum usually with spines, a spine adjacent to the antenniferous tubercle, the pronotum with spines, and a tubercle on the anterior part of the mesepisternum (Maldonado 1976a). Aristathlus does not have spines next to the antenniferous tubercles, on the pronotum, and the connexivum, and the mesepisternum is flat. Aristathlus is further distinguished from the aforementioned genera by the bifid median apical process of the dorsal phallothecal sclerite (see Figs 6C, F, arrows) and the enlarged, paired endosomal processes (Fig. 6A-F). Redescription: Male: Medium sized (table 1), elongate with parallel margins. Coloration: Black with yellow or pale yellow markings on pronotum and hemelytra (Figs 1A, B). Vestiture: Silver, delicate, long, dense setae on head, thorax, mesoscutum, and metanotum; sparse bristlelike, dark setae on legs; short, dark, and bristlelike setae on hemelytra. Other structures with vestiture as indicated below. Structure: Head: Elongate, slightly shorter than length of pronotum; anteocular portion about 1.7 times as long as postocular one (Fig. 2B); postocular portion gently curved in dorsal view towards the neck (Fig. 2B); interocular sulcus deeply impressed, curved in dorsal view (Figs 2A, B); ocellus elevated, located closer to eye than to other ocellus (Fig. 2A); mandibular plate very small, nearly obscured by antenniferous tubercle; maxillary plate large, quadrangular, concave medially (Fig. 2A); clypeus nearly straight, slightly convex at middle; gena about as large as maxillary plate; gula nearly flat; labrum triangular; labial segments: second (first visible) stout, reaching anterior margin of eye, third (second visible) nearly as stout as second, the longest, reaching the neck, fourth (third visible) the shortest; eyes nearly hemispherical in lateral and dorsal views, neither reaching dorsal nor ventral margin of head in lateral view, ommatidia not protruding (Figs 2A, B); antennal segments: scapus about 3-4 times longer than head, glabrous (Fig. 1A); pedicel about a third of scapus
3.20 3.10 3.40 3.30
A. regalis Male Male Female Female
21.20 20.90 22.10 22.10
3.30 3.45
3.60 3.70 3.40 3.45
4.45 4.35
1.15 1.00 1.00 1.00
1.05 1.00
1.55 1.50 1.58 1.64
1.53 1.65
0.80 0.75 0.78 0.86
0.85 0.80
3.75 3.65 3.80 3.70
4.35 4.50
1.25 1.30 1.35 1.33
1.15 1.20
6.30 6.20 7.25 6.80
5.05 -
LenHd LenPrn LenScut WidHd IntOcDi WidPrn WidScut Ant1
TotLen A. imperatorius Male 24.35 Female 25.40
2.10 2.30 2.04
1.75 -
Ant2
4.70
4.00 -
Ant3
2.04
-
Ant4
1.40 1.35 1.35 1.35
1.35 1.40
Lab2
2.00 1.85 2.00 2.00
1.85 2.15
Lab3
0.55 0.55 0.55 0.50
0.50 0.55
Lab4
Table 1. Measurements of A. imperatorius and A. regalis. Abbreviations: TotLen: total length; LenHd, length of head; LenPrn, length of pronotum; LenScut, length of scutellum; WidHd, width of head; IntOcDi, interocular distance; WidPrn, width of pronotum; WidScut, width of scutellum; Ant1-4, antennal segments 1-4 (i.e., scapus, pedicel, basiflagellomere, distiflagellomere); Lab2-4, labial segments (visible) 2-4.
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 89
90 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
length, subequal in diameter to scapus, with dense, short, bristlelike, black setae, and at least six trichobothria on apical half (Fig. 2H, arrows); basiflagellomere about 0.85 times shorter than scapus, slightly stouter than pedicel, with dense very short golden setae; distiflagellomere the shortest. Thorax: Pronotum slightly longer than wide (Figs 1A, B); disc of anterior lobe globose, mostly glabrous with linear areas of dense setae (Figs 2C, D), longitudinal sulcus deeply impressed, reaching anterior margin of pronotum, anterior lateral angles blunt or produced as a small tubercle; posterior lobe wider at posterior margin, disc flat, paired elevated ridges on anterior portion of disc connecting with globose disc of anterior lobe (Fig. 2D), posterior margin curved, humeral angles broadly curved, gently sloping laterally to transverse sulcus; scutellum (mesoscutum) triangular, broadly blunt apically, excavated medially near anterior margin and laterally on each side, produced as an elevated Y-shaped cuticle; metanotum produced dorsally as a blunt tubercle (Fig. 2D, arrow); propleuron with procoxal cavity slightly produced laterally, coxal suture deeply impressed, dorsal margin with a fovea; mesepisternum with sparse setae (Fig. 2E), more dense on anterodorsal margin (Fig. 2F), usually these setae with waxy secretions; anterior margin of mesepisternum flat, without a raised tubercle (i.e., “plica” of authors) (Fig. 2F); mesepimeron small, concave; suture between mesepimeron and metepisternum sinuate (Fig. 2E), mesothoracic spiracle large, ovoid (Fig. 2G), with inner cuticle modified as acute trichia (Fig. 2G, inset); metepisternum with dorsal carina strongly or barely impressed; metepimeron small, with posterior margin strongly
A
B
Fig. 1: Habitus digital photographs. A. Aristathlus imperatorius, male [Monzon Valley, Peru]. B. A. regalis, male [Sinop, Brazil].
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 91
A
B
C
D
E
F
G
H
Fig. 2: Aristathlus imperatorius, male. SEM. A. Head, right, dorsolateral view. B. Head, dorsal view. C. Pronotum, right, lateral view. D. Pronotum, scutellum, and metanotum, right, dorsolateral view. Arrow points to protuberant metanotum. E. Meso- and metapleura, right, lateral view. F. Detail mesepisternum next to posterior margin of pronotum. G. Mesothoracic spiracle, with insert of detailed inner cuticle. H. Pedicel, lateral view of apical half. Arrows point to insertions of trichobothria.
92 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
concave; prosternum triangular, stridulatory sulcus with transverse fine ridges, apex of sulcus reaching middle of anterior coxa, areas lateral to stridulatory sulcus nearly flat; mesosternum slightly concave medially on anterior half, distal half strongly concave and narrowing apically with margins carinate; metasternum slightly convex. Hemelytra: Long, surpassing the abdomen by about half the length of the membrane; margins parallel (Figs 1A, B); corium with cell between cubitus and postcubitus nearly pentagonal (Fig. 1A). Legs: Fore, middle, and hind coxae globose, slightly constricted apically; trochanters nearly quadrangular; coxae and trochanters densely setose on ventral surface; fore femur elongated, nearly cylindrical, stout, slightly swollen at base, ventrally with dense, medium-sized bristlelike setae, dorsally with long, sparse, bristlelike setae; middle femur with greater diameter at middle than basally or apically, shorter than fore femur, ventrally with dense short bristlelike setae; hind femur cylindrical, slightly curved in dorsal view, barely longer than fore femur, constricted subapically, with sparse short bristlelike setae and few medium-sized bristlelike setae; fore tibia stout, lesser in diameter than fore femur, gently curved, apically bulbous, with dense, short and long bristlelike setae on all surface, setae on apex of tibia pale; middle tibia very similar in length and structure to anterior tibia but less setose; hind tibia longer than middle tibia, similar in structure to middle tibia; tarsi on all three legs with three segments, first tarsal segment the shortest, first and second combined as long as third, all segments densely setose on ventral surface; claw of pretarsus with large basal tooth, acute apically, deeply incised, parempodial setae divergent, apparently expanded apically. Abdomen: Elongate, margins parallel; sternites with incomplete, deeply impressed sutures between them, ending each one in a rugose area; spiracle 1 laterally on tergite 1, spiracles 2-7 medially on respective sternite (Fig. 3A), spiracles circular (Fig. 3B). Genitalia: Pygophore elongate ovoid (Figs 4, 5B, G), slightly expanded laterad at level of insertions of parameres (Fig. 4), ovoid in posterior view (Figs 5A, F), slightly constricted in basal third (Figs 5B, G), with a median process on its posterior margin (Figs 5A, B, F, G); single median process of posterior margin directed dorsad, wide or narrow, with apex carinated and reclined (Figs 5C, D, H, I); eighth sternite; strongly concave medially (Figs 4A, C, 5B, G); paramere elongate, sinuate in lateral view, setose apically (Figs 5E, J); phallus with articulatory apparatus with basal plates strongly curved (Figs 6A, B, D, E, bp), dorsally fused (Fig. 6D) or not (Figs 6A); basal plate bridge strongly sclerotized; dorsal phallothecal sclerite ovoid (Fig. 6A, pht) or elongate ovoid (Fig. 6D), strongly sclerotized, with a median, bifid apical process (Figs 6A, D, mp), posterior margin convex or gently concave (Figs 6A, D); base of the struts fused, elongate (Fig. 6D) or not (6A, bstr); basal part of endosoma with paired, dorsal processes (Figs 6A, D, ep), stout, strongly sclerotized and curved laterally, projecting beyond apex of endosoma (Figs 6B, C, E, F), apically notched; apex of endosoma with numerous small, triangular, sclerotized processes (Figs 6B, C, E, F, end), apparently forming dorsal and ventral sacs. Female: Similar to male in coloration, vestiture, and structure, except the following. Structure: Thorax: Anterior lobe of pronotum less elevated than in male. Hemelytra: Longer than in male. Legs: Hind tibia curved ventrally, slightly flattened, and enlarged
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 93
subbasally (Fig. 3E). Abdomen: Spiracle 8 on paratergite 8 (Fig. 3D). Genitalia: Syntergite 9/10 nearly vertical, part corresponding to tergite 10 protuberant, with ventral margin slightly concave (Fig. 3C); first gonacoxa wide (Fig. 3D); first gonapophysis narrow, reaching basal third of first gonacoxa (Fig. 3D); gonoplac paired, not projecting posteriorly (Fig. 3D). Measurements: As in Table 1. Discussion: The type specimens of A. imperatorius and A. regalis were not located. Putshkov & Putshkov (1985) listed in their catalog the type depositories of several spe-
A
B
C
E D Fig. 3: Aristathlus imperatorius, male. SEM. A. Sternites 6 and 7, lateral view. B. Spiracle on sternite 3. Aristathlus regalis, female. C. Syntergite 9/10,posterior view. D. Apex of abdomen, lateroventral view. E. Left, hind tibia. Scales as indicated. Abbreviations: 1gcx, first gonacoxa; 1gpo, first gonapophysis; gpl, gonoplac; ptg 8, paratergite 8; sp 8, spiracle 8; st 6-7, sternites 6 and 7; syn 9/10, syntergite 9/10.
94 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
cies, although they did not cited any information regarding the two species of Aristathlus. The types are neither at the Finnish Museum of Natural History in Helsinki (L. Huldén, pers. com.) nor at the Muséum National d’Histoire Naturelle in Paris (E. Guilbert, pers. com.), where Bergroth deposited some of his types. Because an extensive search for the types has not been carried out in other European collections, and because the identity of those species is not in doubt, we are not designating in this paper neotypes for both Aristathlus species.
A Aristathlus imperatorius
C
D Aristathlus regalis
Fig. 4: Pygophore. A-B: Aristathlus imperatorius. A. Ventral view. B. Dorsal view. C-D: Aristathlus regalis. C. Ventral view. D. Dorsal view.
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 95
Males and females have resinous sticky substances on all tibiae, a fact that already Bergroth (1913) noted. We do not know the origin of this substance, but it is probably produced by the insects and secreted by glandular pores in the tibiae as documented for the anterior tibia of Zelus luridus Stål (Weirauch 2006). An unidentified species of Aristathlus in French Guiana was associated with Aparisthmium cordatum (Euphorbiaceae) (Bérenger & Pluot-Sigwalt 1997), another probable source of the resinous substances on the legs.
C
D
A
E
Aristathlus imperatorius
B
H
I
F
J
G
Aristathlus regalis
Fig. 5: A-E: Aristathlus imperatorius. A. Pygophore, posterior view. B. Pygophore and sternite eight, lateral view. C. Median process of pygophore, lateral view. D. Median process of pygophore, posterior view. E. Right paramere, lateral view. F-J: Aristathlus regalis. F. Pygophore, posterior view. G. Pygophore and sternite eight, lateral view. H. Median process of pygophore, lateral view. I. Median process of pygophore, posterior view. J. Right paramere, lateral view.
96 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
Bergroth (1913) compared the particular structure of the flattened and expanded hind tibia in the females with that of Ixopus Bergroth. This structure also resembles that of Graptoclopius Stål and some species of Notocyrtus [e.g., N. dorsalis (Gray), N. camelus Stål, N. clavipes (Fabricius), see Carvalho & Costa (1993)]. The function of this structure is unknown. Species of Aristathlus are probably mimetic of wasps with metallic wings (Bergroth 1913). Distribution: Aristathlus is found exclusively in tropical areas of South America (Fig. 7). Aristathlus imperatorius and A. regalis are sympatric in their distribution, sometimes co-occurring in the same locality.
Aristathlus imperatorius BERGROTH, 1913 Bergroth 1913: 241 [new species]; Wygodzinsky 1949: 36 [catalog]; Putshkov & Putshkov 1985: 26 [catalog]; Maldonado 1990: 166 [catalog]; Gil-Santana 2007: 61 [distribution, color variation].
Diagnosis: Distinguished by the black hemelytra without yellow markings (Fig. 1A); mesosternum medially pale brown; metasternum brown; middle femur swollen at middle (Fig. 1A); head and thorax densely setose (Figs 1A, 2A, B); metepisternum with dorsal carina strongly impressed; basal plates of articulatory apparatus not fused dorsally (Fig. 6A); base of the struts short (Fig. 6A); dorsal phallothecal sclerite ovoid (Fig. 6A); median, apical process of dorsal phallothecal sclerite strongly acute (Figs 6A, C, arrow). Redescription: Male: Coloration: Head: Dark brown; gula pale brown. Thorax: Pronotum with anterior lobe dark brown, paler on areas with dense setae; posterior lobe dark brown with an U-shaped pale yellow marking from the posterior margin of the pronotum and humeral angles projecting anteriorly to the transverse sulcus (Fig. 1A); raised cuticle of scutellum pale yellow; metanotum apically pale yellow; posterior portion of anterior coxal cavity and adjacent posterior pleural portion of pronotum pale yellow, remaining area dark brown; dorsal area of coxal suture of middle and posterior legs pale brown; stridulatory sulcus pale yellow; sternites brown, central area of mesosternum pale brown. Hemelytra: Black with metallic blue hues. Legs: Dark brown; tibiae apically darker; tarsi black. Abdomen: Tergites and laterotergites black; sternites and laterosternites red yellow. Genitalia: Pygophore and parameres black. Structure: Thorax: Anterolateral angles of pronotum blunt; dorsal carina of metepisternum strongly impressed. Legs: Middle femur strongly expanded medially (Fig. 1A). Genitalia: Basal plates of articulatory apparatus not fused dorsally (Fig. 6A); base of the struts short (Fig. 6A); dorsal phallothecal sclerite ovoid (Fig. 6A); median, apical process of dorsal phallothecal sclerite strongly acute (Figs 6A, C, arrow); dorsal endosomal processes not strongly stout (Figs 6A, B, C). Female: Similar in structure and coloration to male. Coloration: Genitalia: Syntergite 9/10 black, adjacent parts of first gonacoxa and gonapophysis black.
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 97
Variability: The pronotal color pattern is variable, ranging from a narrow, median U-shaped yellow marking, to a wider marking reaching the humeral angles (Gil-Santana 2007). The metanotum is sometimes black, not yellow. Distribution: Aristathlus imperatorius was originally described from Cayenne, French Guiana (Bergroth 1913, Maldonado 1990). Gil-Santana (2007) recorded this species from Brazil (Amazonas, Mato Grosso, and Pará). Specimens from Colombia and Peru represent new country records (Fig. 7). Material examined: BRAZIL, 1♂, Amazonas, Manaus, 14 nov 1955, Elias & Roppa leg. (MNRJ); 1♂, Mato Grosso, Diamantino, Arinos River [14° 22’ S - 56° 07’ W], 16 nov 2002, E. Furtado leg. (MNRJ); 1♀, Pará, Óbidos, may 1956, F. Oliveira leg., [ex] Campos Seabra collection (MNRJ); 1♀, [no specific locality], Mus. Goeldi 17/10, D01 / “Aristathlus imperatorius det. Wygod.” (AMNH). COLOMBIA, 1♂, Meta, Río Ocoa, selva, 100 m, 20 may 1945, L. Richter (ICN). PERU, 1♂, [Huanuco], Tingo María, Monzón Valley, 23 sep 1954, E.L. Schlinger & E.S. Ross collectors / “Aristathlus imperatorius det. Wygod.” (AMNH).
end ep mp pht bstr bp
A
B
D
E
mp
C
F Aristathlus imperatorius
Aristathlus regalis
Fig. 6: Phallus, nearly completely everted. A-C: Aristathlus imperatorius. A. Dorsal view. B. Ventral view. C. Lateral view. D-E: Aristathlus regalis. D. Dorsal view. E. Ventral view. F. Lateral view. Scales as indicated. Abbreviations: bp, basal plates; bstr, base of struts; end, sclerotized processes of the apex of the endosoma; ep, lateral endosomal processes; mp, median apical process of the dorsal phallothecal sclerite; pht, dorsal phallothecal sclerite. Arrows points to the difference between the mp in the two species.
98 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
Aristathlus regalis BERGROTH, 1913 Bergroth 1913: 241 [new species]; Wygodzinsky 1949: 36 [catalog]; Putshkov & Putshkov 1985: 26 [catalog]; Maldonado 1990: 166 [catalog]; Gil-Santana 2007: 61 [distribution].
Diagnosis: Distinguished by the yellow clavus (Fig. 1B); transverse yellow band on corium anterior to the membrane (Fig. 1B); mesosternum medially yellow, metasternum yellow; middle femur not greatly expanded at middle, nearly cylindrical (Fig. 1B); head and thorax not densely setose (Fig. 1B); metepisternum with dorsal carina not strongly impressed; basal plates of articulatory apparatus fused dorsally (Fig. 6D); base of the struts elongate (Fig. 6D); dorsal phallothecal sclerite elongate ovoid (Fig. 6D); median, apical process of dorsal phallothecal sclerite rounded, not acute (Figs 6D, F, arrow). Redescription: Male: Coloration: Head: Dark brown; base of maxillary plate pale yellow; apex of gena pale; gula yellow; postocular portion with faint, longitudinal yellow line. Thorax: Anterior lobe of pronotum dark brown, apex of disc and longitudinal sulcus yellow, lateral margin with narrow yellow line; posterior lobe dark brown; with a U-shaped yellow marking, posterior margin of pronotum yellow; scutellum yellow, metanotum yellow; anterior coxal cavity yellow and adjacent posterior pleural portion of pronotum forming a yellow longitudinal area, remaining area dark brown; meso and metapleura yellow; coxal cavities of middle and hind legs pale yellow; stridulatory sulcus yellow, prosternal lateral areas dark brown; mesosternum medially yellow, laterally dark brown; metasternum yellow. Hemelytra: Dark brown; clavus yellow; apical half of corium yellow. Legs: Dark brown, nearly black, except for trochanters ventrally yellow. Abdomen: Yellow. Genitalia: Apical half of pygophore with inverted U-shaped black marking; parameres dark brown. Structure: Thorax: Anterolateral angles of pronotum produced as a small tubercle; dorsal carina of metepisternum not raised. Legs: Middle femur not strongly swollen medially (Fig. 1B). Genitalia: Basal plates of articulatory apparatus fused dorsally (Fig. 6D); base of the struts elongate (Fig. 6D); dorsal phallothecal sclerite elongate ovoid (Fig. 6D); median, apical process of dorsal phallothecal sclerite rounded, not acute (Figs 6D, F, arrow); dorsal endosomal processes strongly stout (Figs 6D, E, F). Female: Similar to male in coloration and structure, except for the following. Coloration: Abdomen: Pale brown. Genitalia: Syntergite 9/10 dark brown. Variability: Some specimens may have a reddish rather than yellow marking on the pronotum. Distribution: Aristathlus regalis was originally described from Cayenne, French Guiana (Bergroth, 1913). Gil-Santana (2007) recorded it from Brazil (Amazonas, Mato Grosso, and Pará). It is recorded for the first time from Colombia and Suriname. Additional localities are given from Brazil (Fig. 7). Material examined: BRAZIL, 1♀, Amazonas, Tabatinga, aug-1955, E. S. Lima leg., [ex] Campos Seabra collection, “Aristathlus cf. regalis Bergroth Wygodzinsky det.”
Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae) 99
A. imperatorius A. regalis Fig. 7: Distribution map of species of Aristathlus.
(MNRJ); 1♂, Pará, Óbidos, Coleção Campos Seabra, 1955, F.M. Oliveira / “Aristathlus cf. regalis Berg. det. Wygodzinsky” (AMNH); 1♂, 2♀♀, Óbidos, 1955, F.M. Oliveira leg., [ex] Campos Seabra collection, “Aristathlus cf. regalis Bergroth Wygodzinsky det.” (MNRJ); 1♂, 1 adult without abdomen, “Para” [without specific locality]/ PR Uhler collection / Aristathlus regalis Bergr. H.G. Barber” (USNM); 1♂, 1♀, Mato Grosso, 12º31’S 55º37’W, Sinop, Oct-1975, M. Alvarenga (AMNH); 1♂, Diamantino, Arinos River [14° 22’ S - 56° 07’ W], 20 oct 1983, Johann Becker leg. (MNRJ). COLOMBIA, 1♂, Amazonas, Leticia, Km 11 via Tarapacá, 100m, Sistematica Animal col. / 1.3.003 (ICN). SURINAME, 1♀, Brownsberg, 720 m, 18 jan 1972, G.F. Mees (RMNH). List of acronyms used in the text: AMNH – American Museum of Natural History, New York, USA. ICN – Instituto de Ciencias Naturales, Universidad Nacional, Bogota, Colombia. MNRJ – Museo Nacional, Rio de Janeiro, Brazil. RMNH – Nationaal Natuurhistorisch Museum, “Naturalis”, Leiden, The Netherlands. USNM – United States National Museum, Smithonian Institution, Washington D.C., USA.
100 D. Forero, H.R. Gil-Santana & P.H. van Doesburg
ACKNOWLEDGEMENTS We are grateful to Randall T. Schuh (AMNH), Thomas J. Henry (USNM), Luiz A. A. Costa (MNRJ), and Fernando Fernández (ICN) for facilitating the examination of the Reduviidae collection under their care. Marianna Teräväinen and Larry Huldén (Finnish Museum of Natural History, Helsinki), and Eric Guilbert (Muséum National d’Histoire Naturelle, Paris) kindly searched for Bergroth’s type specimens. Christiane Weirauch (University of California, Riverside) and Dávid Rédei (Hungarian Natural History Museum) kindly reviewed the manuscript and provided useful comments.
РЕЗЮМЕ Преописан е неотропичният род Aristathlus BERGROTH, 1913. Представени са дигитални фотографии на дорзалния хабитус на двата изследвани вида: A. imperatorius BERGROTH и A. regalis BERGROTH. Избрани морфологични структури са документирани със микроснимки под сканиращ микроскоп. За първи път са документирани мъжките гениталиии чрез дигитални микрофотографии и чрез рисунки. Представени са нови данни за разпространението на видовете от род Aristathlus в Южна Америка.
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Carrera Z. & E. Osuna (1996). Morfología de la genitalia externa masculina de la tribu Harpactorini (Reduviidae: Harpactorinae). Parte 1a.: Morfología general del falo. — Boletín de Entomología Venezolana N.S., 11(1): 1–9. Chandral S., R.L. Sinazer & S.P. Kumar (2005). Influence of prey on the development, reproduction and size of the assassin bug, Sphedanolestes minusculus Bergroth (Heteroptera: Reduviidae), a potential biological control agent. — Journal of Entomological Research, 29(2): 93–98. Coscarón M.C., E. Barrera & H. Brailovsky (1999). Redescription of Sava Amyot & Serville 1848 (Heteroptera: Reduviidae: Harpactorinae). — Proceedings of the Entomological Society of Washington, 101(4): 827–831. Davis N.T. (1966). Contributions to the morphology and phylogeny of the Reduvioidea (Hemiptera: Heteroptera). Part III. The male and female genitalia. — Annals of the Entomological Society of America, 59(5): 911–924. Davis N.T. (1969). Contribution to the morphology and phylogeny of the Reduvioidea. Part IV. The Harpactoroid complex. — Annals of the Entomological Society of America, 62(1): 74–94. Elkins J.C. (1954a). A new American Harpactorine [sic] genus (Hemiptera, Reduviidae). — Texas Reports on Biology and Medicine, 12(1): 39–48. Elkins J.C. (1954b). A synopsis of Atrachelus (Hemiptera, Reduviidae). — Proceedings of the Entomological Society of Washington, 56(3): 97–120. Elkins J.C. (1962). A new Harpactorinae genus (Hemiptera: Reduviidae). — Journal of the Kansas Entomological Society, 35(4): 360–363. Elkins J.C. (1969). A new genus of Hemipteran wasp mimics (Reduviidae, Harpactorinae). — Journal of the Kansas Entomological Society, 42(4): 456–457. Elkins J.C. & P. Wygodzinsky (1957). A new neotropical Harpactorine genus (Reduviidae, Hemiptera). — Revista Brasileira de Biologia, 17(2): 203–210. Evenhius N. (2007). Insect and spider collections of the world web site. — Available from: http:// www.bishopmuseum.org/bishop/ento/codens-r-us.html. Date of last revision: November 2, 2007. (Accessed November 2007). Gil-Santana H.R. (2007). New records of Reduviidae (Hemiptera: Heteroptera) from Brazil. — Zootaxa, 1390: 59–68. Grundy P.R. (2007). Utilizing the assassin bug, Pristhesancus plagipennis (Hemiptera: Reduviidae), as a biological control agent within an integrated pest management programme for Helicoverpa spp. (Lepidoptera : Noctuidae) and Creontiades spp. (Hemiptera : Miridae) in cotton. — Bulletin of Entomological Research, 97(3): 281–290. Grundy P.R. & D.A. Maelzer (2000). Assessment of Pristhesancus plagipennis (Walker) (Hemiptera: Reduviidae) as an augmented biological control in cotton and soybean crops. — Australian Journal of Entomology, 39(4): 305–309. Grundy P.R. & D.A Maelzer (2003). Towards the on-farm conservation of the assassin bug Pristhesancus plagipennis (Walker) (Hemiptera: Reduviidae) during winter using crop plants as refuges. — Australian Journal of Entomology, 42(2): 153–158. Grundy P.R., D.A. Maelzer, A. Bruce & E. Hassan (2000). A mass-rearing method for the assassin bug Pristhesancus plagipennis (Hemiptera: Reduviidae). — Biological Control, 18(3): 243–250. Hart E.R. (1972a). A systematic revision of the genus Zelus Fabricius (Hemiptera: Reduviidae). — Ph.D. dissertation, Dep. Entomology Texas A&M University: xviii + 1–595., 2 vols.
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Hart E.R. (1972b). A new species of the genus Atopozelus Elkins, with a review of the genus. — Journal of the Kansas Entomological Society, 45(3): 392–396. Hart E.R. (1975). A new species of Ischnoclopius Stål, with notes on the systematic position of the genus (Hemiptera: Reduviidae). — Proceedings of the Entomological Society of Washington, 77(4): 419–425. Hart E.R. (1986). Genus Zelus Fabricius in the United States, Canada, and Northern Mexico (Hemiptera: Reduviidae). — Annals of the Entomological Society of America, 79(3): 535–548. Hart E.R. (1987). The genus Zelus Fabricius in the West Indies (Hemiptera: Reduviidae). — Annals of the Entomological Society of America, 80(2): 293–305. Hijmans R., L. Guarino, A. Jarvis, & R. O’Brien (2007). DIVA-GIS. [software program]. — Available at: http://www.diva-gis.org. Hussey R.F. & J.C. Elkins (1955). Review of the genus Doldina Stål (Hemiptera: Reduviidae). — Journal of the Florida Academy of Sciences, 18(4): 261–278. Ishikawa T., W. Toriumi, W. Susila & S. Okajima (2007). Sycanus aurantiacus (Hemiptera: Heteroptera: Reduviidae), a new harpactorine species from Bali, Indonesia, with brief notes on its biology. — Zootaxa, 1615: 21–27. Jahnke S.M., L.R. Redaelli, L.M.G. Diefenbach & C.F.S. Efrom (2003). Distribuição espacial de posturas de Cosmoclopius nigroannulatus Stål (Hemiptera: Reduviidae) em Nicotiana tabacum L. (Solanaceae). — Neotropical Entomology, 32(1): 123–126. Lent H. & P. Wygodzinsky (1979). Revision of the Triatominae (Hemiptera: Reduviidae) and their significance as vectors of Chagas’ disease. — Bulletin of the American Museum of Natural History, 163(3): 123–520. Maldonado C. J. (1976a). The genus Heza (Hemiptera: Reduviidae). — Journal of Agriculture of the University of Puerto Rico, 60(3): 403–433. Maldonado C. J. (1976b). Three new species of Castolus and a key to the species (Hemiptera: Reduviidae). — Proceedings of the Entomological Society of Washington, 78(4): 435–446. Maldonado C. J. (1983). Concerning new and old species of Heza (Hemiptera: Reduviidae). — Journal of Agriculture of the University of Puerto Rico, 67: 407–418. Maldonado C. J. (1987). Orbella, a new Harpactorinae genus (Hemiptera: Reduviidae). — Journal of Agriculture of the University of Puerto Rico, 71(2): 193–198. Maldonado C. J. (1990). Systematic catalogue of the Reduviidae of the World. — Caribbean Journal of Science, Special publication No. 1, University of Puerto Rico, Mayagüez, Puerto Rico: 1–694. Maldonado C. J. (1992). A new Harpactorinae genus from French Guiana, northern South America (Heteroptera: Reduviidae). — Caribbean Journal of Science, 28(1-2): 38–40. Maldonado C. J. (1993). Redescription of Nesocastolus nigrocoris (Fracker and Bruner) and notes on the genus (Heteroptera: Reduviidae: Harpactorinae). — Proceedings of the Entomological Society of Washington, 95(4): 562–563. Maldonado C. J. & D.J. Carpintero (1993). Redescription of the Harpactorinae genus Sosius Champion 1899, with the description of a new species (Heteroptera: Reduviidae). — Proceedings of the Entomological Society of Washington, 95(2): 223–227. Nishi A., T. Imamura, A. Miyanoshita, S. Morimoto, K. Takahashi, P. Visarathanonth, R. Kengkanpanich, M.E.H. Shazali, & K. Sato (2004). Predatory abilities of Amphibolus venator (Klug) (Hemiptera: Reduviidae), a predator of stored-product insect pests. — Applied Entomology and Zoology, 39(2): 321–326.
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Putshkov V.G. & P.V. Putshkov (1985). A catalogue of assassin-bug genera of the world (Heteroptera, Reduviidae). — Published by the authors, Kiev: 1–137. (Introduction in Ukranian) Rios N.E.& H.L. Bart, Jr. (2005). GEOLocate. Georeferencing software for Natural History Collections. — Version 2.13 [software]. Available at: http://www.museum.tulane.edu/ geolocate/default.aspx. Scudder G.G.E. (1959). The female genitalia of the Heteroptera: morphology and bearing on classification. — The Transactions of the Royal Entomological Society of London, 111(14): 405–467. Stål C. (1872). Enumaratio Reduviinorum americae. Enumeratio Hemipterorum. Bidrag till en förteckning öfver alla hittils kända Hemiptera, Jemte Systematiska meddelanden, 2. — Kongliga Svenska Vetenskaps-Akademiens Handlingar, 10(4): 1–159. Weirauch C. (2005). Pretarsal structures in Reduviidae (Heteroptera, Insecta). — Acta Zoologica, 86: 91–110. Weirauch C. (2006). Observations on the sticky trap predator Zelus luridus Stål (Heteroptera, Reduviidae, Harpactorinae), with the description of a novel gland associated with the female genitalia. — Denisia, 50: 1169-1180. Weirauch C. & D. Forero (2007). Kiskeya palassaina, new genus and new species of Saicinae (Heteroptera: Reduviidae) from the Dominican Republic. — Zootaxa, 1468: 57–68. Wygodzinsky P. (1947). Sobre um novo gênero de Harpactorinae do Brasil, com notas sobre os gêneros Harpactor Laporte e Erbessus Stål (Reduviidae, Hemiptera). — Revista de Entomologia, 17(3): 401–417. Wygodzinsky P. (1948). El género “Diarthrotarsus” Bergroth, 1905 (Harpactorinae, Reduviidae, Hemiptera). — Acta Zoológica Lilloana, 6: 201–213. Wygodzinsky P. (1949). Elenco sistemático de los reduviformes americanos. — Instituto de Medicina Regional, Universidad Nacional de Tucumán, Argentina, Monografia No. 1: 1–102.
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Grozeva & N. Simov (Eds) 2008 A S. revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 105 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 105-120. © Pensoft Publishers Sofia–Moscow
A revision of Leprosoma BAERENSPRUNG, 1859 (Heteroptera: Pentatomidae) D.A. Gapon Zoological Institute of Russian Academy of Sciences, Universitetskaya nab. 1, 199034 Saint Petersburg, Russia. E-mail:
[email protected]
ABSTRACT Two species groups (“inconspicuum” and “reticulatum”) are separated in the Palaearctic genus Leprosoma Baerensprung. The inner and outer ectodermal structures of male and female genitalia are described. Leprosoma inaequale Horváth is synonymized with L. tuberculatum Jakovlev. A new species, L. tenuimarginatum, from Kazakhstan, Turkmenistan, Uzbekistan and Tadjikistan is described in the “reticulatum” species group. Keywords: Aedeagus, Heteroptera, Leprosoma, new species, Palaearctic Region, Pentatomidae.
INTRODUCTION Leprosoma Baerensprung is a Palaearctic genus of the tribe Ventocorini, subfamily Podopinae, family Pentatomidae. It can be easily distinguished from other genera of Ventocorini on the basis of following characters. On the one hand, Leprosoma differs from Ventocoris and Vilpianus in narrower scutellum, dilated connexivum and acute anterior angles of plates on anterior margins of propleurites, on the other hand, it differs from other genera of the tribe in long scutellum reaching apex of abdomen (scutellum of Tshingisella, Asaroticus, Hybocoris, Putonia and Crypsinus slightly extending beyond anterior margin of abdominal tergite VII at the outside). According to Rider (2006), Leprosoma comprises 6 species. In the present paper, a new species of this genus is described from Central Asia and a new synonymy is established. This work is based on the
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extensive material from the collection of the Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZISP), some specimens from the Muséum National d’Histoire Naturelle, Paris (MNHN), and the collection of Dr. Linnavuori (Raisio, Finland). The aedeagi are described in completely inflated state. For the method of inflation and the nomenclature of aedeagal structures see Gapon (2001) and Konstantinov & Gapon (2005), respectively.
Genus Leprosoma BAERENSPRUNG, 1859 Description. Body (Figs 1-4, 6, 7) moderately large, broadly oval, rather weakly convex, yellowish or light or dark brown. Surface asperous, covered reticilately sculptured, without distinct punctures. Head somewhat inclined, about as long as wide or slightly wider than long. Preocular part narrowed and bent downwards apically, with more or less deep lateral incisures. Juga convex, much longer than tylus, converging before it. Tylus narrowed apically. Eyes spherical, moderately large. Ocelli closer to anterior margin of pronotum than to eye margins. Bucculae strongly arcuately convex before middle, each without tooth on anterior part, strongly decreasing caudad. Rostrum reaching middle of metacoxae. Prothorax. Anterior margin of pronotum with thin elevation, lateral margins more or less strongly concave, without rib. Anterior pronotal angles each with small denticle. Humeri slightly to strongly projecting beyond lateral margins of hemelytra. Posterior angles of pronotum shortly rounded. Disc in anterior portion with a small tubercle on each side and a deep furrow (cicatrice) just posterior to it. Up to three transverse carinae developed on pronotum: thin anterior carina just posterior to cicatrices; long, more or less sinuous posterior carina in the middle of pronotum, and shorter intermediate carina sometimes present before the posterior one. Anteror margins of propleurites form short,
1
2
Figs 1-2: Leprosoma, dorsal habitus of L. inconspicuum from Russia, Sarepta (1) and L. stali from Armenia, Erivan (2).
3
4
Figs 3-4: Leprosoma, dorsal habitus of L. tuberculatum, holotypus (3) and L. tenuimarginatum nov. sp., holotypus (4).
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 107
L. inaequale L. tuberculatum
Fig. 5: Ratio of width of abdomen/width of pronotum of L. inaequale and L. tuberculatum.
plates extending ventrad. Anterior angles of these plates acute. Mesothorax. Scutellum narrow, reaching or (in most females) almost reaching apex of abdomen, leaving connexivum and most of corium exposed. Lateral margins of scutellum in basal part subparallel, straight or slightly concave in middle. Apex of scutellum rounded, linguiform. Impressions in basal corners of scutellum barely visible. Basal triangular elevation of scutellum quite high, transformed distally into narrowing carina not reaching apex of scutellum. Inner margins of pleurites before mesocoxae with rectangular plates directed mediad; posterior margins of pleurites without strip of porous cuticle. Metathorax. Stink glands opening in antero-medial part of small, roundish, rather deep impressions; a small tubercle present lateral to gland aperture. Sterna of all thoracic segments with a deep median groove. Legs pale with more or less large dark spots. Meso- and metacoxae with tubercles curved caudad. Abdomen. Connexivum broad, concolorous with rest of dorsum or sometimes darker. Laterotergites smooth or with uneven, wide and low transverse elevation in posterior part. Posterior angles of abdominal segments with tubercles varying in size from small to very large. Ventral surface of abdominal base without tubercle or longitudinal depression. Abdominal venter pale, with small and very shallow pale punctures. Pygophore (Figs 17-20) somewhat wider than long, calyciform at base, slightly 6 7 widened in distal part. Latero-apical anFigs 6-7: Leprosoma, dorsal habitus of L. gles pointed, slightly protruded laterad. reticulatum from Egypt (5) and L. olcesii from Dorsal infolding of pygophoral rim long, Morocco (6).
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strongly elevated, with shallow notch. Latero-apical margins smooth (Figs 8, 11, 13-15) without tubercles or denticles). Ventro-apical margin with very shallow, wide, median notch. Ventral infolding of pygophoral rim perpendicular to ventral wall of pygophore or slightly elevated, long or short. Ventral wall smooth, without punctation, convex in distal part. Parandria small, roundish, as long as wide. Paramere (Figs 9, 12, 16). Basal plate small, roundish. Body of paramere rather long. Sensory processus not developed. Hypophysis long, rather wide in posterior part, narrowed toward apex; its outer margin strongly sclerotized, with a row of very small obtuse denticles. Aedeagus (Figs 21-26). Phallobase slightly wider than long; its ventral processes long, thin, somewhat spaced; dorsal connectives short; capitate processes small, oval. Theca not subdivided into apical and basal parts, moderately long and wide, widest distad of middle; ventral wall almost straight; dorsal wall convex; base with moderately large membranous ventral tubercles. Conjunctiva small. Its paired ventro-lateral lobes with two pairs of branches: lateral and medial. Apex of conjunctiva very short, weakly convex. Dorsal wall forming at base a low elevation bearing two narrow, rather long, pointed sclerotized processes directed towards base of aedeagus (Fig. 26). Longitudinal bands of median penal plates widely spaced at base, converging distad, covered by ventro-lateral lobes and subparallel in their ventral part. Apical processes small, almost nearly contiguous. Vesica narrow, short, tubular. Genitalia of female. First gonocoxae slightly wider than long, almost flat, covered with short setae along caudal and mesial margins. Median genital plate rather long, narrowed caudad, with some short setae on anterior corners. Paratergites IX moderately wide, broadly rounded apically, distinctly projecting beyond dorsal margin of proctiger. Paratergites VIII large, with lateral margins straight, posterior margins arcuately or sub-angularly convex, and mesial margins concave. Bridge between paratergites VIII rectangular, very short, broadly rectangular. Vestiges of first and second gonapophyses as rather large membranous folds. Triangulum large. Gynatrial sclerites. Parabasal sclerite (Fig. 27) wide, rather weakly sclerotized, truncate on anterior end. Basal sclerites broadly triangular, almost touching parabasal sclerite near posterior angles. Outer angles of basal sclerites elongate. Ring sclerites rather small, fused with basal sclerites. Annular sclerite somewhat ovate. Conoid sclerite slightly abbreviated. Spermatheca. Proximal part of spermathecal duct (Figs 28-30) extending slightly beyond anterior margin of abdominal sternum VII. Medial part of duct very short (equal in length to sternum VII) or long, extending beyond anterior margin of sternum III or IV. Proximomedial part of duct (membranous dilation) without sharp constriction on proximal end. Distomedial part (sclerotized invaginated rod) of duct considerably widened before posterior (according to position) end. Visible distal part of duct appreciably longer than pump. Capsule of spermatheca large, spherical, with one or two short processes.
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 109
12 8
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Figs 8-26: Leprosoma, ventro-apical margin of pygophore, ventral view (8, 11, 13, 14, 15); caudal view of paramere (9, 12, 16); parandrium (10); dorsal (17, 19) and caudal (18, 20) views of pygophores; ventral (21-25) and dorsal (26) views of aedeagus of L. inconspicuum (8-10, 21), L. stali (22), L. tuberculatum (11, 12, 17, 18, 23), specimens previously placed in L. inaequale (13), L. tenuimarginatum nov. sp. (14, 19, 20, 24), and L. olcesii (15, 16, 25, 26). Symbols: ap. c – apex of conjunctiva; d. inf – dorsal infolding of rim of pygophore; d. pr – dorsal processes of conjunctiva; l. br – lateral branches of conjunctival ventro-lateral lobes; m. br – medial branches of conjunctival ventro-lateral lobes; m. p. p. – medial penal plates; prd – parandria; v. inf – ventral infolding of rim of pygophore; ves – vesica.
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tr con. s an. s r. s
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29
30
Figs 27-30: Leprosoma, inner ectodermal genitalia of L. inconspicuum, ventral view (27); spermatheca of L. tenuimarginatum nov. sp. (28), L. reticulatum (29) and L. olcesii (30). Symbols: an. s. – annular sclerite of gynatrium; b. s. – basal sclerites of gynatrium; con. s. – conoid sclerite of gynatrium; d. p. – visible distal part of spermathecal duct; dm. p. – distimedial part of spermathecal duct; gp. 1, 2 – gonapophyses 1, 2; m. p. – medial part of spermathecal duct; pb. s. – parabasal sclerites of gynatrium; pm. p. – proximomedial part of spermathecal duct; r. s. – ring sclerites of gynatrium; tr. – triangulum.
Key to species 1(4).
2(3).
Body without incrustation resembling hoarfrost. Head wider than long. Longitudinal carinae on frons obscure or absent. Humeri not incrassate and not separated from pronotal disc by impression. Cicatrices pale. Anterior transverse carina very low, obliterated laterally, often absent or vestigial. Scutellum broad, 0.58-0.65 times as wide as long; tubercles at its base small, rounded ................ ....................................................................... The “inconspicuum” species group Tubercles in anterior portion of pronotum small. Connexivum thin (in strongly lateral view), its laterotergites smooth, with very fine punctation. Tubercles on posterior angles of abdominal segments very small .......Leprosoma inconspicuum
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 111
3(2).
Tubercles in anterior portion of pronotum large. Connexivum thicker (in strong lateral view), its laterotergites with rough punctation and transverse elevation, sometimes strongly smoothed. Tubercles on posterior angles of abdominal segments usually rather large .......................................................... Leprosoma stali 4(1). Body covered with incrustation resembling hoarfrost. Head as long as wide. Longitudinal carinae on frons distinct. Humeri incrassate, separated from pronotal disc by impression. Cicatrices black. Anterior transverse carina high. Scutellum rather narrow, 0.50-0.57 times as wide as long, tubercles at its base large, clear longitudinal .......................................................The “reticulatum” species group. 5(10). Humeri long, strongly projecting beyond lateral margins of hemelytra. 6 (9). Body length 5.25-6.55. Intermediate pronotal carina present. Posterior transversal carina strongly and smoothly curved. Elevations on connexival laterotergites moderately convex, tubercles on posterior angles of abdominal segments moderately large. Medial part of spermathecal duct long, extending beyond anterior margin of abdominal sternum V. 7(8). Humeri broadly or rectangular rounded, directed laterad. Pronotal transverse carinae distinct. Ventral infolding of rim of pygophore very long, perpendicular to ventral wall of pygophore ............................Leprosoma tuberculatum (in part) 8(7). Humeri looking like horns curved forward. Pronotal transverse carinae obsolete and masked by incrustation. Ventral infolding of rim of pygophore very narrow and slightly elevated .................................. Leprosoma tenuimarginatum nov. sp. 9(6). Body length 6.00-6.80. Intermediate pronotal carina absent. Posterior carina weakly and sharply curved. Elevations on connexival laterotergites strongly convex, tubercles in posterior corners of abdominal segments very large. Medial L. inconspicuum L. stali L. tuberculatum, specimens with long humeri L. tuberculatum, specimens with short humeri specimens previously placed in L. inaequale L. tenuimarginatum
Fig. 31: Distribution of Leprosoma species.
112 D.A. Gapon
part of spermathecal duct as long as abdominal sternum VI or slightly extending beyond its anterior margin ....................................................... Leprosoma olcesii 10(5). Humeri short, slightly projecting beyond lateral margins of hemelytra. 11(12). Humeri directed laterad. Medial part of spermathecal duct long, extending beyond anterior margin of abdominal sternum V. Distributed in Russia, Turkey, Transcaucasia, Central Asia and China ...........Leprosoma tuberculatum (in part) 12(13). Humeri weakly curved forward. Medial part of spermathecal duct short, slightly extending beyond anterior margin of abdominal sternum VII. Distributed in North Africa, Israel, Syria, Iran....................................... Leprosoma reticulatum
The “inconspicuum” species group Diagnosis. Body (Figs 1, 2) not covered with thin incrustation resembling hoarfrost. Head slightly wider than long. Longitudinal carinae on frons obscure or absent. Humeri projecting slightly beyond lateral margins of hemelytra, broadly rounded, not incrassate and not separated from pronotal disc by impression. Cicatrices pale. Anterior transverse carina very low, obliterated laterally, often absent or vestigial. Intermediate carina absent. Lateral parts of posterior carina nearly straight, only slightly bent medially. Scutellum relatively broad, 0.58-0.65 times as wide as long; tubercles at its base small, rounded. Pygophore and male genitalia. Latero-apical angles of pygophore rounded. Dorsal infolding of rim of pygophore narrow. Latero-apical margins of pygophore slightly convex, thickened, curved inside in the middle of length, covered with setae directed mediad Ventral infolding of pygophoral rim short, elevated, weakly depressed, with a wide notch. Ventro-apical margin of pygophore with wide, rather deep, smoothly rounded notch (Fig. 8). Inner and outer margins of paramere hypophysis parallel in nearly entire length in apical (caudal) view; outer margin broadly rounded apically (Fig. 9). Medial branches of ventro-lateral lobes of conjunctiva long and wide; lateral branches rather narrow, not curved apically (Figs 21, 22). Apex of conjunctiva rather long, tapered. Female genitalia. Caudal margins of first gonocoxae very weakly curved or straight. Medial part of spermathecal duct extending beyond anterior margin of abdominal sternum III. This species group includes L. inconspicuum and L. stali.
Leprosoma inconspicuum BAERENSPRUNG, 1859 (Figs 1, 8-10, 21, 27)
Material examined (ZISP). Without locality labels, 6.VII.? ( Jakovlev coll.) , 1♂, 1♀, 2♂♂; UKRAINE, Crimea: Eupatoria ( Jakovlev coll.), 1♂; Agarmysh Hill, 4, 11, 15.VI.1906 (V. Jakovlev coll.), 3♀♀; 25-26, 29.VI, 2, 4, 6, 7.VII.?, 7♂♂, 5♀♀, without date, 2 ♀♀ (Al. Kiritshenko); Staryi Krym, 5.VI.1906 ( Jakovlev coll.), 2♂♂; Kerch, 20.IV.1901 (Kiritshenko coll.), 1♂; Donetsk Prov.: Svyatogorskaya, 3-4.VII.1940 (Lukjanovich), 1♀;
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 113
Veliko-Anadol (forestry), 18.V.1895 (Kiritshenko coll.), 1♀; Lugansk Prov.: Provalie, 16.VI.1933 (Talitsky), 1♀; RUSSIA, Rostov Prov.: Tsymla, 12.VI.1924, 1♂; Voronezh Prov.: Kamennaya Step, 11 km S Talovaya, 12.VI.1942 (Stark), 1♀; Stavropol Terr.: Petrovskoe [= Svetlograd], 8.V.1825, 1♀; Volgograd Prov.: Sarepta ( Jakovlev coll.), 2♂♂, 1♀, (Becker), 2♂, 1♀, 2, 21.V.1909 (Glazunov), 2♂♂, 2♀♀; Nikolaevka [=Nikolaevsk], 15.X, 5.XI.1942 (Bogachev), 2♂♂, 1♀; Astrakhan Prov.: Bogdo Hill ( Jakovlev coll.), 1♂; Saratov Prov.: Saratov, 9, 15.IX.1937 (Lukjanovich), 4♂♂, 2♀♀; Dyakovka, 25 km S Krasnyi Kut, 24.VI.1961 (Kerzhner), 1♂; Orenburg Prov.: Buzuluksky bor (reserved area), 2.V., 4.VII, 7, 21.VIII, 6.X.1941 (Tchistovsky), 4♂♂, 3♀♀; Orenburg, 18.V.1924, 27.VII.1924 (A. Ivanov), 2♀; Verkhnedneprovka, 18.V, 4, 10, 11, 13.VI, 4, 21-27.VII.1932, 20-25.V, 12, 20-25.VIII.1934, 31.V, 15, 21.VI.1935 (Zimin), 6♂♂, 9♀♀; Ashche-Butak, 27.V, 24.VII, 2, 10, 11, 13, 14, 23, 29.VIII, 1.IX.1933 (Zimin), 11♂♂, 12♀♀; Elshanka, 27.V, 24, 29.VII, 10, 11, 14, 17.VIII, 1.IX.1933 (Zimin), 4♂♂, 6♀♀; riv. Donguz, 5, 25.VIII.1933 (Zimin), 1♂, 1♀; Spassk, riv. Bolshoy Ik, 30.III, 15.VI.1933 (Rysakov), 1♂, 2♀♀; Novouspenka, 5.IX.1955 (Tchistovsky), 1♂; Dagestan: Petrovsk [= Makhachkala], 2.V.1925 (Kiritshenko), 3♂♂, 4♀♀; Temir-khan-shura [= Buynaksk], 16.V, 22, 23.VI.1925 (A. Kiritshenko), 3♂♂; Chernyi Rynok [= Kochubey], 19.V.1925 (Kiritshenko), 1♀; GEORGIA: Benara, 19 km W Akhaltsykhe, 8.VI, 7, 8, 10, 12, 19.VII.1949 (Kiritshenko), 8♂♂, 9♀♀; ARMENIA: Kushu-Darash near Gokcha [= Sevan] lake, 27.VI.1902 (Elachich), 1♀; Sevan lake, 15.X.1932 (Schelkovnikov), 1♀; KAZAKHSTAN, West Kazakhstan Prov.: Khanskaya Stavka [= Urda], Ryn-peski, 26.V-16.VI.1887 (Plushchevsky), 1♂, 3♀♀; station Dzhanybek, 28.VI.1961 (Kerzhner), 1♀; Petrov, 5.IX.1949 (Kiritchenko), 1♀; Kalmykovo, 12.VI.1909 (Uvarov), 1♂; Yanvartsevo, 29.VI, 3, 4, 9, 15.IX.1949 (Kiritshenko), 2♂♂, 4♀♀, 29.VI.1949 (Arnoldi), 1♂, 1♀, 10, 15.VI.1949, 4.V.1950 (Romadina), 2♂♂, 2♀♀, 14.V.1950 (Grunin), 1♂, 2♀♀; Aktyubinsk Prov.: Aktyubinsk, 28.V.1957 (Mitiaev), 1♂; Biatty-Kuzha, upper riv. Emba, 24.VII.1926 (Burachek), 1♀; station Dzhurum [= Zhuryn], 25.VII.1933 (Formozov), 1♀; Almaty Prov.: Topolevka, Dzhungarsky Alatau, 7.VII.1957 (Kerzhner), 1♂; Sarkand, Dzhungarsky Alatau, 6, 9.VI.1957 (Kerzhner), 2♀♀; KYRGYZSTAN: Frunze [= Bishkek], 3-7.V.1930 (Shnitnikov), 1♂; Belovodskoe, 18.VIII.1931 (Zimin), 1♂; valley of riv. Karabalty, 20.VII.1931 (Zimin), 1♀. Diagnosis. Surface of body rather weakly roughened. Tubercles in anterior portion of pronotum small. Posterior transverse carina higher than anterior, almost straight laterally parts, sometimes slightly curved caudad and slightly interrupted. Tubercles in anterior corners of scutellum very small. Connexivum thin (in strongly lateral view); laterotergites smooth, with very fine punctation. Tubercles on posterior angles of abdominal segments very small. Pygophore as wide as long. Dorsal infolding of rim of pygophore very narrow. Lateral walls of pygophore parallel in distal part. Parandria with evenly convex, not elevate outer margin and slightly concave inner margin (Fig. 10). Female genitalia. Medial part of spermathecal duct extending beyond middle of abdominal sternum IV, sometimes reaching its anterior margin. Measurements (mm) and ratios. Total length 4.85–6.50, width of pronotum 3.20–4.30, width of abdomen 3.25–4.10, ratio head length/head width 0.86–0.96, ratio
114 D.A. Gapon
scutellum width/scutellum length 0.58–0.64. Length of antennal segments I–V: 11–13 : 17–21 : 11–15 : 10–15 : 25–28. Distribution. Southern Europe, Algeria, Russia (south of European territory and West Siberia), Turkey, Iraq, Iran, Georgia, Armenia, Azerbaijan, Kazakhstan, Kyrgyzstan. Biology. The host plants are Brassicaceae, particularly Lepidium.
Leprosoma stali DOUGLAS & SCOTT, 1868 (Figs 2, 22)
Material examined (ZISP). TURKEY: Anatolia, Dikmen-dagh, 11.IV.1934 ( Jenjourist coll.), 1♂; 20.4 km S of Gumushane, 17.VI.2006 (Korotyaev), 1♀; ARMENIA: Erivan, 18.V.1938 (No. 201-938) (Ter-Minasian & Richter), 1♀, 20.V.1947 (Borkhsenius), 1♂, 31.X.1931 (Korinek), 1♀, 6.VII, 3,8.VIII.1935, 25.IX.1937 (Ter-Minasian & Richter), 3♂♂, 2♀♀, 30.III, 4,16.IV.1936 (Richter), 5♀♀; Dzhervezh, 7.IV.1936 (Richter), 2♂♂, 12♀♀; AZERBAIJAN: Talysh Mts, Kosmalian, 19.V.1909 (Kiritshenko coll.), 1♀; IRAN: S slopes of Elburz, Tadzhrish, 25-30.VI.1928 (Siyazov), 1♂; Doab, 14.VI.1914 (Kiritshenko), 1♀; Shachkuh superior, 16, 19-22, 29-30.VI, 2-3.VII.1914 (Kiritshenko), 12♂♂, 7♀♀; Shachkuh-poion, 3, 4.VII.1914 (Kiritshenko), 4♂♂, 1♀. Diagnosis. Surface of body strongly roughened. Tubercles in anterior portion of pronotum larger than in L. inconspicuum. Posterior pronotal transverse carina more strongly curved caudad in lateral parts than in L. inconspicuum. Tubercles in anterior angles of scutellum slightly larger than in L. inconspicuum. Connexivum thicker (in strong lateral view). Laterotergites with rough punctation and transverse elevation, sometimes strongly smoothed. Tubercles on posterior angles of abdominal segments usually slightly larger than in L. inconspicuum. Pygophore slightly wider than long; its lateral walls divergent in distal part. Dorsal infolding of rim of pygophore wider than in L. inconspicuum. Inner margins of parandria straight. Female genitalia. Medial part of spermathecal duct usually extending beyond anterior margin of sternum IV, but sometimes only slightly extending beyond its middle. Measurements (mm) and ratios. Total length 4.95–6.05, width of pronotum 3.10–3.80, width of abdomen 3.35–3.95, ratio head length/head width 0.90–0.93, ratio scutellum width/scutellum length 0.63–0.65. Length of antennal segments I–V: 12–13 : 17–21 : 12–14 : 11–13 : 25–27. Distribution. Recorded from Egypt, Asian part of Turkey, Cyprus, Israel, Syria, Iraq, Iran, Armenia and Azerbaijan (Fig. 31).
The “reticulatum” species group Diagnosis. Body (Figs 3, 4, 6, 7), especially dorsal surface, covered with incrustation resembling hoarfrost. Head as long as wide. Longitudinal carinae on frons distinct. Humeri
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 115
incrassate, from short to very long, separated from pronotal disc by impression. Cicatrices black. Anterior transverse carina high. Intermediate carina present (absent in L. olcesii). Scutellum rather narrow, 0.50-0.57 times as wide as long, tubercles at its base large, clear longitudinal. Pygophore and male genitalia. Latero-apical angles of pygophore acute. Dorsal infolding of rim of pygophore long and wide, strongly elevated, with shallow notch. Latero-apical margins of pygophore not thickened, not convex and not curved inside. Ventral infolding of pygophoral rim long or short. Hypophysis of paramere gradually narrowed along its whole length (Figs 12, 16). Medial branches of ventro-lateral lobes of conjunctiva short and narrow; lateral branches rather wide, curved apically (Figs 23-25). Outer walls of lateral branches of ventro-lateral lobes forming an obtuse angle in frontal view. Apex of conjunctiva short, weakly convex. Female genitalia. Medial part of spermathecal duct (Figs 28-30) either long (extending beyond anterior margin of abdominal sternum V) or very short (as long as abdominal sternum VII or slightly longer). This species group includes L. tuberculatum, L. tenuimarginatum nov. sp., L. reticulatum and L. olcesii.
Leprosoma tuberculatum JAKOVLEV, 1874 Leprosoma inaequale Horváth 1911, nov. syn. (Figs 3, 11-13, 17, 18, 23)
Type material examined (ZISP). Holotype of L. tuberculatum: ♂, Russia, Astrakhan Prov., “Bogdo [Hill]”, “k. V. Jakovleva”, “Leprosoma tuberculatum Jak. V. Jakowlew det.”, “Holotypus ♂ Leprosoma tuberculatum Jak.”. Lectotype of L. turkestanicum: ♂, Uzbekistan, Karaul-Bazar, 13.IV.1927 (Zolotov); paralectotypes: station Ziaddin, 29.V.1927 (Zolotov), 3♀♀. Additional material examined (ZISP). RUSSIA, Astrakhan Prov.: Seroglazinka, 1♂; Volgogard Prov.: Elton lake, 5.VI.1961 (Emeljanov & Kerzhner), 1♂, 2♀♀; Orenburg Prov.: Elshanka, 24.VII.1933 (Zimin), 2♂; ARMENIA: Erivan, 3, 8.VII.1935, 26, 30.VIII.1936 (Richter & Ter-Minasian), 2♂♂, 3♀♀, 31.X., 1.XI.1931 (Korinek), 1♂, 1♀, 18.V.1938 (No. 201-938) (Ter-Minasian & Richter), 1♀; near Erivan (Stekolnikov), 1♂; Dzhervezh, 29.IX.1946 (Richter), 1♂; Sovetashen [= Nubarashen], 27.V.1938, 1♀; AZERBAIJAN: Nakhichevan, 15.VI.1933 (Znoiko), 1♂; Disar near Ordubad, 20.VII.1933 (Znoiko), 1♀, 2, 5.V.1934 (Ter-Minasian), 3♀; Elisavetpol [= Gyandzha], 24.I.1905, 6.VIII.1906 ( Jakovlev coll.), 3♂♂, 4♀♀; IRAN: S slopes of Elburz, Tadzhrish, 25-30. VI.1928 (Siyazov), 1♂, 1♀; Charbag, 4.V.1914 (Kiritshenko), 2♂♂, 1♀; Shakhrud [= Emamshahr], 23, 26, 27, 29, 31.V, 1-3, 5-7, 11.VI.1914 (Kiritshenko), 12♂♂, 13♀♀; Shakhkukh [Shachkuh superior], 15, 16, 18-22, 25, 26, 29.VI, 1-5.VII.1914 (Kiritshenko), 22♂♂, 33♀♀; Shachkuh-poion, 3-5.VII.1914 (Kiritshenko), 6♂♂, 5♀♀; Kazvin, 16.III.1935 ( Jenjouriste), 1♀; KAZAKHSTAN, West Kazakhstan Prov.: Dzhanybek, 26.VI.1961 (Kerzhner), 1♂, 1♀; Kalmykovo, 17.VI.1909 (Uvarov), 1♂, 1♀; Atyrau Prov.: Inder lake, 23, 24.VI.1951 (Steinberg), 1♂; UZBEKISTAN: Shirabad, 27-28.V., 2, 4.VII.1912 (Kir-
116 D.A. Gapon
itshenko), 12♂♂, 4♀♀; Tutak, 31.IX.1911 (Kiritshenko), 1♀; Kyzyl-Kum, Ayakguzhumdy, 29.IV.1965 (Emeljanov), 2♂; Kyzyl-Kum, 20 km N Ayakuzhumdy, 17.VI.1966 (Kerzhner), 1♂; Kyzyl-Kum, 70 km S Tamdy-Bulak, 1.V.1965 (Kerzhner), 1♂, 1. Diagnosis. Body moderately large. Humeri broadly or rectangular rounded, long or shortened, directed laterad. Tubercles in anterior portion of pronotum small. Intermediate transverse carina short, slightly curved caudad. Posterior transverse carina strongly sinuous. Tubercles on posterior angles of abdominal segments large. Laterotergites of connexivum each with distinct transverse elevation. Pygophore (Figs 17, 18) slightly wider than long. Iits ventro-apical margin with shallow, wide, rounded-rectangular notch (Figs 11, 13) and almost straight, slightly concave lateral parts. Ventral infolding of rim of pygophore very long, concave, perpendicular to ventral wall of pygophore. Parandria with convex, elevated, strongly sclerotized outer margin and straight or concave, membranous inner margin. Outer margin of paramere hypophysis convex just before apex (Fig. 12). Female genitalia. Caudal margin of first gonocoxae convex near lateral angles and slightly concave near medial angles. Medial part of spermathecal duct extending beyond anterior margin of abdominal sternum V. Measurements (mm) and ratios. Total length 5.30–6.55, width of pronotum 3.35–4.25, width of abdomen 3.60–4.20, ratio head length/head width 0.98–1.02, ratio scutellum width/scutellum length 0.50–0.55. Length of antennal segments I–V: 12–13 : 18–21 : 11–13 : 11–14 : 23–28. Distribution. Recorded from South of Russian European territory, Asian part of Turkey, Iran, Afghanistan, Armenia, Azerbaijan (new record), Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Kyrgyzstan, northwestern China, but some of these records (definitely those from Turkmenistan and Tajikistan) belong to L. tenuimarginatum. Specimens with long humeri are found in Astrakhan and Orenburg Provinces an in west Kazakhstan; specimens with shortened humeri distributed in the rest part of its natural habitat (Fig. 31). Biology. The host plants are Brassicaceae. L. inaequale Horváth (1911) was described from Armenia (Araxis valley). According to the original description, this species is characterised by the following main characters: abdomen wider than pronotum; lateral margins of pronotum distinctly sinuate in anterior part; humeri broadly rounded, slightly protruding forward; base and sides of scutellum and connexivum blackish; total length 6 mm; width of pronotum 3.75 mm; width of abdomen 4 mm. A large series of specimens of L. inaequale from Armenia and Azerbaijan in the collection of ZISP shows a gradual transition between the characters of L. inaequale and L. tuberculatum. The most important character, ratio of width of abdomen/width of pronotum, is presented in Fig. 5. On the basis of these data, L. inaequale is synonymised with L. tuberculatum. L. turkestanicum Zolotov 1928 was synonymised with L. tuberculatum by Kerzhner (1964). This variety is characterised by the short humeri and short tubercles on posterior angles of abdominal segments and resembles specimens previously placed in L. inaequale.
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 117
Leprosoma tenuimarginatum nov. sp. (Figs 4, 14, 19, 20, 24, 30)
Holotype: ♂, TAJIKISTAN, Pendzhikent, valley of river Zaravshan, 23.XI.1943 (Kiritshenko) (ZISP). Paratypes (ZISP). KAZAKHSTAN, Kzylorda Prov.: valley of riv. Syrdarya, Tartugay, 18.IV.1937 (Lukjanovich), 1♂; Zhambyl Prov.: valley of riv. Talas, foothills of Alatau, VI.1908 (Lebedev), 2♂♂; Mirzoyan [= Taraz], 2-4.V.1937 (Lukjanovich), 1♀; Muyunkum, Kargaly-kul lake, 24.V.1910 (Kiritshenko), 1♂; Qaraganda Prov.: Bet-PakDala, 17.VII.1960 (Asanova), 1♀; Almaty Prov.: Karatala, Kapal, 10.V.1930 (Kirshenblat), 1♀; Kyzylkum, Dort-kuduk [well], 24.III.1914 (Peltz), 1♀; Zhambylskaya Prov.: Kuyuk ( Jakovlev coll.), 1♂; TURKMENISTAN: Turkmenbashi [= Krasnovodsk], 13,14.VI.1899 (Oshanin coll.), 1♀, 7.IV.1914 (Kiritshenko), 1♂; Trans-Caspi (König), 1♀; Ashkhabad ( Jakovlev coll.), 1♂, 3.IV.1922 (Shestoperov), 1♀; 24.III.1933 (Vlasov), 1♂; Kara-Kala, 2.V.1957 (Medvedev), 1♂; Imam-baba, 5.IX.1931 (Frolova), 1♂; Annau, 10.VIII.1914, 1♂; UZBEKISTAN: Chernyayevo [= Khavast], 12-14.VI.1900, 1♀; Tashkent ( Jakovlev coll.), 4♂♂, 4♀♀; station Golodnaya Step [= Gulistan], 15.IV.1903 ( Jacobson), 1♀; Changyr near Khatyrchi, 11, 13, 15.VI.1930 (Zimin), 5♂♂, 7♀♀; Yargak near Khatyrchi, 16.VI.1928 (Zimin), 1♀; Kurgan-tube [= Kurgantepa], 10.VII.1938 (Popov), 1♀; Karzhantau [mountain range], 6.V.1939 (Obukhova), 1♀; Skobelev [= Fergana], 6, 25.III, 7.V.1910 (Shestoperov), 2♂♂, 2♀♀; Gorchakovo, 1.XII.1921 (Shestoperov), 1♂; Chimgan, 15.VII.1920 (Shestoperov), 1♂, 1♀; Dargom [canal] near Samarkand, 16.III.1914 (Blagoveshensky), 1♀; Agalyk, 24.VII.1914, 1917 (Blagoveshensky), 5♂♂, 1♀; station Yakkobag, 12.IX.1911 (A. Kiritshenko), 1♀; Tutak-ata, 31.IX.1911 (A. Kiritshenko), 3♂♂, 1♀; Chapan-ata, 15.IV.1912 (A. Kiritshenko), 2♂♂; Termez, 18.V.1912 (Kiritshenko), 18.V.1912, 1♂; Mts. Aktau near Tamdy, Kyzylkum, 29.IV, 2.V.1966 (Kerzhner), 2♂♂, 3♀♀; 20 km N Ayakguzhumdy, Kyzylkum, 26.IV.1965 (Nartshuk), 1♀; TAJIKISTAN: Pendzhikent, valley of riv. Zaravshan, 20, 23, 28, 29.XI.1943 (Kiritshenko), 4♂♂, 5♀♀; Khurmi, 3.XII.1943 (Kiritshenko), 1♂, 1♀; Tash-bulak – Mazor, Rengan-Tau, IV.1943 (Kiritshenko), 2♀; Dzhar-bulak, Rengan-Tau, 27.IV.1943 (Kiritshenko), 1♀; Dzhilikul, riv. Vakhsh, 29.V.1944, 1♂, 1♀; Kvak – Kondara, valley of riv. Varzob, 1.VII.1943, 2, 4.V.1944 (Kiritshenko), 2♂♂, 1♀; Obi-safed, riverhead of Kondara, 3.V.1944 (Kiritshenko), 1♂, 1♀; Iskander-kul lake, 23.VII.1947 (Kiritshenko), 1♀; riv. Iskander-darya, near source, 22, 24.VII, 2, 5, 7, 9, 11, 22.VIII.1947 (Kiritshenko), 5♂♂, 7♀♀; riv. Khozor-mech, 14, 19.VIII.1947 (Kiritshenko), 1♂, 1♀; riv. Sary-tag, 21.VII.1947 (Kiritshenko), 1♂, 2♀♀; Vorukh, beginning of Kekhskoe ravine, 11.VII.1912 (Tolbeck), 1♂. Diagnosis. Distinguished from L. tuberculatum by the humeri strongly projecting beyond lateral margins of hemelytra and looking like horns curved forward. Pronotal transverse carinae often obsolete and masked by incrustation. Ventro-apical margin of pygophore with oblique, convex lateral parts (Fig. 14). Ventral infolding of rim of pygophore very narrow and slightly elevated (Figs 19, 20). Parandria rounded; their outer
118 D.A. Gapon
half sclerotized, crescent-shaped, inner half almost membranous, without notch. Outer walls of lateral branches of ventro-lateral lobes of conjunctiva more broadly rounded than in other species of this group (Fig. 24). Female genitalia. Caudal margins of first gonocoxae almost straight. Medial part of spermathecal duct extending beyond middle or anterior margin of abdominal sternum IV (Fig. 30). Measurements (mm) and ratios. Total length 5.25–6.40, width of pronotum 3.30–4.25, width of abdomen 3.25–3.95, ratio head length/head width 0.98–1.00, ratio scutellum width/scutellum length 0.50–0.53. Length of antennal segments I–V: 11–13 : 17–21 : 11–12 : 10–12 : 23–26. Etymology. This species is named to denote very narrow ventral infolding of rim of pygophore. Distribution. Kazakhstan, Turkmenistan, Uzbekistan, Tajikistan, mainly in foothills.
Leprosoma reticulatum (HERRICH-SCHAEFFER, 1851) (Figs 6, 28)
Material examined. Egypt: Fajum (U. Sahlb[erg]), 1♀ (R. Linnavuori collection). Diagnosis. Body rather small. Humeri short, slightly projecting beyond lateral margins of hemelytra, rounded rectangular, very weakly curved forward; pronotum as wide as abdomen. Intermediate pronotal carina present; posterior carina strongly and smoothly curved. Laterotergites of connexivum with rather high elevations. Tubercles on posterior angles of abdominal segments rather small. Pygophore and male genitalia not studied. Female genitalia. Genital plates of female and gynatrial sclerites very similar to those of L. tuberculatum. Medial part of spermathecal duct short, slightly extending beyond anterior margin of abdominal sternum VII. Capsule of spermatheca spherical, without processes. Measurements (mm) and ratios (data of VIDAL (1949) are given in brackets). Total length 5.25 (5.00-6.00), width of pronotum 3.50 (3.50-4.00), width of abdomen 3.50 (3.50-4.00), ratio head length/head width 1.02, ratio scutellum width/scutellum length 0.51. Length of segments I-V of antennae: 12:18(22):12(14):11(14):22(30). Distribution. Recorded from Morocco, Mauritania, Libya, Egypt, Israel, Syria, Iran, but some of these records may refer to L. olcesii (Fig. 31).
Leprosoma olcesii FAIRMAIRE, 1886 (Figs 7, 15, 16, 29)
Material examined. Without geography designation ( Jakovlev coll.), 2♀ (ZISP); Morocco: (Puton coll.), 2♂♂, 2♀♀ (MNHN); 1898 (Noualhier coll.), 2 ♀ (MNHN); 1900 (Vaucher), 1♀ (ZISP); Lalla ST, VI.1914 (de Bergevin coll.), 1 ♂, 1♀ (ZISP). Diagnosis. Body rather large. Humeri long, rounded rectangular, rather curved forward. Intermediate pronotal carina absent. Posterior carina curved less strongly, but
A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae) 119
more sharply than in L. reticulatum. Elevations on connexival laterotergites strongly convex, tubercles in posterior corners of abdominal segments very large. Pygophore about as long as wide; its ventro-apical margin with very shallow trapezoid notch and slightly oblique, straight lateral parts (Fig. 15). Ventral infolding of rim of pygophore and parandria very similar to those of L. tuberculatum. Parameres and aedeagus as in L. tuberculatum. Ventro-lateral lobes of conjunctiva forming wider obtuse angle in ventral view. Female genitalia very similar to those of L. reticulatum. Medial part of spermathecal duct as long as abdominal sternum VI or slightly extending beyond its anterior margin (Fig. 29). Measurements (mm) and ratios. Total length 6.00-6.80, width of pronotum 4.35-4.80, width of abdomen 3.85-4.50, ratio head length/head width 0.97-1.02, ratio scutellum width/scutellum length 0.54-0.57. Length of segments I-V of antennae: 1214:21-22:12-14:13-15:26-30. Distribution. Recorded from Morocco (Fig. 31). L. olcesii was synonymized with L. reticulatum by Puton (1886) and restored by Horváth (1911). Other discriminating characters of L. olcesii, additional to the listed above, are given in the original description of L. olcesii and in Vidal’s (1949) key (head apex more broadly rounded, juga with deeper lateral incisures, posterior pronotal carina higher, posterior part of pronotum and all scutellum rougher), but these are not always diagnostic. Fairmaire (1886) considered L. olcesii to be merely a local variety of L. reticulatum with differences caused by the larger body size. The ratio of width of the pronotum to width of the abdomen is the character providing a dichotomy between these two species in keys. Specimens from Morocco examined by me show the following variability of this ratio: 1.0 (2 specimens), 1.11, 1.12 (2 specimens), 1.13, 1.16. Most likely, L. olcesii is not a separate species, judging by the co-existence of forms with short and long humeri in L. tuberculatum and continuity of distribution ranges of L. olcesii and L. reticulatum. However, the material examined by me is not sufficient for conclusion.
ACKNOWLEDGEMENTS With great pleasure I dedicate this work to outstanding entomologist Dr. M. Josifov in recognition of his important contribution to study of the fauna and classification of Heteroptera. I am very grateful to Prof. I.M. Kerzhner (St. Petersburg) for the help in preparation of this paper, Dr. J. Péricart (Saint-Lunaire), Dr. E. Gilbert (Paris), Dr. R. Linnavuori (Raisio) and Dr. V. Rinne (Turku) for the loan of the material.
РЕЗЮМЕ Палеарктичният род Leprosoma BAERENSPRUNG е разделен на две групи видове (“inconspicuum” and “reticulatum”). Описани са вътрешните и външни ектодермални структури на мъжките и женски гениталии. Leprosoma inaequale HORVÁTH е синонимизи-
120 D.A. Gapon
ран с L. tuberculatum JAKOVLEV. В групата “reticulatum” е описан нов вид L. tenuimarginatum от Казахстан, Туркменистан, Узбекистан и Таджикистан.
REFERENCES Fairmaire L. (1886). Notes sur quelques Hémiptères du Maroc. — Revue d’Entomologie, 4: 351–355. Gapon D.A. (2001). Inflation of heteropteran aedeagi using microcapillaries (Heteroptera: Pentatomidae). — Zoosystematica Rossica, 9(1): 157–160. Horváth G. (1911). Hemiptera nova vel minus cognita e regione palaearctica II. — Annales Historico-Naturales Musei Nationalis Hungarici, 9: 573–610. Kerzhner I.M. (1964). Materials on the synonymy of shield bugs (Heteroptera Pentatomoidea) in the fauna of the USSR and adjacent countries. — Entomologicheskoe Obozrenie, 43: 363–367 (In Russian). Konstantinov F.V. & D.A. GAPON (2005). On the Structure of the Aedeagus in Shield Bugs (Heteroptera, Pentatomidae): 1. Subfamilies Discocephalinae and Phyllocephalinae. — Entomologicheskoe Obozrenie, 84(2): 334–352. (In Russian; translated in Entomological Review, 85(3): 221–235). Puton A. (1886). Catalogue des Hémiptères (Hétéroptères, Cicadines et Psyllides) de la faune paléarctique, 3me édn. — Blanc-Hardel, Caën: 1–100. Rider D.A. (2006). Pentatomidae. — In: Aukema B. & Ch. Rieger (Eds). Catalogue of the Heteroptera of the Palaearctic Region. Ponsen & Looijen, Wageningen, The Netherlands, 5: 233–414. Vidal J.P. (1949). Hémiptères de l’Afrique du Nord et des pays circum-méditerranéens. — Mémoires de la Société des Sciences Naturelles du Maroc, 48: 1–238.
S. Grozeva & N. Simov (Eds)species 2008in Slovenia during the Holocene 121 Survival of the endemic Hemiptera ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 121-128. © Pensoft Publishers Sofia–Moscow
Survival of the endemic Hemiptera species in Slovenia during the Holocene * A. Gogala Slovenian Museum of Natural History, Prešernova 20, p.p. 290, SI-1001 Ljubljana, Slovenia. E-mail:
[email protected]
ABSTRACT The endemic Hemiptera species of Slovenia are found in the karstic south-west of the country. Three endemic Heteroptera and one Auchenorrhyncha species are distributed only in karstic mountains and the Kras plateau. As they live in grassland habitats, but the prevailing vegetation in the past was shown to be forest, a question of their survival during Holocene comes into mind. The existence of small grassland refuges in the south slopes at the peaks and ridges of the mountains is proposed. Such small grassland refuges could not be detected by the pollen analysis in distant deposits. In these refuges, endemic species could survive and evolve from the times of glaciations and soon after, when steppes were widely distributed for the last time, to the present. Keywords: endemic species, relicts, Hemiptera, Heteroptera, Auchenorrhyncha, Slovenia.
INTRODUCTION Slovenia is a relatively small country, but its biodiversity is high due to very diverse geographic regions converging in it. There are Alps in the North and Dinaric mountains to the South, Pannonian influences in the East, and Mediterranean influences in the South-West. The submediterranean region is especially reach as 95% of all of Slovenian species live in the southwestern part of Slovenia, mainly in the karst region covering some 10,000 km2 (Mrši * This paper is dedicated to Michail Josifov, a prominent expert on the Balkan Heteroptera, on the occasion of his 80th birthday.
122 A. Gogala
1997). Most of the endemic insect species found in Slovenia are underground dwellers of karst caves and other underground habitats. The majority of these species are beetles of the families Carabidae and Cholevidae. Among the insect groups which do not penetrate into underground habitats, however, endemic species are rare as all surface habitat types continue across borders. We know only three endemic Heteroptera species, one of them lives also in the neighboring territory of Italy. All three of them live in the Kras (Karst) plateau and/ or the karstic mountains nearby. The endemic Auchenorrhyncha species are also confined to the SW part of Slovenia with submediterranean climate and vegetation. They are part of the larger east Adriatic endemism, rich in unique species, and live also in neighboring areas of Croatia or Italy. One species, Jassargus bicorniger (Then) seems to be limited to an area between the Karst edge in the South and the southern slopes of the Trnovski Gozd mountains in the North (Seljak 2004), just like the endemic Heteroptera species. The Kras (Karst) is a calcareous plateau near the Gulf of Trieste, which gave the name to all karst phenomena in the world. It was largely deforested in early history by people who raised sheep and goats and was long known as a windy, stony desert. Nowadays the land is largely left unmanaged, so that the region is in a natural process of reforestation.
ENDEMIC HETEROPTERA SPECIES Halticus henschii Reuter, 1888 Described by Reuter (1888) on basis of the material from Gorica (Gorizia), it is species with the widest distribution of the three endemics. It is present at the edge of the Trnovski Gozd and Nanos mountains, the Kras plateau and at the Karst edge in Slovene Istria (Gogala 2006). Its foodplant is Genista sericea, where I collected and observed it. Its closest relative, Halticus puncticollis Fieber, is distributed in the Balkans, Ukraine, Caucasus and Kazakhstan (Kerzhner & Josifov 1999).
Dimorphocoris saulii Wagner, 1965
Fig. 1: Halticus henschii Reuter, 1888
This brachypterous species was collected first by Luciano Sauli in the mountain Vremščica (1027 m), mentioned in the descriptions by Wagner (1965, 1969) by its Italian name Mt. Auremiano. This karstic mountain is still the only place where it is known from. It can be found there (800 – 930
Survival of the endemic Hemiptera species in Slovenia during the Holocene 123
†
‡
Fig. 2: A couple of Dimorphocoris saulii Wagner, 1965
m a.s.l.) only for a few weeks in June or July, on grass which was identified as Bromus erectus condensatus (Gogala 1992). It is possible that other grass species are also its foodplants. Related species were later described from Italy and the closest locality where one of them (Dimorphocoris servadeii Tamanini) was found, is Mt. Catria on the other side of the Adriatic (Tamanini 1982). Another Italian species (D. marci), found in the central Appennins, was described by Rizzotti Vlach (1998). A related species was discovered also in Greece: Dimorphocoris sari was found on Mt. Taigetos on Peloponnese and described by Linnavuori (1992). All of these species are restricted to a single or a few mountain habitats.
Platycranus boreae Gogala, 2002 I found this species approximately 800 m a.s.l. in the mountain ridge of Lipnik above the Karst edge in Slovene Istria (Gogala 2002). This is still its only known locality, although the same ridge continues across the border to Croatia and I am sure the species lives also there. The locality is one of the largest remains of grasslands in the Slovene karst, but P. boreae lives only at the top of the exposed ridge, where its foodplant, Genista sericea, grows in large aggregations. The closest relative of this small species is probably P. jurineae Putshkov from the Ukraine.
124 A. Gogala
Fig. 3: A couple of Platycranus boreae Gogala, 2002
ECOLOGICAL DEMANDS OF THE ENDEMICS The endemic Hemiptera species live in the open, steppe-like habitats, on grasses or other plants. This type of habitat is endangered today because of natural reforestation as a consequence of diminishing use of the land for pastures. Largest grassland remains exist in the south-exposed slopes of the mountains, where strong bora wind potentiates the effects of summer drought and physiological drought in winter when the ground is frozen. Trees grow also there, but much more slowly and many die or grow only at the side opposite to the prevailing wind. According to the prevailing view all grasslands in the karstic region are man-made and forest is the natural vegetation of the area. How could then the endemic species survive as they cannot live in the forest? A hypothesis was postulated by botanists that seeds of steppe plants were transported in animal fur from the South during migrations of livestock to new grazing lands. The insect species could follow, but what about the endemics? We do not know of their populations in the South, although they could still exist unnoticed. Much more credible is a hypothesis which states that small refuges for grassland flora and fauna existed in the karstic mountains also before people came to the area. Two of the endemic Heteroptera species live on Genista sericea which grows in the
Survival of the endemic Hemiptera species in Slovenia during the Holocene 125
most stony, exposed ridges, in the plant association Genisto sericeae-Seslerietum juncifoliae, together with the grass Sesleria juncifolia (= S. tenuifolia) and other thermophilous and xerophilous plant species of Illyrian or Mediterranean distribution (Kaligari 1997). The areas where this plant association grows were probably never overgrown by forest due to the stony calcareous ground, inclination to the south and windy conditions (the bora wind).
HISTORY OF THE VEGETATION Sediments older than 500 years, useful for pollen analyses, did not accumulate on the karstic ground. The nearest localities where such investigations were done are at the Adriatic coast in Slovene Istria. The investigations in the Škocjanski zatok near Koper and the Sečovlje salt pans showed the development of the Holocene forest vegetation from approximately 7000 to 500 years b.p. (Culiberg 2005). In the oldest sediments pollens of Fagus, Abies, Quercus and Carpinus were found. Such forests thrive today in much higher elevations in the interior. Soon the degradation of the forest began by people. Pollen of tree species became rarer around the year 1000 a.d. when herbaceous and grassland vegetation proliferated. The Karst was deforested in the 15th and 16th century and at the end of the 18th century only bare rock remained in many places. Investigations showed that the natural vegetation in the area was forest of the associations Abieto-Fagetum and Querco- Fig. 4: The meadows on Mt. Vremščica (1027 Carpinetum. What we do not know, m) are home to the endemic heteropteran bug however, is the situation at the peaks Dimorphocoris saulii, which is wingless and thus and ridges of the karstic mountains. Is it cannot fly. The nearest location where a related possible that small grassland refuges re- species, D. servadeii, was discovered, is Mt. mained there from the times when whole Catria in the Italian Apennines, on the other side of the Adriatic sea. The shallow northern part area was covered by steppe vegetation? of the Adriatic (in lighter color) was dry during The existence of relict endemic species the glaciations and a common ancestor of both in meadows at the karstic peaks speaks in species probably also lived in the dry northern favor of this hypothesis. Such situation is Adriatic sea bed during the glaciation periods also indicated by names of many peaks in of the Pleistocene, when the steppes were much the karstic area like Golič (the naked) and more widely distributed.
126 A. Gogala
Pleša (bald one), although we do not know how old these names are. Small grassland refuges could not be detected by the pollen analysis in distant deposits. During the last glaciation period (22,000 – 14,000 years b.p.) cold and dry climate caused that much of southern Europe was covered by dry, almost semidesert steppe vegetation (Adams 1997). The sea level was much lower than today and the shallow northern Adriatic was dry during that time. There was no barrier between the mountains Vremščica and Catria, where related Dimorphocoris species live today. In that time a common ancestor of both species could live in a broad area where steppe vegetation was distributed. When ice caps melted after warming of the climate around 13,000 years b.p., steppes replaced tundra in a great part of Europe as the trees needed much more time for spread Fig. 5: Stony grassland on Mt. Lipnik, the and regrowth. Colder climate returned habitat of Platycranus boreae. again in younger dryas (11,000 – 10,000 years b.p.) when the steppes thrived again in much of southern Europe for the last time. Then the forest reoccured but was more open than at present until about 8000 years b.p. The clearing of the forests for agriculture began soon after 7000 years b. p. Only for about 1000 years the steppe flora and fauna had to survive in the supposed refuges in the mountains to spread again to the man-made pastures. Some endemic Heteroptera species, however, never spread out of their mountain refuges and can be found today only there. The mountains of Trnovski Gozd, Nanos and Vremščica form a barrier today for the warm air from the coast and are dividing the submediterranean vegetation on their south-orienting slopes from the Central European-type vegetation in the interior. During the last glaciation the same barrier was a southern limit for the arctic fauna. North of this barrier in Postojna the remains of marmots, arctic hares and reindeer, animals which live in the tundra, were found. South of this barrier, only cave bears, cave lions, deer and roe lived (Pohar 1992). We can conclude that tundra-type vegetation thrived North of this mountain barrier and steppe-like vegetation to the South. That is the cause for the endemic Hemiptera species, relics of the old Mediterranean steppe fauna, not to appear North of this mountain barrier, where they could not survive the glaciations. They are found only in the Kras and surrounding mountains.
Survival of the endemic Hemiptera species in Slovenia during the Holocene 127
CONCLUSIONS The endemic Hemiptera species of Slovenia live in grassland habitats and are confined to the south-western part of Slovenia with submediterranean vegetation. The three endemic Heteroptera and one Auchenorrhyncha species are found only in the Kras plateau and mountains in its fringe. They are bound to the limestone ground which does not keep moisture well and is prone to summer droughts. The pallinological research showed that forest is a natural vegetation in the area. The endemic species are relics from the times before the glaciations, probably from the tertiary. When the climate was cooler and drier and when steppes covered much of South Europe, they could spread wider and populate new territories. When forests spread to their habitats, however, they survived only in the mountain refuges. They could not populate man-made pastures later from anywhere else than from these grassland refuges, which were most probably situated at the southern slopes at the top of mountain peaks and ridges and the majority of the endemic species never left their mountains. They are still found only there. The grassland refuges were probably maintained by the prevailing bora wind, which blows in the Kras especially strong in winter, desiccates ground and potentiates physiological drought for plants when the ground is frozen as well as summer drought which usually appears in the area. We should not overlook also the potential role of fire. It is a common feature in the summer today, but is usually stopped soon by firemen. In the times before people populated the area fire did not broke out so often, but when it did, nothing stopped its spread. The bora wind successfully accelerates the spread of fire. Fire burns trees and soil which is then eroded away. In such a way, the vegetation is returned to an earlier succesive phase. The habitats of endemic species and the species themselves are endangered today by spread of the forest vegetation and by plans for wind power plants which would be build in the mountain ridges, exact places where the endemic species live.
РЕЗЮМЕ Ендемичните за Словения видове Hemiptera са намерени в югозападната карстова част на страната. Tри ендемични вида Heteroptera и един вид Auchenorrhyncha са описани само за карстовите планини и платото Крас. Тъй като тези видове живеят в тревисти хабитати, появили се на места с преобладаващи в миналото гори, възниква въпросът как те са оцелели през Холоцена. Допуска се съществуването на малки тревни рефугиуми по южните склонове на върховете и по билата на планините. Наличието на такива малки тревни рефугиуми е невъзможно да бъде уловено чрез поленов анализ на древни отложения. В такива рефугиуми ендемичните видове биха могли да оцелеят и еволюират по времето на заледяванията, скоро след това, когато степите са били широко разпространени за последен път в района, и чак до наши дни.
128 A. Gogala
REFERENCES Adams J.M. (1997). Global land environments since the last interglacial. — Oak Ridge National Laboratory, TN, USA. http://www.esd.ornl.gov/ern/qen/nerc.html Culiberg M. (2005). Paleobotanične raziskave na Krasu, Paleobotanic Research on the Kras (Classical Karst). — In: Mihevc, A. (Ed.): Voda in življenje v kamniti pokrajini, Water and life in a rocky landscape. Založba ZRC, Ljubljana, 149-154. Gogala A. (2002). Platycranus (Genistocapsus) boreae nov. sp. from Slovenia (Hemiptera: Heteroptera: Miridae: Orthotylinae). — Acta Entomologica Slovenica, 10: 13-20. Gogala A. (2006). Heteroptera of Slovenia, III: Miridae. — Annales, Annals for Istrian and Mediterranean Studies, Series historia naturalis, 16 (1): 77-112. Gogala M. (1992). Zgodba o dvoličnikih Dimorphocoris (A tale of the Dimorphocoris bugs). — Proteus, Ljubljana, 55: 59-61. Kaligari M. (1997). Rastlinstvo Primorskega krasa in Slovenske Istre – travniki in pašniki (Vegetation of karst of Primorje and Slovene Istra – meadows and pastures). — Zgodovinsko društvo za južno Primorsko, Znanstveno raziskovalno središče republike Slovenije Koper, 1-111. Kerzhner I.M. & M. Josifov (1999). Miridae. — In: Aukema, B. & Ch. Rieger: Catalogue of the Heteroptera of the Palaearctic Region, Volume 3, Cimicomorpha II. Amsterdam. Linnavuori R.E. (1992). The lateralis group of the genus Dimorphocoris Reuter of Greece and the Middle East (Heteroptera, Miridae, Halticini). — Entomologica Fennica, 3: 215-222. Mrši N. (1997). Biotska raznovrstnost v Sloveniji, Slovenija – vroča točka Evrope, Biotic Diversity in Slovenia, Slovenia – the hot spot of Europe. — Ministrstvo za okolje in prostor, Uprava RS za varstvo narave, Ljubljana, 1-129. Pohar V. (1992). Poseljenost Kraškega roba v kvartarju (Population of the Karst edge in the quaternary). — Proteus, 54: 266-268, Ljubljana. Reuter O.M. (1888). Hémiptères-Hétéroptères des environs de Gorice (Illyrie). — Revue d'Entomologie, 7: 57-61. Rizzotti Vlach M.R. (1998). Una nuova specie di Dimorphocoris dell'Appennino centrale (Heteroptera Miridae). — Bollettino della Società Entomologica Italiana, 130: 17-25. Seljak G. (2004). Contribution to the knowledge of planthoppers and leafhoppers of Slovenia (Hemiptera: Auchenorrhyncha). — Acta Entomologica Slovenica, 12 (2): 189-216. Tamanini L. (1982). Dimorphocoris servadeii n. sp., degli Appennini. — Memorie della Società Entomologica Italiana, 60 (1981): 335-341. Wagner E. (1965). Über die Gattung Dimorphocoris Reuter, 1891. — Reichenbachia, 5: 135-156. Wagner E. (1969). Über Dimorphocoris saulii E. Wagner, 1965. — Bollettino della Società Entomologica Italiana, 99/101: 77-80.
S. Grozeva & N. Simov (Eds)josifovi 2008 nov. sp. (Heteroptera, Tingidae) 129 A new species of lacebug fromTajikistan, Catoplatus ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 129-132. © Pensoft Publishers Sofia–Moscow
A new species of lacebug fromTajikistan, Catoplatus josifovi nov. sp. (Heteroptera, Tingidae) V.B. Golub Voronezh State University, Universitetskaya Pl. 1, 394006 Voronezh, Russia. E-mail:
[email protected]
ABSTRACT Catoplatus josifovi nov. sp. from southern spurs of Gyssar Range of Tajikistan is described and illustrated. Keywords: Catoplatus josifovi nov. sp., Heteroptera, Tajikistan, Tingidae.
INTRODUCTION In this paper, Catoplatus josifovi nov. sp. is described as twelfth known Palaearctic species of Catoplatus Spinola. The paper is based on the material of the Zoological Institute, St. Petersburg, Russia. This paper is dedicated to the famous researcher of Heteroptera and remarkable person Dr. Michail Josifov on the occasion of his 80th birthday.
RESULTS Catoplatus josifovi nov. sp. (Figs 1, 2). HOLOTYPE ♂: Tajikistan, N of Stalinabad (now Dushanbe), headwaters of the river Kondara, Obi-safed tract, 3 V 1944 (Kiritshenko). PARATYPES: 1♂, 2♀♀, Tajikistan, 35 km N of Stalinabad (now Dushanbe), headwaters of the Kondara River, Kvak tract, alt. 2000 m, 27 VIII 1937, 6 VI 1943
130 V.B. Golub
Fig. 1: Catoplatus josifovi nov. sp., ♂, general view. Scale bar = 1 mm.
(Kiritshenko); 1♀, Tajikistan, N of Stalinabad (now Dushanbe), valley of the Varzob River, gorge Kondara, 5 VI 1944 (Kirjanova); 1♂, 2♀♀, Tajikistan, N of Stalinabad (now Dushanbe), Varzob River, Kondara-Kvak, 5 VI 1943, 4 V 1944 (Kiritshenko). Holotype and Paratypes deposited in collection of the Zoological Institute, St. Petersburg, Russia. Distribution. Southern spurs of Gyssar Range, Western Tajikistan. Description. Body elongated, 2.57-2.7 (♂♂), or 2.4-2.51 (♀♀) times as long as wide. Dorsal colour dirty yellow, usually rather light. Head black, covered with light bloom, most developed along inner margin of eye; head rather broad, 2.43-2.67 (♂♂) or 2.2-2.7 (♀♀) as wide as long, with only one pair of small frontal spines having dark basis and light apex, directed obliquely forward and
Fig. 2: Catoplatus josifovi nov. sp., paranotum, Scale bar = 0.5 mm.
A new species of lacebug fromTajikistan, Catoplatus josifovi nov. sp. (Heteroptera, Tingidae) 131
downward; spines parallel, often conniving or even contacting with each other. Occipital spines absent. Antennae thick, thickness of their segment I equal 0.14 mm, moderately long, segments I touching or almost touching by their bases; antennae almost wholly brownish yellow or light yellow, only apex of segment IV blackish. Antennae covered with extremely short, light, pressed hairs which are a little longer on segment IV. Ratio of length of antennal segment III to width of head 1.07-1.18 (♂♂) or 0.93-1.09 (♀♀). Antenniferous tubercles short, their apex curved toward medial line of head. Pronotum with strongly convex disc and three very low longitudinal carinae without areolae; carinae almost deleted in the highest place of disc. Hood (or else vesicula) roof-shaped, low, with slightly prominent towards anterior margin. Paranota reflexed and pressed to sides of pronotal disc, rather wide (see sideways), with two rows of angular areolae along almost all their length, only in the same base and in apex with one row, ♀♀ not rarely having several areolae of the third row in the middle part of their length. Macropterous form. Hemelytra with rather small areolae. Exterior margin of hemelytra sinuate in the anteapical part. Costal area very narrow, carinate and without of areolae in the most part of its length, slightly dilated in the base of hemelytron and near of its apex and here with one row of very small and not always distinct areolae. Subcostal area biseriate in almost all its length, with one row only in its base and apex. Discoidal area with 6-7 (♂♂) or 7 (♀♀) irregular rows of areolae in the widest part. Sutural (inner) area with 10-11 (♂♂) or 11-12 (♀♀) irregular rows of areolae in the widest part. Body ventrally brownish yellow, places brown. Legs rather long, almost wholly light dirty yellow, not darker than body and antennae, only the apices of tarsi are blackish. Femora tibiae not very thick, approximately as thick as antennal segment III or slightly thinner. Fore tibiae 1.45; middle tibiae 1.5-1.55; hind tibiae 1.8 times longer than head width. Fore tibiae 1.2; middle tibiae 1.3; hind tibiae 1.5 longer than antennal segment III. Measurements (in mm). Body length 3.8-4.1, body width 1.48-1.62, pronotum width 1.25-1.35, head length 0.21-0.23 (♂♂) or 0.21-0.26 (♀♀), head width 0.53-0.56 (♂♂) or 0.56-0.59 (♀♀), vertex width 0.26-0.29. Length of antennal segments (I-IV): 0.18-0.19; 0.16-0.18; 0.6-0.66(♂) or 0.51-0.61(♀); 0.26-0.29. Length of tibiae: fore 0.81; middle 0.86; hind 1.0. Length of tarsi: fore 0.26; middle 0.27; hind 0.29 mm. Etymology. I am very glad to dedicate this species to the famous heteropterologist Michail Josifov. Diagnosis. The new species is similar to Catoplatus hilaris Horváth, 1906 in the very narrow, carinae like costal area, its light, brownish yellow antennae (except apex of segment IV, and its two rows of areolae of subcostal area of hemelytra. The latter species is distributed in Transcaucasia (Armenia, Georgia) and Mediterranean area (Péricart 1983; Péricart & Golub 1996). C. hilaris distinguishes from the C. josifovi in having 4-5 head spines (2-3 frontal and 2 small occipital), shorter as a rule antennal segment III (0.4-0.58 mm), narrower as a rule head (0.48-0.55 mm), narrower paranota with two rows of areolae only in their fore half, the lack of a deep depression in the anteapical part of exterior margin of hemelytron.
132 V.B. Golub
The very narrow costal area of hemelytra of C. hilaris has nevertheless 1 row of areolae in its whole its length. Body of C. hilaris is usually narrower (1.3-1.52 mm).
ACKNOWLEDGEMENTS The author is indebted to Professor I.M. Kerzhner for the opportunity to study materials of the Zoological Institute, St. Petersburg, Russia. The work was supported by the Russian Foundation for Basic Research (grants 05-04-49089-a, 05-04-49917-a, 08-04-00212-a).
РЕЗЮМЕ Описан и илюстриран е Catoplatus josifovi nov. sp. от южните склонове на Хисарския хребет в Западен Таджикистан.
REFERENCES Horváth G. (1906). Synopsis Tingitidarum Regionis palaearcticae. — Annales Historico-Naturales Musei Nationalis Hungarici, 4: 1-117. Péricart J. (1983). Hémiptères Tingidae euro-méditerranéens. — Faune de France, 69: 1-620. Péricart J. & Golub V.B. (1996). Superfamily Tingoidea Laporte, 1832. — In: B. Aukema & Chr. Rieger (Eds). Catalogue of the Heteroptera of Palaearctic Region. Vol. 2. Cimicomorpha I Wageningen: 3-83.
S. Grozeva & N.1909 Simov 2008 A new species of Euchilofulvius Poppius, from(Eds) Malaysia (Heteroptera: Miridae: Cylapinae) 133 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 133-137. © Pensoft Publishers Sofia–Moscow
A new species of Euchilofulvius POPPIUS, 1909 from Malaysia (Heteroptera: Miridae: Cylapinae) J. Gorczyca Department of Zoology, University of Silesia, Bankowa 9, 40-007 Katowice, Poland. E-mail:
[email protected]
ABSTRACT A new species of the Cylapinae genus Euchilofulvius is described on the basis of specimens collected in Malaysia. A key to the species of the genus is given, and the parameres of the new species and those of E. antennatus Gorczyca are presented. Keywords: Cylapinae, Euchilofulvius, Heteroptera, Miridae, new species, taxonomy.
INTRODUCTION The genus Euchilofulivus has been established by Poppius (1909) for a single species E. tibialis, described on the basis of a single female collected on Mentawei Island. For a long time this was the only report on this genus and species. Several years ago I revised the genus Euchilofulvius, described two new species from India and Sumatra, and provided new data on the distribution of Euchilofulvius tibialis Popp (Gorczyca 1998). In the following years new species were described from Sulawesi, the Solomon Islands and Japan (Gorczyca 1999, 2002; Yasunaga & Miyamoto 2006). Up to now six species of the genus Euchilofulvius have been described and one has been synonymised (Gorczyca 2002; Yasunaga & Miyamoto 2006). This genus was erroneously divided into two subgenera and further into two complexes (Gorczyca 1999, 2002). Within the material deposited in the collection of Naturhistorisches Museum in Vienna I found three specimens representing a new species of the genus Euchilofulvius from Malaysia. The description of the new species is given below.
134 J. Gorczyca
TAXONOMY Euchilofulvius josifovi nov. sp. Type material: Holotype ♀: [Malaysia W., Kelantan, Road between Kampong Raja and Gua Musang 1400-1700 m (Ladang Pandrak), 1-28. iv 2006, 4º63` N - 101º45` E/4º88` N - 101º95` E., Cechovsky Petr leg.], paratype ♂ the same data as holotype, paratype ♀: [Malaysia West, Pahang Cameron Higlands, Tanah Rata, 1200-1500m, 3. Ii – 19. Ii. 2005, Cechovsky Petr leg.]. Housed in Naturhistorisches Museum in Vienna, Austria. Etymology. This species is named for the prominent heteropterist Professor Michail Josifov in honour of his inestimable contribution to the Heteroptera. Diagnosis. This species can be easily distinguished from E. zdzislawi Gorczyca and other representatives of the genus by a longitudinal pale stripe on scutellum and head with pale swellings, as well as by the shape of parameres (Figs 1-4). Description. Male. Body dark brown covered with pale scale-like setae. Length of the body 3.60 mm, width 1.28 mm, head dark brown with five slightly paler swellings covered with pale, scale-like setae. Length of the head 0.60 mm, width 0.62 mm, diameter of eye 0.20 mm. The first antennal segment relatively thick, brown, covered with short setae, second segment almost cylindrical, only slightly thicker in apical part. Third and fourth segments very short, third segment pale, fourth darker. Length of the antennal segments in mm: 0.45: 1.12: 0.27: 0.30. Rostrum brown, thin, reaching at least mesocoxae, length of the individual segments unmeasurable in the specimen examined. Pronotum dark brown, calli slightly raised, posterior margin paler, the posterior lobe of pronotum with two pale, longitudinal stripes perpendicular to the posterior margin. Length of pronotum 0.65 mm, length of the anterior margin 0.45 mm, the lateral margins 0.70 mm, the posterior margin 1.15 mm. Mesoscutum dark brown with paler stripe on sides, scutellum dark brown with short, pale, longitudinal stripe in the middle. Apex of scutellum pale. Hemelytra brown with pale patches, clavus unicoloured, dark brown, corium with large pale patch contiguous with median fracture, embolium and cuneus slightly chestnut, there are small pale patches above the cuneus, apex of cuneus pale. Membrane dark grey, venation dark, slightly marked, major cell triangular, minor cell hardly visible. Body dark brown ventrally, abdomen tinged with red. Coxae and trochanters invisible in the specimen examined. Femora pale at base then dark brown with a wide, pale ring near the apex, apex dark brown with pale patch tinged with red. Tibiae brown at base, pale brown at the apex, tarsi very short, slim, two-segmented, claws toothed subapically. Aedeagus membranous, parameres as on Figures 1, 3. I compared the parameres of the new species with those of E. antennatus because genitalia of this species have never been pictured. Female. Similar to male but bigger, pale stripes and patches on the body more contrasting than in male. First and second antennal segments covered with scale-like setae,
A new species of Euchilofulvius Poppius, 1909 from Malaysia (Heteroptera: Miridae: Cylapinae) 135
pale, longitudinal stripe on scutellum reaching mesoscutum. Embolium and cuneus more distinctly tinged with red than in male, apex of cuneus only slightly paler. Length of the body 3.70 – 3.80 mm, width 1.40-1.47 mm, length of head 0.55 mm, width 0.60 mm, diameter of eye 0.12 mm. Length of the antennal segments in mm: 0.45: 1.05 (third and fourth segments broken in the specimens examined). Length of the individual rostral segments unmeasurable in the specimens examined. Length of pronotum 0.67 mm, anterior margin 0.50 mm, lateral margins 0.75 mm, posterior margin 1.18-1.20 mm.
1
2
3
4
0.1 mm
Figs 1-2: left parameres, 3-4 right parameres, Figs 1, 3: Euchilofulvius josifovi nov. sp.; 2, 4 - E. antennatus Gorczyca
136 J. Gorczyca
Key to the species of Euchilofulvius POPPIUS 1. 2. 3. 4. 5. 6. -
Length of the body more than 3.50 mm ..............................................................2 Length of the body less than 3.50 mm.................................................................4 Scutellum with a pale longitudinal stripe in the middle reaching mesoscutum in female, shorter in male, head with five pale swellings ................. E. josifovi nov. sp. Scutellum without a pale longitudinal stripe in the middle or with a very short stripe in female, never reaching mesoscutum, head without pale swellings ..........3 Posterior lobe of pronotum unicoloured, pale brown, tibiae brown, length of the male body more than 4.0 mm .......................................... E. zdzislawi Gorczyca Posterior lobe of pronotum darkened on sides, tibiae paler in the apical part, length of the male body less than 4.0 mm ........... E. lepidocerus Yasunaga & Miyamoto Antennae, tibiae and hemelytra pale .....................................E. pallens Gorczyca Antennae, tibiae and hemelytra at least partly darkened ......................................5 Anterior lobe of pronotum pale brown................................E. carinatus (Poppius) Anterior lobe of pronotum dark brown, almost black (sometimes calli paler) ......6 Tibiae pale or tinged with red ........................................ E. antennatus Gorczyca Tibiae pale at base than dark brown and contrasting pale in the apical part ......... ..................................................................................................E. tibialis Poppius
ACKNOWLEDGEMENTS I would like to express my sincere thanks to Herbert Zettel (Naturhistorisches Museum Wien, Vienna, Austria) for the loan of the specimens, Snejana Grozeva and Nikolay Simov (Bulgarian Academy of Sciences) who kindly invited me to contribute to this volume and Krystyna Warchał (University of Silesia) for her help and improvements in the language.
РЕЗЮМЕ На базата на индивиди, събрани в Малайзия, е описан нов вид от род Euchilofulvius (Cylapinae). Предложен е ключ за определяне на видовете от рода. Представени са парамерите на новия вид и на E. antennatus GORCZYCA.
REFERENCES Gorczyca J. (1998). A revision of Euchilofulvius Poppius (Heteroptera: Miridae: Cylapinae). — European Journal of Entomology, 95: 93-98.
A new species of Euchilofulvius Poppius, 1909 from Malaysia (Heteroptera: Miridae: Cylapinae) 137
Gorczyca J. (1999). On the Euchilofulvius-complex (Heteroptera, Miridae, Cylapinae). — International Journal of Invertebrate Taxonomy Genus, 10(1): 1-12. Gorczyca J. (2002). A new species of the genus Euchilofulvius Poppius (Hemiptera: Heteroptera: Miridae) from the Solomon Islands. — Polskie Pismo Entomologiczne, 71: 297-299. Poppius B. (1909). Zur Kentnis der Miriden-Unterfamilie Cylapina Reut. — Acta Societatis Scientiarum Fenniceae, 37(4): 1-46. Yasunaga T. & S. Miyamoto (2006). Second report on the Japanese cylapine plant bugs (Heteroptera, Miridae, Cylapinae), with descriptions of five new species. In: Rabitsch W. (Ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von ERNST HEISS. — Denisia 19: 721-735.
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S. Grozeva & N. (Eds) 2008 OnSimov some threatened Heteroptera from the Iberian fauna 139 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 139-158. © Pensoft Publishers Sofia–Moscow
On some threatened Heteroptera from the Iberian fauna M. Goula1, M. Costas2, S. Pagola-Carte3, M. Baena4, T. López2, A. Vázquez2, F. Gessé1, J. Ribes5 & E. Ribes5 1
Dpt. Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Avda Diagonal 645, 08028 Barcelona, Spain. E-mail:
[email protected],
[email protected] 2 Dpto. de Zoología y Antropología Física, Facultad de Biología, Universidad Complutense de Madrid, C./ José Antonio Novais, 2, 28040 Madrid, Spain. E-mail:
[email protected] 3 Azpeitia 3, 7 D, E-20010 Donostia, Gipuzkoa, Spain. E-mail:
[email protected] 4 Plaza Flor del Olivo 4, Bloq. 7, 1º B, 14001 Córdoba, Spain. E-mail:
[email protected] 5 València 123-125, ent., 3a., 08011 Barcelona, Spain. E-mail:
[email protected],
[email protected]
ABSTRACT The authors were in charge of selecting Heteroptera species under threat in Spain, in order to publish a Red List of Invertebrates, launched by the Spanish Ministry of Environment in 1998 in the frame of the National Inventory of Biodiversity. Twenty-three bug species were suggested. Nine of them, in the category of “Vulnerable” according to the UICN criteria, were finally included in the Red List. The remaining fourteen also deserve to be considered as threatened for different reasons. In this paper, a summary report for each of those fourteen species, concerning their biology, distribution and threatening factors is presented. Species included are: Pachycoleus pusillimus (Sahlberg, 1870) (Dipsocoridae), Rhagovelia (Rhagovelia) nigricans nigricans (Burmeister, 1835) (Veliidae), Acalypta suturalis (Puton, 1879) (Tingidae), Isometopus intrusus (HerrichSchaeffer, 1835), Bryocoris pteridis (Fallén, 1807), Ribautocapsus bruckii (Reuter, 1879), Roudairea crassicornis (Puton & Reuter, 1886) (Miridae), Anthocoris visci Douglas, 1889 (Anthocoridae), Rhynocoris (Rhynocoris) lineaticornis (Reuter, 1895) (Reduviidae), Metatropis rufescens (Herrich-Schaeffer, 1835) (Berytidae), Leptodemus minutus ( Jakovlev, 1874), Auchenodes costalis (Lethierry, 1877) (Oxycarenidae), Trichaphanus fuentei (Puton, 1894) (Rhyparochromidae) and Leptoceraea femoralis (Horváth, 1897)
140 M. Goula et al.
(Rhopalidae). All of them are under threat due to human activities, and their survival includes the protection of their respective habitats. Legislation to promote protection, as well as environmental education and information of the public, are indispensable to achieve the success in species conservation. Keywords: True bugs, Heteroptera, Iberian fauna, protection, threatened fauna.
INTRODUCTION In 1998, the Spanish Ministry of Environment launched the National Inventory of Biodiversity, to give compliment to article number 7 of the Biological Diversity Agreement. This article 7 conveys to identify and monitor those organisms that deserve to be protected, and utilized in a sustainable way. The Inventory was conceived as a series of independent projects, one for each biological group. An Atlas and a Red Book (Red List) were planned to be published for each biological group. The Atlas provides the information on the distribution of the species, and the Red Book compiles the information on the conservation status of those species. After the vertebrates and plants inventories have been already developed and successively published, invertebrates were tackled. Arthropods and terrestrial molluscs were the first groups taken, as the existence of their respective Societies (Asociación Española de Entomología, Sociedad Española de Malacología) do much facilitate the joint work of more than a hundred specialists. The CIBIO (Iberoamerican Centre of Biodiversity) and the University Institute of Research placed in the University of Alicante were in charge of the coordination of all projects. Concerning arthropods, the invertebrate habitats included in the Habitat Directive (Galante & Verdú 2000; Ramos et al. 2001) and the hundred arthropod species included in the Spanish National Catalogue of Threatened Species were the ground precedents to the National Inventory of Biodiversity. Main criteria to select species were those of the IUCN version 3.1 (2001), where rarity, habitat singularity and degree of threaten are taken into account, among others. As a result of the whole process, the group working on Heteroptera made a proposal of 23 threatened species. Nine of them were finally included in the Red List of Invertebrates of Spain, under the threat category “Vulnerable”. The species were the mirids (Plant bugs) Leptopterna pilosa Reuter, 1889, Orthotylus (Orthotylus) siuranus Wagner, 1964, Orthotylus (Pachylops) blascoi J. Ribes, 1991, Parahypsitylus nevadensis E. Wagner, 1957 and Solenoxyphus minor Wagner, 1969, the reduviids (Assassin bugs) Collartida tanausu J. Ribes, Oromí & E. Ribes, 1997, Polytoxus siculus (A. Costa, 1842) and Vibertiola cinerea (Horváth, 1907) and the lygaeid (Seed bug) Tropidothorax sternalis sternalis (Dallas, 1852). Details on their biology, habitat and distribution may be consulted in Verdú & Galante (2006). It is worth to say that total number of threatened arthropod species included in the list was 177, and that the Heteroptera were considered into account for the first time in Spain as threatened organisms. In this paper we introduce the 14 Heteroptera species that also
On some threatened Heteroptera from the Iberian fauna 141
deserve to be considered under threat for different reasons, although the present state of knowledge does not allow to include them in the Red List of Invertebrates of Spain. At present, Heteroptera are absent from the European conservation treatises or agreements. However, Heteroptera have been included in several Red Lists, as seen in the following examples. In Great Britain, 143 Heteroptera species were included in the British Red Data Book concerning insects (Shirt 1987). In Slovenia, 30 species were included in the Red List of endangered Heteroptera of that country (Gogala 1992). In Germany, 303 Heteroptera species were listed as threatened, including those already extinct, as part of a Red List of threatened animals in Germany (Günther et al. 1998). In the Slovakian Republic, 31 heteroptera species have been mentioned in different endangerment categories (Baláž et al. 2001). In the Czech Republic, the Red List includes 260 Heteroptera species (Kment & Vilímová 2005).
LIST OF SPECIES The fourteen species considered are exposed in taxonomic order. The scheme in the Catalogue of the Heteroptera of the Palaearctic region has been chosen (Aukema & Rieger 1995). The only exception is with “lygaeids” (Lygaeidae sensu lato), for which the taxonomic scheme of Henry (1997) has finally been adopted. For each of them, a survey of the most important data on their biology, distribution and threatening factors is included. Also, specific measures of conservation are suggested. In figure 1 the distribution of the fourteen species in Spanish provinces is stated. In each distribution area paragraph, name of the province is in brackets.
Dipsocoromorpha Dipsocoridae Pachycoleus pusillimus (SAHLBERG, 1870) Distribution area. Pachycoleus pusillimus is a European species (Kerzhner 1995). In the Iberian Peninsula, it has been only mentioned in the central zone. It is included in the Regional Catalogue of threatened species of wild fauna and flora from the Community of Madrid as species to be protected (Catálogo in press). The first record was from Guadalix de la Sierra (Madrid), 1♂ (Costas et al. 1993). The other Iberian records are from Miraflores de la Sierra (Madrid), UTM 30TVL4813, 3-VI-1977, R. Outerelo leg. 1♂, on wet-moss (in a pine forest); Laguna El Campillo, Rivas-Vaciamadrid (Madrid), UTM 30TVK5864, 28-XI-2004, M. Á. Vázquez leg., 1♂ and 11-III-2005, M. Á. Vázquez leg., a nymph, in withered leaves. Habitat and biology. Pachycoleus Fieber, 1860 species live under stones and in the slopes of streams and rivers, in the interstitial spaces with high humidity degree (Tamanini 1979; Wagner 1967). Štys (1990) identifies the habitat of Pachycoleus, which
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prefers humid zones with Sphagnum or similar mosses, but without abundant mud; usually Pachycoleus is associated to the presence of other Heteroptera of the genera Hebrus Curtis, 1833, Saldula Van Duzee, 1914, and Chartoscirta Stål, 1868. They predate on other small Arthropoda. Phenology of P. pusillimus is unknown. The four captures of P. pusillimus in the Iberian Peninsula occured by means of Berlese method. In June 1977, the sample came from a sandy slope of a dry stream; in October 1990, the sample was a moss collected on a pine roots near a stream, in November 2004 the species was extracted from wet litter of Populus; and in March 2005 a nymph was obtained also from wet litter of Populus. Threatening factors. Pachycoleus pusillimus may be threatened by several factors.The most important is the misuse, overuse and contamination of rivers and streams, as a consequence of massive enlargement of many towns of the Madrilenian Mountains by the construction of residential areas. Canalization or deflection of streams decrease their natural zones. The increasing spent of water conducts to a reduction of water volume in rivers and streams under the minimum ecological volume, increasing the ecological impact. Finally the existence of uncontrolled spills to the bodies of water and in its banks low the habitat quality. Measures of conservation. The conservation of P. pusillimus is closely tied to the conservation of its habitat. The recommended measures must include the protection of this habitat. For example, it is necessary to preserve streams slopes, and/or to guarantee the minimum volume of water in streams and rivers in order to keep their optimal quality levels. Also, to regulate the use of water providing guarantee of the minimum phreatic level. Canalizations of streams, coating of stream slopes and margins with cement concrete, PVC or similar, modifications of the courses of streams and the throwing of solid or liquid spills, should be avoided. Most important is also the protection of bank vegetation, including mosses, herbs and forests.
Gerromorpha Veliidae Rhagovelia (Rhagovelia) nigricans nigricans (BURMEISTER, 1835) Distribution area. Rhagovelia (R.) nigricans nigricans has been mentioned from Morocco, Tunisia, Egypt, Israel, Yemen, Lebanon and Cyprus, and doubtfuly from Saudi Arabia (Andersen 1995; Baena et al. 1994). An only Iberian locality is known: Guadalete River (Cádiz), 31-X-1988, A. Gallardo leg.; II -VI-90 and IX-90, M. Baena leg. (Baena et al. 1994). This until present only Iberian population seems numerous and stable by the presence of adults and nymphs. Habitat and biology. The species lives in running waters, whenever enough volume and speed of water, low salinity and temperatures between 12 and 23ºC throughout the year are provided; heavy grain substrate, stone blocks, gravel and sands, with scarce vegetation composed by Tamarix africana, Typha dominguensis, Nerium oleander, Scirpus sp. and Salix sp. are preferred.
On some threatened Heteroptera from the Iberian fauna 143
Threatening factors. Rhagovelia nigricans nigricans can be threatened by the degradation of its habitat, by water pollution or reduction of its volume. Conservation measures. It is necessary to directly conserve and maintain the habitat of R. nigricans nigricans. Then, it is needed to prevent the uncontrolled spills, to avoid the alteration of water volumes, not to make deflections of the water channels or their canalization and to protect the margins with native vegetation.
Cimicomorpha Tingidae Acalypta suturalis (PUTON, 1879) Distribution area. Acalypta suturalis is known from Spain and southern France, generally around the Pyrenees (Péricart & Golub 1996). In Spain it has been recorded from Irun (Gipuzkoa) (Puton 1879) and from a collection in Can Tunis (= Casa Antúnez), a littoral Barcelona city area (Barcelona), dated 3.VII.1914, A. Codina leg. (Codina 1915; Sánchez 1920). This Barcelona record is quite dubious. J. Ribes et al. (2004) report Codina’s and Sánchez’s works, whilst Péricart (1983) reports only Codina’s. Habitat and biology. This is a brachypterous species, living on mosses, in the Pyrenees area, probably in turfs and moist woods. Adults are captured from April to September. Nymphs are unknown. Threatening factors. Acalypta suturalis may be threatened by the effect of several activities which affect to its habitat and host plant. Mosses are intensively collected for ornamental purposes, especially in Christmas time. Mosses do not tolerate walking on them, so that it may be impacted by frequentation in the Pyrenees due to mountain tourism, or by wood extractions. Also, the opening of mountain roads that facilitates the vehicle entrance to areas unaccessible until recent also contributes to accelerate the degradation of environment. Water stress caused by long periods with no rain is a threat to the survival of A. suturalis due to drying of mosses. Conservation measures. Increasing touristic activities in the mountains have to be regulated. On the other hand, building of new roads, even for timber activities, should be restricted, as well as circulation of four wheeled vehicles as a good dissuasive method to approach people everywhere in the mountain. The purpose of all those measures is to keep certain areas untouched, specially woods, leaving them to evolve by themselves.
Miridae Isometopus intrusus (HERRICH-SCHAEFFER, 1835) Distribution area. Isometopus intrusus is present in central and southern Europe, and also in Turkey (Kerzhner & Josifov 1999). In the Iberian Peninsula it has been mentioned from El Montecillo (Pozuelo de Calatrava, Ciudad Real) (De la Fuente
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1920), Prades (Tarragona) (Goula & J. Ribes 1995), Santa Coloma (Andorra) (Gessé et al. 1997) and Arraia-Maeztu (Araba) (Pagola-Carte et al. 2004). Habitat and biology. This species lives on the bark of apple trees and other deciduous trees like Tilia sp., Salix sp. ( Josifov 1974; Wagner & Weber 1964) and Fraxinus sp. ( Jordan 1941). It has been collected on the branches of Quercus ilex (De la Fuente 1920) and Q. pyrenaica (Pagola-Carte et al. 2004) and it has been also found in pistachio’s crops in the Anatolian Peninsula (Yanik & Yücel 2001). The nymphs can be found in May and the adults from June to August. It is an agile predator that feeds on aphids (Eriosoma Leach, 1818) and other small arthropods (Wagner & Weber 1964). Threatening factors. This species is considered rare although it is widely distributed all over Europe. The possible fragmentation and reduction of its habitat can cause damages for the maintenance of its populations. The forests where I. intrusus is found are progressively being replaced by other plant species, like plane tree, or occupied by farming and real state constructions. Contamination (pesticides, industrial or urban spills) also contributes to the degradation of its habitat. Conservation measures. The conservation of Isometopus intrusus must be addressed specifically towards the preservation of the habitat that provides shelter to it. It is very important to preserve wide areas big enough to guarantee the maintenance of the characteristics of this habitat, particularly the river banks forests, since they are especially fragile. Agriculture and urban or tourist developments do not imply the complete degradation of the environment. When growing fruit trees suitable for I. intrusus, the use of selective pesticides that do not harm the wild fauna is recommended, as well as keeping some of them in the marginal zones only for preservation, not for production. It cannot be neglected that I. intrusus contributes to the depredation of aphids and other small arthropods, so its presence in crops can be developed within integrated or biological pest control programs.
Bryocoris pteridis (FALLÉN, 1807) Distribution area. Bryocoris pteridis is widely distributed in middle Europe and extends to the Far East of Russia (Kerzhner & Josifov 1999). It has been recently found to the south of the Pyrenees, in the Basque Country (Pagola-Carte et al. 2004, 2005). It is so far only known from a certain number of localities (Errenteria, Hernani, Irun and Larraul) of the north-eastern part of the province of Gipuzkoa. Habitat and biology. Bryocoris pteridis is an inhabitant of damp and shady forests undergrowths, where it lives on several fern species (Wagner 1974a; Ehanno 1987a; Wachmann et al. 2004). It usually shares the habitat and plant hosts with another Bryocorinae, Monalocoris (M.) filicis (Linnaeus, 1758). The Iberian specimens have been collected on Dryopteris filix-mas (Dryopteridaceae), almost always together with M. (M.) filicis. Despite the parallelisms with that other Bryocorinae, in the Basque Country B. pteridis is less frequent, as stated for France (Ehanno 1987b). B. pteridis is univoltine
On some threatened Heteroptera from the Iberian fauna 145
and overwinters as egg. Adults have been observed at the beginning of summer. Typically, males are macropterous and females are brachypterous. A marked preference for plants with well-developed sporangia has been stated (Pagola-Carte et al. 2004). Threatening factors. Iberian populations of B. pteridis constitute the western and southernmost limit of its distribution range. In addition to the intrinsic handicap of female brachypterism and the associated low dispersion rates which prevent rapid colonization of new localities, several factors are identified as threats for the species in the Iberian Peninsula. B. pteridis is considered to be highly sensitive to forest disturbances. The removal of the habitat as a side effect of the wood industry activities, the plantation of coniferous species which preclude the continuity of the undergrowth with ferns, and the use of the forest for cattle or recreational activities are thought to be major problems. It is also presumably that climate warming could affect the habitat of this and other insect species that, given their ecological requirements, are at the edges of their range in the northern Iberian Peninsula. Conservation measures. The conservation of B. pteridis is closely linked to the protection of its habitat. A proper forest management is needed in the areas with known populations of the mirid, as well as in those of presumed presence. Particular attention should be paid to the herbaceous level of the forest undergrowth, avoiding the use of those areas for livestock grazing or sheltering. The opening of new trails or roads should also be controlled. In addition, local measures should be taken to regulate the impact of recreational activities (fishing, hunting, walking, etc.) on the habitats near paths and rivers.
Ribautocapsus bruckii (REUTER, 1879) Distribution area. Ribautocapsus bruckii has been recorded from France, Portugal, Spain and Algeria (Kerzhner & Josifov 1999). In Spain, it had been recorded by Reuter (1879), under a vague location “Hispania”, with no more details. Recently, more specimens have been collected by M. Baena and J.M. Vela in Andalusia, Guéjar Sierra (Granada), in early summer. Habitat and biology. Ribautocapsus bruckii lives in uncultivated, dry and sunny habitats (Wagner 1974b), in lowest levels of herbs (Ehanno 1987a). Perrier (in Wagner & Weber 1964) mentions Arctostaphylos uva-ursi and Echium vulgare as host plants. Streito & Matocq (2006) collected R. brucki with a sweeping net. Adults are present in June and July (Wagner 1974b). M. Baena found the species living in Artemisia sp. Threatening factors. Ruderal or anthropogenic areas frequented by R. bruckii are usually not taken into account, but they are under the threat of certain activities related to agriculture: stubble burning, that fragments and reduces the habitat in time and space, the use of unspecific pesticides, destruction of margins and boundaries due to agriculture intensification. Other threats to the unspecific habitat of R. bruckii are clearing of underwood to prevent wood fires, or of ditches to maintain roads, and increasing urbanization in wasted lands.
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Conservation measures. Conservation of the species requires information and environmental education of both the public and the environmental managers, so that they become conscious of the value of those unspecific and apparently banal habitats along the roads, of the uncultivated lands, or the undergrowth, that are the refugees for species like R. bruckii. Less emblematic habitats are sometimes very difficult to preserve, as they are considered with no ecological value and unthreatened. Only an attitudinal change in the sense of increasing the positive appreciation of such habitats, may be the key to include them in management decisions.
Roudairea crassicornis PUTON & REUTER, 1886 ( = Roudeirea [sic] launaeae Wagner, 1975; Wagner (1975, 1976); syn. J. Ribes in J. Ribes et al. 1997) Distribution area. Roudairea crassicornis is an Eremian species described from Tunisia, also recorded from Spain, Algeria, Saudi Arabia and Iraq (Kerzhner & Josifov 1999). Spanish records are very scarce, based on a few specimens collected in Playa del Saladar beach, belonging to Arenales del Sol (Alacant, País Valencià) ( J. Ribes & Sauleda 1979) and Retuerta de Pina, in Los Monegros steppic region (Zaragoza, Aragón) ( J. Ribes et al. 1997). Habitat and biology. Very little is known on R. crassicornis biology. In Alacant it lives on the Asteraceae Launaea fragilis (=L. resedifolia) within the Ammophilion plant community, between May and June (in 1973 and 1974). It is worth to stress the homochromy between the mirid and the flower buds of its host plant ( J. Ribes & Sauleda 1979). In 1993 in Los Monegros, where L. fragilis also thrives, it was collected by light trap. Threatening factors. In Alacant, beach tourism is the main negative impact for the species since it involves urban development in the coast, together with extremely high frequentation rates during long periods each year. All that is harmful for the typical flora and, hence, the habitat of R. crassicornis. In Los Monegros, the major threatening factor does not only concern this mirid, but the whole biota. The ancestral xeric features of the area of Retuerta de Pina may become seriously affected by the irrigation-intensive farming, which is being developed in recent years. Damages to the open forests of Juniperus thurifera and their associated communities could soon be irreversible. Conservation measures. The conservation measures necessarily involve the protection of the species habitat. In general, the aim is to minimize the impact of human activities. In the beaches of Alacant, legal actions have to be designed to conserve some patches of littoral areas including the features required by R. crassicornis; buildings, traffic and human frequentation would be prevented there. In the continental xeric area Los Monegros (Zaragoza), quite enough wide areas of Juniperus forests should be kept protected from irrigation-intensive farming, in order to provide with shelter R. crassicornis and a vast set of other interesting species present in the area and similarly sensitive to the same threats.
On some threatened Heteroptera from the Iberian fauna 147
Anthocoridae Anthocoris visci DOUGLAS, 1889 Distribution area. Anthocoris visci is present in Belgium, Czech Republic, France, Great Britain, Germany, Luxemburg, Macedonia, the Netherlands, Spain, and Asian part of Turkey, and doubtfully in former Yugoslavia (Péricart 1996), so that its distribution area is smaller than the area of its host plant, Viscum album. Spanish records are all only Catalan: Les, Val d’Aran (Lleida), 26.II.1914, L. Hilaire leg., Sánchez (1920); Les, Val d’Aran (Lleida), 26-IV-1914, Codina (1915); Montgat (Barcelona), Torre Bueno (1912), reported also in Gómez Menor (1956) and Péricart (1972). J. Ribes et al. (2004) include both records from Montgat and Les, as well as two more citations from the Natural Park of Collserola, erroneously attributed to J.Ribes & E.Ribes (2001), that of course have to be discarded. A more recent collection of a very small female, under the usual species size, at Els Torms, Garrigues (Lleida), 23.VI.1968. J.Ribes leg. et coll., confirms the presence of A.visci in Catalonia. Habitat and biology. Anthocoris visci lives exclusively on mistletoe, Viscum album, where it probably preys on Cacopsylla visci (Curtis, 1835) (Psyllidae). A. visci has been abundantly collected on mistletoe of apple and almond trees. Eggs are laid under the epidermis of the inner face of very young leaves of mistletoe, either isolated or in little groups (2-3 eggs). Only the operculum protrudes outside the leaf. Adults are collected from spring to autumn. Maybe they hibernate under the bark of mistletoe host plant. Threatening factors. Anthocoris visci is thought to be vulnerable because its host plant, V. album, may be locally very abundant, but it is considered a rare species generally speaking. Mistletoe is intensively collected for ornamental purposes, specially, in Christmas time. Also, woods parasited by mistletoe are not left abandoned, but managed, threatening the survival of A. visci. Reduction on mistletoe distribution may also occur by unappropiated forest management, as is reforestation to thick densities providing a too shady environment not suitable to V. album, or by introduction of allochtonous shade trees not susceptible to mistletoe. Conservation measures. The conservation measures are directly correlated to factors which negatively affect mistletoe. Extraction of mistletoe should be regulated to avoid abusive practice. Also, a change in the perception of old or parasitized woods is needed, that should be understood as part of the wood cycle. The preservation of untouched woody areas where the parasite could install and spread would be required. Also in fruit trees areas, some of the old trees could be spared in the margins, giving support to parasitation. The aim is to provide an stable population of mistletoe in which A. visci may complete its cycle during long periods of time. A. visci is catalogued as “Notable B” in Great Britain (Kirby 1992), and as endangered, with unknown status, in Germany (Günther et al. 1998).
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Reduviidae Rhynocoris (Rhynocoris) lineaticornis (REUTER, 1895) Distribution area. Rhynocoris lineaticornis is known from Spain, Algeria, Morocco and Chad (Putshkov & Putshkov 1996). The only Spanish record is the capture of two males in Fuente del Pozuelo, near to the locality of Grazalema (Cádiz) (Baena 1985). Habitat and biology. Populations of R. lineaticornis are usually scarce all over the distribution area. The two macropterous males of Grazalema were captured beating on Mediterranean bush vegetation. The brachypterous females must live on the soil, where are more difficult to be captured. Probably, hibernation occurs in the last nymphal stages, as it happens in other Rhynocoris Hahn, 1833 species. Threatening factors. The two threatening factors for R. lineaticornis are bush fires and wood reforestation. It has to be taken into account that Mediterranean bush is very susceptible to fires, and human frequentation by trekking in the wild may increase fire thread. On the other hand, Mediterranean bush is a stage in vegetation succession to Mediterranean woods. Reforestation may be promoted by administration, affecting to the natural conditions of the habitat of R. lineaticornis. Conservation measures. Survival of R. lineaticornis is linked to the conservation of the Mediterranean bush vegetation. With this aim, it is necessary to enhance the persistence of this transitional stage of vegetation by avoiding or reducing reforestation. Also, as the locality where the species was found is very close to a very frequented area of “Pinsapar of Grazalema”, general information about habitat with examples of threatened plant and animal species must be given to inform the visitors and to avoid fires.
Berytidae Metatropis rufescens (HERRICH-SCHAEFFER, 1835) Distribution area. Metatropis rufescens is widely distributed in northern and central Europe (Péricart 2001b). It is less frequent towards the southern parts of the continent, being virtually absent from the Mediterranean area. Considered to be a rare species (Péricart 1984), it is found in quite good numbers when especifically searched in its habitat, natural or almost natural forest areas with base-rich substrates and sufficient humidity, and therefore it is a good indicator species of preservation of particular forest communities (Sprynar & Kment 2005). As to the Iberian Peninsula, it has been very scarcely recorded from some northern regions: Catalonia, Fogars de Montclús (Barcelona), J. Ribes (1990); Basque Country, Hernani (Gipuzkoa), Pagola-Carte et al. (2003), and Turtzioz (Bizkaia), Pagola-Carte & J. Ribes (2007). Habitat and biology. Metatropis rufescens mainly lives on Circaea spp., a glandularhairy Onagraceae. The three known Iberian populations were found on that plant. The berytid host plant can be found in humid environments receiving low light levels, and
On some threatened Heteroptera from the Iberian fauna 149
this habitat is usually shared with a peculiar hemipteran fauna, such as several species of Miridae Bryocorinae (Péricart 1984; J. Ribes 1990). Adults overwinter under moss or barks, always under conditions of high humidity, and reappear in spring. Oviposition takes place at the beginning of summer. Adults of the new generation live at the end of summer and in autumn. Threatening factors. Metatropis rufescens is highly dependent on the availability of its host plant and habitat. The species is threatened by the reduction or loss of humid environments. Iberian populations, the southernmost distribution localities, are rather isolated and probably with a reduced genetic flow. Among the threats to the habitat, it has to be emphasized some forestry activities involving aggressive methods of timber extraction as well as plantation of dense stands of coniferous trees, which prevent any herbaceous undergrowth; and the cleaning of road and trail edges, where C. lutetiana frequently thrives. Conservation measures. The conservation of M. rufescens relies on the protection of its habitat. A proper forest management is needed to protect the herbaceous level of the forest undergrowth, including the edges of trails and roads, particularly where C. lutetiana grows. Indiscriminate clearings should be avoided. According to experience in the Czech Republic (Petr Kment, pers. comm.), recreational activities (fishing, hunting, walking, etc.) may be allowed whenever performed in an extensive and disperse way. M. rufescens is included in the Red List of Czech Republic, under the category “near threatened” (Kment & Vilímová 2005).
Oxycarenidae Auchenodes costalis (LETHIERRY, 1877) Distribution area. Auchenodes costalis has a disjoint distribution. Its most western border is the Iberian Peninsula, and its most eastern one is the Balkan Peninsula (Péricart 2001a). The Spanish records are: Badajoz, Uhagón leg., col. MNCN compiled in Péricart (1999a); Malagón (Ciudad Real), De la Fuente leg., col. MNCN, compiled in Péricart (1999a); Pozuelo de Calatrava (Ciudad Real), Sienkiewicz (1964) compiled by Péricart (1999a); Alpedrete (Madrid), 18-VI-1991, M. Á. Vázquez leg., col. Universidad Complutense de Madrid, 1 specimen on Poaceae, compiled in Péricart (1999a); Santa Helena ( Jaén), IV-1925, Lindberg. leg., compiled in Péricart (1999a). Most part of all those records are very old. The species is included in the Regional Catalogue of threatened species of wild fauna and flora from the Community of Madrid (Catálogo in press). Habitat and biology. Auchenodes costalis is found in mountain rocky steppes, as well as in sand dunes on the sea coast, and it is reputed as a xerophilous species. It lives on Matricaria (Asteraceae) and on lichens. Adults have been captured from April to September, and nymphs have been recorded in June (Péricart 1999a). The capture of a male in Alpedrete took place on Poaceae.
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Threatening factors. Auchenodes costalis is threatened by the factors that affect their habitat and its host plants. In addition, it is not a very abundant species, with scarce and scattered populations. Burning of stubbles reduce the habitat of the species and makes difficult the continuity of its life cycle. Also the elimination of plants from trial edges reduces the mobility of the species. In wooded lands, clearings of the underwood as prevention of fires may reduce the density and the richness of plant species that host A. costalis. Finally, the use of non selective pesticides in agriculture, can directly reduce the populations of A. costalis when its habitat is next to agricultural zones. Conservation measures. The conservation of A. costalis is linked to the conservation of its habitat. For example it is necessary to control stubble burning, since the scrubs and the edges are the refuge of many species of Heteroptera and other insects. Also, in road improvement and clearing of underbrush, the conservation of edge vegetation apt to A. costalis and to many other equally interesting species attached to that habitat has to be recommended. Also the use of selective pesticides in extensive agriculture has to be implemented.
Leptodemus minutus ( JAKOVLEV, 1874) Distribution area. Leptodemus minutus is largely distributed in the southernmost area of the Palaearctic region, extending to Sudan (Péricart 2001a). It has been mentioned from a scarce number of European localities. In Spain, it is known by a short number of specimens attracted to light in Córdoba and Baena (Baena in litteris; Péricart 1999a). Habitat and ecology. Péricart (2001a) states that L. minutus lives on sandy soils, in desert climates. It may also be collected at 4000 m (Atlas range, Morocco). As host plants, Artemisia inodora, Plantago indica and Aristida scoparia hav been recorded. In Spain, the species was collected in light traps, so that its habitat is unknown. It is speculated that maybe it frequents dry and sunny places, where grasses are abundant, as wasted lands, field margins, road and trail borders, among others. Threatening factors. Leptodemus minutus may be threatened by the effect of several activities which may affect its habitat and host plants. Moreover, it is a scarce species, and their populations are rare and scattered. The threat against the species is increased by the fact that its habitat is considered not to be of special ecological interest, or menaced. However, this habitat may be disturbed by different human activities related to agriculture development (burn of stubbles, use of pesticides, agricultural intensification in general), which cut the insect life cycle in time and space. The clearing of undergrowth to prevent fires, or the removing of weeds for the conservation of roads may also negatively affect L. minutus habitat and populations both directly and indirectly. In the last times, urbanization of wasted lands near villages is an increasing threat to this species. Conservation measures. The conservation of L. minutus is largely interferred by activities that very often are considered desirable and necessary. In agriculture, the use of selective pesticides and the conservation of stream banks and field margins could help
On some threatened Heteroptera from the Iberian fauna 151
to maintain L. minutus living conditions. In general, the reserve of part of L. minutus habitat near woods, villages or cultivated areas, keeping it undisturbed, may contribute to the survival of the species.
Rhyparochromidae Trichaphanus fuentei (PUTON, 1894) Distribution area. Trichaphanus fuentei shows a disjoint distribution, being the Iberian Peninsula the most western end and Iran the most eastern end (Péricart 2001a). The species is included in the Regional Catalogue of threatened species of wild fauna and flora from the Community of Madrid (Catálogo in press). In Spain, it has been recorded from Laroles to the Puerto de la Ragua (Almería), V-1981, Remane leg. compiled in Péricart (1999b); Pozuelo de Calatrava (Ciudad Real), male lectotype, Péricart (1999b); Fuencaliente (Ciudad Real), Péricart (1999b); Castril (Granada), Péricart (1999b); Valdemoro (Madrid), 31-III-1974, S. Pérez Minocci leg., 1 ex. (under bark) and 31-III-1974, R. Outerelo leg. (under stone), Péricart (1999b); Sierra de las Nieves, Ronda (Málaga), Péricart (1999b). Most of those are old records. Habitat and biology. Trichaphanus fuentei has been captured under stones, and under barks and shrubs of different plant species, in not very high woods. T. fuentei is a large-sized, always macropterous species. Some of its biological characteristics, as its egg stage, are unknown. Threatening factors. Trichaphanus fuentei is threatened by the effects of diverse activities that negatively affect to its habitat. Most important threats for bark niche are burning of stubbles, clearing of underbrush to prevent fires, or of edge vegetation along trails and roads to maintain them. The type and degree of threat on the lapidicolous habitat depends on its location. In the vicinity of towns, the space may be wanted for industrial or real estate urbanization. In rural zones, agricultural use of land, including spraying of non selective pesticides, may interfere with the living conditions of T. fuentei. Conservation measures. The conservation of T. fuentei is closely tied to the conservation of its habitat. It is necessary to control the burning of stubbles and the clearing of the underbrush and road edges, to keep some areas with untouched vegetation where T. fuentei may thrive. The implementation of selective pesticides has also to be recommended. The excessive growth of urban nuclei needs to be regulated.
Rhopalidae Leptoceraea femoralis (HORVÁTH, 1897) Distribution area. Leptoceraea femoralis is a Irano-Mediterranean species (Dolling 2006) with a short number of records. Most of the Spanish records are quite old: Pozuelo de Calatrava (Ciudad Real), De la Fuente (1899), specimens out of location; Almagro
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(Ciudad Real), De la Fuente (1920), specimens out of location; Cartagena (Murcia), a couple, col. MNCN, Vázquez (1985); Conil (Cádiz), col. M. Baena, Moulet (1995). Habitat and biology. This species has been associated to Aeluropus littoralis; this Poaceae is a halophylous hemicryptophyte that needs humidity and grows only in natural habitats. De la Fuente (1920) points out that this species is associated to wetlands. Other data of their biology are not known. Threatening factors. Leptoceraea femoralis can be threatened in many ways, by alteration of its habitat. The banks of streams, ponds and brooks it inhabits are under several threatening factors. Grazing and frequentation by livestock can contribute to the deterioration of the habitat. On the other hand, the maintenance of the phreatic level and environmental humidity, and of water quality, can be seen highly harmed by the abusive extraction or inadequate uses of water. In addition, the generally flattened topography of the humid zones makes them susceptible to drying and urbanization, depending on its geographic situation next to inhabited nuclei. Finally, the attraction of natural humid zones for ecological tourism can put them under the risk of an excessive, undesirable human impact, either direct or indirect. Conservation measures. The conservation of L. femoralis is closely tied to the conservation of its habitat. It is necessary not to alter the phreatic levels, by regulating the use of water, so that these zones stay in their optimal levels. Also an excessive pressure by livestock or by agricultural uses must be avoided as much as possible. The human frequentation due to leisure has also to be controlled. 3, 5, 10 10 4 8 8, 10
7 4
1, 11, 13
11
4, 11, 13, 14 12
2, 9, 14
13
11 6, 13
14 13
Fig. 1: Distribution in Spain of the fourteen species considered in the article. Lines correspond to province limits. Numbers in the provinces indicate the species that have been collected in each one, according to the following code: 1, Pachycoleus pusillimus; 2, Rhagovelia nigricans nigricans; 3, Acalypta suturalis; 4, Isometopus intrusus; 5, Bryocoris pteridis; 6, Ribautocapsus bruckii; 7, Roudairea crassicornis; 8, Anthocoris visci; 9, Rhynocoris lineaticornis; 10, Metatropis rufescens; 11, Auchenodes costalis; 12, Leptodemus minutus; 13, Trichaphanus fuentei; 14, Leptoceraea femoralis.
On some threatened Heteroptera from the Iberian fauna 153
DISCUSSION The fourteen species considered present different characteristics in regard to their habitats, to their threats and to the specific recommended measures of protection. Most of them are threatened by human activities. However, the conservation of those species may be achieved by the protection of their respective habitats. Several of those species are linked to rivers and water streams, as they are aquatic or live on the vegetation or in the soil interstices in the banks. Main threats come from misuse or overuse of water, water pollution and bank dirt, and from canalization which implies alteration of stream bottom and banks. In this case, conservation of the habitat means to keep an appropriate flow and quality of water, and to preserve structure and vegetation of banks. Another group of species live in mountain or forest habitats. Main threat is human frequentation facilitated by the opening of mountain roads. Main reasons of frequentation may be either leisure, timber industry works, or wood management. When the habitat is placed in a dry area, clearing of the underwood for fire prevention negatively affect to the habitat conservation. On the other hand, the fire itself may be a menace. In all those cases, the reservation of undisturbed patches of land could be a good measure to assure the survival of the species. A certain number of species are linked to very specific habitats, as is A. visci and the mistletoe, or I. intrusus and old trees. Others, on the contrary, live in habitats considered not emblematic at all, as are ruderal or herbaceous dry habitats. Main threats are some human activities considered necessary or even desirable, as fire prevention, agricultural management or road preservation are. Lack of appreciation for old, parasitized woods, or for abandoned lands, does not help to the preservation of those habitats. Again, the preservation of undisturbed patches of land, together with the promotion of respect for those apparently not interesting habitats, could be key measures for the survival of the insect. Summarizing, the protection of threatened species and their habitat involves several actions. On the one hand, environmental education and information of the public is most important, as a way to promote respect behaviour in the wild, which helps to its preservation. On the other, administration is the only statement, at local, regional or national level, that may promote legislation to protect habitats and species, making compatible their survival with human interests. Private attitudes and public actions are the two faces of the coin, and the success in reducing threats to the species is built on the harmony of both.
ACKNOWLEDGEMENTS We are very grateful to our colleague Juan Antonio Arce for his always opportune commentaries and suggestions.
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РЕЗЮМЕ През 1998 г. на авторите на тази статия бе възложено от Испанското министерство на околната среда (в рамките на Националната Инвентаризация на Биоразнообразието) да изберат застрашени видове от разред Heteroptera от Испания, с цел да бъде публикуван «Червен списък» на безгрьбначните животни. Бяха предложени двадесет и три вида. Девет от тях от категорията «Уязвими», съгласно критериите на UICN, са вече включени в «Червения списък» на безгрьбначните животни. Останалите видове по различни причини също заслужават да бъдат разглеждани като застрашени. В тази статия има кратко описание на всеки от тези 14 вида, което включва кратко описание на тяхната биология, разпространение, и факторите, които ги застрашават. Включените видове са: Pachycoleus pusillimus (SAHLBERG, 1870) (Dipsocoridae), Rhagovelia (Rhagovelia) nigricans nigricans (BURMEISTER, 1835) (Veliidae), Acalypta suturalis (PUTON, 1879) (Tingidae), Isometopus intrusus (HERRICH-SCHAEFFER, 1835), Bryocoris pteridis (FALLÉN, 1807), Ribautocapsus bruckii (REUTER, 1879), Roudairea crassicornis (PUTON & REUTER, 1886) (Miridae), Anthocoris visci DOUGLAS, 1889 (Anthocoridae), Rhynocoris (Rhynocoris) lineaticornis (REUTER, 1895) (Reduviidae), Metatropis rufescens (HERRICH-SCHAEFFER, 1835) (Berytidae), Leptodemus minutus (JAKOVLEV, 1874), Auchenodes costalis (LETHIERRY, 1877) (Oxycarenidae), Trichaphanus fuentei (PUTON, 1894) (Rhyparochromidae) и Leptoceraea femoralis (HORVÁTH, 1897) (Rhopalidae). Всички тези видове са застрашени от човешката дейност, и оцеляването им изисква защита на техните хабитати. Опазването на тези видове изисква задължителна подкрепа чрез законодателни мерки, природозащитно образование и информиране на общественността, за да имат успех мероприятията по опазване и консервация на застрашените видове.
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S. Grozeva & N. Simov (Eds) 2008 Pentatomidae, Pentatominae) 159 Review of Parachinavia Roche (Hemiptera, ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 159-169. © Pensoft Publishers Sofia–Moscow
Review of Parachinavia ROCHE (Hemiptera, Pentatomidae, Pentatominae) * J. Grazia1 & C.F. Schwertner2 1
Universidade Federal do Rio Grande do Sul (UFRGS), Instituto de Biociências, Departamento de Zoologia, Av. Bento Gonçalves 9500, Prédio 43435, 91501-970 Porto Alegre, RS, Brazil. E-mail:
[email protected] 2 Universidade Federal do Rio Grande do Sul (UFRGS), Instituto de Biociências, Departamento de Zoologia, Av. Bento Gonçalves 9500, Prédio 43435, 91501-970 Porto Alegre, RS, Brazil. E-mail:
[email protected]
ABSTRACT Parachinavia Roche, monotypic until the present work, is reviewed and broadened with the inclusion of Acrosternum prunasis (Dallas). The genus is redescribed, including the genitalia of both sexes. Among green stink bugs, Parachinavia is distinguished by oval body, small to medium size, color pale green to ochraceous, punctures concolorous and regularly distributed, ocelli on a pale callosity, gonocoxites 8 conspicuously inflated, gonocoxites 9 with anterior margin concave and posterior margin slightly convex, and secondary thickenings of gonapophyses 9 in 1+1 small rounded areas. Diagnostic characters for the species, as well as illustrations for several morphological structures, are provided. Keywords: Afrotropical Region, Afrotemperate Region, Heteroptera, Parachinavia creola, Parachinavia prunasis, Nezarini.
INTRODUCTION The genus Parachinavia was described by Roche (1977) in a review of Pentatomidae of the granitic island of Seychelles. The author mentioned that three male specimens collected *
Contribution N° 529 of the Department of Zoology, UFRGS.
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on Mahé do ‘… not fit into any Acrosternum group of genera…’ (Roche 1977, p. 564), and proposed the new genus and new species, Parachinavia creolea Roche. In a tentative tribal classification of Pentatominae, Rider (2005) included Parachinavia in the tribe Nezarini. Schwertner (2005), studying the phylogenetic relationships of a group of green stink bug genera, established Parachinavia as the sister-group of genus Acrosternum, the two genera sharing at least three synapomophies. Our study of type-material of both genera had allowed the recognition of Acrosternum prunasis (Dallas, 1851), a continental African species, as congeneric to P. creolea. A new combination, Parachinavia prunasis nov. comb., is proposed here in. Gerlach et al. (2005) extended the distribution of P. creolea to Silhoutte island, this species being endemic to Seychelles. Therefore, the new combination here proposed extends the geographical distribution of Parachinavia into the continental area of Afrotropical region and to the Afrotemperate region (Morrone 2002). In this work, Parachinavia is redescribed and, for the first time, a detailed description of the genitalia of both sexes is presented. Diagnostic characteristics of the species are given. We dedicate this paper to Michail Josifov, in recognition of his contribution to the study of the Heteroptera.
MATERIAL AND METHODS Material was examined from the following collections (acronyms and curators in parentheses): American Museum of Natural History, New York, USA (AMNH, R.T. Schuh); Museé Royal d’Afrique Centrale, Tervuren, Belgium (MRAC, U. Dall’Asta); Museum Nationale d’Histoire Naturelle, Paris, France (MNHN, D. Pluot-Sigwalt); National Museum of Natural History, Washington D. C., USA (NMNH, T.J. Henry); The Natural History Museum, London, UK (BMNH, M. Webb,). Dissection of the internal genitalia followed Grazia et al. (2006); terminology for genitalia followed mainly Dupuis (1970) for males and females, and Baker (1937) and Schaefer (1977) for males. Drawings were made with camera lucida mounted on a Leica MZ125 stereomicroscope; photographs were taken with a Microptics-USA photographic system.
RESULTS AND DISCUSSION Parachinavia ROCHE Parachinavia Roche, 1977:564 (nov. gen.). Type species: Parachinavia creolea Roche
Diagnosis. Recognized among green stink bugs by oval body, small to medium size, color pale green to ochraceous in preserved specimens, punctures concolorous and regularly
Review of Parachinavia Roche (Hemiptera, Pentatomidae, Pentatominae) 161
distributed, ocelli on a pale callosity but body without calloused areas; unique apomorphies among the group include the following characters of the female genitalia: gonocoxites 8 conspicuously inflated, posterior margin of gonocoxites 9 slightly convex, anterior margin concave; secondary thickenings of gonapophyses 9 in 1+1 small caps. Distinguished from Acrosternum by several morphological characters (see comments below). Description. General color pale green to ochraceous in preserved specimens; apical half of antennal segment III and most of segments IV and V reddish. Dorsal and ventral surfaces with concolorous punctures regularly distributed, coarse; callosity areas absent. Broad, oval, of medium size (9 to 12 mm). Head triangular, juga as long as tylus. Ocelli on a pale callosity; lateral margins of juga slightly concave. Antennifer tubercle not seen from dorsal side; lateral spine of antennifer tubercle obsolete, ventral spine absent. Antennae five segmented; first segment not surpassing apex of head, antennal segment increasing in length from I to V; segment IV and V subequal. Bucculae low, uniform in height, margins subparallel and evanescent near base of head. Rostrum just attaining or surpassing metacoxae; segment I not surpassing bucculae; segment II shorter than III and IV combined. Pronotum trapezoidal, humeral angles not developed; antero-lateral margins rectilinear. Scutellum almost as broad as long at base, apex surpassing level of fourth connexival segment; parafrenal lobe about two and a half size of postfrenal lobe. Hemelytra: corium surpassing anterior half of connexival segment VI; membrane hyaline, sometimes with green speckles. Connexivum more or less exposed, postero-lateral angles slightly pronounced, tiny black spot on posterior angle of urosternite present. Mesosternum conspicuously carinate; metasternum longitudinally hexagonal, longer than wide, feebly sulcate. Ostiole of metapleural scent gland elliptical, developed in a long, narrow ruga, evanescent at apex, extending to the ¾ of the metapleura; mesopleural evaporatorium wide, occupying almost half of sclerite; metapleural evaporatorium also wide, occupying most of sclerite. Legs immaculate, tibia dorsally sulcate. Abdominal spine short, blunt, scarcely attaining metacoxae. Spiracles concolorous; lateral trichobothria placed along spiracular line. Abdominal midline with scattered concolorous punctures. Male genitalia (Figs 4-19). Pygophore quadrangular, postero-lateral angles not developed, round. Genital cup opening dorso-posterior. Superior process (sp) of genital cup present. Dorsal rim (dr) sinuous at middle, scarcely projected over segment X; lateral thirds of dorsal rim with 1+1 spines (sdr). Ventral rim (vr) of pygophore shallowly concave at middle (me); infolding of ventral rim (ivr) wide on dorsal view, simple, inflated laterad to inferior ridge. Inferior ridge (ir) not visible in ventral view. Segment X perpendicular in relation to the sagittal plane of pygophore, quadrangular in outline; dorsal surface uneven at basal half, apical half without processes. Parameres (pa) uniramous, laterally flat; apical part of parameres (ap) more or less wide and latero-basally directed; hairy process at base of paramere (bp) present. Phallotheca (ph) tubular, more than four times as long as vesica (v). Vesica short, straight, with 1+1 processes (pv); processes subequal in length to vesica and obscured by conjunctiva (cj) when not extended.
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Female genitalia (Figs 20-21). Posterior margin of segment VIII in open U along gonocoxites 8 (gc8). Laterotegites 8 (la8) as long as laterotergites 9 (la9); spiracles (s) present on basal angle of laterotergites 8. Gonocoxites 8 conspicuously inflated, posterior margin slightly convex; sutural margins convex, juxtaposed only at middle. Surface of basal 2/3 of laterotergites 9 concave. Posterior margin of gonapocoxites 9 (gc9) slightly concave, anterior margin concave, prolonged anteriorly in 1+1 short arms; secondary thickenings of gonapophyses 9 (sg9) in 1+1 small caps on anterior margin; chitinellipsen (ch) present. Ductus receptaculi (dre) before vesicular area (va) almost as long as ductus receptaculi after vesicular area; in this portion, reeled and strongly inflated before anterior flange (af ). Internal ductus of vesicular area reeled at basal fourth. Capsula seminalis (cs) as long as pars intermedialis (pi), assymetrical and with one projection anteriorly bent. Comments. Parachinavia was described for P. creolea, in comparison with Acrosternum heegeri, based on three male specimens (Roche 1977). Among green stink bugs, Acrosternum and Parachinavia represent a monophyletic group, sharing the following synapomorphies (Schwertner 2005): antennifer tubercle obsolete, not visible in dorsal view; mesosternum conspicuously carinate; ductus receptaculi after vesicular area reeled and strongly inflated before anterior flange; capsula seminalis asymmetrical and with one projection anteriorly bent. Parachinavia can be readily distinguished from Acrosternum by punctures apparently coarser and less dense, scutellum almost as broad as long at base, postero-lateral angles of pygophore not developed, superior process of genital cup conspicuous, dorsal rim only slightly projecting over segment X, lateral third of dorsal rim with 1+1 spines, ventral rim of pygophore shallowly concave at middle; infolding of ventral rim simple, slightly inflated laterad to inferior ridge, and wide in dorsal view; inferior ridge not visible in ventral view; segment X with dorsal surface uneven at basal half, apical half of segment X without processes, internal ductus of vesicular area reeled at basal fourth, and capsula seminalis as long as pars intermedialis. Distribution. Widely distributed in the Afrotropical and Afrotemperate regions, including the west-Pacific islands of Seychelles.
Parachinavia creolea ROCHE (Figs 1-11, 22) Parachinavia creolea Roche, 1977:565 (nov. sp.); Gerlach et al. 2005:107 (distribution). Type-locality: Seychelles.
Male. Rostrum just attaining metacoxae (Fig. 3). Measurements (mm±standard deviation): Total length 9.26±0.29, total width 5.20±0.38, head length 1.93±0.11, head width 2.58±0.11, interocular distance 1.48±0.04, interocelar distance 0.88±0.04, head length in front of eyes 0.93±0.04, antennal segment I 0.58±0.02, antennal segment II 0.98±0.07, antennal segment III 1.28±0.02, antennal segment IV 1.43±0.09, antennal segment V 1.42±0.07, rostral segment I 0.73, rostral segment II 1.33, rostral segment III
Review of Parachinavia Roche (Hemiptera, Pentatomidae, Pentatominae) 163
0.93, rostral segment IV 0.97, pronotum length 1.67±0.28, pronotum width 5.47±0.38, scutellum length 3.33±0.00, scutellum width 3.33±0.19. Pygophore: excavation of ventral rim occupying more than median third of this structure (Figs 4, 6). Parameres: apical portion almost as wide as basal portion; apical margin obtusely rounded; apical process inconspicuous (Fig. 7-8). Female. Unknown. Comments. Roche (1977) mentioned that this species was apparently rare, and until recently it was known only from type material. More recently, Gerlach et al. (2005) extended its distribution. Distinguished from P. prunasis nov. comb. by shorter rostrum, ampliate excavation of ventral rim of pygophore, and narrower parameres, obtusely rounded at apex.
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Figs 1-6: Parachinavia creolea, holotype ♂. 1-3, Habitus in dorsal, ventral and lateral views; 4-6, pygophore in dorsal, posterior and lateral views (dissected and partially damaged).
164 J. Grazia & C.F. Schwertner bp
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Figs 7-11: Parachinavia creolea, holotype ♂. 7-8, Left paremere in latero-medial and latero-external views; 9-11, Phallus in dorsal, ventral and lateral views (aa- articulatory apparatus; ap- apical portion of paramere; bp-basal portion of paramere; cj- conjunctiva; dsd- ductus seminis distalis; dsp- ductus seminis proximalis; er- ejaculatory reservoir; pc- processus capitati; ph- phallotheca; v- vesica).
Holotype ♂ with the following labels (pygophore dissected): [Seychelles, Mt Fleuri, Mahe, 19.III.1972, P.J.L. Roche//Holotype//7178//Parachinavia creolea gen. & nov. sp., P.J.L. Roche det. 1975] (BMNH). Paratype ♂ with the following labels (pygophore dissected): [Seychelles: Mahe, Mt Fleuri, 10.I.1972, P.J.L. Roche //Paratype//6712//Parachinavia creolea gen. & nov. sp., P.J.L. Roche det. 1975] (MRAC). Distribution (Fig. 22). Seychelles, known from Mahé (Mt. Fleuri) and Silhoutte islands. Parachinavia prunasis (DALLAS) nov. comb. (Figs 12-22) Raphigaster prunasis Dallas, 1851: 279 (nov. sp.). Nezara prunasis: Distant, 1900:392 (list); Hesse, 1926: 36 (distribution). Nezara klugi Schouteden, 1909:59 (taxonomy and distribution). nov. syn. Nezara (Acrosternum?) prunasis: Kirkaldy, 1909: 121 (taxonomy and distribution). Nezara conspersa Schouteden, 1909:59 (syn. by Linnavuori, 1972). Acrosternum prunasis prunasis: Linnavuori, 1972:429 (taxonomy and distribution); 1975:99 (distribution), 1982a:25 (distribution). Acrosternum prunasis klugi: Linnavuori, 1972:429 (taxonomy and distribution); 1975:99 (distribution). nov. syn. Acrosternum (Acrosternum) prunasis: Linnavuori, 1982b:142 (taxonomy and distribution). Type-locality: Congo
Review of Parachinavia Roche (Hemiptera, Pentatomidae, Pentatominae) 165
Male. Rostrum surpassing metacoxae. Measurements (mm±standard deviation): Total length 9.62±0.33, total width 5.66±0.38, head length 1.86±0.06, head width 2.63±0.08, interocular distance 1.61±0.05, interocellar distance 1.02±0.02, head length in front of eyes 0.86±0.04, antennal segment I 0.46 ±0.10, antennal segment II 0.99±0.09, antennal segment III 1.23±0.11, antennal segment IV 1.41+0.13, antennal segment V 1.33±0.04, rostral segment I 0.78±0.07, rostral segment II 1.34±0.12, rostral segment III 1.17±0.14, rostral segment IV 0.96±0.11, pronotum length 1.74±0.10, pronotum width 5.82±0.26, scutellum length 3.60±0.22, scutellum width 3.60±0.17. Pygophore: median excavation of ventral rim occupying less than median third of this structure (Figs 12, 14). Parameres: apical portion wider than basal portion; apical margin somewhat truncate, apical process conspicuous (Figs 15-16). Female. Rostrum same as male. Measurements (mm±standard deviation): Total lenght 10.90±0.82, total width 6.27±0.45, head length 2.01±0.11, head width 2.80±0.17, interocular distance 1.76±0.11, interocellar distance 1.15±0.07, head length in front of eyes 0.92±0.06, antennal segment I 0.50±0.09, antennal segment II 1.02±0.15, antennal segment III 1.24±0.25, antennal segment IV 1.39±0.05, antennal segment V 1.35, rostral X
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Figs 12-19: Parachinavia prunasis nov.comb. 12-14, Pygophore in dorsal, lateral and ventral views; 15-16, Left paramere in latero-medial and latero-external views; 17-19, Phallus, with vesica expanded, in dorsal, ventral and lateral views; articulatory apparatus omitted (cj- conjunctiva; drdorsal rim; er- ejaculatory reservoir; ir – inferior ridge; ivr- infolding of ventral rim; me- median excavation of ventral rim; pa- paramere; ph- phallotheca; pv- processus vesicae; sdr- spine of dorsal rim; ps- superior processes of dorsal rim; v- vesica; vr- ventral rim; X- segment X).
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segment I 0.82±0.11, rostral segment II 1.36±0.10, rostral segment III 1.14±0.09, rostral segment IV 0.95±0.06, pronotum length 1.89±0.12, pronotum width 6.38±0.42, scutellum length 4.05±0.38, scutellum width 4.02±0.28. Genitalia as described for the genus. Comments. Dallas (1851) described this species in Raphigaster, subgenus Nezara, based on a female collected from Congo. Stål (1876) included R. prunasis in his ‘Inserti Pentatomidarum Loci Systematici’ list, while Kirkaldy (1909) expressed doubt as to its position within Nezara. Distant (1900) and Hesse (1926) listed the species in Nezara, and Linnavuori (1972, 1975, 1982a, 1982b) included it in Acrosternum. Schouteden (1909) described Nezara conspersa and Nezara klugi, and compared them to Nezara miliaris Klug. Linnavuori (1972) included all these species in Acrosternum, established N. conspersa as a junior synonymy of A. prunasis, and considered A. klugi as a subspecies of A. prunasis; he also compared A. prunasis with A. millierei (Mulsant & Rey) and, according to him, the two differ mainly in the structure of male genitalia. In fact, this difference among A. prunasis and Acrosternum spp. corresponds to the diagnostic character proposed by Roche (1977) to establish Parachinavia. Linnavuori (1972) proposition of A. prunasis klugi was based just on head length; the variation observed in the specimens studied did not support this hypotheses. Therefore, A. prunasis klugi is synonymized with P. prunasis comb.nov. dre
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21 Figs 20-21: Parachinavia prunasis nov. comb. 20- genital plates; 21- Ectodermal ducts and laterotergites, gonocoxites an gonapophyses of ninth segment (af- anterior flange; atv- anterior thickenings of vaginal intima; ch- chitinellipsen; cs- capsula seminalis; dre- ductus recepatuculi; g9- gonapophyses 9; gc8- gonocoxites 8; gc9- gonocoxites 9; la8- laterotergites 8; la9- laterotergites 9; pf- posterior flange; pi- pars intermedialis; s- spiracles; tg9- thickenings of gonapophyses 9; va- vesicular area; VII- segment VII; X- segment X).
Review of Parachinavia Roche (Hemiptera, Pentatomidae, Pentatominae) 167
Parachinavia prunasis can be distinguished from P. creolea by the longer rostrum, the narrower excavation of ventral rim of pygophore and the somewhat wider parameres, truncate at apex. Holotype: ♀ [a. Congo] (BMNH). Other specimens examined: 1♀ [Kordofan, Tendelti, UmmRuwaba, Sudan, 25.I.1963, Linnavuori] (AMNH); 1♂ [Sudan, Equatoria, Tambura-Wau road, 25-26. IV.1963, Linnavuori] (AMNH); 1♂ [Legon, A.D., Ghana, 15.XI.68, D. Leston//U.V.] (AMNH); 1♂ [Legon, Ghana, 20.XI.1968, D. Leston//U.V. trap] (AMNH); 1♀ [Tron, Ghana, 14.X.1967, D. Leston] (AMNH); 1♂ [Olokemepi, Ibadan, Nigeria//Bridwell collection //Acrosternum (A.) prunasis, Rider det. 1989] (NMNH); 1♂ [Botswana: Xugana, Okavango swamp, 19°05’S 23°06’E, 3200ft, 1-12 December, 1975, W. Carter//collected at Black light] (NMNH); 1♀ [Pretoria, Transvaal Meientpes Kop., 25.Nov.1967, JA & S Slater, T.Schuh] (AMNH); 1♂ [Rostenburg, U. SO. Afr., XII.1961, A.L. Capener] (AMNH); 1♀ [Umtentweni, Natal, VII.1953, A.L. Capener] (AMNH); 1♀ [Museum Paris, Transvaal, HammanSkraal, E. Simar, Coll. Noualhier 1898] (MNHN); 1♀ [Uitenhage, Dunbroby, Rev. O’Neil//prunasis Dall.//N°8//Distant Coll. 1911-383] (BMNH); 1♀ [Museum Paris, Cape Town, E. Simon, coll. Noualhier 1898] (MNHN); Distribution (Fig. 22). Widely distributed in the continental area of Afrotropical region (Sudan, Ghana, Ivory Coast, Nigeria, Somalia, Congo, Botswana, Mozambique) and Afrotemperate region (South Africa).
P. creolea P. prunasis (material examined in this work) P. prunasis (Linnavuori 1972, 1975) P. prunasis (Linnavuori 1982a) P. prunasis (Linnavuori 1982b) P. prunasis (Schouteden 1909)
Figs 22: Distribution of Parachinavia spp.
168 J. Grazia & C.F. Schwertner
ACKNOWLEDGEMENTS To the curators of the listed collections, especially M. Webb (BMNH) and U. Dall’Asta (MRAC) for the loan of type-material. Our sincere thanks to Randall T. Schuh, for assistance and facilities during the stay of CFS at AMNH, where this study was partially carried out. This work was supported by grants of Conselho Nacional de Pesquisa (CNPq) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) to the authors.
РЕЗЮМЕ Moнотипичният досега род Parachinavia ROCHE е ревизиран и в него е включен Acrosternum prunasis (DALLAS). Родът е преописан, включени са данни за гениталиите на двата пола. От останалите пентатомиди със зелена окраска, Parachinavia се отличават с овално тяло, с малки или средни размери, с бледо зелено до охрено оцветяване, пунктировката едноцветна и равномерна, оцелите на светли мазолести издатини, 8 гонококсити значително издути, 9 гонококсити с преден влъбнат и заден изпъкнал ръб, и вторично одебеляване на на гонококситите 9 в 1+1 малки кръгли простанства. Представени са диагностичните белези за видовете; някои морфологични структури са илюстрирани.
REFERENCES Baker A.D. (1937). A study of the male genitalia of Canadian species of Pentatomidae. — Canadian Journal of Research, 4: 148–220. Dallas W.S. (1851). List of the specimens of hemipterous insects in the collection of the Brithsh Museum. — Trustees of the British Museum, London: 1–368+pls. 1–11. Distant W.L. (1900). Rhynchotal notes IV. Heteroptera: Pentatominae (part). — Annals and Magazine of Natural History, (7) 5:420–435. Dupuis C. (1970). Heteroptera. — In: Tuxen S.L. (ed.): Taxonomist’s glossary of genitalia in insects, 2nd ed. Munksgaard, Copenhagen: 190–209. Gerlach J., M. Madl & P. Matyot (2005). Review of the Pentatomidae of Seychelles Hemiptera, Heteroptera). — Phelsuma, 13: 104–108. Grazia J., C.F. Schwertner & A. Ferrari (2006). Description of five new species of Chinavia Orian (Hemiptera, Pentatomidae, Pentatominae) from western and northwestern South America. — In: Rabitsch W (Ed.): Denisia 19. Biologiezentrum der Oberösterreichischen Landesmuseen, Linz: 423–434. Hesse A.J. (1926). A list of the Heteropterous and Homopterous Hemiptera of South West Africa. — Annals of the South African Museum, 23: 1–177+2 plates. Kirkaldy G.W. (1909). Catalogue of the Hemiptera (Heteroptera). 1- Cimicidae. — Felix L. Dames, Berlin: xxxix+1–392.
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Linnavouri R. (1972) Studies on African Pentatomidae. — Arquivos do Museu Bocage, 3: 415–434. Linnavuori R. (1975). Hemiptera of the Sudan, with remarks on some species of the adjacent countries 5. Pentatomidae. — Boletim da Sociedade Portuguesa de Ciências Naturais, 15: 5–128. Linnavuori R. (1982a). Some hemipterous fauna from Somalia and Ethiopia. — Monitore zoologico italiano, 16: 1-33. Linnavuori R. (1982b) Pentatomidae and Acanthosomidae of Nigeria and the Ivory Coast, with remarks on species of the adjacent countries in West and Central Africa. — Acta Zoologica Fennica, 163: 1–176. Morrone J.J. (2002). Biogeographical regions under track and cladistic scrutiny. — Journal of Biogeogaphy, 29: 149–152. Rider D. (2005). Pentatomodeia Home page. North Dakota State University, Fargo. Available at http://www.ndsu.nodak.edu/ndsu/rider /Pentatomoidea/. Acess in: 01.IX.2005. Roche P.J.L. (1977). Pentatomidae of the granitic islands of Seychelles (Heteroptera). — Revue Zoologique africaine, 91: 558–572. Schaefer C.W. (1977). Genital capsule of the trichophoran male (Hemiptera: Heteroptera: Geocorisae). — International Journal of Insect Morphology & Embryology, 6: 277–301. Schouteden H. (1909). Catalogues raisonnés de la Faune entomologique du Congo Belge. Hémiptèrés, fam. Pentatomidae. — Annales du Museé du Congo Belge. Zoologie, Serie 2c (3), 1: 1–85. Schwertner C.F. (2005). Filogenia e classificação dos percevejos-verdes do grupo Nezara Amyot & Serville, 1843 (Hemiptera, Pentatomidae, Pentatominae). — DSc Thesis, Universidade Federal do Rio Grande do Sul, Porto Alegre: xv+1–246. Stål C. (1876). Enumeratio Hemipterorum V. — Kongliga Svenska Vetenskaps-Akademiens Handlingar, 14: 1–162.
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S. Grozeva & N. Simov (Eds) 2008 species (Heteroptera, Miridae) 171 Cytotaxonomy of two Cremnocephalus ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 171-179. © Pensoft Publishers Sofia–Moscow
Cytotaxonomy of two Cremnocephalus species (Heteroptera, Miridae) * S. Grozeva1 & N. Simov2 1 2
Institute of Zoology, Tzar Osvoboditel 1, 1000 Sofia, Bulgaria. E-mail:
[email protected] National Museum of Natural History, Tzar Osvoboditel 1, 1000 Sofia, Bulgaria. E-mail:
[email protected]
ABSTRACT The karyotype of Cremnocephalus alpestris Wagner, 1941, 2n=30 (28+XY) and C. albolineatus Reuter, 1875, 2n=28 (26+XY) is studied. The original information on the behaviour of the chromosomes, and on the amount (small), distribution (telomeric), and content (mainly AT-rich) of the C-heterochromatin is provided. The nucleolus organizer regions (NORs) are localized on the sex chromosomes. Present karyological study shows that the karyotype can be used as a reliable taxonomic character to identify these two species. Such study should be applied to the rest three Cremnocephalus species, which are morphologically very close and show variable diagnostic characters. Keywords: Heteroptera, Miridae, Cremnocephalus, male achiasmatic meiosis, cytotaxonomy.
INTRODUCTION The genus Cremnocephalus includes five species widespread in Western Palaearctic (Kerzhner & Josifov 1999; Magnien 2000), associated with coniferous trees as Abies, Pinus, Picea, Larix and Pseudotsuga (Wagner 1974; Rieger 1983; Magnien 2000; *
This paper is dedicated to our colleague and teacher Dr. M. Josifov in ocassion of his 80th anniversary and in recognition of his remarkable contribution to the taxonomy and faunistics of Palaearctic Heteroptera.
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Wachman et al. 2004; Gossner et al. 2005). Some of the recently described species of Cremnocephalus (Rieger 1983; Magnien 2000) are very similar in their morphological characters, for instance Cremnocephalus kariae Rieger, 1983 and C. alpestris Wagner, 1941 (Magnien 2000). By examination of C. alpestris specimens from Bulgaria was detected that some of the diagnostic characters used for differentiation of above listed species, as drawing on clavus and especially d/c ratio (Magnien 2000), vary in different level even among the specimens from the same locality and the same tree species. The aim of the authors was to study the karyotypes and other cytogenetic characters of different Cremnocephalus species, and to evaluate these characters as taxonomical markers for species discrimination.
MATERIAL AND METHODS About 30 males of Cremnocephalus alpestris Wagner, 1941 and over 40 males C. albolineatus Reuter, 1875 were collected from different coniferous trees in Bulgarian mountains (Table 1) and fixed in 3:1 fixative (96% ethanol - glacial acetic acid mixture). Species identification is done after Magnien (2000). The species identity of C. alpestris specimens with variable characters is confirmed by male and female genitalia’s structures after Magnien (2000). Table 1. List of localities of collected specimens Species Cremnocephalus albolineatus Reuter, 1875
Karyotype 2n= 8(26+XY)
Cremnocephalus alpestris Wagner, 1941
2n=30(28+XY)
Locality West Rhodopes, 1400m, hut “Martziganitsa”, on Pinus sylvestris, 2.07.2004 West Rhodores, 1200m, Dobrostan village, on Pinus nigra, 2.07.2004 Pirin mts, Bossovo brdo, on Pinus nigra, 16.06.04 Kresna, Cool bath, on Pinus nigra, 14.06.04 Pirin mts, 1335m, Predela, on Abies alba, 2.04.2004 Pirin mts, 1335m, Predela, on Picea abies, 8.07.2004 Pirin mts, hut ”Bynderitsa”, on Pinus peuce, 18.08.2004 Rila mts., Semkovo, on Abies alba, 8.07.04 Central Balkan, 1700m, Tsarichina Reserve, on Abies alba, 1.08.2004
The gonads were squashed in a small drop of 45% acetic acid. The cover slips were removed by dry ice technique. Slides were dehydrated in fresh fixative (3:1) and air dried. Part of the preparations were stained by Schiff-Giemsa method of Grozeva & Nokkala (1996) to study the number and the behaviour of the chromosomes, whereas for the other slides different method was applied. The DNA binding fluorochromes,
Cytotaxonomy of two Cremnocephalus species (Heteroptera, Miridae) 173
GC-specific chromomycin A3 (CMA3) and AT-specific 4’-6’ diamino-2-phenylindole (DAPI) were used to reveal molecular organization of the chromatin according to the method of Schweizer (1976) and Donlon & Magenis (1983), with minor modifications by Kuznetsova et al. (2001). Chromosomes spreads were analysed using Laborlux 12 (Leitz, Wetzlar, Germany) with Olympus C 5060 digital camera; fluorochrome labelled slides were studied using the fluorescent microscope Dialux 22 (Leitz, Wetzlar, Germany). The fluorescence pictures were documented on Kodak 160 ASA slide colour film.
RESULTS Cremnocephalus albolineatus REUTER, 1875 2n=28 (26+XX/XY) Spermatogonial metaphases of this species consist of 26 autosomes, gradually decreasing in size, very large X and small Y sex chromosomes (Fig. 1a). This stage shows the X chromosome with a degree of condensation comparable with that of the autosomes. The Y chromosome is one of the smallest chromosomes while the X chromosome is several times larger than the Y and the largest chromosome in the set (Fig. 1b). The X chromosome is more condensed in mitotic stages. The condensation stage revealed 13 autosomal bivalents consisted of two parallel aligned chromosomes, connected usually by one or two collochores, and the positively heteropycnotic X and Y sex chromosomes, which were situated either spatially well separated (Fig. 2) or together as one body (Fig. 3) till the prometaphase (PM) (Fig. 4). In PM (Fig. 5) and metaphase I (MI) (Fig. 6), the 13 bivalents could be observed, while sex chromosomes could not be distinguished among them. The sex chromosomes underwent equitional separation in anaphase I and segregation in AII. Thus at the radial second metaphase (MII), it was easy to recognize the asymmetric pseudobivalent of sex chromosomes (Fig. 7). In some telophase II plates a large chromosome (probably the X) was observed to lag (Fig. 8).
Cremnocephalus alpestris WAGNER, 1941 2n=30 (28+XY) Spermatogonial metaphases were not found. Nuclei at condensation stage revealed 14 autosomal bivalents, consisted of two autosomes without chiasmata, connected however by one or two collochores (Fig. 9); and a positively heteropycnotic body of the sex chromosomes (X and Y). In early (Fig. 10) and mature (Fig. 11) metaphase I (MI), sex chromosomes could be distinguished among the autosomal bivalents. At the second metaphase (MII), autosomes formed clear ring and showed radial like arrangement. The sex chromosomes formed asymmetric pseudobivalent in the centre of the plate (Fig. 12).
174 S. Grozeva & N. Simov
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Figs 1-8: Cremnocephalus albolineatus: Fig. 1: Spermatogonial metaphase, plate (a), karyogram (b); Figs 2-3: Condensation stage, the sex chromosomes and separately (2) or together (3); Fig. 4: Late condensation stage; Fig. 5: Prometaphase I (PMI); Fig. 6: Metaphase I (MI); Fig. 7: Metaphase II (MII); Fig. 8: Anaphase II (AII). Figs 9-15: Cremnocephalus alpestris: Fig. 9: Condensation stage; Fig. 10: Prometaphase I (MI); Fig. 11: Metaphase I (MI); Fig. 12: Metaphase II (MII); Figs 13-15: Fluorochrome staining: Condensation stage after CMA3 staining (13), Prometaphase I after DAPI staining (14-15). Bar equals 10µm.
Cytotaxonomy of two Cremnocephalus species (Heteroptera, Miridae) 175
The Y chromosome was one of the smallest chromosomes while the X chromosome was larger and seemed to be the largest in the set. When CMA 3 -stained, condensation stages showed bright sex chromatin bodies and one or two signals on their surface (Fig. 13). Positive signals of CMA3 revealed GC-rich areas on sex chromosomes, and indicated the NOR position on the X or Y chromosomes. DAPIstaining revealed bright signals on one of the telomeres of two larger bivalents. Some positional polymorphism of one of these bivalents was observed. The signal of every homolog was found to be on the adjacent (Fig. 14) or on the opposite ends of the homologs (Fig. 15). The examination of C. alpestris specimens from Bulgaria showed that some of the diagnostic characters used for differentiation of Cremnocephalus species, namely drawing on clavus and d/c ratio (Fig. 16) varied even among the animals from the same locality and the same tree species (Picea abies). For example the d/c ratio in specimens of C. alpestris collected in one locality on Picea abies at 1600 m a.s.l. on Vitosha Mts., varied from 0.45 to 0.68. After Magnien (2000) the value of d/c ratio is <0.55 for C. alpestris and >0.55 for C. kariae. At the same time, any notable difference in the karyotype of the morphologically variable specimens collected from one locality or even from one tree species (Abies alba, Picea abies or Pinus peuce) was observed.
c
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Fig. 16: Cremnocephalus alpestris – Variation of drawing on clavus among specimens from locality Zlatnite Mostove 1600 m a.s.l. in Vitosha Mt., Bulgaria, collected on Picea alba.
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DISCUSSION Untill the present time about 200 (less then 4%) species from 6 mirid subfamilies have been karyologically studied (Ueshima 1979; Nokkala & Nokkala 1986a, b; Grozeva 2003; Grozeva et al. 2006; Grozeva & Simov 2008). Although the chromosome numbers of the mirid species range from 13 to 80, however the majority of species display the modal chromosome number for the family is 2n = 34 (XY), described in 79 species (about 46% of species studied). In the tribe Hallodapini (the subfamily Phylinae) the karyotype of only two species, Hallodapus linnavuori Miyamoto, 1966 with 2n=26(24+XY) (Muramoto 1973), Orectoderus obliquus Uhler, 1876 with 2n=34(32+XY) (Akingbohungbe 1974), have been so far known. Two Cremnocephalus species from the same tribe studied in the present paper display different chromosome numbers, 2n = 28(26+XY) in C. albolineatus and 2n = 30(28+XY) in C. alpestris. The data available, few as they are, suggest the variety of chromosome numbers in the tribe that provides a good impetus to further karyotaxonomical studies of the group. In the species studied, the male meiosis is achiasmatic and of collochore type, as in other Miridae (Nokkala & Nokkala 1986a; Grozeva at al. 2006), as well as in Cimicidae (Grozeva & Nokkala 2002). Till now, two basic types of achiasmatic meiosis have been reported in the Heteroptera. By the descriptions of Nokkala & Nokkala (1986a) and Kuznetsova et al. (2007) in the most widespread type, the alignment of homologous chromosomes attained early in meiosis remained externally unchanged until the beginning of anaphase, whereas in the other type, after synapsis, the so-called collochores are formed, and homologous chromosomes are connected with each other at specific positions via these tenacious structures. Within the Heteroptera, the “alignment” type occurs in the families Saldidae, Micronectidae, Anthocoridae, Microphysidae, and Nabidae (Nokkala & Nokkala 1983; Nokkala & Nokkala 1984; Nokkala & Nokkala 1986a, b; Kuznetsova & Maryaska-Nadachowska 2000; Nokkala & Grozeva 2000; Kuznetsova et al. 2004; Grozeva et al. 2008), whereas the “collochore” type is encountered in the cimicomorphan families Miridae and Cimicidae (Nokkala & Nokkala 1986a; Grozeva & Nokkala 2002). The pattern of meiosis, especially the behaviour of the sex chromosomes, varies between species of the Heteroptera. In majority of heteropteran species and in all Miridae species studied, the autosomal bivalents segregate reductionally during the first meiotic division, whereas the sex chromosomes divide equationally at anaphase I, associate in second meiotic division showing “touch-and-go pairing” and segregate reductionally at anaphase II (Ueshima 1979; Nokkala & Nokkala 1986c). The same behaviour of sex chromosomes was observed in two Cremnocephalus species form the present study. The true bugs characteristically display small amount of heterochromatin in their karyotypes with both telomeric and interstitial localization and variously distribution in different species (Grozeva & Nokkala 2003; Grozeva et al. 2004). The analysis of constitutive heterochromatin content and distribution in C. alpestris revealed the
Cytotaxonomy of two Cremnocephalus species (Heteroptera, Miridae) 177
presence of conspicuous DAPI-positive bands (indicating an excess of AT base pairs) of the terminal location in two large bivalents with some positional polymorphism in one of these bivalents. The CMA-bright bands (indicating an excess of GC base pairs) corresponding with NORs has also been reported in other heteropteran species (González-García et al. 1996; Rebagliati et al. 2003; Kuznetsova et al., 2007). The NORs are usually localized on the sex chromosomes (Grozeva et al. 2004), and this is true for C. alpestris. Thus, present karyological study proves that C. alpestris and C. albolineatus can be distinguished very well by their different chromosome numbers. The study of further Cremnocephalus species is needed to learn if chromosome markers are reliable taxonomic characters for systematics of the group.
РEЗЮМЕ Изучен е кариотипът на два вида Cremnocephalus: C. alpestris WAGNER, 1941 (2n=30, 28+XY) и C. albolineatus REUTER, 1875 (2n=28, 26+XY). Получена е оригинална информация за поведението на хромозомите, както за количество (малко), състава (предимно от АТповтори) и разпределението (теломерно) на хетерохроматина. Ядърцевият организатор (NOR) е разположен на половите хромозоми. Настоящото кариологично изследване показва, че кариотипът може да бъде надежден белег за разграничаване на тези два вида. Такова изследване би било полезно да се проведе и за останалите три вида от рода, които са морфологично много близки, и с вариабилни диагностични белези.
REFERENCES Akingbohungbe A. E. (1974). Chromosome numbers of some North American mirids (Heteroptera: Miridae). — Canadian Genetics and Cytology, 16: 251-256. Donlon T.A. & R.E. Magenis (1983). Methyl green is a substitute for distamycin A in the formation of distamycin A/DAPI C-bands. — Human Genetics, 65: 144-146. González-García J.M., C. Antonio, J.A. Suja & J.S. Rufas (1996). Meiosis in holocentric chromosomes: kinetic activity is randomly restricted to the chromatid ends of sex univalents in Graphosoma italicum (Heteroptera). — Chromosome Research, 4: 124-132. Gossner M., A. Gruppe & U. Simon (2005). Aphidophagous insect communities in tree crowns of Norway spruce (Picea abies L.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco). — Journal of Applied Entomology, 129 (2): 81-89. Grozeva S. (2003). Karyotype of three endemic Mediterranean Miridae species (Heteroptera) from Bulgaria. — Acta Zoologica Bulgarica, 55 (1): 53-59. Grozeva S., V. Kuznetsova & S. Nokkala (2004). Patterns of chromosome banding in four nabid species (Heteroptera, Cimicomorpha, Nabidae) with high chromosome number karyotypes. — Hereditas, 140: 99-104.
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Grozeva S. & S. Nokkala (1996). Chromosomes and their meiotic behaviour in two families of the primitive infraorder Dipsocoromorpha (Heteroptera). — Hereditas, 125: 31-36. Grozeva S. & S. Nokkala (2002). Achiasmatic male meiosis in Cimex sp. (Heteroptera, Cimicidae) — Caryologia, 55(3): 189-192. Grozeva S. & S. Nokkala (2003). C-heterochromatin and extra (B) chromosome distribution in six species of the Nabis (Heteroptera, Nabidae) with the modal male karyotype 2n = 16 + XY. — Folia biologica (Kraków), 51(1/2): 13-21. Grozeva S., S. Nokkala & N. Simov (2006). First evidence of sex chromosomes prereduction in male meiosis in the Miridae bugs (Heteroptera). — Folia Biologica (Krakow), 54(1-2): 9-12. Grozeva S., S. Nokkala & N. Simov (2008). Achiasmatic meiosis in two Micronecta species (Heteroptera, Nepomorpha, Micronectidae). — Comparative cytogenetics, 2(1) (in press). Grozeva S. & N. Simov (2008). Cytogenetic Studies of Bryocorinae true bugs (Heteroptera, Miridae). — Acta Zoologica Bulgarica 60 (1): (in press). Kerzhner I.M. & M. Josifov (1999). Family Miridae Hahn, 1833. — In: Aukema, B. & C. Rieger (Eds) Catalogue of the Heteroptera of the Palaearctic Region. The Netherlands Entomological Society, Amsterdam, Vol. 3: 1-576. Kuznetsova V.G., S. Grozeva & S. Nokkala (2004). New cytogenetic data on Nabidae (Heteroptera: Cimicomorpha), with a discussion of karyotype variation and meiotic patterns, and their taxonomic significance. — European Journal ofEntomology, 101: 205-210. Kuznetsova V.G., S. Grozeva, J. N. Sewlal & S. Nokkala (2007). Cytogenetic characterization of the endemic of Trinidad, Arachnocoris trinitatus Bergroth: the first data for the tribe Arachnocorini (Heteroptera: Cimicomorpha: Nabidae). — Folia Biol. (Krakow), 55 (1/2): 17-26. Kuznetsova V.G. & A. Maryaska-Nadachowska (2000). Autosomal polyploidy and male meiotic pattern in the bug family Nabidae (Heteroptera). — Journal of Zoological Systematics and Evolutionary Research, 38: 87-94. Kuznetsova V. G., M. Westendorff & S. Nokkala (2001). Patterns of chromosome banding the sawfly family Tenthredinidae (Hymenoptera, Symphyta). — Caryologia, 54: 227-233. Magnien P. (2000). Revision du genre Cremnocephalus Fieber, 1860; description de deux nouvelles especes du sud de la France et de Calabre; note sur les genitalia femelle (Heteroptera, Miridae). — Nouvelle Revue Entomologique (N.S.), 17 (1): 51—67 Muramoto N (1973). A chromosome study in eighteen Japanese Heteropterans (in Japanese). — La Kromosomo, 91: 2896-2905. Nokkala S. & S.Grozeva (2000). Male meiosis of achiasmatic type in Myrmedobia coleoptrata (Fn.) (Heteroptera, Microphysidae). — Caryologia, 53 (1): 5-8. Nokkala S. & C. Nokkala (1983). Achiasmatic male meiosis in two species of Saldula (Saldidae, Hemiptera). — Hereditas, 99: 131-134. Nokkala S. & C. Nokkala (1984). Achiasmatic male meiosis in the Heteropteran genus Nabis (Nabidae, Hemiptera). — Hereditas, 101: 31-35. Nokkala S. & C. Nokkala (1986a). Achiasmatic male meiosis of collochore type in the heteropteran family Miridae. — Hereditas, 105: 193-197. Nokkala S. & C. Nokkala (1986b). Achiasmatic male meiosis in Anthocoris nemorum (L.) (Anthocoridae, Hemiptera). — Hereditas, 105: 287-289.
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Nokkala S. & C. Nokkala (1986c). The mechanisms behind the regular segregation of autosomal univalents in Calocoris quadripunctatus (Vil.) (Miridae, Hemiptera). — Hereditas, 105: 199-204. Rebagliati P., A.G. Papeschi & L.M. Mola (2003). Meiosis and fluorescent banding in Edessa meditabunda and E. rufomarginata (Heteroptera: Pentatomidae: Edessinae). — European Journal of Entomology, 100: 11-18. Rieger C. (1983). Ein neuer Cremnocephalus aus Griechenland (Heteroptera, Miridae). — Nachrichtenblatt Bayerischen Entomologen, 32: 75-77. Schweizer D. (1976). Reverse fluorescent chromosome banding with chromomycin and DAPI — Chromosoma, 58: 307-324. Wachmann E., Melber A. & J. Deckert (2004). Wanzen 2. — Die Tierwelt Deutschlands 75: 1-294. Wagner E. (1974). Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Teil 2. — Entomologische Abhandlungen Staatliches Museum für Tierkunde in Dresden, 39 Suppl. ii+421. Ueshima N. (1979). Hemiptera II: Heteroptera. — In: Bernard J. (ed.): Animal Cytogenetics. Vol. 3. Insecta. B. John, Berlin, Stuttgart: 1-113.
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S. Grozeva &Spain: N. Simov (Eds)michaili 2008 sp. nov. (Heteroptera: Tingidae) 181 A new tingid species from Southern Dictyonota ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 181-184. © Pensoft Publishers Sofia–Moscow
A new tingid species from Southern Spain: Dictyonota michaili nov. sp. (Heteroptera: Tingidae) * H. Günther Eisenacher Straße 25, D-55218 Ingelheim, E-mail:
[email protected]
ABSTRACT A new species of Tingidae from Southern Spain is described. Dictyonota michaili nov. sp. was collected on Retama sphaerocarpa (L.) Boiss. (Fabaceae) in the desert like region of Tabernas, province of Almeria. Its characters are described and compared with those of other species of Dictyonota in Europe. Keywords: Dictyonota michaili nov. sp., Tingidae, Spain, Retama sphaerocarpa.
Dictyonota michaili nov. sp. Material studied Holotypus ♂, 19.05.1997 surrounding of Tabernas, „Minihollywood“, on Retama sphaerocarpa (L.) Boiss. U. Koschwitz leg., in the collection of the Naturhistorisches Museum Mainz (NHMM). Paratypes 2 ♂, 19.05.1997 Tabernas, U. Koschwitz leg., 1 ♀, Tabernas, 19.06.2005, H. Günther leg. All deposited in the NHMM. The host plant of Dictyonota michaili nov. sp., Retama sphaerocarpa (L.) Boiss. (Fabaceae), is a common shrub in the region round Tabernas, province of Almeria, Southern Spain (Fig. 1) (Sagredo 1987). *
I dedicate the new species to my friend and colleague Michail Josifov, Sofia, on the occasion of his 80th birthday.
182 H. Günther
Inspite of that, the new species of Dictyonota is very rare. Besides the three specimens collected in 1997 by U. Koschwitz, only one female could be recorded in 2005. It was the only record during three further excursions in the region. Description of the new species (Figs 2, 3) Body length ♂ 3.0 – 3.3 mm; ♀ 3.0 mm; breadth 2.4 – 2.8 mm. Body yellow and brown with bright, opaque cells. Head black, curved and distinctly punctuated with two short, pale occipital and two frontal spines. Occipital spines prominent obliquely upward and forward. Behind eyes, a neck is present. Antennae slender. Segments one to three brown, segment four black. Total length of antennae 1.4 mm, length of the segments one to four 0.15, 0.15, 0.85 and 0.25 mm. Third segment long, covered with small tubercles, leaving visible the surface of the segment. By this characteristic, D. michaili nov. sp. distinguishes from most all other European Dictyonota species with a short, thickened third antennal segment, which is totally covered and masked by tubercles with bristles (Péricart 1983, p. 144). Pronotum proximally black, distally brown. Paranota bearing a row of six to seven large cells, occasionally anterior cells divided and therefore arranged in two rows. Pronotal ampoule very low, not reaching the height of median pronotal carina in lateral view, markedly lower than in all other European Dictyonota species. The three pronotal carinae low, each with a row of eight to nine small cells. Cells in the middle of carinae with very small lumen or even totally closed. By this character, D. michaili nov. sp. distinguishes from D. oblita, the only Dictyonota species in Europe with a long, unmasked second antennal segment. (Péricart Fig. 1: Retama sphaerocarpa (L.) Boiss., the host plant of 1983, p.159, 161). Dictyonota michaili nov. sp.
A new tingid species from Southern Spain: Dictyonota michaili sp. nov. (Heteroptera: Tingidae) 183
Fig. 3: Dictyonota michaili, Pronotum lateral, photo G. Strauß
Fig. 2: Dictyonota michaili, total, photo G. Strauß
Hemelytra with bright, opalescent cells of different sizes separated by robust brown veins. Costal area (lame costale, Randfeld) with 18 (± 1) cells. Posterior five to six cells of medium size; anterior five to six cells large, accompanied with two small cells between them. Subcostal area (aire subcostale, Seitenfeld) with three rows of cells, two rows of small cells on outer side, one row of large cells on inner side. Discoidal area (aire discoidale, Mittelfeld) with two or three markedly large, opalescent cells on outer side, two or three cells of medium size, also opalescent on inner side, and three rows of cells of different sizes in the middle. Sutural area (aire suturale, Schlußfeld) with six to seven rows of cells of different sizes. Tibiae and tarsi yellow; femora brown.
DISCUSSION Dictyonota michaili nov. sp. belongs to the group of Dictyonota species with long and slender antennae, especially with a long third antennal segment. Its surface is not totally covered with tubercles and therefore visible. There exists only one other Dictyonota species with a similar characteristic in Europe, Dictyonota oblita Péricart, 1981. This species is smaller (2.5 – 2.75 mm), it distinguishes from D. michaili nov. sp. by the form of the cells in the pronotal carinae, they are distinctly visible and of equal sizes.
184 H. Günther
In contrast to Dictyonota michaili nov. sp. and D. oblita, all other European Dictyonota species have shorter and thicker antennae with a shorter and totally or nearly totally masked third antennal segment. For the group of Dictyonota oblita, D. gracilicornis Puton, 1874 and further nonEuropean species Péricart (1983) gives Chenopodiaceae as food plants. The group of Dictyonota strichnocera Fieber, 1844, D. fuliginosa A. Costa, 1853, D. marmorea Baerensprung, 1858 and others live on Fabaceae, as does Dictyonota michaili nov. sp.
ACKNOWLEDGEMENTS I thank my friend Dr. U. Koschwitz, Eppenbrunn, who let to me the three specimens of Dictyonota michaili nov. sp., collected by him in 1997 and my friend Gerhard Strauß, Biberach, who made the excellent photos of Dictyonota.
РЕЗЮМЕ Описан е нов вид от сем. Tingidae от Южна Испания. Dictyonota michaili nov. sp. e събран на Retama sphaerocarpa (Fabaceae) в пустинен район в Taбeрнас, провинция на Aлмерия. Описани са особеностите на вида и са сравнени с тези на други видове от род Dictyonota в Европа.
REFERENCES Péricart J. (1983) Hémiptères Tingidae Euro-Méditerranéens. – Faune de France 69: Paris: 1-618. Sagredo, R. (1987) Flora de Almeria. Plantas Vasculares de la Provincia. – Diputacion Provincial de Almeria: 1-557.
S. Grozeva & N.ofSimov (Eds) 2008 A new species Pilophorus from the Far East (Heteroptera, Miridae) 185 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 185-188. © Pensoft Publishers Sofia–Moscow
A new species of Pilophorus from the Far East (Heteroptera, Miridae) I.M. Kerzhner Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia
ABSTRACT Pilophorus josifovi nov. sp. is described from Primorsk Territory of Russia and North Korea. The species is closely related to P. erraticus Lnv., P. pseudoperplexus Jos., and P. okamotoi Miy. & Lee. Keywords: Heteroptera, Miridae, Pilophorus, new species, Russia, Korea.
INTRODUCTION The genus Pilophorus is represented in the Far East by about 15 species (Kerzhner & Josifov 1999), some of them very similar. The new species described below was misidentified by Josifov (1987) and partly by Kerzhner (1988) as P. erraticus Lnv.
Pilophorus josifovi nov. sp. Pilophorus erraticus (non Linnavuori, 1962): Josifov 1987: 120; Kerzhner 1988: 72 (part., male from Vinogradovka). Holotype: ♂, Russia, Primorsk Territory, Vinogradovka, 6.VIII.1929 (A.N. Kiritshenko), Zoological Institute, St. Petersburg, Russia.
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Paratypes: 2♀♀, North Korea, Hangjang-pukdo, Džikha-ri, 16 km S of Čhongdžin, on Ulmus, 31.VIII.1970 (M. Josifov), Institute of Zoology, Sofia, Bulgaria and collection of E. Heiss, Innsbruck, Austria. Description. Elongate, macropterous, 3.55 (♀) to 3.58 (♂) times as long as width of pronotum. Pubescence of dorsum golden, very short, appressed. Head brownish black in upper half, red in lower half. Hind margin of head with several erect dark setae, these about half as long as those in P. erraticus. Vertex 1.6 times as wide as eye. Antennal segment 1 red with narrowly yellowish base and yellow ventral side; segment 2 red proximally and gradually becoming dark reddish brown in apical third, gradually and moderately thickened toward apex (15 times as long as wide), not clavate; segment 3 white in basal half and black in apical half; segment 4 black with narrowly white base. Length of antennal segments (1–4): 0.31–0.33, 1.47–1.57, 0.57–0.64, 0.50 mm. Rostrum yellow, black apically, extending to fore margin of middle coxae. Pronotum brownish black, weakly shining, with almost indistinct transverse striation, 1.25–1.30 times as long as wide (measured perpendicular to surface of pronotum!), concave laterally. Scutellum brownish black, with two longitudinal spots of silvery scales. Clavus and most of corium pale brown, dull; exocorium behind distal silvery band and cuneus darker and shiny. Corium with two silvery bands formed of scales; distal band crossing clavus without shift. Cuneus with band of silvery scales on fore margin. Membrane smoky brown, dull, with all margins (except base) paler and shiny. Ventral side of body dark brown, with a line of silvery scales on hind margin of mesothorax and two spots of silvery scales near base on each side of abdomen. Fore and hind coxae and all trochanters yellowish white; middle coxae brownish. Fore femora pale yellow, reddish in central part. Middle and hind femora and all tibiae red. Hind tibiae distinctly curved, with black bristles in apical 5/6. Tarsi yellow, with blackish brown segment 3. Parameres as in Figs 1–3. Phallotheca (Fig. 4) with a long pointed apex. Vesica (Fig. 5) slender; its lateral process short, pointed, with two lateral teeth directed oppositely. Body length 4.4 mm. Comparison. The new species is similar to the East Palaearctic P. erraticus Linnavuori, 1962 (= P. alni Josifov, 1987), P. pseudoperplexus Josifov, 1987 (= P. oculatus Kerzhner, 1988), and P. okamotoi Miyamoto & Lee, 1966, but these species have all femora and usually also antennal segment 1, pale portion of segment 2, and tibiae yellow. In addition, in P. erraticus antennal segment 2 is wider and slightly clavate; in the other two species the hind tibiae are not curved, P. pseudoperplexus has antennal segment 3 blackened only at the extreme apex (incorrectly shown in Kerzhner 1988, fig. 212), and in P. okamotoi the vertex is about twice or more than twice as wide as the eye and antennal segment 3 is half as long as segment 2. The male genitalia of the new species, especially the phallotheca and vesica, are also distinctive. Etymology. The new species is named after famous Bulgarian heteropterist Michail V. Josifov, my old friend and colleague, on the occasion of his 80th birthday. We met both in Russia and Bulgaria, participated in a joint field trip to Tajikistan and Kyrgyzstan, and coauthored papers about Mongolian and Korean Heteroptera and in the Miridae
A new species of Pilophorus from the Far East (Heteroptera, Miridae) 187
1 2
4
3
5
Figs 1-5: Pilophorus josifovi nov. sp., holotype. 1, left paramere; 2–3, right paramere; 4, phallotheca; 5, vesica.
volume of the Palaearctic catalogue. Collaboration with Misha was always pleasant and productive.
ACKNOWLEDGEMENTS I am thankful to Nikolay Simov (Sofia) for important information. The collection of Zoological Institute, St. Petersburg, is supported by Rosnauka for UFC no. 2-2.20.
РЕЗЮМЕ Pilophorus josifovi nov. sp. е описан от Русия и Северна Корея, граничния район Приморск. Видът е близък до P. erraticus LNV., P. pseudoperplexus JOS., и P. okamotoi MIY. & LEE.
REFERENCES Josifov M. (1987). Einige neue Miriden aus Nordkorea (KDVR) (Heteroptera). — Reichenbachia, 24: 115–122.
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Kerzhner I.M. (1988). Novye i maloizvestnye poluzhestkokrylye nasekomye (Heteroptera) s Dal’nego Vostoka SSSR [New and little-known heteropterous insects (Heteroptera) from the Far East of the USSR]. — USSR Academy of Sciences, Far East Centre, Vladivostok, (1987): 1–83. (In Russian). Kerzhner, I.M. & Josifov, M. (1999). Cimicimorpha II. Family Miridae. — Aukema, B. & Rieger, Chr. (Eds). Catalogue of the Heteroptera of the Palaearctic Region, The Neteherlands Entomological Society, Amsterdam, 3: 1–576.
Grozeva & N. Simov (Eds) 2008 Review of theS. family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 189 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 189-213. © Pensoft Publishers Sofia–Moscow
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia P. Kment1 & Z. Jindra2 1
Department of Entomology, National Museum, Kunratice 1, CZ-148 00 Praha & Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44 Praha 2, Czech Republic. E-mail:
[email protected] 2 Department of Plant Protection, Faculty of Agrobiology, Food and Natural Resources, Czech Agricultural University, CZ-165 21 Praha 6, Czech Republic. E-mail:
[email protected]
ABSTRACT Review of the nine species of Nerthra Say, 1832 (Hemiptera: Heteroptera: Gelastocoridae: Nerthrinae) occurring in south-eastern Asia west of Wallace line is given. The male of N. nieuwenhuisi Todd, 1957, the 4th instar larva of N. asiatica (Horváth, 1892), and 3rd, 4th and 5th instar larvae of N. indica (Atkinson, 1889) are described, and illustrated for the first time. Nerthra indica = N. arunachalensis Thirumalai, 1998 is proposed as a new synonymy, and the variability of N. indica is briefly discussed. First exact records are given for N. nieuwenhuisi from Sarawak (Malaysia), and N. indica from Fujian (China), Laos and Vietnam. Keywords: Gelastocoridae, Nerthra, larva, Oriental region, China, India, Indonesia, Laos, Malaysia, Nepal, Vietnam, faunistics
INTRODUCTION Toad bugs (Gelastocoridae) are a remarkable group of ‘aquatic’ bugs (Nepomorpha) which are derived from aquatic ancestors and and have become secondarily terrestrial (Hebsgaard et al. 2005). This small family is divided into two subfamilies, Gelastocorinae and Nerthrinae; Gelastocorinae, include the genera Gelastocoris Kirkaldy, 1897
190 P. Kment & Z. Jindra
and Montandonius Melin, 1929, inhabit wet margins of water bodies in America, from southern Canada in the north to Argentina in the south (e.g., Todd 1955, 1961a; Nieser 1975; Schnack & Estévez 1979; Štys & Jansson 1988). The Nerthrinae includes two genera. The fossil Cratonerthra Martins-Neto, 2005 (with two species) was recently described from Early Cretaceous sediments of Santana Formation in Ceará state, south-eastern Brazil (López Ruf et al. 2005). The only recent genus, Nerthra Say, 1832 (sensu Todd 1955), currently includes 91 valid species (Kment, in prep.). However Cassis (2006) and Cassis & Silveira (2006), have proposed a splitting of the genus based on a phylogenetic analysis of Nerthra species to include five genera within the current concept of Nerthra sensu lato. Nerthra species are distributed in tropical and subtropical regions on all continents (except for Europe). However, the most speciose fauna of Nerthra is found in America, Australia, and Malesia (e.g., Todd 1955, 1961a; Cassis & Silveira 2001, 2002, 2004, 2006; Andersen & Weir 2004; Chen et al. 2005). Only one species, N. grandicollis (Germar, 1837) is restricted to the Afrotropical region including Madagascar (Todd 1955, 1961a). Another nine species of Nerthra are known from the south-eastern Asia, west of Wallace’s line (Todd 1961b). Nerthra species inhabit various littoral (wet banks of fresh waters, sea coasts) as well as truly terrestrial habitats, such as banana plantations (Kevan 1942), open heathland or forests (Cassis & Silveira 2001). Some species are thought to be nocturnal, hiding during daytime under stones, plant debris or in humid sand or mud. They are predators of various small invertebrates (e.g., Kevan 1942, Todd 1955, Chen et al. 2005). We have the pleasure to dedicate this paper to Michail Josifov, an outstanding Bulgarian heteropterist, on the occasion his 80th birthday.
MATERIAL AND METHODS Dry-mounted specimens were studied under a binocular stereomicroscope MBS-10. Measurements were made with an ocular micrometer. For the study of male genitalia, specimens were softened in hot water with admixture of 70% ethanol. The line-drawings were prepared from photographs taken by an Olympus Camedia C-5060 camera attached to an Olympus SZX9 binocular microscope. Base maps were downloaded from On-line Map Creation http://www.aquarius.geomar.de/omc/. The following codens of the collections are used throughout the paper: BMNH Natural History Museum, London, United Kingdom; MCSN Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy; MMBC Moravian Museum, Brno, Czech Republic; MNHN Muséum National d’Histoire Naturelle, Paris, France; NHMW Naturhistorisches Museum, Wien, Austria; NMEG Naturkundesmuseum, Erfurt, Germany; NMPC National Museum, Praha, Czech Republic;
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 191
NZSI
National Zoological Collection, Zoological Survey of India, Calcutta, West Bengal, India; OXUM University Museum of Natural History, Oxford, United Kingdom; PBPC Petr Baňař private collection, Brno, Czech Republic; RMNH Nationaal Natuurhistorische Museum, Leiden, Netherlands; SEMC Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA; USNM United States National Museum of Natural History, Washington, D.C., USA; VSPC Vít Socha private collection, Pardubice, Czech Republic; ZJPC Zdeněk Jindra private collection, Praha, Czech Republic; ZMAS Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia; ZMUH Zoologisches Museum, Universität von Hamburg, Germany.
REVIEW OF THE SPECIES Nerthra asiatica (HORVÁTH, 1892) (Figs 1, 5, 8) Mononyx asiaticus Horváth, 1892: 136 (description, type locality). Holotype: female, ‘China: Flumen Poi-ho (G. N. Potanin)’ [= Sichuan, Gar Qu (= Pai Ho River)] (ZMAS) (see Kiritshenko 1926 and Polhemus 1995). Mononyx asiaticus: Oshanin (1909): 956 (catalogue, distribution); Oshanin (1912): 89 (catalogue, distribution); Kiritshenko (1926): 226 (taxonomy, synonymy); Kiritshenko (1930): 435 (distribution); Wu (1935): 559 (catalogue, distribution); Hoffmann (1933): 250 (catalogue: China); Hoffmann (1941): 44 (catalogue, distribution). Nerthra asiatica: Todd (1955): 349, 410–411, 472–473 (redescription, figure, key, catalogue, distribution); Todd (1957): 154 (variability, faunistics); Todd (1961a): 466 (catalogue, distribution); Todd (1961b): 93 (faunistics); Nieser & Chen (1992): 5–6 (comparative note); Polhemus (1995): 24 (catalogue, distribution); Thirumalai (1998): 192 (key); Bal & Basu (2003): 542–544 (key, distribution, faunistics). Nerthra asiaticus: Hua (2000): 214 (checklist: China). Mononyx grossus Montandon, 1899a: 395, 398. Syntypes: China, ‘Thibet, Mou-pin (A. David 1870)’ [= Sichuan, Baoxing (= Mu-p’ing)] & ‘Tchouen (A. David 1875)’ [not identified] (MNHN & coll. Montandon → BMNH, 1 syntype in SEMC) (see Todd 1955 and Polhemus 1995) Mononyx grossus: Distant (1906): 16 (redescription, comparative note, faunistics); Oshanin (1909): 956 (catalogue, distribution); Oshanin (1912): 89 (catalogue, distribution); Kiritschenko (1926): 226 (taxonomy, synonymy); Wu (1935): 559 (catalogue, distribution).
Material examined. CHINA: Sichuan, Huatan env. (Shpshin), ca 50 km SW of Yáan, 29°47‘N 102°41‘E, 16.vii.1995, 1 female, M. Trýzna & O. Šafránek lgt. (ZJPC); Jiulonggou env., near Dayi, 70 km W of Chengdu, 28.vi.–2.vii.1995, 1 larva, Z. Jindra lgt.
192 P. Kment & Z. Jindra
(ZJPC). INDIA: Assam, Noa Dihing, 1 female, Chennell [lgt.], Distant coll., 1911–383, ‘grossus Montand.’ (BMNH). Variability of adults. Todd (1961b) gave the following measurements: body length 11.0-12.2 mm, pronotum width 8.0-8.1 mm, and abdomen width 8.0-8.5 mm. The two examined females in this work are within these ranges, except the pronotum width of the female from Noa Dihing (former measurements belong to the female from India: Noa Dihing, latter to the female from China: Huatan env. (Fig. 1)): total body length 11.3 / 11.6 mm, head width 4.3 / 4.6 mm, inter-ocular width 2.5 / 2.6 mm, pronotum length 3.1 / 3.2 mm, pronotum width 7.3 / 8.0 mm, and abdomen width 8.2 / 8.5 mm, respectively (see also Table 1). Description of 4th instar larva (Fig. 5). Body uniformly brown, covered with pale clavate setae; dorsal surface partly covered with muddy crust. Head with five small tubercles in front of eyes, one pair laterally on each side and one medially; one flat elevation on frons and two slightly higher elevations on vertex between eyes. Pronotum flattened laterally, elevated medially; lateral margins not expanded, straight and paralell in posterior 2/3, bent towards eyes in anterior 1/3, weakly sinuate before anterior angles; poste1 2 rior margin straight. Mesonotum slightly depressed laterally and medially, lateral margins not expanded, straight and paralell; wing pads reaching half length of metanotum. Metanotum with strong depressions sublaterally and submedially, distinctly elevated medially. Abdominal tergites convex, decreasing towards connexivum; lateral margin 3 4 of connexivum turned upwards, forming distinct groove between Figs 1–4: Habitus of adults. 1 – Nerthra asiatica tergites and connexivum; lateral (Horváth, 1892), female from China, Huatan env. margin of connexival segments (11.6 mm); 2 – N. indica (Atkinson, 1889), female from India, Peeling env. (9.5 mm); 3 – N. lobata (Montandon, almost straight, bearing clavate 1899), female from Indonesia, Gugul (10.5 mm); 4 – N. setae in posterior half, posteronieuwenhuisi Todd, 1957, male from Malaysia, Mt. lateral angles slightly prominent f rom the connexival outline, Malang (10.2 mm) (Photo: Jan Macek).
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 193
which seems to be serrate. Measurements (see also Table 2). Differential diagnosis of larva. The studied larva was not found in association with adults of N. asiatica, however, we consider the similarity of head tubercles and shape of thorax as a basis for their identification. The larva of N. asiatica differs from larvae of N. 5 6 indica, in the following characters: Head with five small tubercles in front of eyes (one pair laterally on each side and one median tubercle); pronotum less elevated medially than in N. indica; lateral pronotal margins not expanded, 7 paralell, almost without sinuosity before anterior angles; metanotum not expanded laterally, paralell; Figs 5–7: Habitus of larvae. 5 – Nerthra asiatica (Horváth, lateral margins of connexival 1892), 4th instar larva from China, Jiulonggou env. (7.1 th larva from segments almost straight, postero- mm); 6 – N. indica (Atkinson, 1889), 5 instar rd Nepal, SW Sauraha (8.1 mm); 7 – N. indica, 3 instar larva lateral angles slightly prominent, from China, Maguan (4.5 mm) (Photo: Jan Macek). and connexival outline serrate. Ecology. Collected at 5000 ft [= 1524 m a.s.l.] in Darjiling district, West Bengal (Todd 1961b). Distribution (Fig. 8). China: Sichuan (Horváth 1892; Montandon 1899a; Todd 1957), ?Tibet (Montandon 1899a). India: Assam (Distant 1906), Sikkim (Bal & Basu 2003), West Bengal (Todd 1961b). Discussion. This species is known only from the females. Todd (1955: 411) noted: ‘In the shape of the pronotum it would seem to be closely related to N. spissa (Distant), but the worker has not seen a female of the latter species or a male of N. asiatica (Horvath), so no definite comparison can be made between these two species at the present time’. Additional material of both species is needed to resolve the possible synonymy of N. asiatica and N. spissa.
Nerthra eximia TODD, 1957 (Fig. 9) Nerthra eximia Todd, 1957: 153, 155–156. Holotype: female, Indonesia, Sumatra, ‘Tanangataloo, Ophir-Sum. [= Mt. Ophir], 1915, A. de Kock’ (RMNH).
194 P. Kment & Z. Jindra
Fig. 8: Distribution map of Nerthra asiatica (Horváth, 1892) (circles), N. spissa (Distant, 1911) (rhombs), and N. unguistyla Todd, 1957 (squares).
Fig. 9: Distribution map of Nerthra indica (Atkinson, 1889) (circles), N. eximia Todd, 1957 (asterisk), N. lobata (Montandon, 1897) (squares), N. nieuwenhuisi Todd, 1957 (crosses), and N. serrata (Montandon, 1897) (rhombs).
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 195
Nerthra eximia: Todd (1961a): 468 (catalogue, distribution); Chen et al. (2005): 414 (catalogue: Malesia, distribution).
Ecology. Unknown. Distribution (Fig. 9). Indonesia (West Sumatra) (Todd 1957). Discussion. This species is only known from the female holotype. Todd (1957) wrote: ‘It is very closely related to the preceding species [i.e. N. nieuwenhuisi] and may subsequently prove to be but a form of that species, but for the present I prefer to describe it as a separate species. This species agrees with N. nieuwenhuisi n. sp. and differs from N. lobata (Montandon), the only species previously reported from Sumatra, by the absence of lateral tumescences of the last visible abdominal sternite, by the strongly elevated scutellum, and by the greatest width of the pronotum being at the level of the transverse furrow. It differs from N. nieuwenhuisi n. sp. by its smaller size, differently shaped lateral margins of the pronotum, and differently shaped lateral margin of the embolium. It should be pointed-out, however, that the two sides of the pronotum of this specimen are not alike, and therefore differences of the shapes of the lateral margins of the pronota of the two species may not be significant in this instance. The facts that these are insular species and from different islands was another factor in my decesion to treat this specimen as a separate species’. Further material is needed to improve our knowledge of N. eximia and to confirm its species status.
Nerthra indica (ATKINSON, 1889) (Figs 2, 6–7, 9–14, 17–20) Mononyx indicus Atkinson, 1889: 345. Lectotype: male, India, Sikkim (OXUM). Designated by Lansbury (1988). Mononyx indicus: Montandon (1899a): 394, 397 (key, redescription, faunistics); Distant (1906): 15 (redescription, distribution); Maxwell-Lefroy (1909): 709 (ecology); Paiva (1919): 372 (ecology, faunistics). Nerthra indica: Todd (1955): 350, 405–406, 466–467 (redescription, figure, taxonomy, key, catalogue, distribution, faunistics); Todd (1957): 152 (variability, faunistics); Todd (1961a): 469 (catalogue, distribution); Todd (1961b): 93–94 (taxonomy, figure, faunistics); Nieser (1977): 298 (faunistics); Todd (1977): 216 (faunistics); Lansbury (1988): 248–250 (lectotype designation, redescription); Nieser & Chen (1992): 5–6, 10 (comparative note, figure, faunistics); Polhemus (1995): 24 (catalogue, distribution); Thirumalai (1998): 189–190, 192 (key, faunistics); Hua (2000): 214 (check-list: China); Bal & Basu (2003): 542–544 (key, diagnosis, distribution, faunistics). Mononyx projectus Distant, 1911: 310–311 (description, figure, type locality). Syntypes(?): India, ‘Assam, Sibsagar (Coll. Distant)’ (BMNH). Mononyx projectus: Todd (1955): 405–406 (taxonomy, synonymy). Mononyx turgidulus Distant, 1911: 311–312 (description, figure, type locality). Syntypes(?): ‘Nepal, Gowchar (Coll. Distant)’ (BMNH).
196 P. Kment & Z. Jindra
Nerthra turgidula: Todd (1955): 406–407 (original description repeated); Todd (1961a): 474 (catalogue, distribution); Todd (1961b): 93–94 (taxonomy, synonymy); Bal & Basu (2003): 542–544 (key, diagnosis, distribution, faunistics). Nerthra arunachalensis Thirumalai, 1998: 190–192 (description, key, figures, type locality), nov. syn. Holotype: male, India, Arunchal Pradesh, Julli village, Near Ganga, Itanagar, 6.viii.1991, D. B. Bastawade coll. (‘The type material is deposited at Arunchal Pradesh Field Station, Zoological Survey of India, Itanagar and will be transfered to National Collections of Zoological Survey of India, Calcutta’ [= NZSI] (Thirumalai 1998)). We did not examine the type.
Material examined. CHINA: NW Fujian, Wuyi Shan Mts., Kuatun, 27°40‘N 117°40‘E, 2300 m a.s.l., 19.iv.1938, 1 larva (instar 3), and 4.vi.1938, 1 male, J. Klapperich lgt. (NMPC); Kuatun, 18.vi.1946, 1 female; 24.vi.1946, 1 male; 25.vi.1946, 1 female; 27.vi.1946, 1 female; 28.vi.1946, 1 larva (instar 5); 2.vii.1946, 1 male; 15.vii.1946, 2 females, Tschung Sen lgt. (NMPC). S Yunnan, ‘30 km SW Tszin’pina [= Pingbian], 370 m a.s.l., 13.vi.1956, 1 male, D. Panfilov lgt. (NMPC); ‘okr. Cheli’ [= Yunjinghong env.], 700 m a.s.l., 20.iv.1957, 1 male, 1 female, D. Panfilov lgt. (NMPC); Yunnan, ‘okr. Binbyanya’ [= Pingbian env.], 700 m a.s.l., 29.vi.1956, 1 male, D. Panfilov lgt. (NMPC); ‘Daveyshan’, okr. Binbyanya’ [= Pingbian env., Daweishan Nature Reserve], 1350 m a.s.l., 23.vi.1956, 1 male, and 1500 m a.s.l., 22.vi.1956, 1 male, D. Panfilov lgt. (NMPC); Maguan [= Anping], 23°04‘N 104°25‘E, 1500–1600 m a.s.l., 25.–26.vi.1994, 2 males, 1 larva (instar 3), V. Kubáň lgt. (NMPC). INDIA: Meghalaya, Nokrek NP, 3 km S of Daribookgiri, 25°27‘N 90°19‘E, 1400 m a.s.l., 26.iv.1999, 1 female, Z. Košťál lgt. (ZJPC); 3 km E of Tura, 25°30‘N 90°14‘E, 500–1150 m a.s.l., 15.–22.v.1999, 1 female, Z. Košťál lgt. (ZJPC). Sikkim, Phodong village env., 3000 m a.s.l., 28.v.2002, 1 female, P. Benda lgt. (PBPC); Tankhola, Tista R[iver], 19.ii.1920, 1 female, H. Stevens lgt., E. L. Todd det. (BMNH); E Sikkim, Gantok [= Gangtok], 5000 ft, 6.v.1924, 1 female, R.W.G. Hingston lgt. (BMNH); Gangtok, 1.–4.v.1984, 1 male, A. Hamet lgt. (ZJPC); Gangtok env., Fambong-Lho forest, 2000–2500 m a.s.l., 8.–15.vii.1997, 1 female, J. Schneider lgt. (ZJPC); W Sikkim, Peeling env., 2100 m a.s.l., 18.–20.vii.1997, 1 female, J. Schneider lgt. (ZJPC). West Bengal, Darjeeling env., 2000–2400 m a.s.l., 4.–7.vii.1997, 1 male, J. Schneider lgt. (ZJPC); Kurseong [= Karsiyang], N.E. Himalayas, 5000 ft, 21.–29.v.[19]06, 1 male, N. Annandale lgt. (BMNH). LAOS: La Casade, 30 km from Xieng Khouang [= Xiangkhoang], 5.i.1918, 1 female, R.V. de Salvaza lgt. (BMNH); Hua Phan province, Phu Phan Mt., 20°12‘N 104°01‘E, 1500–1800 m a.s.l., 20.–31.v.2007, 1 male, local collector lgt. (NMPC); Louangnamtha province, Namtha → Muang Sing, 21°09‘N 101°19‘E, 900–1200 m a.s.l., 5.–31.v.1997, 1 male, 1 female, V. Kubáň lgt. (MMBC). NEPAL: Kimpti Kola, 2000 m a.s.l., 31.x.1961, 1 female, Gruber lgt. (NMPC). Narayani province, Chitwan district, Sauraha, 20.–25.v.1992, 1 male, I. Jeniš lgt. (ZJPC); Royal Chitwan National Park, 7.5 km of NW Sauraha, Rapti riverside, 27°34‘N 84°24‘E, 180 m a.s.l., 19.vi.2005, 1 female, M. Hartmann lgt. (NMEG); Sauraha SW, Rapti river, bank, 27°34‘80“N 84°29‘49“E, 18.iv.2000, 1 larva (instar 5), A. Weigel lgt. (NMEG). VIETNAM: Tonkin, Huong By, May 1916, 1 male, R.V. de
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 197
Salvaza lgt. (BMNH). Vinh Phuc province, Tam Dao, 3.–11.vi.1985, 1 male, 1 female, and 27.v.–2.vi.1986, 1 male, 1 larva (instar 4), V. Švihla lgt. (NMPC); Tam Dao, 890 m a.s.l., 18.v.2007, 1 male, 2 females, V. Socha lgt. (NMPC, VSPC). Variability of adults. Males (n = 20): For measurements see Table 1. In 17 males the pronotum was wider than maximal abdomen width (ratios (mm): 7.1 : 6.7, 7.0 : 6.6, 6.8 : 6.3, 6.7 : 6.3, 6.6 : 6.4, 6.6 : 6.3 (3x), 6.5 : 6.2, 6.5 : 6.1, 6.4 : 6.3, 6.4 : 6.2 (2x), 6.3 : 6.0, 6.3 : 5.9, 6.0 : 5.9, 5.8 : 5.6), while in three remaining males the ratio was equal (6.6 : 6.6, 6.3 : 6.3, 6.1 : 6.1). Females (n = 20): For measurements see Table 1. In eight females the pronotum was wider than maximal abdomen width (ratios 7.6 : 7.4, 7.5 : 7.4, 7.5 : 7.1, 7.4 : 7.3, 7.4 : 7.1, 7.2 : 7.1, 6.9 : 6.6, 6.8 : 6.7), in four females the ratio was equal (7.8 : 7.8, 7.6 : 7.6, 6.8 : 6.8, 6.7 : 6.7), and in eight females the pronotum was narrower than abdomen (ratios 7.5 : 7.6, 7.1 : 7.6, 7.0 : 7.1 (2x), 6.6 : 6.8, 6.6 : 6.7, 6.4 : 6.7, 6.4 : 6.6). Body brown, with variable yellowish, grayish or reddish colouring, often obscured by muddy crust. We found extreme morphological variability in N. indica, especially body sculpture, and outlines of the pronotum, hemelytra, and abdomen. In particular, we did not find two specimens with exactly the same shape of pronotum. Moreover, in 23 of 40 examined specimens the lateral margins of the pronotum were markedly asymmetrical (see Figs 2, 10–13). On the other hand, all males shared the same paramere shape (Figs 17–19). In all investigated females the outline of the ovipositor was the same and the ventral submarginal tumescences on the last visible abdominal sternite were absent. Description of 5th instar larva (Fig. 6). Body brown, thorax laterally and legs pale brown; body covered with sparse pale clavate setae; upper surface partly covered with muddy crust. Head with three tubercles in front of eyes (one high, conical on each side, one smaller, conical to flat medially); two low elevations on vertex between eyes. Pronotum flattened laterally, strongly elevated medially; lateral margins rounded and expanded laterally, deeply incised before anterior angles, bearing sparse clavate setae; posterior margin bisinuous (slightly convex laterally and medially, slightly concave submedially). Table 1. Nerthra asiatica (Horváth, 1892), N. indica (Atkinson, 1889), and N. nieuwenhuisi Todd, 1957 – measurements of adults.
N. asiatica female (n = 2) N. indica male (n = 20) female (n = 20) N. nieuwenhuisi male (n = 2)
Total body length (mm)
Head width (mm)
Interocular width (mm)
Pronotum length (medially) (mm)
Maximal pronotum width (mm)
Abdomen width (across connexivum) (mm)
11.3–11.6
4.3–4.6
2.5–2.6
3.1–3.2
7.3–8.0
8.2–8.5
8.5–9.4 9.2–10.7
3.7–4.3 3.9–4.4
1.8–2.1 1.8–2.2
2.5–2.9 2.6–3.3
5.8–7.1 6.4–7.8
5.6–6.7 6.6–7.8
10.2–10.5
4.2–4.3
1.9–2.0
2.7–3.0
7.0–7.2
6.6–6.7
198 P. Kment & Z. Jindra
Mesonotum laterally flattened, medially elevated and projected posteriorly, bearing three distinct tubercles (one apically, two subapically) and shallow depression among them. Mesonotal wing pads large, flattened, expanded laterally, reaching anterior margin of abdominal tergite III, bearing sparse row of pale clavate setae on lateral margin. Metanotum anteriorly covered by mesonotum, posterior margin straight, metanotal wing pads completely covered by mesonotal ones. Abdominal tergites convex, decreasing towards connexivum; connexivum with lateral margins slightly turned upwards, forming shallow
10
11
12
13
14
15
16
Figs 10–16: Variability of pronotum and hemelytra outline. 10–13. Nerthra indica (Atkinson, 1889): 10–11 – males from China, Maguan; 12 – female from India, Fambong Lho forest; 13 – female from India, 3 km E of Tura. 14 – N. arunachalensis Thirumalai, 1998, male, holotype (according to Thirumalai 1998). 15–16 – N. nieuwenhuisi Todd, 1957, males from Malaysia, Mt. Malang.
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 199
groove between tergites and connexivum; lateral margin of connexival segments rounded medially, bearing long clavate setae; connexival outline thus undulating. Tergite I medially projected posteriorly and elevated, bearing dense group of stout, black clavate setae on each side of the projection (in one specimen missing, probably rubbed off ); however, dorso-abdominal scent gland (DAG) ostioles not apparent. No trace of DAG ostioles observed on remaining tergites. Measurements: see Table 2. Description of 4th instar larva. Generally similar to instar 5, if not stated otherwise. Pronotum: posterior margin only slightly bisinuous. Mesonotum laterally flattened, medially sligthly elevated and projected posteriorly, without tubercles, but with shallow depression subapically; posterior margins slightly bisinuous; mesonotal wing pads not developed, however, lateral margins flattened and expanded laterally. Metanotum narrower than mesonotum, laterally flattened but not expanded, wing pads not developed. Tergite I medially projected posteriorly and elevated, dense group of stout, black clavate setae on each side of the projection not present (possibly damaged). Measurements: see Table 2. Description of 3rd instar larva (Fig. 7). Generally similar to instar 5, if not stated otherwise. Head with three small tubercles in front of eyes (one laterally on each side, one medially; the lateral ones not much higher than median one). Pronotum: posterior margin slightly convex. Mesonotum depressed laterally, lateral margins expanded and rounded; posterior margin almost straight, slightly convex medially. Metanotum narrower than mesonotum, laterally narrowly flattened, not expanded. Tergite I medially projected
Head width (mm)
Inter-ocular width (mm)
Head heigth (mm)
Thorax length (mm)
Pronotum length (mm)
Pronotum width (mm)
Mesonotum width (mm)
Abdomen length (mm)
Abdomen width (mm)
Metatibia length (mm)
N. ASIATICA 4th instar Sichuan: Jiulonggou N. INDICA 3rd instar Yunnan: Maguan Fujian: Kuatun 4th instar Vietnam: Tam Dao 5th instar Fujian: Kuatun Nepal: Sauraha
Total body length (mm)
Table 2. Nerthra asiatica (Horváth, 1892) and N. indica (Atkinson, 1889) – measurements of larvae.
7.1
3.3
1.6
1.6
3.3
1.8
5.2
5.5
3.2
5.3
2.5
4.5 4.7
2.4 2.5
1.1 1.1
1.2 1.2
2.0 2.2
1.3 1.4
3.7 3.8
3.8 4.0
2.1 2.2
3.2 3.7
1.7 1.3
6.1
3.1
1.4
1.6
2.9
1.9
5.0
5.2
2.6
4.8
2.4
7.4 8.1
3.4 3.6
1.6 1.8
1.7 2.1
3.5 3.6
2.1 2.3
5.6 6.1
5.9 6.4
3.1 3.5
5.7 6.2
3.1 3.3
200 P. Kment & Z. Jindra
posteriorly and elevated, dense group of stout, black clavate setae on each side of the projection not present (possibly damaged). Measurements: see Table 2. Differential diagnosis of larva. The studied larvae were found in association with adults of N. indica. They differs from larvae of N. asiatica in the following characters: head with three tubercles in front of eyes (one laterally on each side, one medially); pronotum strongly elevated medially, lateral margins expanded laterally, rounded, deeply incised before anterior angles; metanotum laterally expanded and rounded; lateral margins of connexival segments rounded medially, connexival outline undulating. Ecology. Maxwell-Lefroy (1909) characterized N. indica as follows: ‘it is found on grass paths, on the soil and under stones, as well as on hard roads; it is not aquatic and is possibly predaceous on small insects’. According to Todd (1961b) it was collected between 600–5000 ft [= 183–1524 m a.s.l.] in Darjiling. Our records from Sikkim and West Bengal extends its altitudinal limit to 3000 m a.s.l. Distribution. Bangladesh (Sylket [= Sylhet] – Todd (1955)). Burma (Lansbury 1988 – no exact record). China: Fujian (this paper), Guangdong (Nieser 1977; Todd 1977), Guangxi (Nieser & Chen 1992), Guizhou (Nieser & Chen 1992), Sichuan (Nieser & Chen 1992), ?Tibet (Nieser & Chen 1992), Yunnan (Nieser & Chen 1992). India: Arunachal Pradesh (Thirumalai 1998), Assam (Todd 1955, 1957; Nieser 1977; Lansbury 1988), Meghalaya (Paiva 1919; Bal & Basu 2003), Sikkim (Montandon 1899a; Todd 1957; Lansbury 1988; Bal & Basu 2003), Tamilnadu (Thirumalai 1998, this outlying record needs confirmation), West Bengal (Montandon 1899a; Todd 1961b). Laos (new record). Nepal (Distant 1906). Vietnam (Todd 1957 – Tonkin, no exact locality; here confirmed). Discussion. Nerthra indica is a highly variable species. Todd (1957) wrote: ‘The specimens from Tonkin differ slightly in the shape of the lateral margin of the pronotum which appears more like the margin of the pronotum of N. lobata (Montandon), but the absence of lateral tumescences on the last visible abdominal sternite of the female and shape of the clasper of the male reveal their relation to N. indica (Atkinson)’. Todd (1961b) gave the following size ranges: ‘Male: Length, 8.2 to 9.1 mm; width of pronotum, 5.5 to 6.7 mm; width of abdomen, 5.7 to 6.4 mm. Female: Length, 8.5 to 10.2 mm; width of pronotum, 6.4 to 7.8 mm; width of abdomen, 6.4 to 8.4 mm’. Further he stated: ‘The specimens under consideration verify the fact that both sexes are extremely variable in the relative widths of the pronotum and abdomen. In some instances the pronotum is wider, in others the abdomen is the wider. Correspondingly, the width of the connexivum is also variable. The species is laso variable in the shape of the lateral margin of the pronotum, but those presently studied have that part less irregular in shape than in those specimens previously studied and in a few instances the specimens from the Darjiling area agree with the illustration of Mononyx turgidulus Distant in the shape of the margin. Therefore, I have placed that name in the synonymy of indica’ (Todd 1961b). Lansbury (1988), redescribing the syntype series, presented the following measurements: ‘Males: 9–9.6 mm long; pronotal width 6.5 mm; abdominal
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 201
17
21
18
19
22
20 23
Figs 17–23: Parameres in different views. 17–19. Nerthra indica (Atkinson, 1889), male from Laos, Namtha → Muang Sing. 20 – N. arunachalensis Thirumalai, 1998, male, holotype (according to Thirumalai 1998). 21–23 – N. nieuwenhuisi Todd, 1957, male from Malaysia, Mt. Malang.
width 6.3–6.4 mm; head width 4–4.1 mm; width between eyes 2 mm. Females: 9.4–10 mm long; pronotal width 6.9–7.1 mm; abdominal width 6.8–7.0 mm; head width 4–4.1 mm; width between eyes 2–2.1 mm.’ Thirumalai (1998) gave measurements of one female from southern India, Tamilnadu: ‘Length 10.0 mm; width of pronotum 7.7 mm; width of abdomen 8.2 mm’. These above mentioned characters well correspond with the observations we made on our material. Thirumalai (1998) described a new species, N. arunachalensis, based on one male (holotype) and one female (paratype) from Arunachal Pradesh, northern India. In his key he distinguished N. arunachalensis and N. indica from N. asiatica and N. spissa by having the ‘lateral margin of pronotum broadly sinuous’. Both species differ in ‘lateral margin of pronotum projecting beyond the base of embolium’ in N. arunachalensis, and ‘lateral
202 P. Kment & Z. Jindra
margin of pronotum projecting not beyond the base of embolium’ in N. indica. Further he wrote: ‘N. arunachalensis is close to N. indica (Atkinson) and N. serrata (Montandon) in general appearance, but differs distinctly in the nature of pronotum [Fig. 14], ventral abdominal segments, and male paramere [Fig. 20], and other characters mentioned in the text’. He gave the following measurements of N. arunachalensis: ‘Male: Length, 8.3 mm., width of the pronotum, 6.2 mm., width of abdomen, 5.9 mm., Female: 9.4 mm., width of the pronotum, 7.1mm., width of abdomen, 6.9 mm. [sic!]’. Thirumalai (1998) listed only following references – Atkinson (1889), Distant (1906, 1911), and Todd (1955), being completely unaware of all papers pointing-out the variability of N. indica (Todd 1957, 1961b; Lansbury 1988; Nieser & Chen 1992), as well as the existence of another endemic Indian species N. unguistyla. We have compared the description, figures and measurements of N. arunachalensis given by Thirumalai (1998) with available series of N. indica and other published data on its variability (see above). This revealed, that N. arunachalensis fits in all aspects (including geographical distribution) within the range of variation of N. indica. On this basis we regard N. arunachalensis to be a junior subjective synonym of N. indica. The shape of male paramere has been represented in previous works as being variable, but we regard this as an artefact of the rendering of the illustrations (Todd 1959, 1961b; Lansbury 1988; Nieser & Chen 1992; Thirumalai 1998). However, we found that by rotating the extracted pygophore with the left paramere exposed, we reconcile our observations with those of the above mentioned authors (see also Figs 17–20).
Nerthra lobata (MONTANDON, 1899) (Figs 3, 9) Mononyx lobatus Montandon, 1899a: 394, 397–398. Syntypes: Indonesia, ‘Sumatra, Pangherang Pisang (E. Modigliani)’ (MCSN, coll. Montandon (currently in BMNH)) & ‘Java’ (NHMW). Nerthra lobata: Todd (1955): 350, 407–409, 466–467 (redescription, figures, key, catalogue, distribution, faunistics); Todd (1957): 153–154 (figure, distribution, faunistics); Todd (1961a): 469 (catalogue, distribution); Nieser (1977): 298 (faunistics); Chen et al. (2005): 414 (catalogue: Malesia, distribution); Buzzetti et al. (2006): 32 (distribution, faunistics).
Material examined. INDONESIA: West Sumatra, Gugul, 12 km W from Padangpanjang, iv.–v.1992, 3 females, local collector lgt. (NMPC, ZJPC). MALAYSIA: Pahang, Sungai Taban, 24.iv.1922, 1 female, H.M. Pendelbury lgt., coll. F.M.S. Museum, E.L. Todd det. (BMNH). Ecology. Unknown. Distribution (Fig. 9). Indonesia: Java (Montandon 1899a), Sumatra (Montandon 1899a; Todd 1955; Nieser 1977; Buzzetti et al. 2006). Malaysia: Pahang (Todd 1957).
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 203
Nerthra macrothorax (MONTROUZIER, 1855) (Fig. 24) Galgulus macrothorax Montrouzier, 1855: 110–111 (description, type localiy). Holotype (probable): Papua New Guinea, Woodlark Is. (MNHN) (see Andersen & Weir 2004; Polhemus (1995) gave the type material as syntype(s)). Scylaecus macrothorax: Stål (1861): 201 (generic placement); Stål (1876): 139 (catalogue, distribution). Peltopterus macrothorax: Stål (1863): 408 (catalogue, distribution); Stål (1865): 173 (comparative note); Stål (1871): 610, 706 (check-list: Philippines); Montandon (1899b): 779–780 (redescription, faunistics); Kirkaldy (1906): 150 (catalogue of genera); Esaki (1928): 75 (faunistics); Ohshima (1933): 410 (faunistics); Sonan (1934): 21–22 (faunistics); Esaki (1936): 43 (faunistics); Hoffmann (1941): 44–45 (catalogue, distribution); Miyamoto (1953): 35 (faunistics); Miyamoto (1954): 28 (faunistics). Nerthra macrothorax: Todd (1955): 348, 414–416, 468–473 (redescription, figures, key, catalogue, distribution, faunistics); Todd (1957): 157 (faunistics); Todd (1959): 63–64, 67, 69–71 (figures, key, comparative note, synonymy, catalogue, ecology, distribution, faunistics); Todd (1960a): 172, 176–177, 192 (figures, key, comparative note, synonymy, catalogue, distribution, faunistic); Todd (1960b): 116 (ecology, faunistics); Todd (1961a): 470 (cata-
Fig. 24: Distribution map of Nerthra macrothorax (Montrouzier, 1855) in south-east Asia and adjacent parts of Malesia and Australia.
204 P. Kment & Z. Jindra
logue, distribution); Popov (1971): 70 (figure); Polhemus (1976): 240 (ecology); Nieser (1977): 299 (faunistics); Todd (1977): 216 (faunistics); Miyamoto & Yasunaga (1989): 156 (check-list: Japan); Cassis & Gross (1995): 86–87 (catalogue: Australia, distribution, faunistics); Polhemus (1995): 24 (catalogue, distribution); Nieser & Chen (1999): 94 (key, distribution); Hua (2000): 214 (check-list: Taiwan); Andersen & Weir (2004): 273, 275, 279–280, 336 (key, figures, distribution); Chen et al. (2005): 414 (catalogue: Malesia, distribution); Nieser & Chen (2005): 308 (diagnosis, ecology, distribution, faunistics); Cassis & Silveira (2006): 145 (distribution, zoogeography); Evenhuis & Polhemus (2006): 4 (check-list: Fiji).
Ecology. According to Todd (1959) it ‘has been found burrowing in rotten Pandanus logs. This activity may be important in the distribution of the species as such debris could easily drift from one island to another through the action of storms and ocean currents’. Todd (1960b) reported additional findings of this species ‘under Pandanus and decaying leaves of Erythrina indica near the sea shore’ on Philippine Islands of Cebu and Negros. Nieser & Chen (2005) observed these toad bugs burrowing in the sand on a beach in the south of Taiwan. The possible transmarine transport with plant debris is a reasonable explanation for its wide distribution throughout the IndoPacific region (e.g., Polhemus 1976). Distribution (Fig. 24). Widely distributed across Indian and Pacific Oceans. South-east Asia: China: Taiwan (Esaki 1928; Sonan 1934; Hua 2000; Nieser & Chen 2005), Lu Dao Island (= Green Island, Kasho To Island) (Todd 1957), Lan Yu Island (= Orchid Island, Koto Sho Island) (Todd 1957). Indonesia: East Kalimantan: Maratua Island (Todd 1957). Japan: Ryukyu Islands: Yaeyama Island, Kikai-Shima Island, Tokara Group (Takajimi Island); Kyushu: Osumi; Shikoku: Satano Misaki [= Ashizuri-misaki] (Esaki 1928, 1933, 1936; Ohshima 1933; Miyamoto 1953, 1954). ?Malaysia: Sabah (North Borneo – Montandon 1899b). Further distribution: Comoro Islands: Mayotte Island; ?Tanzania; Indonesia: Aru Island, Buru, Flores, Halmahera, Irian Jaya, Kei Islands [= Kepulauan Kai], Postiljon [= Sablana] Islands, Sulawesi; Philippines: Biliran, Cebu, Leyte, Negros, Panay; Mariana Islands; Caroline Islands; Australia: Northern Territory; Papua New Guinea: incl. New Britain and other adjacent islands; Solomon Islands; Fiji; Tonga; Samoa (e.g., Montandon 1899b; Esaki 1928; Todd 1955, 1957, 1959, 1960a, 1960b, 1961a; Nieser 1977; Cassis & Gross 1995).
Nerthra nieuwenhuisi TODD, 1957 (Figs 4, 9, 15–16, 21–23) Nerthra nieuwenhuisi Todd, 1957: 153–155. Holotype: female, Indonesia, ‘Borneo, Boven, (upper) Mahakkam River, 1894, Borneo Exped., Dr. Nieuwenhuis’ (RMNH). Nerthra nieuwenhuisi: Todd (1961a): 471 (catalogue, distribution); Chen et al. (2005): 414 (catalogue: Malesia, distribution).
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 205
Material examined. MALAYSIA: Sarawak, Mt. Malang, 2500 ft, 1 male, B. Mjoberg lgt., Ex F.M.S. Museum, B.M. 1955–354 (BMNH); Malang, April 1904, 1 male, Ex F.M.S. Museum, B.M. 1955–354 (BMNH). Description of male (Fig. 4). Colouration: Ground colour orange-brown, scutellum, median elevation on anterior part of pronotum and head between eyes with blackish spots, abdominal sternites (except their orange-brown outer margins), apices of femora, entire tibiae and tarsi blackish; membrane of the same colour as rest of hemelytra, or slightly darker. The structure of the two examined males fits well to description of female by Todd (1957). We found the following structural differencies: lateral margins of pronotum variable, from almost regularly rounded (Fig. 15) to median and posterior portions nearly straight (Fig. 16); lateral margin of embolium only slightly expanded laterally, regularly rounded (Figs 4, 15–16); and connexivum less prominent than in female, only five abdominal segments visible. Paramere as in Figs 21–23. Measurements (n = 2): Body length 10.2 / 10.5 mm; head width 4.2 / 4.3 mm; inter-ocular width 1.9 / 2.0 mm; head heigth 2.4 / 2.5 mm; pronotum length (medially) 2.7 / 3.0 mm; (maximal) pronotum width 7.2 / 7.0 mm; (maximal) abdomen width (across connexivum) 6.7 / 6.6 mm, respectively (see also Table 1). Pronotum slightly wider than abdomen (ratios 7.2 : 6.7 and 7.0 : 6.6). Differential diagnosis. This species was previously known only from the female holotype (Todd 1957). Todd (1957: 155) wrote: ‘It is slightly larger than N. asiatica (Horváth), from which it may be easily separated by the dilated margin of the embolium, distinctive lateral margin of the pronotum, and proportionally longer hind legs. The size, shape of the lateral dilation of the embolium, and the shape of the lateral margin of the pronotum will separate this species from the other species of the grandicollis group.’ According to Todd (1957: 156) N. nieuwenhuisi is ‘very closely related’ to N. eximia ‘and may subsequently prove to be but a form of that species’. This species is differentiated fom other south-eastern Asian species of the N. grandicollis-group, as follows: i) from N. asiatica it differs in the shape of pronotum, outer margin of embolium laterally expanded, and the number and shape of tubercles on the head; ii) from N. spissa and N. unguistyla it mainly differs in the shape of the left paramere, and iii) from N. unguistyla in its welldeveloped hemelytral membrane. According to external characters and the shape of the paramere (if known), N. nieuwenhuisi is most similar to N. indica, N. lobata, N. serrata, and N. eximia. From these species it differs in the combination of the following characters: body orange-brown, apices of femora, entire tibiae and tarsi blackish; a combination not observed in any other studied Nerthra species from south-eastern Asia. Furthermore, N. nieuwenhuisi is similar to N. eximia in that the scutellum is strongly elevated, distinguished from N. indica in which the scutellum is moderately elevated, and from N. lobata, wherer the scutellum is distinctly depressed basally. Nerthra nieuwenhuisi can be seperated from the other species by the less expanded lateral margin of embolium, the shape of lateral margin of pronotum, and abscence of lateral submarginal tumescences ventrally on the last visible abdominal sternite of female. The latter, are well developed in N. lobata, and
206 P. Kment & Z. Jindra
are also absent in N. indica and N. eximia, and by the shape of the left paramere. The left paramere (Figs 21–23) is very similar to that of N. indica (Figs 17–19) and N. lobata (see Todd (1955): 467), but differs from them being slightly shorter and wider basally. The female of N. nieuwenhuisi should be distinguishable also by its large size (holotype: body length 12.5 mm, pronotum width 8.5 mm, abdomen width 8.3 mm) (see Todd 1957). Ecology. Unknown. Distribution (Fig. 9). Indonesia: Kalimantan Timur (Todd 1957). Malaysia: Sarawak (new record).
Nerthra serrata (MONTANDON, 1897) (Fig. 9) Mononyx serratus Montandon, 1897: 365–366 (description, type localiy). Syntypes (both sexes): Burma, ‘Carin Chebà, Carin Ghecù’ [‘Type localities Carin Ghecú and Carin Cheba are now known to be in that section of Burma between the Salwin and Sittang rivers, east and northeast of Toungoo.’ (Todd 1957)] (MCSN, ZMUH, coll. Montandon (? currently in BMNH)). Mononyx serratus: Montandon (1899a): 394, 396 (key); Distant (1906): 15 (redescription, figure, faunistics). Nerthra serrata: Todd (1955): 408–410 (original description repeated, comparative note); Todd (1957): 152 (note on type localities); Todd (1961a): 473 (catalogue, distribution); Thirumalai (1998): 192 (comparative note).
Ecology. Unknown. Distribution (Fig. 9). Burma (Montandon 1897; Distant 1906; Todd 1957). Discussion. Montandon (1899a) gave an extensive description of the species; however, he did not compare it with any other species, stating only : ‘La forme très dilatée du pronotum, le lobe de la marge élytrale et les sinuosités du connexivum permettront de reconnaître facilement cette espèce.’ [= Very enlarged shape of pronotum, lobe of the elytral margin, and the sinuosity of connexivum enable an easy identification of this species]. Distant (1906) diagnosed the species and provided a habitus figure of a syntype (probably male). Todd (1955) repeated the original description in verbatim with statement: ‘The writer has not seen this species or at least has not been able to recognize it’. Nevertheless, Todd (1955: 408–409) discussed the status of N. serrata as follows: ‘It may be that N. serrata (Montandon) will prove to be this species [= N. lobata] in which case Nerthra lobata (Montandon) would fall as a synonym since it was described two years after the former species. Doctor R.L. Usinger examined the clasper of the type of N. serrata (Montandon) and is of the opinion that it is the same as my drawing of the clasper of N. lobata (Montandon). Montandon’s description of N. serrata states that the pronotum laterally is very dilated and that the segments of the connexivum are concave. None of the specimens I have examined are of this nature. For that reason and because both species were described by Montandon, this writer prefers to retain both species
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 207
until more specimens can be examined.’ (Todd 1955). So far, the types of N. serrata have never been reexamined and redescribed, and identity of the species is unclear.
Nerthra spissa (DISTANT, 1911) (Fig. 8) Mononyx spissus Distant, 1911: 312–313 (description, figure, type locality). Lectotype: male, India, Assam, ‘Sibs. [= Sibsagar] 6257’, ‘Distant coll 1911–383’ (BMNH). Designated by Nieser & Chen (1992). Mononyx spissus: Chakrabarty et al. (1994): 2, 24–25 (diagnosis, faunistics). Nerthra spissa: Todd (1955): 350, 411–412, 466–467 (redescription, figure, key, catalogue, distribution, faunistics); Todd (1961a): 473 (catalogue, distribution); Nieser & Chen (1992): 6 (lectotype designation, figure); Thirumalai (1998): 190, 192 (key, faunistics).
Ecology. Unknown. Distribution (Fig. 8). India: Arunachal Pradesh (Chakrabarty et al. 1994; Thirumalai 1998), Assam (Todd 1955; Nieser & Chen 1992). Discussion. This species is so far known from the males only. For further comments on its status see Discussion under N. asiatica.
Nerthra unguistyla TODD, 1957 (Figs 8, 25) Nerthra unguistyla Todd, 1957: 152–154. Holotype: male, South India, Mayavaram, 8.x.1945, P.S. Nathan lgt. ( J.C. Lutz collection, currently in USNM). Nerthra unguistyla: Todd (1961a): 474 (catalogue, distribution); Todd (1977): 217 (faunistics).
Material examined. INDIA: Tamilnadu, Coimbatore, ii.1950, 1 male, 1 female; 1.iv.1950, 1 female; iv.1950, 1 female, P.S. Nathan lgt. (NMPC). Ecology. Unknown. Distribution (Fig. 8). India: Pondicherry (Todd 1977), Tamilnadu (Todd 1957). Discussion. This species diff ers substantially from all remaining species of N. grandicollis species-group lacking the hemelytral membranes (Todd 1957).
Fig. 25: Habitus of Nerthra unguistyla Todd, 1957, female from India, Coimbatore (10,4 mm length) (Photo: Jan Macek).
208 P. Kment & Z. Jindra
[Nerthra grandicollis (GERMAR, 1837)] Mononyx grandicallis [sic!]: Hua (2000): 214 (checklist: China – Hongkong).
Discussion. Afrotropical species (e.g., Todd 1959, 1961a). The record from Hongkong (Hua 2000) is apparently erroneous.
DISCUSSION At present, nine species of Nerhra are known from south-eastern Asia west of Wallace’s line. Todd (1955) synonymized all previously described genera of Nerthrinae with Nerthra, establishing eight informal species-groups. To this, Cassis & Silveira (2004) add a ninth species-group to accomodate the endemic Western Australian species N. tuberculata Montandon 1899. Two of these species-groups occur also in south-eastern Asia, the N. rugosa (Desjardins, 1837) species-group and the N. grandicollis species-group. The N. rugosa species-group, comprised of four species, is distributed in coastal areas of tropical and subtropical regions (especially in eastern Pacific) (see Todd 1955; Cassis & Silveira 2006), is represented by single species – N. macrothorax. The remaining eight species belong to the N. grandicollis species-group, including nine species: N. grandicollis (widely distributed in Afrotropical region and Madagascar), N. asiatica (south-eastern China, northern India), N. eximia (Sumatra), N. indica (widely distributed from south-eastern China to southern India, Laos, and Vietnam), N. lobata ( Java, Sumatra, Malayan Peninsula), N. nieuwenhuisi (Borneo), N. serrata (Burma), N. spissa (northern India), and N. unguistyla (southern India). Most families of Nepomorpha and Gerromorpha are species rich in south-eastern Asia, with many newly described species every year (see e.g. Chen et al. 2005). In contrast, the diversity of Gelastocoridae in this region is apparently low. Taking the possible synonymies of N. spissa with N. asiatica and N. serrata with either N. lobata or N. indica into account, the number of described species from south-eastern Asia is less diverse, and we have found no evidence of new species. However collections of toad bugs from southeast Asia are limited, and additional survey, especially from Burma, Thailand, Cambodia, Malayan Peninsula, Borneo, Sumatra, and southern India, are needed to determine this hypothesis. The larval morphology of Gelastocoridae is poorly studied and the identity of the larvae of most species remains unkown. So far, all five instars of only two species were described in detail and illustrated – Nearctic Gelastocoris oculatus oculatus (Fabricius, 1798) (Hungerford 1919, 1922; Brown & McPherson 1994) and Neotropical Nerthra ranina (Herrich-Schaeffer, 1853) (Estévez & Schnack 1978). Further, only Melin (1929) shortly described and illustrated larvae of two species of Gelastocoris and four species of Nerthra from South America, and Cassis & Silveira (2001) gave descriptions of larvae of five Australian species belongig to the N. alaticollis (Stål, 1854) species-group.
Review of the family Gelastocoridae (Heteroptera: Nepomorpha) of south-eastern Asia 209
ACKNOWLEDGEMENTS We are very obliged to Mick Webb (BMNH, London, United Kingdom), Igor Malenovský (MMBC, Brno, Czech Republic), Petr Baňař (Brno, Czech Republic), and Vít Socha (Pardubice, Czech Republic) for loan or gift of the material. Jan Macek (National Museum, Praha, Czech Republic) kindly made the photographs, and Denise Wyniger (NaturMuseum Luzern, Switzerland) and Gerry Cassis (University of New South Wales, Sydney, Australia) made some important comments on an early stage of the manuscript.
РЕЗЮМЕ Направена е ревизия на 9 вида от род Nerthra SAY, 1832 (Hemiptera: Heteroptera: Gelastocoridae: Nerthrinae), срещащи се в Югоизточна Азия, западно от Уолесовата линия. Описани са мъжките на N. nieuwenhuisi TODD, 1957, четвърта възраст ларва на N. asiatica (HORVÁTH 1892), трета, четвърта и пета възраст ларви на N. indica (ATKINSON, 1889), и са илюстрирани за първи път. Предложена е нова синонимия на Nerthra indica (ATKINSON, 1889) = N. arunachalensis THIRUMALAI, 1998 и накратко е обсъдена изменчивостта на N. indica. За първи път са съобщени с точни находища на N. nieuwenhuisi от Саравак (Малайзия), и N. indica от Фудзян (Китай), Лаос и Виетнам.
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Hungerford H.B. (1919). The biology and ecology of aquatic and semiaquatic Hemiptera. — Kansas University Science Bulletin, 11: i–iii (colour pls.) + 3–328 + 337–341 (index). Hungerford H.B. (1922). The life history of the toad bug Gelastocoris oculatus Fabr. (Gelastocoridæ). — Kansas University Science Bulletin, 14: 145–171. Kevan D.K.M. (1942). Some observations on Mononyx nepaeformis (Fabricius 1775), a toad-bug (Mononychidae, Hemipt.-Heteropt.). — Proceedings of the Royal Entomological Society of London, Series A, 17: 109–110. Kiritshenko A.N. (1926). Beiträge zur Kenntnis palaearktischer Hemipteren. — Konowia, 5: 218–226. Kiritshenko A.N. (1930). Nauchnye rezul’taty Entomologicheskikh ekspeditsiy Zoologicheskogo Muzeya v Ussuriyskiy kray. IV. Hemiptera cryptocerata. (Résultate scientifiques des Expéditions entomologiques du Musée Zoologique dans la région d’Oussouri. IV. Hemiptera cryptocerata). — Ezhegodnik Zoologicheskogo Muzeya Akademii Nauk SSSR, 1930: 431–440 (in Russian, Latin diagnoses of new species and French title). Kirkaldy G.W. (1906). List of the genera of the pagiopodous Hemiptera-Heteroptera, with their type species, from 1758 to 1904 (and also of the aquatic and semi-aquatic Trochalopoda). — Transactions of the American Entomological Society, 32: 117–156, 156a, 156b. Kment P. (in prep.). Catalogue of the toad bugs (Heteroptera: Gelastocoridae) of the World. Lansbury I. (1988). Gelastocoridae (Hem.-Het.) in the Hope Entomological Collections, University Museum, Oxford. — Entomologist’s Monthly Magazine, 124: 243–250. López Ruf M., P. Pérez Goodwyn & R.G. Martins-Neto (2005). New Heteroptera (Insecta) from the Santana Formation, Lower Creatceous (Northeastern Brazil), with description of a new family and new taxa of Naucoridae and Gelastocoridae. — Gaea, 1: 68–74. Maxwell-Lefroy H. (1909). Indian Insect Life. — Thacker, Spink & Co., Calcuuta and Simla: 1–786. [Not seen, fide Todd (1955)]. Melin D. (1929). Hemiptera from South and Central America. I. (Revision of the genus Gelastocoris and the American species of Mononyx). — Zoologiska Bidrag (Uppsala), 12: 151–194. Miyamoto S. (1953). [Tokara expedition]. — Nymph, 2: 33–41 (In Japanese). [Cited according to Todd (1957, 1961a)]. Miyamoto S. (1954). [Collecting records from Tokara Islands]. — Shin Konchû, 7(1–2): 28–34 + 4 pls (In Japanese). [Cited according to Todd (1957, 1961a)]. Miyamoto S. & T. Yasunaga (1989). Hemiptera-Heteroptera. — In: Hirashima Y. (Ed.): Nihon-san Konchu Somokuroku. (A Checklist of Japanese Insects). Vol. 1. Kyushu University Entomological Laboratory, Fukuoka: 151–188. (In Japanese). Montandon A.L. (1897): Viaggio di Leonardo Fea in Birmania e regioni vicine. LXXV. Hemiptera Cyrptocerata. — Annali del Museo Civico di Storia Naturale di Genova (2), 17: 365–377. Montandon A.L. (1899a): Hemiptera Cryptocerata. S. fam. Mononychinae. Notes et descriptions d’espèces nouvelles. I-ére partie. —Buletinul Societaţii de Sciinţe din Bucureşti-Româna, 8: 392–407. Montandon A.L. (1899b): Hemiptera Cryptocerata. S. fam. Mononychinae. Notes et descriptions d’espèces nouvelles. 2-ére partie. —Buletinul Societaţii de Sciinţe din Bucureşti-Româna, 8: 774–780. Montrouzier P. (1855). Essai sur la fauna de l’île de Woodlark ou Moiou. — Annales de la Société Linnéene de Lyon (2), 7: 1–114. [Not seen, fide Todd (1955)].
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Nieser N. (1975). The water bugs (Heteroptera: Nepomorpha) of the Guyana Region. — Studies on the Fauna of Suriname and Other Guyanas, 16: 1–310 + 24 pls. Nieser N. (1977). Gelastocoridae in the Zoologisches Museum der Humboldt-Universität zu Berlin (Heteroptera). — Deutsche Entomologische Zeitschrift (N. F.), 24: 293–303. Nieser N. & P.-P.Chen (1992). Notes on Gelastocoridae and Ochteridae (Heteroptera) with the description of five new species. — Storkia, 1: 2–13. Nieser N. & P.-P.Chen (1999). Sixteen new species of Nepomorpha (Heteroptera) mainly from Sulawesi (Indonesia). — Tijdschrift voor Entomologie, 142: 77–123. Nieser N. & P.-P.Chen (2005). The water bugs (Hemiptera: Nepomorpha and Gerromorpha) of Vanuatu. — Tijdschrift voor Entomologie, 148: 307–327. Ohshima H. (1933). [Collecting trip to the Yaeyama and Miyako Islands]. — Botany and Zoology, Theoretical and Applied, 1(4): 529–539 (In Japanese). [Cited according to Todd (1957, 1961a)]. Oshanin B. (1909). Verzeichnis des Palaearktischen Hemipteren mit besonderer Brücksichtigung ihrer Verteilung im Russischen Reiche. I. Band. Heteroptera. III. Lieferung. Ende des Bandes. — Ezhegodnik Zoologicheskago Muzeya Imperatorskoy Akademii Nauk, 14 (Prilozhenie): 587–1087. Oshanin B. (1912). Katalog der paläarktischen Hemipteren (Heteroptera, HomopteraAuchenorrhynchaund Psylloidea). — R. Friedländer & Sohn, Berlin: 1–187. Paiva C.A. (1919). Rhynchota from the Garo Hills, Assam. — Records of the Indian Museum, 16: 349–377 + pls. xxxiv–xxxvi. Polhemus J.T. (1976): Water-boatmen of saline waters (Hemiptera: Corixidae). — In: Cheng L. (Ed.): Marine insects. North-Holland Publishing Company, Amsterdam – Oxford & American Elsevier Publishing Company, New York: 263–289. Polhemus J.T. (1995). Family Gelastocoridae Kirkaldy, 1897 – toad bugs. — In: Aukema B. & Ch. Rieger (Eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1. Enticocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam: 23–25. Popov Yu.A. (1971). Istoricheskoe razvitie poluzhestkokrylykh infraotryada Nepomorpha (Heteroptera). [Historical development of the hemipterous infraorder Nepomorpha (Heteroptera)]. — Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSSR, 129: 1–230 + 9 pls (In Russian). Schnack J.A. & A.L. Estévez (1979). Monografia sistematica de los Gelastocoridae de la Republica Argentina (Hemiptera). — Limnobios, 1(9): 346–380. Sonan J. (1934). [On Peltopterus macrothorax (Montrouzier)]. — Transactions of the Natural History Society of Formosa, 24: 21–22 (In Japanese). [Cited according to (Todd 1957, 1961a)]. Stål C. (1861). Nova methodus familias quasdam Hemipterorum disponendi. — Öfversigt af Kungliga Vetenskapsakademiens Förhandlingar, 18(4): 195–212. Stål C. (1863). Verzeichniss der Mononychiden. — Berliner Entomologische Zeitschrift, 7: 405–408. Stål C. (1865). Hemiptera Africana. Tomus tertius. — Officina Norstedtiana, Holmiæ: 1–276. Stål C. (1871). Hemiptera insularum Philippinarum. – Bidrag till Philippinska öarnes Hemipter-fauna. — Öfversigt af Kungliga Vetenskapsakademiens Förhandlingar, 27(7) (1870): 607–776 (In Swedish and Latin). Stål C. (1876). Enumeratio Hemipterorum. Bidrag till en förteckning öfver alla hittills kända Hemiptera, jemte systematiska meddelanden. 5. — Kongliga Svenska Vetenskapsakademiens Handlingar, 14(4) (1875): 1–162.
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Štys P. & A. Jansson (1988). Check-list of recent family-group and genus-group names of Nepomorpha (Heteroptera) of the world. — Acta Entomologica Fennica, 40: 1–44. Thirumalai G. (1998). The genus Nerthra (Say) from India with a description of a new species from Arunachal Pradesh, North Eastern India (Gelastocoridae: Hemiptera: Insecta). — Records of the Zoological Survey of India, 96 (1997): 189–193. Todd E.L. (1955). A taxonomic revision of the family Gelastocoridae (Hemiptera). — University of Kansas Science Bulletin, 37: 277–475. Todd E.L. (1957). Five new species of Gelastocoridae with comments on other species (Hemiptera). — Proceedings of the Entomological Society of Washington, 59: 145–162. Todd E.L. (1959). The Gelastocoridae of Melanesia. — Nova Guinea, New Series, 10: 61–95. Todd E.L. (1960a). The Gelastocoridae of Australia (Hemiptera). — Pacific Insects, 2: 171–194. Todd E.L. (1960b). Notes on Nerthra macrothorax (Montrouzier) (Hemiptera: Gelastocoridae). — Proceedings of the Entomological Society of Washington, 62: 116. Todd E.L. (1961a). A checklist of the Gelastocoridae (Hemiptera). — Proceedings of the Hawaiian Entomological Society, 17: 461–476. Todd E.L. (1961b). Notes on some toad bugs (Hemiptera: Gelastocoridae) from India. — Proceedings of the Biological Society of Washington, 74: 93–94. Todd E.L. (1977). Distributional and nomenclatural notes on some Gelastocoridae in the collection of the Bernice P. Bishop Museum (Hemiptera). — Proceedings of the Biological Society of Washington, 90: 214–217. Wu Ch.-F. (1935). Family Gelastocoridae.— In: Wu Ch.-F.: Catalogus Insectorum Sinensium. Volume II. Fan Memorial Institute of Biology, Peiping: 559.
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Grozeva & N.ofSimov (Eds)to2008 Review of Omocoris Lindberg, 1930 S. and a description a new genus accommodate Eurycolpus dimorphus... 215 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 215-230. © Pensoft Publishers Sofia–Moscow
Review of Omocoris LINDBERG, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus WAGNER 1961 (Heteroptera: Miridae: Phylinae) F.V. Konstantinov Department of Entomology, Faculty of Biology and Soil Sciences, St. Petersburg State University, Universitetskaya nab. 7/9, St. Petersburg 199034, Russia. E-mail:
[email protected]
ABSTRACT The genus Omocoris Lindberg is revised and O. unicolor nov. sp. is described from Uzbekistan. A new genus, Josifovius, is erected to accommodate the type species Eurycolpus dimorphus Wagner, 1961. Omocoris erythrophtalmus Carapezza, 1997 is considered as a species incertae sedis. Illustrations of the male and female genitalia, tarsus and pretarsus, photographs of the dorsal habitus, hosts and distributional records of discussed species are provided. Keywords: Heteroptera, Miridae, Phylini, Omocoris, Josifovius, new genus, new combination, distribution.
INTRODUCTION Omocoris Lindberg, 1930 is a small genus currently containing three species known from South Mediterranean or Central Asian regions. The present paper provides revised generic assignments for the species currently placed in the genus and a description of a new species found in the collection of Zoological Institute, Russian Academy of Sciences. A revised diagnosis and redescriptions for the genus and known species are given. Examination of O. dimorphus (Wagner, 1961) concludes that a new genus is required to accommodate this species properly.
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Bar code labels were attached to the specimens and are referred to as unique specimen identifiers (USIs). Generally each USI label corresponds to a single specimen; however, some USI labels correspond to two or three specimens if several specimens are mounted on one pin. As a way of accessing additional information, such as color photographs, specimens dissected, notes, collecting method, and specimens photographed for specimens examined in the Planetary Biodiversity Inventories Project on Plant Bugs and the present paper please refer to the www.discoverlife.org website. The original locality data are given in square brackets, if different from the currently existing toponyms (see specimens examined). All measurements are in millimeters (see Table 1). All scale bars are 0.05 mm. All specimens examined in the course of this study, including types, are retained at the Zoological Institute, St. Petersburg. This paper is dedicated to Michail Josifov in recognition of his inestimable contribution to our knowledge of the Palearctic Heteroptera, particularly the Miridae.
TAXONOMY Omocoris LINDBERG, 1930 Omocoris Lindberg, 1930: 20-21 (nov. gen.). Type species by original designation: Omocoris parallelus Lindberg, 1930 (= Oncotylus cunealis Reuter, 1904; placed by Carvalho (1952: 62) in synonymy with Eurycolpus; restored by Kerzhner & Muminov 1964: 46-47). Oncotylista Kiritshenko, 1959: 108-109 (nom. nud.). Omocoris: Wagner, 1975: 267-269 (description and key to spp.).
Diagnosis: Recognized by the strong sexual dimorphism, viz. the male being long, slender, and parallel-sided (Figs 1, 3), while the female is broadly ovate, strongly brachypterous, with the fused corium and clavus, absent cuneus and membrane, and apex of hemelytron reaching at most posterior margin of abdominal tergum III (Figs 2, 4). Antennal segment I long, nearly equal in length to pronotum, at least twice as long as interocular distance in both sexes. Dorsum with simple dark setae; tibial spines dark. Sensory lobe of the left paramere with flattened, obtuse, apically bifid apex (Figs 23, 24). Vesica with subapical secondary gonopore, two straight apical blades, and more or less developed series of spicules located between secondary gonopore and base of the blades (Figs 7-10). Most similar among the Palearctic phyline genera to Eurycolpus Reuter, 1875 and Oncotylus Fieber, 1858 in the structure of the pretarsus with a comparatively large pulvillus which is apically unattached to the claw, the general appearance, coloration, body proportions of the male, and the elongate head. Separated from Oncotylus by the strongly brachypterous female, the distinctly concave pronotal margins in the male, long antennal segment I, comparatively short pulvillus, and the different structure of the left paramere
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 217
and vesica. The male of O. viridiflavus (Goeze, 1778) is similar to the male of Omocoris species in the long, slender and parallel-sided body with thin, long legs, somewhat concave pronotal margins, and the comparatively long antennal segment I. However, the female of this species as well as all its congeners are macropterous; additionally the male has a single-coned vesica, and the left paramere has a long and smoothly tapering sensory lobe. Eurycolpus differs from Omocoris in the presence of a subapical notch on the claw
Table 1. Measurements, mm Species Body O. cunealis ♂ (N=5)
CunClyp
Length Pronotum
AntSeg1
AntSeg2
Head
Width Pronotum
InterOcDi
Mean SD Range Min Max Mean SD Range Min Max
6.40 0.38 1.00 5.80 6.80 5.34 0.38 0.80 5.00 5.80
5.58 0.37 0.87 4.98 5.85 2.92 0.21 0.48 2.70 3.18
0.76 0.05 0.13 0.70 0.83 0.76 0.04 0.10 0.70 0.80
0.86 0.08 0.15 0.78 0.93 0.94 0.02 0.05 0.90 0.95
2.09 0.17 0.40 1.90 2.30 1.97 0.11 0.23 1.88 2.10
1.02 0.03 0.07 0.98 1.05 1.10 0.07 0.18 1.05 1.23
1.52 0.05 0.12 1.48 1.60 1.26 0.06 0.13 1.20 1.33
0.37 0.02 0.04 0.35 0.39 0.45 0.02 0.04 0.43 0.46
Mean SD Range Min Max Mean SD Range Min Max
5.48 0.16 0.40 5.30 5.70 4.78 0.24 0.60 4.50 5.10
4.74 0.14 0.33 4.58 4.90 0.44 0.25 0.58 0.00 0.58
0.69 0.05 0.13 0.60 0.73 0.71 0.03 0.08 0.68 0.75
0.65 0.04 0.08 0.63 0.70 0.68 0.05 0.13 0.63 0.75
1.65 0.10 0.25 1.55 1.80 1.54 0.10 0.25 1.38 1.63
0.98 0.02 0.05 0.95 1.00 1.04 0.01 0.03 1.03 1.05
1.35 0.09 0.20 1.25 1.45 1.18 0.02 0.05 1.15 1.20
0.36 0.02 0.04 0.34 0.38 0.43 0.02 0.05 0.40 0.45
J. dimorphus ♂ (N=2) Mean Range Min Max ♀ (N=2) Mean Range Min Max
4.60 0.20 4.50 4.70 3.65 0.30 3.50 3.80
4.10 0.20 4.00 4.20 2.83 0.25 2.70 2.95
0.53 0.00 0.53 0.53 0.50 0.05 0.48 0.53
0.45 0.00 0.45 0.45 0.35 0.00 0.35 0.35
1.58 0.00 1.58 1.58 1.10 0.05 1.08 1.13
0.78 0.01 0.78 0.79 0.76 0.08 0.73 0.80
1.08 0.00 1.08 1.08 1.01 0.08 0.98 1.05
0.29 0.00 0.29 0.29 0.38 0.00 0.38 0.38
♀ (N=5)
O. unicolor ♂ (N=5)
♀ (N=5)
218 F.V. Konstantinov
(Figs 31-32), the macropterous female, the mixed vestiture composed of silver and dark setae, antennal segment I shorter than interocular distance, the thin and apically pointed sensory lobe of the left paramere, and the vesica strongly twisted with a very long and thin single apical blade, and secondary gonopore distant from the apex. Description: Male: COLORATION (Figs 1, 3): Dorsum and venter pale, greenish yellow; labium somewhat darkened apically; all femora densely covered with minute rounded pale brown spots, rarely uniformly pale; tibiae always uniformly pale; cuneus pale or reddish. SURFACE AND VESTITURE: Dorsum smooth, shining; hemelytra weakly granulate; entire dorsum with dense, dark, adpressed to semierect, simple setae; venter covered with reclining pale simple setae and sparse dark setae; all legs and antennae with dense, dark, semiadpressed, simple setae half as long as those on hemelytra; tibial spines dark; apex of antennal segment I and femora with long dark spine-like setae somewhat
1
3
2
4
5
6
Figs 1-4: Dorsal habitus of Omocoris and Josifovius spp.: 1 – 2, O. cunealis: 1 – ♂ (AMNH_PBI 00149570), 2 – ♀ (AMNH_PBI 00149626); 3-4, O. unicolor: 3 – ♂ (AMNH_PBI 00149580), 4 – ♀ (AMNH_PBI 00149626); 5-6, J. dimorphus: 5 – ♂ (AMNH_PBI 00149574), 6 – ♀ (AMNH_PBI 00149576).
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 219
thinner than tibial spines; spine-like setae on femora sparse, mainly distributed along fore margin of fore femora, hind margin and apices of middle and hind femora. STRUCTURE: Body strongly elongate, parallel-sided, total length 5.3-6.8. Head: Elongate, frons sloping and distinctly projecting beyond anterior margin of eyes; clypeus prominent and visible from above; eyes large; antennae rather long, with segment I as long as or slightly shorter than pronotum; labium reaching but not surpassing hind coxae. Thorax: Pronotum trapeziform, elevated posteriorly, with weakly defined calli, concave margins, rounded and posterior angles somewhat protruding laterally; metathoracic scent-gland evaporatory area long and narrow, wedge-shaped. Legs: Slender; hind femora rather thin and long, surpassing apex of abdomen; tarsal segment II slightly longer than segment III (Fig. 28); claw (Fig. 29) rather long, thin, apically curved; pulvillus large, flaplike, reaching midpoint of claw curvature, apically free, not attached to the claw. GENITALIA: Genital segment: Rather large, about 0.4 of abdomen, gradually tapering, with broadly rounded apex, without distinctive ornamentation. Parameres: Right paramere lanceolate, of typical phyline shape, comparatively long (Fig. 26); left paramere (Figs 23, 24, 27) with very broad, apically bifid sensory lobe, apical process thin and strongly curved apically. Apex of theca: Shape typical of many phylines (Fig. 25). Vesica: Comparatively long and thin, S-shaped, apically with two straight blades of unequal length and more or less developed series of spicules between secondary gonopore and base of blades; secondary gonopore subapical, with well developed sculpture (Figs 7-10). Female: Body (Figs 2, 4) short, stout, strongly brachypterous, total length 4.55.8. COLORATION, SURFACE AND VESTITURE: As in male. STRUCTURE: Head: Similar to that of male but with distinctly smaller eyes and more convex frons. Thorax: Not elevated posteriorly; pronotum and scutellum almost flat in lateral view; pronotum trapeziform to nearly square, with straight margins; posterolateral angles not protruded. Hemelytra: Strongly brachypterous; lateral margins broadly convex; corium and clavus fused; cuneus and membrane absent; apex of hemelytron rounded, reaching at most posterior margin of abdominal tergum III. Legs: Not as elongate as in male, hind femora almost reaching apex of abdomen. Abdomen: Broad and deep, 1.8-1.9 times as broad as basal width of pronotum. GENITALIA: As in Figs 12-13, with more or less triangular sclerotized rings; posterior wall simple, with minute, hardly visible and evenly distributed rows of spinules (Fig. 16); vestibulum narrow, S-shaped, irregularly sclerotized (Fig. 15). Distribution: Central Asia (Fig. 41). Discussion: Omocoris was described by Lindberg (1930) to accommodate a single species O. parallelus. Although the tribal affiliation was not discussed in the description, the genus was originally placed in orthotylines. Carvalho (1952) revealed the close affinity of the genus with Eurycolpus and considered the latter to be a senior synonym of Omocoris. The generic status of Omocoris was subsequently restored by Kerzhner & Muminov (1964), who provided diagnoses for both genera based on the vestiture, length of the antennal segment I, degree of the sexual dimorphism, and the male genitalic structures, especially on the number of apical processes of the vesica and the shape of the sensory
220 F.V. Konstantinov
lobe of the left paramere. Kerzhner & Muminov also synonymized O. paralellus with Oncotylus cunealis Reuter, 1904 and transferred the latter to Omocoris. O. erythrophtalmus is described by Carapezza (1997) on the basis of a male and female collected in Tunisia, and is so far known only from the type series. The species does not fit the diagnosis of Omocoris presented above in all respects except in the structure of the pretarsus. The following combination of characters clearly indicates that the placement of O. erythrophtalmus in Omocoris is unsatisfactory: antennal segment I short, not longer than interocular distance; vestiture and tibial spines pale; the legs comparatively short; the females macropterous; the sensory lobe of the left paramere long, apically pointed. The characters listed above are shared by a number of Palaearctic phyline genera. The species resembles Eurycolpus in external appearance and pretarsal structure, but lacks the subapical notch on the claw, a character, according to my observations, shared by all representatives of Eurycolpus. The correct assignment of O. erythrophtalmus evidently would not become possible without examination of the vesica, which is missing in the holotype, the only known male specimen (Carapezza 1997). The female genitalia were found to be of no value at the generic level in Oncotylus, Omocoris, Eurycolpus, Josifovius and other related genera. Due to these reasons, I herein declare Omocoris erythrophtalmus Carapezza, 1997 as a species incertae sedis.
Omocoris cunealis (REUTER, 1904) Figs 1, 2, 8, 10, 12, 23, 41 Oncotylus cunealis Reuter, 1904: 7, 14. Omocoris parallelus Lindberg, 1930: 21 (syn. Kerzhner & Muminov, 1964: 46). Oncotylista (nom. nud.) cunealis Reuter: Kiritshenko, 1959: 108-109
Diagnosis: Easily distinguished from O. unicolor by the larger size, reddish cuneus in the male, longer hemelytron in the female always reaching posterior margin of the abdominal tergum III, comparatively robust vesica with the well developed twofold subapical series of teeth, narrow and weakly sclerotized rings of the bursa copulatrix. Description: Male: COLORATION (Fig. 1): Greenish yellow; femora apically with minute pale brown spots or uniformly pale; cuneus and adjacent medioapical margin of corium pale reddish; outer margin of cuneus greenish yellow; membrane uniformly smoky pale brown, sometimes with indistinct brown patches. SURFACE AND VESTITURE: As in generic description. STRUCTURE: Body 4.0 – 4.5 × as long as width of pronotum; total length 5.8-6.8. Head: Elongate; vertex 1.0-1.2 × as wide as eye; antennal segment I 1.0-1.2 × as long as pronotum; antennal segment II 1.3-1.4 × as long as basal width of pronotum, 1.9-2.2 × as long as width of head. Thorax: Pronotum 1.9-2.1 × as wide as long; tarsus and claw as in generic description. GENITALIA: Genital segment, theca and right paramere as in generic diagnosis; left paramere (Fig. 23) almost flat apically
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 221
with slightly bifid sensory lobe; vesica larger and more robust than in O. unicolor, strongly sclerotized, with well developed, twofold series of spicules running from projection distal to secondary gonopore (Figs 8, 10). Female: Body 4.0-4.4 × as long as width of pronotum; total length 5.0-5.8. COLORATION: Uniformly greenish yellow, rarely femora with indistinct minute pale brown spots apically. SURFACE, VESTITURE AND STRUCTURE: As in generic description. Head: Vertex 1.2-1.5 × as wide as eye; antennal segment I 1.2-1.3 × as long as pronotum; antennal segment II 1.5-1.6 × as long as basal width of pronotum, 1.71.9 × as long as width of head. Thorax: Pronotum 1.5-1.7 × as wide as long; hemelytra reaching posterior margin of abdominal tergum III (Fig. 2). GENITALIA: As in Fig. 12, with weakly sclerotized, narrow, triangular sclerotized rings and narrow, S-shaped vestibulum. Host plant: O. cunealis was collected from Malcolmia sp. (Brassicaceae), annual, biennial, or perennial herb distributed in Central and South-West Asia, China, and Mediterranean region. Distribution: Known from Iran, Turkmenistan, Tajikistan, and Uzbekistan (Fig. 41). Material examined: Holotype of O. cunealis: IRAN: Khorasan: Kafar Qal’eh [Kyafirkala] on Gerirud River, E Khorasan, 36.4731°N 60.7553°E, 460 m, 09 Apr
7
8 9 10
Figs 7-10 : Vesica of Omocoris spp.: 7-8, in ventral view: 7 – O. unicolor (AMNH_PBI00149620), 8 – O. cunealis (AMNH_PBI00149561); 9-10, in lateral view: 9 – O. unicolor (AMNH_ PBI00149583), 10 – O. cunealis (AMNH_PBI00149547).
222 F.V. Konstantinov
1898, Zarudny, ♂ (AMNH_PBI 00149514). TAJIKISTAN: Daryakul [Dar’ya-Kul’] Lake, left bank of Vakhsh River, 37.31°N 68.50917°E, 21 Mar 1944, A. N. Kiritshenko, 3♂ (AMNH_PBI 00149516, AMNH_PBI 00149520-AMNH_PBI 00149521), 3♀ (AMNH_PBI 00149515, AMNH_PBI 00149517-AMNH_PBI 00149518), 11 larvae (AMNH_PBI 00149519, AMNH_PBI 00149522-AMNH_PBI 00149531); 20 Mar 1944, A. N. Kiritshenko, 2♂ (AMNH_PBI 00149537, AMNH_PBI 00149538), 5 larvae (AMNH_PBI 00149532-AMNH_PBI 00149536). Peschany Pass, 8 km N Bura-tau, 37.21888°N 68.44277°E, 10 Apr 1948, Shchetkin, 2♂ (AMNH_PBI 00149551, AMNH_PBI 00149552), 2♀ (AMNH_PBI 00149550, AMNH_PBI 00149553). TURKMENISTAN: Foothills 13 km SW Gyzylarbat [Kizyl-Arvat], 38.878°N 56.161°E, 25 Apr 1952, Ilyichev, 1♂ (AMNH_PBI 00149542). Mts 12 km SW from Gyzylarbat [Kizyl-Arvat], 38.889°N 56.175°E, 29 Apr 1952, Steinberg, 1♂ (AMNH_PBI 00149541); 22 Apr 1952, K.G. Romadina, 2♂ (AMNH_PBI 00149539, AMNH_PBI 00149540). Toutly, 35 km NE of Gyzylarbat, 39.18333°N 56.53333°E, 20 Apr 1952, Ilyichev, 1♂ (AMNH_PBI 00149544); 20 Apr 1952, K.G. Romadina, 1♂ (AMNH_PBI 00149543). Uzboy, 25 km W of Yaskhan, 39.681°N 55.282°E, 28 Apr 1952, Slepyan, 3♂ (AMNH_PBI 00149545-AMNH_PBI 00149547). UZBEKISTAN: 20 km N of Ayakguzhumdy, 40.739°N 63.747°E, 17 Jun 1966, I. M. Kerzhner, Malcolmia sp. (Brassicaceae), 22♂ (AMNH_PBI 00149554-AMNH_PBI 00149572), 19♀ (AMNH_PBI 00149554-AMNH_PBI 00149569, AMNH_PBI 00149573). Termez [Buchara mer. = former Bukhara Chanate], 37.21666°N 67.26666°E, Apr 1913, A. N. Kiritshenko, 2♂ (AMNH_PBI 00149548, AMNH_PBI 00149549).
Omocoris unicolor nov. sp. Figs 3, 4, 7, 9, 13, 15, 16, 19, 25-27, 28, 29, 41
Diagnosis: Differs from O. cunealis in the smaller size, uniformly pale greenish hemelytron usually with small pale brown spots in male, shorter hemelytron reaching at most posterior margin of the abdominal tergum II in female, thin vesica with a straight and hardly recognizable series of subapical spicules, and broadly rounded sclerotized rings of the bursa copulatrix. Description: Male: COLORATION (Fig. 3): Uniformly pale, greenish yellow; all femora, hemelytra, rarely base of pronotum densely covered with minute rounded pale brown spots; these spots usually more or less reduced on corium; rarely hemelytra uniformly pale; membrane pale, rarely slightly infuscate, outer margin with brown stripe extending from cells to apex of wing. SURFACE AND VESTITURE: As in generic description, but venter of fore coxa with dark spine-like setae in addition to those on antennal segment I and femora. STRUCTURE: Body elongate, parallel sided, 3.7-4.3 × as long as width of pronotum; total length 5.3-5.7. Head: Elongate; vertex 1.10-1.25 × as wide as eye; antennal segment I 0.9-1.0 × as long as pronotum; antennal segment II 1.2-1.3 × as long as basal width of pronotum, 1.55-1.85 × as long as width of head.
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 223
Thorax: Pronotum 1.8-2.1 × as wide as long; tarsus as in Fig. 28, claw as in Fig. 29. GENITALIA: Theca as in Fig. 25; right paramere as in Fig. 26; left paramere (Figs 24, 27) distinctly bifid sensory lobe apically; vesica thin and slightly sclerotized, with single, straight, and in some specimens bearly visible series of spicules (Figs 7, 9). Female: Body short, 3.8-4.3 × as long as width of pronotum; total length 4.5-5.1. COLORATION: As in male, brown spots on hemelytra faint to absent. SURFACE AND VESTITURE: As in generic description. STRUCTURE: Head: Vertex 1.3-1.5 × as wide as eye; antennal segment I 0.9-1.1 × as long as pronotum; antennal segment II 1.2-1.4 × as long as basal width of pronotum, 1.3-1.6 × as long as width of head. Thorax: Pronotum 1.6-1.7 × as wide as long; apex of hemelytron rounded, reaching to about posterior margin of abdominal tergum II (Fig. 4). GENITALIA: As in Fig. 13, with well sclerotized and comparatively rounded sclerotized rings, vestibulum as in Fig.
11
14
12
13
15
16
17
Figs 11-17: Female genital structures of Omocoris and Josifovius: 11-13, bursa copulatrix and associated structures in dorsal view: 11 – J. dimorphus (AMNH_PBI 00149577), 12 – O. cunealis (AMNH_PBI 00149515), 13 – O. unicolor (AMNH_PBI00149609); 14-15, vestibulum in anterior view: 14 – J. dimorphus (AMNH_PBI 00149577), 15 – O. unicolor (AMNH_PBI00149585); 1617, sculpture of the ventral wall of bursa copulatrix: 16 – O. unicolor (AMNH_PBI00149585), 17 – J. dimorphus (AMNH_PBI 00149577).
224 F.V. Konstantinov
15, ventral wall of bursa copulatrix simple, without any armament, with fine sculpture as in Fig 16. Host plant: Adults and larvae of the species were collected from Hypecoum pendulum var. parviflorum (Kar. & Kir.) Cullen (Fumariaceae). This annual herb is known from Central Asia, Turkey, Iran, Afghanistan, and Pakistan. Single occurrence on Malcolmia sp. is considered a sitting record. Distribution: Uzbekistan (Fig. 41). Etymology: unicolor is the Latin adjective meaning “of a single color”, referring to the absence of contrasting color pattern on the dorsum of this new species. Material examined: Holotype: UZBEKISTAN: Aktau, Tamdytau, 40.739°N 63.747°E, 29 Apr 1966, I. M. Kerzhner, ♂ (AMNH_PBI 00149622). Paratypes: UZBEKISTAN: 18 km N of Ayakguzhumdy, 40.919°N 63.747°E, 18 Apr 1966, I. M. Kerzhner, Hypecoum pendulum var. parviflorum (Fumariaceae), 1♀ (AMNH_PBI 00149589). 20 km E of Dzhankel’dy, 40.85°N 63.573°E, 19 Apr 1966, I. M. Kerzhner, Malcolmia sp. (Brassicaceae), 1♂ (AMNH_PBI 00149582). 20 km N of Ayakguzhumdy, 40.739°N 63.747°E, 17 Jun 1966, I. M. Kerzhner, Hypecoum pendulum var. parviflorum (Fumariaceae), 8♂ (AMNH_PBI 00149594-AMNH_PBI 00149599, AMNH_PBI 00149601-AMNH_PBI 00149602), 11♀ (AMNH_PBI 00149591-AMNH_PBI 00149592, AMNH_PBI 00149594-AMNH_PBI 00149599, AMNH_PBI 00149601), 2 larvae (AMNH_PBI 00149593, AMNH_PBI 00149593). 30 km N of Ayakagytma, 40.935°N 64.498°E, 25 Apr 1966, I. M. Kerzhner, 1♀ (AMNH_PBI 00149632). Aktau, Tamdytau, 40.739°N 63.747°E, 02 May 1966, G.S. Medvedev, 3♂ (AMNH_PBI 00149612, AMNH_PBI 00149620-AMNH_PBI 00149621); 01 May 1966, G.S. Medvedev, 1♂ (AMNH_PBI 00149611), 2♀ (AMNH_PBI 00149609, AMNH_PBI 00149610); 02 May 1966, I. M. Kerzhner, 10♂ (AMNH_PBI 00149615, AMNH_PBI 00149617-AMNH_PBI 00149619), 6♀ (AMNH_PBI 00149613-AMNH_PBI 00149614, AMNH_PBI 00149616, AMNH_PBI 00149616, AMNH_PBI 00149626-AMNH_PBI 00149627); 29 Apr 1966, I. M. Kerzhner, 3♂ (AMNH_PBI 00149623-AMNH_PBI 00149625), 4♀ (AMNH_PBI 00149628-AMNH_PBI 00149631). Ayakguzhumdy, E of Dzhankel’dy [Dzhangildy], 40.739°N 63.747°E, 18 Apr 1966, I. M. Kerzhner, 2♂ (AMNH_PBI 00149583, AMNH_PBI 00149600), 10♀ (AMNH_PBI 00149584-AMNH_PBI 00149588, AMNH_PBI 00149600), 2 larvae (AMNH_PBI 00149590, AMNH_PBI 00149590); 23 Apr 1966, G.S. Medvedev, 4♂ (AMNH_PBI 00149603-AMNH_PBI 00149605), 14♀ (AMNH_PBI 00149603AMNH_PBI 00149608). Between Dzhankel’dy and Ayakguzhumdy, 40.792°N 63.529°E, 22 Apr 1966, I. M. Kerzhner, 3♂ (AMNH_PBI 00149580, AMNH_PBI 00149581), 3♀ (AMNH_PBI 00149579, AMNH_PBI 00149581). Kyzylkum Sands, 10 km SW of Turtkuduk, 40.643°N 64.184°E, 24 Jun 1966, I. M. Kerzhner, 3♂ (AMNH_PBI 00149636), 3♀ (AMNH_PBI 00149637). Kyzylkum Sands, Besapan, 41.5°N 64.5°E, 28 Apr 1966, I. M. Kerzhner, 5♂ (AMNH_PBI 00149634, AMNH_PBI 00149634, AMNH_PBI 00149634-AMNH_PBI 00149635, AMNH_PBI 00149635), 2♀ (AMNH_PBI 00149633, AMNH_PBI 00149635).
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 225
18
19
23
22
20
24
21
25
26
27
Figs 18-27: Male genitalia: 18-19, vesica in lateral view: 18 – Josifovius dimorphus (AMNH_PBI 00149574), 19 – Dasycapsus cunealis (AMNH_PBI 00155719); 20-21, vesica in ventral view: 20 – J. dimorphus (AMNH_PBI 00149574), 21 – D. cunealis (AMNH_PBI 00155719); 22-24: left paramere, view in dorsal view: 22 – J. dimorphus (AMNH_PBI 00149574), 23 – Omocoris cunealis (AMNH_PBI 00149547), 24 – O. unicolor (AMNH_PBI 00149583); 25-27, O. unicolor (AMNH_PBI 00149583): 25 – apex of theca, 26 – right paramere, 27 – left paramere.
226 F.V. Konstantinov
Josifovius nov. gen. Type species: Eurycolpus dimorphus Wagner, 1961
Diagnosis: Recognized by the pale yellow coloration with somewhat darkened claval comissure, simple pale vestiture, dark tibial spines, pulvillus reaching to half the length of the claw and with the extreme apex not attached to the claw, submacropterous female, and J-shaped vesica with a small, apical secondary gonopore and series of square dentations running on the distal half of the vesica. Most similar to Eurycolpus Reuter in the color pattern, body proportions, vestiture, and comparatively short rostrum reaching the middle coxae. Eurycolpus distinguished from Josifovius by the presence of subapical notch on the claw (Figs 30-32), subbrachypterous females, antennal segment I considerably shorter than distance between the eyes, and entirely different, long and thin, usually strongly twisted vesica with a long and gradually tapering apical process and secondary gonopore located far from the apex. The series of peculiar square dentations on the vesica also occur in some Tinicephalus Fieber, 1858 and several clearly unrelated phylines, namely in many species of Moissonia Reuter, 1894, Chrysochnoodes rufus Wagner, 1959, and Megalodactylus macularubra (Mulsant & Rey, 1852). Several species of Tinicephalus further resemble Josifovius in having submacropterous females, similar coloration, and uniformly pale vestiture. However, the former genus differs from Josifovius in the body proportions, vesica with long, typically two or even three lobed apical process, and pretarsus with claw strongly curved apically, pulvillus large, closely adhered to claw with the apex almost reaching tip of claw (Figs 37, 38). The apically located secondary gonopore is reminiscent of two Palaearctic Phylini genera: Dasycapsus Poppius, 1912 and Acrotelus Reuter, 1885. However, both genera have a very short, robust vesica twice as long as the phallobase (Fig. 19), with large secondary gonopore, and a large pulvillus almost reaching the apex of the claw (Figs 35, 36, 39, 40). Description: Male: COLORATION (Fig. 5): Uniformly pale yellow, apex of labium darkened; femora apically with indistinct, small, rounded, pale brown spots, rarely uniformly pale. Outer parts of clavus and corium usually paler, whitish; claval comissure somewhat darkened, saturated yellow to pale brown; membrane whitish, with brown edging along outer cell and brown kidney-shaped spot extending from cells to apex of wing. SURFACE AND VESTITURE: Dorsum smooth, shining; whole body with silver, adpressed, simple setae; tibial spines dark; antennal segment I with single spine-like setae on medial surface. STRUCTURE: Body elongate, parallel-sided. Head: Short, declivent, weakly projecting beyond anterior margin of eyes; clypeus visible from above; eyes large; antennae relatively long; labium reaching to apex of middle coxae. Thorax: Pronotum trapeziform, with weakly defined calli; metathoracic scent-gland evaporatory area relatively short, oval. Legs: Slender; hind femora rather thin and long, surpassing apex of abdomen; tarsal segment II slightly longer than segment III (Fig. 33); claw (Fig. 34) thin, gradually curved; pulvillum flaplike, reaching to half length of claw, with extreme
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 227
apex free, not attached to the claw. GENITALIA: Genital segment of moderately size, about 0.3 of abdomen, conical, without distinctive ornamentation; theca L-shaped, of shape typical of many Phylinae, lacking distinctive morphological features; left paramere with gradually tapering sensory lobe and thin, smoothly curved apical process (Fig. 22); right paramere small, typical Phyline; vesica J-shaped (Figs 18, 20), thin, with a series of square, apically dilative dentations running from middle to apex of vesica; secondary gonopore small, located at extreme apex of vesica, with distinctive ornamentation. Female: COLORATION, SURFACE AND VESTITURE: As in male. STRUCTURE: Body elongate oval, submacropterous. Head: Longer than in male, projecting
28
29
31 30 32
34 33
35
37
39
36
38
40
Figs 28-40: Hind tarsus and pretarsus: 28-29 – Omocoris unicolor (AMNH_PBI 00149583), 3031 – Eurycolpus flaveolus (AMNH_PBI 00146819), 32 – E. aureolus (AMNH_PBI 00146764), 33-34 – Josifovius dimorphus (AMNH_PBI 00149575), 35-36 – Dasycapsus cunealis (AMNH_PBI 00155719), 37-38 – Tinicephalus hortulans (AMNH_PBI 00154012), 39-40 – Acrotelus pilosicornis (AMNH_PBI 00147807).
228 F.V. Konstantinov
well beyond anterior margin of eyes, with distinctly smaller eyes and more convex frons. Hemelytra submacropterous (Fig. 6), surpassing anterior margin of abdominal tergum VIII; cuneus broadly triangular, with apex reaching middle of abdominal tergum VII. Legs: Not as elongate as in male; hind femora not reaching apex of abdomen. Abdomen: Rather broad, 1.5-1.6 times as broad as basal width of pronotum. GENITALIA: As in Fig. 11, sclerotized rings rounded, more or less straight medially, posterior wall simple, without any armament, with fine sculpture as in Fig 17, vestibulum long, twisted, narrow and almost entirely membranous (Fig. 14). Host plant: Unknown. According to Carapezza (1997), the species was generally collected by sweeping on Asteraceae in desert habitats. Distribution: Originally described from Egypt and later recorded from Algeria (Eckerlein & Wagner 1965), Libya (Eckerlein & Wagner 1970), Tunisia, and Israel (Carapezza 1997). Etymology: The genus is named after Michail Josifov in recognition of his substantial contributions to our knowledge of heteropteran systematics. The gender is masculine. Discussion: Eurycolpus dimorphus Wagner, 1961 was transferred to Omocoris by Wagner (1975) evidently due to the submacropterous females and the antennal segment I somewhat longer than in Eurycolpus. However, in the type species of Omocoris females are strongly brachypterous, with fused corium and clavus, and absent cuneus and membrane, while the antennal segment I is much longer than in both sexes of O. dimorphus. Combination of other characters, including vestiture, structure of the head, pretarsus, sensory lobe of the left paramere and peculiar vesica also clearly contradicts with the current placement of O. dimorphus.
Omocoris cunealis Omocoris unicolor
Fig. 41: Distribution of Omocoris spp.
Review of Omocoris Lindberg, 1930 and a description of a new genus to accommodate Eurycolpus dimorphus... 229
Based on a preponderance of morphological evidence, I have to conclude that neither Omocoris, nor any other phyline genus, can adequately accommodate O. dimorphus and the new monotypic genus Josifovius is erected to place it.
Josifovius dimorphus (WAGNER, 1961) nov. comb. Figs 5, 6, 11, 14, 17, 18, 20, 22, 33, 34 Eurycolpus dimorphus Wagner, 1961: 318-320 Omocoris dimorphus: Wagner, 1975: 268-269
Diagnosis: Recognized by the characters given in the generic diagnosis. Description: Male: COLORATION, SURFACE, VESTITURE AND GENITALIA: As in generic description. STRUCTURE: Body 4.2-4.4 × as long as width of pronotum; total body length 4.5-4.7. Vertex 1.15-1.20 × as wide as eye; antennal segment I 0.9 × as long as pronotum; antennal segment II 1.5 × as long as basal width of pronotum and 2.0 × as long as width of head; pronotum 2.05 × as wide as long. Female: COLORATION, SURFACE, VESTITURE AND GENITALIA: As in generic description. STRUCTURE: Body 3.6 × as long as width of pronotum; total body length 3.5-3.8. Vertex 1.8-2.1 × as wide as eye; antennal segment I 0.7 × as long as pronotum; antennal segment II 1.1 × as long as basal width of pronotum, 1.4-1.5 × as long as width of head; pronotum 2.0 × as wide as long. Material examined: LIBYA: Kseia, 32.581°N 14.041°E, 02 May 1961 - 03 May 1961, Eckerlein, 2♂ (AMNH_PBI 00149574, AMNH_PBI 00149575), 2♀ (AMNH_ PBI 00149576, AMNH_PBI 00149577).
ACKNOWLEDGEMENTS I am greatly indebted to Dr I. M. Kerzhner (Zoological Institute, Russian Academy of Sciences), and Dr. Michael D. Schwartz (ECORC, Agriculture and Agri-Food, Ottawa, Canada) for comments on the manuscript. This paper is a contribution to the NSF Planetary Biodiversity Inventory grant (PBI) DEB-0316495. The equipment used during the work was partly funded by the Russian Foundation for Basic Research, project No 0504-48179 and “Leading scientific schools” project No. НШ 7130.2006-4. The collection of Zoological Institute, St. Petersburg, is supported by Rosnauka for UFC no. 2-2.20.
РЕЗЮМЕ Ревизиран е род Omocoris LINDBERG и е описан нов вид O. unicolor nov. sp. от Узбекистан. Описан е новият род Josifovius, с типов вид Eurycolpus dimorphus WAGNER, 1961. Omocoris
230 F.V. Konstantinov
erythrophtalmus CARAPEZZA, 1997 се разглежда като species incertae sedis. Представени са илюстрации на мъжките и женски гениталии, тарсуса и претарсуса, фотографии на дорзалния хабитус, както и данни за разпространението на видовете и растенията гостоприемник.
REFERENCES Carapezza A. (1997). Heteroptera of Tunisia. — Naturalista Siciliano, (4) 21. Suppl. A: 1-331. Carvalho J.C.M. (1952). On the major classification of the Miridae (Hemiptera) (With keys to subfamilies and tribes and a catalogue of the world genera) . — Anais da Academia Brasileira de Ciencias, 24: 31-110. Eckerlein H. & E. Wagner (1965). Ein Beitrag zur Heteropterenfauna Algeriens. — Acta Faunistica Entomologica Musei Nationalis Pragae, 11: 195-243. Eckerlein H. & E. Wagner (1970). Die Heteropterenfauna Libyens. — Acta Entomologica Musei Nationalis Pragae , 38 (1969): 155-194. Kerzhner I.M. & N.N. Muminov (1964). On some little-known capsid bugs (Heteroptera, Miridae) from Middle Asia. – Izvestiya Akademii Nauk Tadzhikskoi SSR, Otdelenie Biologicheskikh Nauk, 1964 (2): 45-49 (in Russian). Kiritshenko A.N. (1959). True bugs (Hemiptera-Heteroptera) of the “Tigrovaya Balka” Nature Reserve. — Trudy Instituta Akademii Nauk Tadzhikskoi SSR, 16(1): 97-110 (in Russian). Lindberg H. (1930). Neue oder wenig bekannte Capsiden aus Turkestan und dem Kaukasus. — Notulae Entomologicae, 10: 18-22. Reuter O.M. (1904). Capsidae persicae a D:o N.A. Zarudny collectae enumeratae novaeque species descriptae. — Ezhegodnik Zoologicheskogo Muzeya Imperatorskoi Akademii Nauk, 9: 5-16. Wagner E. (1961). Ein weiterer Beitrag zur Miriden-Fauna Aegyptens (HemipteraHeteroptera). — Bulletin de la Société Entomologique d’Egypte, 45: 315-322. Wagner E. (1975). Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera), Teil 3. — Entomologische Abhandlungen herausgegeben vom Staatlichen Museum für Tierkunde Dresden, 40, Suppl.: i-ii + 1-483.
N.northern Simov (Eds) 2008 alboacuminatus josifovi nov. ssp. 231 A new subspecies S. of Grozeva Lygaeidae & from Iran: Raglius ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 231-234. © Pensoft Publishers Sofia–Moscow
A new subspecies of Lygaeidae from northern Iran: Raglius alboacuminatus josifovi nov. ssp. (Heteroptera, Lygaeidae) * R.E. Linnavuori1 & E.Heiss2 1 2
Saukkokuja 10, FIN-21220 Raisio, Finland. E-mail: rauno.linnavuori@kolumbus.fi Research Entomologist, Tiroler Landesmuseum, Josef-Schraffl-Strasse 2 a, A-6020 Innsbruck, Austria. E-mail:
[email protected] (corresponding author)
ABSTRACT A new subspecies of the Lygaeidae taxon Raglius albocuminatus (Goeze, 1778), R. alboacuminatus josofovi nov. ssp. is described from northern Iran. Keywords: Heteroptera, Lygaeidae, Raglius, new subspecies, Iran.
INTRODUCTION The article is based on material of Raglius alboacuminatus (Goeze, 1787) collected by the authors in the Gilan and Golestan provinces in northern Iran, which shows distinct differences to the nominate taxon of widespread European distribution and is recognized and described as a new subspecies.
*
The authors dedicate this paper to their friend and excellent heteropterist Michail Josifov, in occasion of his 80th birthday, recognizing his valuable contributions to the taxonomy and faunistics of this order.
232 R.E. Linnavuori & E.Heiss
TAXONOMY Raglius alboacuminatus josifovi nov. ssp. (Figs 1, 3) Type material: Iran. Gilan province: Damashk- Bareh Sar, male holotype, male and female paratype, 27.VII.2002; Deylaman, female paratype, 18-20.VII.1996; DeylamanBarasar, female paratype, 23.VII.1996; Rustam Abad- Salan Sar, 53 paratypes (males and females) 24.VII.1996, 8-9.X.2000, Linnavuori leg.; Golestan province: Golestan N.P., Jahan Nama Prot. Area, 36°40’ N, 54°18’ E, 1700-1800m under loose bark of Quercus castaneifolia, 72 paratypes (males and females), 20-22.X.2003, Heiss leg.; Golestan N.P., Almeh 37°21’N, 56°07’ E, 1700m, 1 female 26-27 X 2003, Heiss leg.; Golestan N.P.,
A
B
C
Fig. 1: Raglius alboacuminatus josifovi nov. ssp.: A- subbrachypterous male; B- macropterous male; C- macropterous female.
A
B
C
Fig. 2: Raglius alboacuminatus alboacuminatus (Goeze, 1778): A- subbrachypterous male; B- macropterous male; C- macropterous female.
A new subspecies of Lygaeidae from northern Iran: Raglius alboacuminatus josifovi nov. ssp. 233
Deras Nu 36°39’N, 54°07’E, 2300m, 1male, 1 female 23 X 2003, Heiss leg. The holotype will be deposited in the future in the National Museums and Galleries of Wales. The paratypes are deposited in the collections of the authors and of B.Aukema, A.Carapezza, H.Günther, J.Péricart, J.Ribes, Ch. Rieger, W.Rabitsch, Zool. Inst. Acad. Sciences Sofia, Zool. Inst. St. Petersburg, Nat. Hist. Mus. Vienna, Prague and Budapest. Description. Length male 5.5-6.5 mm, female 6.5-6.75 mm. Head black, eyes brown. Antennae black, with 2nd segment and apical part of 1st yellowish. Rostrum dark brown. Anterior part of pronotum black, basal part yellow- brown, slightly darker medially, elevations of basal angles pale, rarely slightly darkened laterally, puncturing brownish. Scutellum black. Hemelytra yellowish brown; clavus with scutellar margin more or less darkened, elevated area between two puncture rows along claval suture whitish; apical part of corium blackish with triangular whitish spot, costal margin pale; membrane blackish with pale apical spot. Abdomen black, lateral margins of connexivum narrowly reddish brown. Under surface black, lateral margins of propleura narrowly, caudal margins broadly whitish yellow; caudal margins of mesopleura narrowly, of metapleura broadly whitish yellow. Legs yellow-brown, extreme tips of tibiae and tarsal segments 2 and 3 slightly darkened. Tibial spines dark brown. Macropterous and subbrachypterous. Body relatively elongate. Head 1.13-1.17 (males) or 1,18-1.27 (females) times as broad as long, 0.53-0.64 (males and females) times as long as basal width of pronotum, finely shagreened and indistinctly punctate, with some erect brown setae, ocular index 3.42-3.9 (males), 3.7-4.1 (females). Antennae long and gracile, proportions between segments 12:29:26:32 (male), 11:26:24:29 (female), 2nd segment 1.0-1.12 (male) or 0.93-1.07 (female) as long as diatone, 0.58-0.68 (male) or 0.53-0.64 (female) times as long as basal width of pronotum. A
D
B
C
E
Fig. 3: Raglius alboacuminatus josifovi nov. ssp.: A- male fore femur and tibia; C- male hind femur; R.alboacuminatus alboacuminatus (Goeze, 1778): B- male fore tibia; D-E left paramer in two positions; Scale 0.5 mm. D-E after Péricart (1998).
234 R.E. Linnavuori & E.Heiss
Rostrum extending to middle coxae. Pronotum 1.31-1.48 times as broad basally as long in middle, distincly broadening caudad, anterior margin punctate, anterior lobe finely shagreened and, like lateral margins, with long erect setae, basal part medially punctate. Scutellum finely punctate. Hemelytra distinctly punctate, membranes in subbrachypterous form extending to 5th tergite. Connexivum and venter with long erect brown setae. Legs appearing long and gracile, fore tibia in male nearly straight (Fig. 3 A), hind femur (Fig. 3 C) with a sharp subapical tooth. Male genitalia as in the nominate form; parameres as in Fig. 3 D-E. Habitat: In mountain meadows. Very common on Stachys sp. in Rustam Abad- Salan Sar in Gilan province. The records from Jahan Nama under bark of Quercus seem to be already aggregations for overwintering in adult stage. In the meadows nearby the dominating plant was Stachys byzantinica, but no specimen was collected from these plants. Etymology: The new subspecies is dedicated to Prof. Dr. Michail Josifov for his excellent work on Heteroptera. Differential diagnosis: The nominate subspecies Raglius acuminatus acuminatus (Goeze, 1778) (Fig. 2) (redescription in Péricart 1998: 271-274), which is widely distributed from Western Europe and the Mediterranean area to Middle Asia, is of very variable colouration. Anyway, it is distinguished by the blackish brown basal lateral spots on the pronotum, the apically somewhat darkened 2nd antennal segments and the dark pattern of the legs: fore femora largely, the other femora apically darkened, apices of tibiae also dark. Moreover the body is somewhat broader. The head is broader, 1.23-1.30 times as broad as long. The antennae are shorter and more incrassate. The pronotum is broader, 1.4-1.6 times as broad basally as long in middle and more parallel-sided. The protibias of males (Fig. 3 B) are gently arched. Since there is practically no individual variability in the large Iranian material, it undoubtedly represents a separate subspecies.
ACKNOWLEDGMENTS The authors are greatly indebted to Mr.Veikko Rinne of the Zoological Museum of the Turku University, Turku, Finland, for the photos given in the paper.
РЕЗЮМЕ Нов подвид на вида Raglius albocuminatus (GOEZE, 1778) (Lygaeidae), R. alboacuminatus josofovi nov. ssp. е описан от Северен Иран.
REFERENCES Péricart, J. (1998). Hémiptéres Lygaeidae euro-méditerranéens. 3. – Faune de France 84 C: I-VI, 1-487.
Grozeva & and N. Simov (Eds)in2008 AlaryS. polymorphism new localities palaearctic Oncocephalus Klug, 1830 235 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 235-240. © Pensoft Publishers Sofia–Moscow
Alary polymorphism and new localities in palaearctic Oncocephalus KLUG, 1830 (Heteroptera, Reduviidae, Stenopodainae)* P. Moulet Museum Requien, 67 rue Joseph Vernet, 84000 Avignon, France. E-mail:
[email protected]
ABSTRACT Description of a brachypterous specimen of Oncocephalus aspericollis; the occurrence of this species in Europe is documented. Description of the sub macropterous morph of O. squalidus and of a macropterous specimen of O. afghanus. O. bipartitus must be cancelled from European fauna. Keywords: Oncocephalus, O. afghanus, O. aspericollis, O. bipartitus, O. squalidus.
INTRODUCTION Oncocephalus Klug, 1830 is the largest genus in the sub family Stenopodainae. Nearly 200 species are known mainly from Palaearctic and Africa, the genus is poorly known in America (Maldonado Capriles 1990). In the Palaearctic area Putshkov & Putshkov (1996) recorded 66 species, and Moulet (2001, 2003, 2005) described some new taxa mainly from the Iran, Saudi Arabia and Yemen. The diagnostic characters of Oncocephalus are summarized as: two strong tubercles on the head, fore femora with one or two row(s) of spines underneath, tarsi 3-segmented. The general colour light brown, beige or light greyish is widely distributed in the genus * This paper, the 5th contribution to palaearctic Stenopodainae, is dedicated to commemorate the 80th birthday of eminent Prof. M. Josifov.
236 P. Moulet
and, at first glance, the species are very close. Often, only ♂ can be distinguished by their genitalia (parameres and pygophore). The wing polymorphism is well-known in both sexes in Oncocephalus although sometimes, in some species, one can discover new alary forms. On my request my colleagues Dr. C. Rieger, Dr. P. Kment (Praga, NMPC) and Dr. D. Redei (Budapest, HNHM) sent me some Reduviidae from their own collection or institutions. This study focuses on Oncocephalus and allows me to make the following remarks. 1) Occurrence of Oncocephalus aspericollis REUTER, 1882 in Europe and description of the brachypterous form. According to Putshkov & Putshkov (1996) O. aspericollis Reuter is only known from Israel and O. arcticeps Puton & Noualhier from the Asian part of Turkey, Iraq, Syria and Israel. According to Linnavuori (1986) the latter is present in Saudi Arabia too but Putshkov & Putshkov (1996) doubt of such an observation. Moulet (2002) studied the types of both species and considered O. arcticeps a synonym of O. aspericollis which ipso facto is distributed from Saudi Arabia to Turkey. In C. Rieger’s collection there is 1♂ from Greece identified as O. aspericollis by G. Seidenstücker. The pygophore of this specimen is quite identical with the one of O. arcticeps figured by Moulet (2002, Fig. 45) and the parameres are the same as those of O. aspericollis (Moulet 2002, Fig. 47). Oncocephalus aspericollis must therefore be considered as a new taxon for Europe and the up to now unknown brachypterous form is described below. Material: 1 ♂; [Kos Island, near Tingaki, 1976 may 19th, leg Schmidt]; in coll. C. Rieger. Adult (Fig. 1). Ground colour light beige to pale yellow. Head 1.5 times longer than width across eyes ; anterior lobe 2.21 times as long as posterior. Second antennal joint 1
2
Fig. 1: Brachypterous form of Oncocephalus aspericollis, habitus. Fig. 2: Sub macropterous form of Oncocephalus squalidus, habitus. Fig. 3: Macropterous form of Oncocephalus afghanus, habitus. Scale bars : 3 mm. All original
3
Alary polymorphism and new localities in palaearctic Oncocephalus Klug, 1830
237
1.8 times as long as first with setae 2.33 times longer than diameter. Anterior lobe of pronotum yellowish, mid line dark, two sub median feebly carinate light stripes continuing on posterior lobe as two little sub lateral lines not surpassing the transverse furrow; lateral edges of anterior lobe thickened; posterior lobe brownish with a second yellow line not reaching posterior margin. Hemelytron 2.5 times as long as scutellum. Underside of fore femora with a row of seven acute teeth and a second of only two teeth. Abdominal tergites pale yellow (nearly whitish) with a median pale grey spot (spots increase in size from forward to backwards); paratergites brown except apex; tip of femora and tibiae more or less dark brown (more than in macropterous forms). Gula brown; venter of abdomen laterally embrowned. Length: 11. 4 mm. Some differences on measurements can be noted in this specimen. In macropterous, the anterior lobe of head is 2.5-3 times longer than posterior and the second antennal joint is 2.15 times as long as the first. One observes that generally macropterous specimen is lighter. 2) Description of the sub macropter form of Oncocephalus squalidus (ROSSI, 1790). Oncocephalus squalidus is a well-known species throughout Western Palaearctic, the Iran (Dispons & Villiers (1967) and pers. obs.) and as far as the Sudan (Putshkov & Putshkov 1996). Nevertheless, up to now, only the macropterous form was known. A quite singular specimen in C. Rieger’s collection can be considered as the sub macropterous form of O. squalidus. Two characters particularly lead to this conclusion. First, the head shape which anterior lobe is clearly widened in front of eyes, only O. squalidus is diagnosed by this feature among the species with only one row of profemoral teeth. Second, the colour of the dorsal connexivum with one large and one little black spots is so far known only in O. squalidus (Moulet 2002, Fig. 31). The little size of this specimen (10.45 mm) is unusual, since the macropterous specimens that I have studied are 13-14.5 mm. long. Material examined: 1 ♂ ; [Syria, Salkhad, april 5th, 1988, on a basaltic plate at 9001200 m high]; in coll. C. Rieger. Adult (Fig. 2). Plain general colour, even without diffuse coloured spots. Head and antennal joints I and II brown (sienna), interocular furrow well developed and deep ; anterior lobe of head 3 times as long as posterior and 0.75 times as wide as head across eyes. Pronotum darkened, anterior and lateral edges yellowish; two stripes on the disc with little round tubercles; mid costulations nearly reaching anterior and posterior margins, the lateral fused with the median ones anterior to the transverse furrow; the latter very well developed and deep; no lateral tubercle. Scutellum blackish, mid line light. Hemelytron beige-brown, the up most lateral margin and the membrane slightly embrowned, veins little pronounced (even on the corium). Legs of general colour; apex of femora, tip of tibiae and sub apical ring very faintly embrowned. Dorsum beige brown. Segments of the dorsal connexivum with a big quadrangular blackish spot forward and a small rounded one backwards. Antennae, legs, dorsum and exocorium with transparent, short and very strong setae. Length: 10.45 mm.
238 P. Moulet
3) Oncocephalus bipartitus HORVÁTH, 1911 cancelled of European fauna. In his study on the Heteropterological fauna of Santorin, Rieger (1995) reported the presence of O. bipartitus on this island. This seemed possible because this species is known to occur in Lebanon and Syria (Putshkov & Putshkov 1996). I have studied this specimen and found that, in fact, it represents the apterous form of O. obsoletus Klug. The main difference between the two species is the ornamentation of the hemelytron: brown with a transverse black stripe in the apterous form of O. obsoletus and lighter without spot or stripe in O. bipartitus. Other differences are found in the colour of the pronotum and the dorsal connexivum. The record of O. bipartitus in Europe is based on a misidentification and this taxon must be removed from the European list of Heteroptera which must be considered as a near eastern species. 4) Description of the macropterous form of Oncocephalus afghanus HOBERLANDT, 1961. In the collection loaned by NMPC there was a little series of O. afghanus Hoberlandt with ♀ type, one ♂ brachypterous and one ♂ macropterous, a form up to now unknown in this species. Material examined: 1 ♂ ; [Parwan-Tal, 4 km W of Gulbahar (Afghanistan, prov. Parwan), 1600 m high, february 18th, 1962, unter Steinen, am Ufer, Kullman leg]. The antennae, rostrum, left legs and right fore tarsus missing. Adult (Fig. 3). Head long (0.88 times as long as the pronotum and 1.54 times as long as width across eyes), black, lightened from base of antennal tubercle to anterior end with faint and shortly setiferous granulations; cephalic furrow behind eyes well developed and deep; posterior lobe higher than anterior and 0.39 times as long as it; eyes voluminous, globose; ocelli large. Pronotum dark brown to blackish; anterior margin lightened; anterior angles short, acute and bent forward ; transverse furrow distinct but not deep. Disc of anterior lobe with some granulations and very little bright spaces, integument mat between spaces. Posterior lobe bell-shaped, posterior margins broadly convex, 1.13 times as long as the anterior lobe; humeral angles feebly obtuse, hardly surpassing the corium laterally; with two lateral yellow stripes almost reaching the posterior margin and two sub median stripes continuing on the anterior lobe. Lateral margins straight, not sinuous at the transverse furrow level, with very short denticulation; no lateral tubercle. Scutellum black, tip long and regularly tapered; mid line and farthest apex somewhat light. Hemelytron swarthy, surpassing abdominal tip, base of exocorium, of discal cell and apex of apical cells very slightly brownish; inferior margin and apex of membrane very light brown, some little spaces decoloured; veins strong and light, Sc vein very strong (nearly carinate). Yellowish dorsal connexivum, a quadrangular black spot in the middle of each segment. Fore femora rather thin, brown-yellowish, tip blackish, a median ring more or less complete, clear and brown (sometimes fused with apical spot); two rows of teeth underneath: the external with three teeth, the internal with seven; fore tibiae yellowish, a broad brown median ring, the base slightly brown. Middle and hind femora yellowish
Alary polymorphism and new localities in palaearctic Oncocephalus Klug, 1830
239
with an indefinite sub median ring and another, sub apical, clear brown-black. Middle and hind tibiae with a dark ring on the apical third and a sub median one hardly darker than the ground colour of legs. Gula, thoracic pleura, sternites and venter black; coxal cavities and mid abdominal carina yellow. Parameres as previously figured (Moulet 2003). Length: 15.3 mm. The macropterous O. afghanus is closely related to O. aspericollis Reuter and O. hierosolymensis Moulet. It can be distinguished by its swarthy colour and the absence of hemelytral spots which exist, though not very clearly, in the other two species; the rings and spots of the legs are distinct in O. afghanus particularly on the middle and hind legs; the membrane in O. afghanus surpasses the tip of the abdomen (hardly longer in the other two taxa). The ornamentation of the pronotum of this form is similar to O. gilvostriatus Moulet although the latter can be separated by the general colour, the very distinct black spots on the hemelytron, the head hardly 1.38 times as long as width across eyes, the distinct pronotal tubercle, the absence of a sub median black ring on the fore and middle tibiae, the membrane hardly longer than the abdomen and the shape of the parameres (Moulet 2005). The differences in this specimen and the one previously described (Moulet 2003) are mainly due to the wing polymorphism.
ACKNOWNLEDGEMENTS It is a great pleasure for me to thank warmly my friends Mrs Pluot-Sigwalt (Paris), P. Kment (Praga) and D. Redei (Budapest) for their help in this study. I am particularly indebted to Dr. C. Rieger for the loan of his collection which brought me the occasion of the present study. Many thanks to the two anonymous reviewers.
РЕЗЮМЕ Описан е брахиптерен индивид на Oncocephalus aspericollis и е потвърдено наличието на този вид в Европа. Описани са субмакроптерни индивиди на вида O. squalidus и макроптерни индивиди от вида O. afghanus. O. bipartitus трябва да бъде премахнат от европейската фауна.
REFERENCES Dispons P. & A. Villiers (1967). Contribution à la faune de l’Iran. 2. Hémiptères Reduviidae. — Annales de la Société entomologique de France, 3: 1067-1085. Linnavuori (1986). Heteroptera of Saudi Arabia. — Fauna Saudi Arabia, 8: 31-197.
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Maldonado Capriles J. (1990). Systematic catalogue of the Reduviidae of the World (Insecta, Heteroptera). — Caribbean Journal, Special edition. Porto Rico : x + 1-694. Moulet P. (2001). Quatre Oncocephalus Klug, 1830 nouveaux et description de la forme macroptère de O. pennatulus (Heteroptera, Reduviidae, Stenopodainae). — Nouvelle Revue d’Entomologie, 18(4): 353-368. Moulet P. (2002). Sur quelques Oncocephalus Klug, 1830 ouest-paléarctique (Hemiptera, Reduviidae, Stenopodainae). — Nouvelle Revue d’Entomologie, 19(3): 273-289. Moulet P. (2003). Contribution à l’étude des Stenopodainae paléarctiques (Heteroptera, Reduviidae). — Nouvelle Revue d’Entomologie, 20(3): 281-297. Moulet P. (2005). Sur quelques Stenopodainae paléarctiques (Hemiptera, Reduviidae). — Bulletin de la Société entomologique de France, 110(1): 31-36. Putshkov P. & V. Putshkov (1996). Family Reduviidae. — In: Aukema B. & C. Rieger (Eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 2 : Cimicomorpha I. The Netherlands Entomological Society, Amsterdam: 148-265. Rieger C. (1995). Die Fauna der Ägäis-Insel Santorin. Teil 9. Heteroptera. — Stuttgarter Beiträge zur Naturkunde, 520: 1-26.
S. Grozeva of &Th N.ailand Simov 2008 The Notonectidae (Heteroptera, Nepomorpha) and(Eds) adjacent areas, with keys for identification... 241 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 241-292. © Pensoft Publishers Sofia–Moscow
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification and five new records for Thailand N. Nieser1, P.-p. Chen2 & P. Leksawasdi3 1
Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands National Museum of Natural History (Naturalis), P.O. Box 9517, 2300RA Leiden, The Netherlands (international correspondence). 3 Department of Biology, Faculty of Sciences, Chiang Mai University, Chiang Mai, Thailand (correspondence within Thailand). 2
ABSTRACT Survey of the Notonectidae from Thailand with keys for males and females and diagnoses to all species known from Continental SE Asia and distributional records for Thailand. Five species are recorded for the first time in Thailand: Anisops exiguus Horváth, A. lansburyi Leong, A. niveus (Fabricius), A. tahitiensis Lundblad and Enithares malayensis Brooks. Additional first records: Anisops exiguus in Indonesia and Laos; A. nigrolineatus Lundblad in Laos and Vietnam and Enithares sinica (Stål) in Laos. Keywords: Notonectidae, keys, Continental SE Asia, Thailand, new records.
INTRODUCTION The family Notonectidae belongs to the infraorder Nepomorpha of the (sub)order Heteroptera or true bugs. Notonectidae swim with their belly up, hence their vernacular name backswimmers. It is a cosmopolitan family with about 350 described species worldwide (Chen et al. 2005), of which 16 have been recorded from Thailand. There are two subfamilies which have different ways of life which will be discussed below.
242 N. Nieser, P.-p. Chen & P. Leksawasdi
In the Old World the dominant genus of the subfamily Anisopinae is Anisops (Fig. 1) with about 130 species of which five species have been recorded from Thailand previously (Nieser 1998). In the Americas the dominant genus of Anisopinae is Buenoa Kirkaldy (1904) which is very similar to Anisops in build and way of life. On the whole results of general biological, morphological and physiological studies obtained in one of these genera will apply to the other too. A second genus of Anisopinae, Paranisops (Fig. 35), was considered an Australian endemic. It was recently found in Thailand where it is represented by two species (Nieser & Zettel 2001; Chen et al. 2002). The dominant genus of the subfamily Notonectinae is Notonecta Linnaeus (1758), which has a cosmopolitan distribution although it is poorly represented in tropical areas. In the Old World tropics Enithares, (Fig. 41) with about 75 species worldwide of which only two occur in the Americas, is the dominant genus of Notonectinae. Five species of Enithares have actually been recorded from Thailand but four more occur in adjacent countries. Most Thai species of backswimmers are only known in the macropterous form. Only a few species are predominantly brachypterous, but even in those a considerable portion of the population is able to fly and thereby to colonize different kinds of stagnant waters including small pools and puddles and artificial ponds. However, some species are more restricted in their habitat choice, e.g. being found nearly always in stagnant waters associated with streams or rivers. This is e.g. the case with most species of Enithares. Two additional genera: Aphelonecta (Fig. 40) and Nychia (Fig. 63), each with one species have been recorded from Thailand. Anisopinae, at least the genera Anisops and Buenoa, have cells associated with tracheoles in the ventral part of the basal half of the abdomen. These cells contain haemoglobin (Hungerford 1922). In Anisops these cells have been studied by Poisson (1926), but the best morphological study on the haemoglobin cells, combined with some basic biological observations, is by Bare (1928) in a species of Buenoa. The function of these haemoglobin cells in Anisopinae has been elucidated by Miller (1964) and Wells et al. (1981). All Notonectidae use oxygen from atmospheric air for respiration. They come to the surface to replenish a store of oxygen, which they use up during a dive. In Anisops most of the oxygen consumed during a dive is derived from the external air store. However, during a dive oxygen is gradually released from the oxyhaemoglobin into the air store enabling the bug to maintain neutral buoyancy during most of a dive. This means that they can be suspended in the water without actively swimming. This makes Anisopinae one of the two types of really planktonic insects, the other type being larvae of Chaoboridae (Diptera). The Notonectinae lack haemoglobin cells and obtain their oxygen from a large store of air, which they carry during a dive, which means that they are considerably lighter than water. They will float against the underside of the surface film when not clinging to a perch under water or actively and vigorously swim downward. Indeed many species of Notonectinae await their prey floating against the surface film. Thanks to sensory organs in their legs they can feel vibrations made by potential prey floating on the surface film. With their large eyes they can observe potential prey under water.
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 243
Notonectidae are strong predators, and many species of Notonectinae are reported to show strong preference for mosquito larvae and pupae (Papáek 2000, 2001). In Thailand some preliminary research into predation of mosquito larvae by a species of Enithares has been carried out (Chowanadisai 1986). Anisopinae are less frequently reported to attack mosquito larvae. This may be partly due to the fact that most observations have been made in temperate regions where Notonecta is dominant. But it has also to do with the fact that Notonectinae often hunt at the underside of the water surface whereas Anisopinae usually hunt deeper in the water where they are less likely to come across mosquito larvae. However, at least Anisops sardeus Herrich-Schaeffer, 1849, which occurs from the Mediterranean eastward to India and Burma, is reported to be an active predator of mosquitoes (Tawfik et al. 1986). Species of Culicidae of which the larvae live in open water can be eliminated entirely from ponds with Notonecta whereas larvae of Culicidae which tend to stay in the vegetation can coexist with Notonecta (Blaustein et al. 2004). Another indication that Notonectidae are important predators of mosquitoes is the fact that some species of mosquitoes tend to avoid oviposition in localities inhabited by backswimmers both Notonecta (Chesson 1984; Blaustein et al. 2004) and Anisops (Eitam et al. 2002). Several fish are reported to prey on backswimmers, specifically Gambusia affinis (Baird & Gerard) (e.g. Miura et al. 1979, 1984) which is sometimes used to diminish mosquito populations in rice-fields. Personal field experience shows that Notonectidae are relatively scarce or absent from ponds with fish unless there is well developed marginal vegetation which provides cover. Older instars often prey on younger instars of their own species. The latter may also seek cover between vegetation, whereas adults stay in the more open water. On the other hand, backswimmers can reach considerable densities in ponds used for the culture of fish, preying on the fry. This has a.o. been reported for two species occurring in Thailand: Anisops bouvieri (Gorai & Chaudhuri, 1962) and A. breddini (Leong, 1962). Judging form the ovipositors, the egg-laying habits are also different between the subfamilies. Notonectinae, notably Enithares have short and blunt ovipositors (first gonapophyses) which are moderately sclerotized and beset with stout spines (Fig. 52). Eggs are glued on the surface of a substrate. Females of Anisops on the contrary have strongly sclerotized and serrate ovipositors with which they deposit their eggs inside plant tissues.
MATERIAL AND METHODS The nomenclature of taxa above the species level has been discussed by Štys & Jansson (1988). This paper is to be consulted for justification and references for the names above species level used. Although this paper focuses on the Thai fauna, species occurring in Peninsular Malaysia and Singapore (together indicated as Malay Peninsula), southern part of Burma, Laos and Cambodia are incorporated in the keys and briefly discussed.
244 N. Nieser, P.-p. Chen & P. Leksawasdi
Specimens collected by P.- p. Chen and/or N. Nieser have been divided between the Nieser collection, Tiel, The Netherlands (NCTN) and the National Natural History Museum Naturalis at Leiden, The Netherlands (RMNH), some reference specimens have been deposited in the Collection of the Entomology Department, Chiang Mai University, Chiang Mai, Thailand (CMUC). In addition specimens from the Natural History Museum at Vienna, Austria (NHMW) have been studied. Measurements are in mm, length is measured from the anterior tip of the head to the posterior tip of the hemielytra, and width is the maximal width of the specimen. For recognition of Notonectidae and most of the terminology used the reader is referred to a general text on entomology or a more specialized text on Heteroptera e.g. Chen et al. (2005) or Schuh & Slater (1995). A few terms more or less typical for Notonectidae are explained below, see also Figures 1, 5, 42, 49, 54, 63. Cephalic projection (= cephalic horn) in some male Anisops the part of the interocular space extending in front of the eyes (Fig. 1). Coxal plate (= ventral metepisternal lobe) lateral lobe ventrally on metasternum at base of connexiva and covering the base of posterior coxa (Fig. 2). Genital capsule see introduction of Enithares. Nodal furrow a short groove running from the costal margin of a hemielytron inward, just proximally of the suture between corium and membrane (Figs 42, 43). For the distance of the nodal furrow to the membranal suture, see introduction of genus Enithares. Parameres see introduction of genus Enithares. Rostral prong a pair of lateral outgrows of the third rostral segment in male Anisops (Fig. 10). Stridulatory comb a ridge with teeth near the base of the fore tibia of male Anisops (Fig. 6). The apex of the comb is towards the posterior (i.e. the concave, flexor) margin of the tibia. Synthlipsis he shortest distance between eyes posteriorly (Fig. 1). Tylus the area of the frons just above the rostrum (Fig. 10); formerly called facial tubercle (Brooks 1951) actually it is the clypeus which is entirely fused with the frons. (Metasternal) xiphus (= metaxiphus) median, more or less arrow head-shaped projection just anteriorly of the hind coxae medially on metasternum (Fig. 49).
Abbreviations asl. = above sea level AT = A. Thanyakam BK = Bomee Kavinsekan brach. = brachypterous CD = C. Duangsupa CMC = Chiang Mai City
CMU = Chiang Mai University Dist. = district ex(x). = specimen(s) KB = K. Boonthavon leg. = collected by Lv. = larva(e)
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 245
macr. = macropterous MFLU = Mae Fah Luang University NN = N. Nieser NP = National Park PC = P.-p. Chen PL = P. Leksawasdi Prov. = province RT = R. Thapa SC = Sombat Chuenchooklin
SU = S. Chumphongphan SP = S. Piyaprichart SW = Siriwat Wongsiri TJ = T. Jongjitvimol TH = T. Jamyanya WJ = W. Jaiyai WU = W. Juntawong WW = W. Wattanachiyingcharoen
ACKNOWLEDGEMENTS The field work during which most of the material reported in this publication was collected by the first two authors in 2001, 2002 and 2004/5 was sponsored by grants from the Uyttenboogaart-Eliasen Stichting. We thank Dr. N. Tuwanon (president of Chiang Mai University) and Dr. C. Kasemset (Director of the Institute for Science and technology Research and Development, Chiang Mai University) for supporting the making of an inventory of water bugs of northern Thailand. Many colleagues helped with work in the field, they are mentioned in the list of abbreviations above. We want to thank especially Sawang Priyaprichart for his extensive logistical support including creating the opportunities to collect in various National Parks and other restricted areas.
SYSTEMATIC PART KEY TO SUBFAMILIES AND GENERA OF NOTONECTIDAE OCCURRING IN INDOAUSTRALIA 1. -. 2.
-. 3.
Hemielytral commissure with a definite hair-lined pit at anterior end (Figs 1, p; 25, pi) Anisopinae ...............................................................................................2 Hemielytral commissure without a hair-lined pit at anterior end Notonectinae.. .............................................................................................................................4 Coxal plates of hind legs bare (Fig. 2), occasionally distally fringed with black hairs. Male fore tibia proximally with a row of stridulatory pegs usually on a stridulatory ridge (Fig. 6) .........................................................................................................3 Coxal plates of hind legs covered with long black hairs (Fig. 3). Males without stridulatory pegs on fore tibia. [Australia and Thailand] ........................Paranisops Antennae three-segmented. Male, rostrum with a pair of prominent lateral prongs (Fig. 10); fore tibia proximally with stridular pegs packed closely to form a comb placed on a stridulatory ridge (Fig. 8). [Palaeo-tropical and -subtropical] ............. ................................................................................................................... Anisops
246 N. Nieser, P.-p. Chen & P. Leksawasdi
-.
4. -. 5. -. 6. -.
Antennae two-segmented. Male, rostrum without lateral prongs; fore tibia proximally with stridulatory pegs clearly separate and not on a stridulatory ridge (Fig. 9) [Australia] ...................................................... Walambianisops Lansbury, 1984 Anterolateral margins of prothorax foveate (Figs 41, 63) .....................................5 Anterolateral margins of prothorax not foveate. [Cosmopolitan but rare in tropical areas] .......................................................................................................Notonecta Mid femur with a pointed protuberance distally (Fig. 5). [Palaeotropical] ............ ................................................................................................................Enithares Mid femur without a pointed protuberance .........................................................6 Eyes in dorsal view caudally holoptic, forming an ocular commissure (Fig. 63) [Palaeotropical] ........................................................................................... Nychia Eyes in dorsal view widely separated caudally (Fig. 40). [SE Asia] ......Aphelonecta
SUBFAMILY ANISOPINAE Anisops SPINOLA, 1837 This genus has been revised by Brooks (1951) who provides a key to species for males. Since then many species have been added, including one described from Melaka (Leong 1963) and one described from Laos (Nieser et al. 2004). In addition the distribution of various species has been found to be much more extensive than reported by Brooks (1951). Synonymy is abbreviated, notably synonyms mentioned by Brooks (1951) are not repeated. Including the four species newly recorded in this paper, there are nine species actually recorded from Thailand. Definite identification of Anisops species is usually possible only for males which have several distinguishing secondary sexual characteristics, notably length and shape of the rostral prong and number and shaped of teeth in the stridulatory comb (see terms used). However, a key to females of Anisops actually found in Thailand has been provided. The easiest character to use for sexing a sample is in the tarsus of the front leg which is one-segmented in males and two segmented in females (Figs 6, 7). A character used in identification of females is the number of spines on the femur of the hind leg which was first used in Buenoa by Truxal (1953). The posterior margin of the hind femur has two rows of spines: one ventrally consisting of numerous small spines and one dorsally consisting of a smaller number of somewhat larger spines (Figs 21-24). The number of spines in the dorsal as well as the ventral row may differ in some closely related species and can be used to separate their females. The colour of Anisops species in dorsal view tends to degenerate into a uniform sordid pale grey or whitish in specimens killed and/or stored in alcohol. Living and freshly killed specimens have a lustrous or pearly appearance due to the shiny translucent wings overlying the abdomen. The abdominal venter is uniformly blackish with median keel and lateral margins yellowish. As killing and keeping specimens in alcohol
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 247
a s
w p c
2
3
4 1 co s p
f
tr
ti
5
ta1 ta2
6
ta
ta3
7
ta
r
cl
8
9
Fig. 1: Anisops kuroiwae, male, length 6.4 mm; a = anterior width of vertex, h = cephalic horn, p = hair lined pit, s = synthlipsis (adapted from Chen et al. 2005). Figs 2-3: Ventral view of the thorax in Anisopinae: 2 = Anisops, 3= Paranisops; c = coxal plate (from Chen et al. 2005). Fig. 4: Larva of Enithares, dorsal view; w = wing pads (adapted from Chen et al. 2005). Fig 5: Middle leg of Enithares, illustrating the various leg segments; cl = claws, co = coxa, f = femur, p = subapical protuberance of femur, ta1, ta2, ta3 = first, second and third tarsal segment, ti = tibia, tr = trochanter (from Nieser, 2004). Figs 6-7: Fore leg of Anisops kuroiwae: 6 = male, 7 = female; s = stridulatory comb, ta = tarsus (adapted from Chen et al. 2005). Figs 8-9: Base of fore tibia in male Anisopinae: 8 = Anisops sp., 9 = Walambiopsis; r = stridulatory ridge with comb (8 adapted from Chen et al. 2005; 9 adapted from Lansbury 1984).
248 N. Nieser, P.-p. Chen & P. Leksawasdi
is common practice in collecting aquatic insects little attention is paid to colour in this genus. However, when one collects Anisops in alcohol it is often worthwhile to note the colour in the field.
KEY TO MALES OF ANISOPS SPECIES IN THAILAND AND ADJACENT REGIONS 1. -. 2. -.
3. -. 4. -. 5. -. 6.
-.
7. -. 8. -. 9. -.
Interocular space anteriorly produced into a cephalic projection (Fig. 1) .............2 Interocular space anteriorly rounded, truncate or indented, not produced into a cephalic projection ...............................................................................................4 Cephalic projection in dorsal view rounded at apex, in lateral view extending in front of eye less by than half the total length of the frons (Figs 1, 13) .................3 Cephalic projection in dorsal view more or less acuminate at apex, in lateral view extending in front of eye by half or more the total length of the frons (Fig. 11) ... .............................................................................................................. A. bouvieri In frontal view, frons diverging ventrally and with a lateral carina over its entire length (Fig. 20) .................................................................................... A. kuroiwae In frontal view frons narrowest halfway between the eyes, with a weak lateral carina in its dorsal part only (Fig. 19) ............................................................... A. nasutus In dorsal view eyes holoptic in posterior half (Figs 25, 26)....................A. breddini In dorsal view eyes not holoptic ...........................................................................5 Tylus medially excavate (Fig. 17) ..........................................................................6 Tylus flat or somewhat swollen, not medially excavate .........................................7 Length less than 5.5 mm. In frontal view, excavation of tylus laterally with single carinae which run up to ventral quarter of eyes and do not meet (Fig. 17); in dorsal view anterior margin of head smooth .......................................................A. niveus Length over 5.5 mm. In frontal view, excavation of tylus laterally with double carinae which run up to halfway the eyes where the outer ones meet in an acute apex (Fig. 18); in dorsal view anterior margin of head sharply indented....... A. ogasawarensis Rostral prong shorter than third rostral segment, originating in its proximal third (Fig. 14) ................................................................................................................8 Rostral prong longer than third rostral segment, originating in its distal third (Fig. 15) ......................................................................................................................12 Synthlipsis one fifth or less the anterior width of vertex; length up to 4.8 mm ..... ............................................................................................................... A. exiguus Synthlipsis about one third or more the anterior width of vertex, length usually over 5.0 mm .........................................................................................................9 Length over 6.5 mm ..........................................................................................10 Small species, length up to 5.0 mm; stridulatory comb on fore tibia with about 10 teeth (Figs 32, 33) ........................................................................... A. brooksianus
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 249
p
16
r
t
10
11
13
12
14
17
15
18
20 19 Figs 10-16: Head of male Anisops spp. in lateral view: 10 = A. barbatus, 11 = A. bouvieri, 12 = A. tahitiensis, 13 = A. kuroiwae, 14 = A. exiguus, 15 = A. lansburyi; 16 = A. brooksianus; p = rostral prong (dotted in A. barbatus), r = rostrum, t = tylus (from Nieser 2004). Figs 17-20: Head of male Anisops spp. in frontal view: 17 = A. niveus; 18 = A. ogasawarensis; 19 = A. nasutus; 20 = A. kuroiwae (17, 19, 20 from Nieser 2004).
250 N. Nieser, P.-p. Chen & P. Leksawasdi
10. Length 8.5 mm or more; tylus swollen, with a pair of tufts of bristles which reach the base of the labrum (Fig. 10).............................................................A. barbatus -. Length up to 8.3 mm; tylus flat of slightly swollen, without tufts of bristles .....11 11. Fore femur distally narrow, its dorsal margin smooth, without subapical impression (Fig. 27); stridulatory comb with about 20 teeth increasing in width from base to apex ..................................................................................................... A. campbelli -. Fore femur broadly rounded distally, its dorsal margin with a wide shallow indentation in distal third (Fig. 28); stridulatory comb with 20-25 teeth, decreasing in width from base to apex ...................................................................... A. occipitalis 12. Synthlipsis very narrow, one seventh or less the anterior width of vertex .............. ............................................................................................................ A. lansburyi -. Synthlipsis one third or more the anterior width of vertex.................................13 13. Along the median longitudinal axis the head is less than one half the length of pronotum; tylus and frons not carinate, in lateral view frons extending in front of the eyes over nearly its entire length; stridulatory comb on fore tibia with about 22 teeth of which the apical 14 are distinctly longer than the basal 8 which are differently orientated (Fig. 30) .......................................................A. nigrolineatus -. Along median longitudinal axis the head is more than one half the length of pronotum; tylus and adjacent part of the frons carinate, in lateral view dorsal half of frons not extending in front of eyes (Fig.16); stridulatory comb of about 25-32 teeth which decrease from base to apex (Fig. 31) .............................. A. tahitiensis
KEY TO FEMALES OF SPECIES OF ANISOPS ACTUALLY RECORDED FROM THAILAND 1. -. 2. -. 3. -. 4. -. 5. -.
In dorsal view eyes holoptic posteriorly (Fig. 25) ..................................A. breddini Eyes not holoptic, synthlipsis at least 0.14 times the anterior width of vertex .....2 Large species, length over 8.0 mm, width over 2.5 mm; synthlipsis very wide, over 0.6 times the anterior width of vertex ...................................................A. barbatus Smaller species, length less than 7.0 mm, width less than 2.0 mm; synthlipsis narrower, 0.5 times the anterior width of vertex or less .............................................3 Synthlipsis less than 0.25 times the anterior width of vertex ...............................4 Synthlipsis over 0.3 times the anterior width of vertex ........................................5 Posterior margin of hind femur with 40-50, rather evenly spaced, spines in ventral row (Fig. 24) ........................................................................................ A. lansburyi Posterior margin of hind femur with 20-30 spines in ventral row which are mostly placed in the distal half of the femur (Fig. 23) ....................................... A. exiguus Along the median longitudinal axis the head is less than one half the length of pronotum .......................................................................................A. nigrolineatus Along the median longitudinal axis the head is two thirds or more the length of pronotum .............................................................................................................6
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 251
6.
Number of spines in dorsal row on posterior margin of hind femur 10-15 (Fig. 22) ........................................................................................................................7 Number of spines in dorsal row on posterior margin of hind femur 20-30 (Fig. 21) ........................................................................................................................8 Along the median longitudinal axis the length of head is two thirds the length of pronotum; body length 5.4-6.3 mm .................................................... A. kuroiwae
-. 7.
21 22
23
24
h pr
31
s
29 pi
25
26 30
27
28
32 33
Figs 21-24: Left hind femur of Anisops spp., illustrating the spines on hind margin: 21 = A. bouvieri, 22 = A. kuroiwae, 23 = A. exiguus; 24 = A. lansburyi; scales 1 mm; 21, 22 dorsal, 23, 24 ventral view. Figs 25-26: Anisops breddini, anterior part of body in dorsal view: 25 = brachypterous form, 26 = macropterous form; h = eyes posteriorly holoptic, pi = hair lined pit, pr = pronotum, s = scutellum (adapted from Nieser 2004). Figs 27-28: Outline of fore femur in male Anisops spp.: 27 = A. campbelli, 28 = A. occipitalis, arrow indicates shallow subapical indentation dorsally (27 adapted from Brooks 1951). Figs 29-32: Stridulatory combs of male Anisops spp.: 29 A. lansburyi, 30 = A. nigrolineatus, 31 = A. tahitiensis, 32 A. brooksianus (from Nieser 2004 and Nieser et al. 2004). Figs 33: Anisops brooksianus, male, fore leg (from Nieser et al. 2004).
252 N. Nieser, P.-p. Chen & P. Leksawasdi
-. 8. -.
Along the median longitudinal axis the length of head is subequal to the length of pronotum; body length 4.8-5.6 mm .........................................................A. niveus Width of head 5.5-6 times the anterior width of vertex, synthlipsis about half the anterior width of vertex ...................................................................... A. tahitiensis Width of head 4 times the anterior width of vertex, synthlipsis about one third the anterior width of vertex ......................................................................... A. bouvieri
Anisops barbatus BROOKS Anisops barbata Brooks, 1951: 387-389. Anisops barbata; Lansbury 1964a: 211. Anisops barbatus; Nieser 2004: 84-85. Material examined. - Thailand: CHIANG RAI Prov. Muang Dist., campus of MFLU, pond in front of C-building, 6.X.2004, artificial pond, recently constructed, margins composed of large stones with a thin layer of mud on top, no vegetation, sampled at edge, leg. PC & NN, CN0435, 1♀; campus of MFLU, pond at Chinese Language and Culture Centre, 30.X.2004, artificial pond, 1.5 m deep, with fountain and sparse lotus plants, leg. PC & NN, CN0454, 12♂ 9♀. NAN Prov., Pua Dist., 5 km NE of Pua town, 19.XI.1994, lotus pond by a bus stop, leg. PC & SP, 2♂ 1♀. Vietnam: 40 km NW An Khe Buon Luoi, 14°10’N 108°30’E, 620-750 m asl., 28.III.1995, leg. Patchulátko & Dembicky, 1♂ (NHMW).
Distribution and habitat. - A widespread species recorded from India through SE Asia (Burma, Southern Shan States; Laos; Malay Peninsula) to Sumatra, Java and SW China (including Taiwan). This species was first recorded from Thailand by Hanboonsong et al. (1996) from Mung River in Ubon Ratchathani province. Judging from our collections this may be a pioneer species colonizing recently created ponds with little or no vegetation. Diagnosis. - Macropterous form. The largest species occurring in Thailand (but see under A. occipitalis). Length, male 8.5-9.3, female 8.0-9.1; width, male 2.7-3.0, female 2.8-3.1. Colour. Head and pronotum pale yellowish, eyes light brown to castaneous. Scutellum pale orange. Abdominal dorsum variably marked with dark brown to blackish, shining through the translucent wings. Width of head about eight times the anterior width of vertex; synthlipsis wide, at least two thirds the anterior width of vertex. Male. In dorsal view, anterior margin of head truncate; width of head seven to eight times the anterior width of vertex; rostral prong slightly shorter than third rostral segment, originating in proximal third of third rostral segment (Fig. 10). Tylus somewhat swollen, with a pair of tufts of bristles which reach the base of the labrum. Fore femur narrowed at apex; stridulatory comb on foretibia consisting of 20-25 comparatively broad teeth. Female. Width of head 5 to six times the anterior width of vertex. Tylus slightly swollen, without tufts of hairs. Brachypterous form not known.
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Comparative notes. - This species is within the Thai fauna recognizable by its size, some small females may be as long as large females of A. occipitalis but they are broader (see under A. occipitalis).
Anisops bouvieri KIRKALDY Anisops bouvieri Kirkaldy, 1904: 116-117. Anisops bouvieri; Brooks 1951: 430-432 (redescription). Anisops bouvieri; Nieser 2004: 85. Material examined.- Thailand: CHIANG RAI Prov. Muang Dist., Boonrod Beverage Company, Mount Phangan, stream, 15.I.1995, leg. PC & Anonymous, C9504, 3♂ 2♀; campus of MFLU, pond at Chinese Language and Culture Centre, 30.X.2004, artificial pond, 1.5 m deep, with fountain and sparse lotus plants, leg. PC & NN, CN0454, 5♂ 2♀; Wiang Chai Dist., Nong Luang lake, 3.XI.2004, large lake, banks partly artificially strengthened with stones, vegetation Eichhornia and Mimosa, bottom muddy with plant debris, water clear to slightly opaque, sampled at edge up to 0.5 m deep, leg. PC, NN, RT & SC, CN0459, 3♂ 2♀ 5Lv.; CHIANG MAI Prov., Chom Thong Dist., Baan Mae Jam Fang Khaw, Ping River, under bridge, 82 Km SW CMC, 3.II.2002, leg. PC, NN, CD & AT, C0227, 1♂ 1♀; at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 1♂ 1♀; LAMPANG Prov., Hang Chat Dis., Thai Elephant Conservation Centre, 30 km N of Lampang City, 400m asl.,11.II.2002, ponds in mountain stream, polluted by elephant waste, leg. PC, NN, CD, AT & WJ, C0233, 2♂ 1♀; SAKON NAKHON Prov., constructed wetland for water treatment near Nong Haan, open pond with marginal vegetation, 23.XI.1995, leg. NN, 1♂ 2♀; KRABI Prov., Ao Luk Dist., P.N. Mountain resort, tiled basin with fountain in front of resort building, on Nymphaea-like plant, 20.XI.2004, leg. PC, NN & SP, CN0489, 2♂, 3♀.
Distribution and habitat. - A widespread species, recorded from India through SE Asia to southern China and New Guinea. In Thailand it has previously been recorded from Bangkok (Brooks 1951) and Chiang Mai (Lansbury 1962). This species seems to have a preference for artificial or otherwise ecologically disturbed ponds with no or sparse vegetation. Diagnosis. - Macropterous form. A medium sized, spindle-shaped, sordid whitish species with its greatest body width somewhat anteriorly of the middle. Length male 6.06.3, female 5.7-6.0, width male 1.5-1.8, female 1.7-1.8. Colour. Sordid whitish overlaid by the shiny translucent wings. Eyes light brown to light grey. Apex of abdomen dorsally black. Width of head slightly more than three times the anterior width of vertex. Posterior margin of hind femur with 23-28 spines in dorsal row becoming slightly smaller and more densely packed distally (Fig. 21); ventral row with 35-50 small spines. Male. In dorsal view the head has a rather long cephalic projection with an acute apex, in lateral view the cephalic projection is extending about half the length of the frons anteriorly of eyes (Fig. 11). Synthlipsis narrow, 0.20-0.25 times the anterior width of vertex; width of head 3.5 times the anterior width of vertex (1.4/0.4). Tylus medially
254 N. Nieser, P.-p. Chen & P. Leksawasdi
deeply excavate, the lateral rims grooved, these grooves running upward to the apex of the cephalic projection. Labrum with three narrow tufts of hairs, one at each basal angle and one at the apex. Rostral prong slightly shorter than third rostral segment, originating in the proximal third of third rostral segment (Fig. 11). Fore femur narrowed at apex. Stridulatory comb on fore tibia consisting of about 12 teeth, which become shorter towards apex. Distally of the tibial comb four to five normal (not clavate) bristles. Female. In dorsal view the head is rounded anteriorly with the vertex very slightly produced. Tylus somewhat swollen. Labrum with some short hairs, not united into tufts as in the male. Synthlipsis about one half the anterior width of vertex. Brachypterous form. Not known. Comparative notes. - Males are at once recognized by the structure of the cephalic projection. In A. sardeus Herrich Schäffer, 1850 distributed from Africa and the Mediterranean to India and Burma, males have a similar cephalic projection. However, the fore tibia of males of A. sardeus have about three distinctly clavate bristles along anterior margin, well visible at a magnification of about 30x. In addition there are only two tufts of bristles placed in the basal angles of the labrum, directed forward in a curve and meeting apically thus forming a loop. Females of A. bouvieri are very similar to those of A. kuroiwae which has been found in the same locality as A. bouvieri in over half our samples. A difference is found in the number of dorsal spines on the posterior margin of hind femur which is 23-28 in A. bouvieri and 11-15 in A. kuroiwae (Figs 21, 22).
Anisops breddini KIRKALDY Anisops breddini Kirkaldy, 1901: 5-6. Anisops breddeni Brooks, 1951: 439-441 (misspelling, redescription). Anisops breddini; Nieser 2004: 85-86. Anisops kempi Brooks 1951: 441-442 (synonymized by Nieser 2004). Material examined. - Thailand:CHIANG RAI Prov.,Muang Dist., Boonrod Beverage Company, Mount Phangan, stream, 15.I.1995, leg. PC & Anonymous, C9504, 1♀ macr.; Chiang Saen Dist., Baan Sop Ruank, polluted pond at Thai side of Golden Triangle, 26.I.2002, leg. PC, CD, AT & WJ, C0214, 1♂1♀ brach.; Muang Dist., Baan Khun Khon, 33 Km SW Chiang Rai City, ponds behind hill tribe house, 23.II.2002, leg. PC, NN, CD, AT, C0243, 2♀ brach.; same, 16.X.2004, leg. PC & NN, CN0444, 2♀ brach.; Campus of MFLU, ponds, 13-22.X.2004, leg. PC & NN, CN0440, CN0446 2♀ brach., 2Lv.; Wiang Chai Dist., Don Sila Subdist., ditch without current along road, width 6m, no aquatic vegetation but at places overgrown with terrestrial plants, 3.XI.2004, leg. PC, NN, RT& SU, CN0458, 2♀ brach., 1♀ macr.;Wiang Chai Dist., Nong Luang lake, 3.XI.2004, large lake, banks partly arificially strengthened with stones, vegetation Eichhornia and Mimosa, bottom muddy with plant debris, water clear to slightly opaque, sampled at edge up to 0.5 m deep, leg. PC, NN, RT & SC, CN0459, 1♀ macr.; CHIANG MAI Prov., Ban Mae Kap Nam Mae To, 98°37’E, 18°51’N, 600m asl., 14.III.1992, leg. F. Malicky, 1♀ macr. (NHMW); Doi Saket Dist., Hui Hong Khrai, 30 Km N CMC, Royal Development Study Centre, 500m asl., big and deep open lake for rearing fish, 1.XII.1994, leg. PC & S. Wongsiri,
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numerous specimens, 8♂ 12♀ brach. collected; San Kamphaeng Dist., Ban Rong Wow Daeng village, 25 Km SE of CMC, two polluted ponds behind a rose farm, 22.I.2002, leg. PC, CD & AT, C0209, 1♂ brach.; Muang Dist., pond beside road near department of Physics, campus CMU, muddy shallow pond, surrounded by palm trees, 13.II.2002, leg. PC & NN, C0234, 1♂ 5♀ brach.; Saraphi Dist., pond at side of highway 11, 14 Km S of CMC, 14.II.2002, leg. PC, NN, CD & AT, C0236, 12♂ 7♀ brach., 1♂ macr., 21Lv.; Muang Dist., stream near entrance of Chiang Mai Zoo, virtually stagnant part at bridge to CMU, bottom soft black mud (0.05-0.1 m thick) on sand, 1♂ macr.; at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 1♂ brach., 1♂ macr.; ponds on campus of faculty of Food-engineering and Agro-industry, CMU, 14.XII.2004, leg. PC & NN, 1♀ brach.; PHAYAO Prov., Muang Dist., pond on campus of Naresuan University, 24.IX.2004, leg. PC, NN & SP, C0432, 4♂ 3♀ brach., 5Lv.; LAMPANG Prov., Amuan Agricultural Experimental and Training Station, pond surrounded by weeds, 10.I.1995, leg. PC & SW, 11♂ 9♀ brach.; Mount Lung Khuntan, pond in Elephant Training Centre, 27.III.2001, leg. PC, SW & BK, C0140, 4♂ brach.; SUKOTHAI Prov., Taleh Luang, large lake, sampled at edge between Mimosa, many snails, 18.IX.2004, leg. PC, NN, SC & WU, CN0417, 11♂ 19♀ brach., 5♂ macr., 9Lv.; Sisaamrong Dist., roadside ditch of flooded agricultural fields near Sukhotai City, 18.IX.2004, leg. PC, NN, SC & WU, C0418, 1♀ macr.; PHITSANULOK Prov., Muang Dist., outskirts of Phitsanulok City, lotus pond at Amarin Hotel, 7.III.2001, leg. PC, SP & WW, C0113, 2♂ brach., 44♂ 33♀ macr., 12Lv.; outskirts of Phitsanulok City, Biological Control Centre, frog rearing pond, 7.III.2001, leg. PC, SP & WW, C0114a, 1♂ brach., 2♂ 4♀ macr.; outskirts of Phitsanulok City, fish pond near agricultural station of Naresuan University, 7.III.2001, leg. PC, SP & WW, C0115, 1♂ brach., 1♂ macr.; Muang Dist., ponds on campus of Naresuan University, 8.IX.2004, leg. PC & NN, 1♀ macr., 1Lv; Phitsanulok City, house of S. Piyaprichart, at light, 9.XI.2004, leg. PC, NN & SP, CN0460, 1♀ macr.; PHICHIT Prov., Nachilaparami Dist., 18 Km S of Pichit City, abandoned rice fields and road ditch, 15.IX.2004, leg. PC, NN, KB & TJ, C0414, 4♂ brach., 14Lv.; NAKHON SAWAN Prov., Muang Dist., Beung Boraphet Lake, canal in marsh near office station of National Park, 15.IX.2004, leg. PC, NN, KB & TJ, CN0412, 4♂ 3♀ brach.; Muang Dist., Sawan park in the centre of Nakhon Sawan City, small lake with edge of Poaceae, which are trimmed and kept neat, 15.IX.2004, leg. PC, NN, KB & TJ, C0413, 1♀ brach.; SAKON NAKHON Prov.,constructed wetland for water treatment near Nong Haan, open pond with marginal vegetation, 23.XI.1995, leg. NN, N9525, 6♂ 7♀ brach., 2♂ macr.; same, overgrown marshy pond, Eichornia, Hydrilla, Ipomoea, Lotus, 23.XI.1995, leg. NN, N9526, 10♂ 8♀ brach., 1♂ 1♀ macr., 3Lv.; UDON THANI Prov., W of Udon Thani City, roadside pond choked with Eichornia, 27.XII.1994, leg. PC et al., C09428, numerous sepcimens; SURIN Prov., Ban Lak Wo, 5m wide stream, 200m asl., 2.I.1997, leg. P. Schwendinger, 1♂ macr. (NHMW); CHON BURI Prov., Khao Khaew Open Zoo, ponds, 6.IV.2001, leg. PC, B. Kavinseksan & S. Leepitakrat, C0157/8, 5♂ 10♀ brach.; KRABI Prov., Ao Luk Dist., P.N. Mountain resort, tiled basin with fountain in front of resort building, on Nymphaea-like plant, 20.XI.2004, leg. PC, NN & SP, CN0489, 1♀, macr.; PHATTALUNG Prov., Muang Dist., Kuttasawan, pond near Taleh Luang, shaded pond in small stand of woods, a Convolvulaceae-like plant growing into the water, no aquatic vegetation, 13.XI.2004, leg. PC, NN & SP, CN0468, 10♂ 6♀ brach., 21♂ 20♀ macr., 25Lv.
Distribution and habitat. - A widespread species recorded from India and Sri Lanka through SE Asia to Java, Sulawesi and New Guinea (Lansbury 1961; Nieser & Chen
256 N. Nieser, P.-p. Chen & P. Leksawasdi
1991). Although this is the commonest species of Anisops in Thailand and Nieser (1998) recorded this species from Thailand in his table there seems to be no previous record with locality data from Thailand. A. breddini is a typical species of man-made ponds and ditches, it is found often between vegetation. Its life history was studied by Leong (1962) in the Malay Peninsula where it is also the commonest species of Anisops (Nieser, 2004). Diagnosis. - Brachypterous form. A medium sized, slender, pale yellow to whitish species, length male 5.7-6.6, female 5.4-6.8; width male 1.2-1.6, female 1.2-1.7. Eyes holoptic posteriorly (Fig. 25); vertex projecting slightly in front of the eyes in dorsal view. Tylus smoothly convex. Width of head about equal to width of pronotum, five to six times the anterior width of vertex. Lateral margins of pronotum parallel (Fig. 25); basal width of pronotum 0.6-0.7 times the posterior width of the pronotum. Hemielytra with reduced membranes; hind wings reduced to club-shaped elongate straps. Males. Rostral prong slightly longer than third rostral segment, second rostral segment with a distal outgrowth posteriorly, which reaches the tip of the rostrum. Fore femur strongly swollen. Stridulatory comb on foretibia consisting of a few irregular teeth. Female. Second rostral segment with a short distal outgrowth posteriorly, which does not reach halfway the third rostral segment. Macropterous form. Length male 5.7-6.6, female 5.4-6.7; width male 1.3-1.6, female 1.4-1.7. Vertex not or hardly projecting beyond anterior margin of eyes. Lateral margins of pronotum slightly diverging posteriorly; humeral width of pronotum slightly larger than width of head (Fig. 26); basal width of scutellum about 0.9 times the humeral width of pronotum. Hemielytra with fully developed membrane. Hind wings fully developed. Other structural characters, including secondary sexual characters of male and female as in brachypterous form. Comparative notes. - This species can be recognized immediately by the posteriorly holoptic eyes in both sexes.
Anisops brooksianus NIESER et al. Anisops brooksianus Nieser, Chen, Leksawasdi, Thanyakam & Duangsupa, 2004: 36-37. Material examined. - Laos: CHAMPASAK Prov., Bolavens Plateau, Paksong, 1260m asl., 27.XII. 1996, leg. P. Schwendinger, 1♂1♀ paratypes.
Distribution. - Only known from the type series from Laos. Diagnosis. - Macropterous form. A small blackish species, length male 4.9, female 5.2-6.1; width male 1.3, female 1.4-1.6. In dorsal view the head is anteriorly truncate with vertex not or very slightly indented, width of head five to six times the anterior width of vertex. Tylus flat, smooth without specialized hairs. Posterior margin of hind femur with about 30 spines in dorsal and about 40 spines in ventral row.
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Male. Synthlipsis 0.3 times the anterior width of vertex. Rostral prong (Fig. 16) about half the length of third rostral segment, originating near proximal end of third segment, apex of rostrum rounded. Stridulatory comb on fore tibia (Fig. 32) consisting of 11 teeth of approximately the same size. Fore tarsus without small pegs on inner surface (Fig. 33). Female. Synthlipsis 0.4 times the anterior width of vertex. Brachypterous form unknown. Comparative notes. - A. brooksianus is similar to A. exiguus and A. lansburyi, which both have a narrower synthlipsis. The male of A. lansburyi has a longer rostral prong; the male of A. exiguus has five small spines on inner surface of fore tarsus, which are lacking in A. brooksianus.
Anisops campbelli BROOKS Anisops campbelli Brooks, 1951: 322-324.
Distribution. - India and Burma (Southern Shan States). We have not seen this species which has not been recorded since its original description. In view of its occurrence in Burma it might occur in NW Thailand, however, this is the best researched area for aquatic Heteroptera of Thailand and so far this species has not been found there. Diagnosis. (after Brooks 1951). - A large, brown to grey species, length, male 7.3-8.1, female 8.1-8.4; width both sexes 2.2-2.5. In dorsal view anterior margin of head rounded with vertex extending slightly in front of eyes; width of head three to four times the anterior width of vertex; synthlipsis about two thirds the anterior width of vertex. Tylus flat. Male. Rostral prong about as long as third rostral segment, originating halfway third rostral segment which has a procumbent tuft of hairs along each side of the dorsal surface. Apex of fore femur distally narrowed (Fig. 27). Stridulatory comb on fore tibia with approximately 18 teeth which increase in size from base to apex. Comparative notes. - Slightly smaller than A. barbatus, males of which have two tufts of bristles on tylus and a stridulatory comb of 20-25 teeth of subequal size. A. occipitalis which is of the same size has the apex of anterior femur broadly rounded (Fig. 28).
Anisops exiguus HORVÁTH, first record Anisops exigua Horváth, 1919: 314. Anisops exigera Brooks, 1951: 447-449 (misspelling, redescription). Anisops exiguus; Nieser 2004: 86. Material examined. - Thailand: CHIANG MAI Prov., Muang Dist., at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open
258 N. Nieser, P.-p. Chen & P. Leksawasdi
vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 6♂ 6♀; PHRAE Prov., roadside pond somewhat downstream of Mae Kam waterfall; 20x10m, water turbid light brown due to suspended silt; sample from edge with shade, some dead plant material haning in the water, bottom very soft mud with some plant debris, depth unknown, 16.XI.1995, leg. NN, N09516, 2♂; KANCHANABURI Prov., Thong Pha Phum NP, stream at Jed Mit, 9.II.2002, leg. S. Mongchoichana, 1♂; KRABI Prov., Ao Luk Dist., P.N. Mountain resort, tiled basin with fountain in front of resort building, on Nymphaea-like plant, 20.XI.2004, leg. PC, NN & SP, CN0489, 9♂ 8♀. First record for Thailand. Laos: Central Laos, Viangchang prov., Phou Khao Khouay NP, Nam Leuk near Tad Leuk waterfall, leg. Schilhammer, 1♂ (NHMW). First record for Laos. Indonesia: Sumatra, stream, 2°47’N, 99°14’E, leg. F. Malicky, 1♂ (NCTN). First record for Indonesia.
Distribution and habitat. - A widespread species recorded from Papua New Guinea, Sumatra, SE Asia and India. Although widespread this species is relatively rarely collected. The few habitat data point to a preference for temporary ponds. Diagnosis. - Macropterous form. A small, sordid white to brown species with greyish brown eyes and abdomen dorsally blackish; length, male 4.3-4.8, female 4.35.3; width, both sexes 1.2-1.4. In dorsal view the head is anteriorly truncate with vertex slightly indented; width of head about six times the anterior width of vertex. Synthlipsis very narrow, 0.12-0.2 times the anterior width of vertex. Posterior margin of hind femur with 18-20 spines in dorsal and 22-31 spines in ventral row. Male. Tylus slightly swollen with a smooth surface. Rostral prong slightly shorter than third rostral segment, originating in proximal third of third rostral segment (Fig.14). Stridular comb of fore femur with 9-11 teeth, which are longest in the middle. Fore tarsus with five small spines on inner surface. Female. Tylus flat. Brachypterous form unknown. Comparative notes. - At first sight very similar to A. lansburyi, with which it has been found together. Males can be recognized by the rostral prong which is short in A. exiguus and long in A. lansburyi (Figs 14, 15); in addition the stridular comb is lacking the one or two very small teeth at apex present in A. lansburyi. Females can be recognized by the spines on hind margin of hind femur especially the ventral row which in A. exiguus has 22-31 spines and in A. lansburyi 38-50 spines (Figs 23, 24). Apart from the number the placing is also different: in A. exiguus the spines are absent proximally and proximally distinctly more widely spaced than distally; in A. lansburyi the spines are more evenly spaced over the entire length of the femur. Males of A. brooksianus, which are of the same size and have a short rostral prong (Fig. 16) differ in having a wider synthlipsis (0.3 times anterior width of vertex) and lacking the row of about five small spines on the inner surface of the fore tarsus, instead there is one somewhat larger spine near base of fore tarsus (Fig. 33). Females can be separated by the synthlipsis which is 0.4 times the anterior width of vertex in A. brooksianus and 0.2 times the anterior width of vertex in A. exiguus.
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Anisops kuroiwae MATSUMURA Anisops kuroiwae Matsumura, 1915: 109. Anisops batillifrons Lundblad, 1933b: 463-464. Anisops batillifrons; Brooks 1951: 420-423 (redescription). Anisops kuroiwai Miyamoto, 1964: 67-68 (synonymy, unjustified emendation of name). Anisops kuroiwae; Nieser 2004: 86-87. Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Boonrod Beverage Company, Mount Phangan, stream, 15.I.1995, leg. PC & Anonymous, C9504, 12♂ 18♀; Mae Fah Luang Dist., hilltribe village 15 Km N of Maesalong village, 29.X.2004, leg. PC, NN & RT, CN0449, 2♂ 2♀; Muang Dist.,campus of MFLU, pond at Chinese Language and Culture Centre, 30.X.2004, artificial pond, 1.5 m deep, with fountain and sparse lotus plants, leg. PC & NN, CN0454, 28♂ 42♀; Wiang Chai Dist., Nong Luang lake, 3.XI.2004, large lake, banks partly artificially strengthened with stones, vegetation Eichhornia and Mimosa, bottom muddy with plant debris, water clear to slightly opaque, sampled at edge up to 0.5 m deep, leg. PC, NN, RT & SC, CN0459, 1♂ 1♀; CHIANG MAI Prov., Muang Dist., at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 22♂ 15♀ ; Chom Thong Dist., Doi Inthanon NP, 70 km SW CMC, Wang Khwaay waterfall, pools in streambed downstream of waterfall, 31.I.2002, leg. NN, CD & AT, C0223, 1♀; Chom Thong Dist., Baan Luang Subdist., swimming pool at little home resort, 16.I.2005, leg. PC & NN, CN0509, 3♀; NAN Prov., Pua Dist., Ban Hon, 5 Km NE of Pua, lotus pond at busstop, 19.XI.1994, leg. PC & SP, C9409, > 50 specimens; PHRAE Prov., roadside pond somewhat downstream of Mae Kam waterfall; 20x10m, water turbid light brown due to suspended silt; sample from edge with shade, some dead plant material hanging in the water, bottom very soft mud with some plant debris, depth unknown, 16.XI.1995, leg. NN, N09516, 13♂ 18♀; LAMPANG Prov., Hang Chat Dist., Thai Elephant Conservation Centre, 30 km N of Lampang City, 400m asl.,11.II.2002, ponds in mountain stream, polluted by elephant waste, leg. PC, NN, CD, AT & WJ, C0233, 7♂ 12♀; SUKOTHAI Prov., Sisaamrong Dist., roadside ditch at flooded agricultural fields, near Sukhotai City, 18.IX.2004, leg. PC, NN, SC & WU, CN0418, 1♂; UBON RATCHATHANI Prov., Warin Chamrap Dist., pond beside Moon River, 25.XII.1994, leg. PC et al., C9423, 1♂, 3Lv. Laos: 10 Km N of Luang Prabang, Mekhong River area, 250m asl., XI.1992, leg. I. Sonsy, 2♂ 1♀ (NHMW); Vietnam: 40 Km NW of An Khe Buon Luoi, 14°10’N, 108°30’E, 620-750m asl., leg. P. Pacholatko & L. Dembicky, 5♂ 4♀ (NHMW).
Distribution and habitat. - A widespread species, from N India through SE Asia to southern China, Japan (Okinawa and Iriomote) and the Philippines. In Thailand it has been recorded from Chiang Mai (Lansbury 1962) and Udon Ratchathani (Hanboongsong et al. 1996). We have seen only specimens from the northern part of Thailand but as it has been recorded from West Malaysia (Nieser 2004) this species will probably occur in the whole of Thailand. A. kuroiwae was mostly found in man made ponds either without vegetation or with mainly marginal vegetation.
260 N. Nieser, P.-p. Chen & P. Leksawasdi
Diagnosis. - Macropterous form (Fig. 1). A medium sized, pale yellow species. Length, male 5.6-6.4, female 5.4-6.3; width male 1.5-1.6, female 1.3-1.8. Head width four to five times the anterior width of vertex. Posterior margin of hind femur with 1015 spines in dorsal and 40-50 spines in ventral row. Male. In dorsal view the head has a cephalic projection with a rounded apex; in lateral view the cephalic projection extends less than half the total length of the frons in front of the eyes (Fig. 13). In frontal view tylus and frons are excavate with two carinae on each side of which the inner ones meet in the apical (dorsal) sixth of the frons (Fig. 20). Rostral prong slightly longer than third rostral segment, originating in proximal third of rostral segment (Fig. 13). Fore femur apically narrowed. Stridulatory comb on fore tibia with about 13 teeth of which the basal six are much longer than the apical seven. Female. In dorsal view the head is rounded anteriorly; synthlipsis slightly less than one third the anterior width of vertex. Tylus flat, not swollen. Brachypterous form unknown. Comparative notes. - Males are easily recognized by the structure of the cephalic horn, see also under A. ogasawarensis. Females are very similar to those of A. bouvieri with which they have several times been found together. A difference is found in the number of dorsal spines on the posterior margin of hind femur, which is 23-28 in A. bouvieri and 11-15 in A. kuroiwae (Figs 21, 22). Females of A. kuroiwae are also similar to females of A. niveus which have about the same number of spines on the posterior margin of hind femur. Females of A. niveus are usually smaller (length 4.8-5.6) than females of A. kuroiwae (length 5.4-6.3). In addition, the median lengths of head and pronotum are subequal in A. niveus females whereas the median length of head is about two thirds the median length of pronotum in A. kuroiwae females.
Anisops lansburyi LEONG, first record Anisops lansburyi Leong, 1963: 101-102. Anisops lansburyi; Nieser 2004: 87. Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Boonrod Beverage Company, Mount Phangan, stream, 15.I.1995, leg. PC, C9504, 1♂ 1♀; CHIANG MAI Prov., Muang Dist., at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 10♂ 8♀; CHON BURI Prov., Khao Khaew Open Zoo, ponds, 6.IV.2001, leg. PC, B. Kavinseksan & S. Leepitakrat, C0158/9, 1♂ 1♀; SONGKHLA Prov., fishing village near The Pha, small trench draining cocos orchards and meadows, water clear brow, hardly brackish although only 200-300 m from the seafront, 28.XI.1995, leg. NN, N9535, 1♂. First record for Thailand.
Distribution. - So far recorded from the Malay Peninsula and Thailand only.
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Diagnosis. - Macropterous form. A small, sordid white species. Length, male 4.55.6, female 4.6-5.9; width male 1.1-1.4, female 1.1-1.5. Width of head about six times the anterior width of vertex. Hind margin of posterior femur with 10-20 spines in dorsal and 40-50 in ventral row (Fig. 24). Male. In dorsal view the head is rounded with anterior margin truncate and vertex slightly indented. Synthlipsis narrow, 0.10-0.15 times the anterior width of vertex. Rostral prong longer than third rostral segment, originating near its apex (Fig. 15). Tylus somewhat swollen, smoothly convex. Tylus and labrum with a few short scattered hairs. Fore femur narrowed at apex. Tibial comb with eight to nine, rather large, teeth of subequal length and one or two very small teeth at its apex (Fig. 29). Female. Except for sexual characters, females of this species are very similar to males. They are on average slightly larger and the synthlipsis is slightly wider than in males: 0.15-0.25 times the anterior width of vertex. Brachypterous form unknown. Comparative notes. - This species is of the same size as A. brooksianus and A. exiguus. Males of these two species are recognized by their short rostral prongs. Females of A. exiguus have 20-30 spines in ventral row on posterior margin of hind femur whereas A. lansburyi has 40-50 spines in this row (Figs 23, 24). Females of A. brooksianus have the synthlipsis 0.4 times the anterior width of vertex. Remarks. - The range of lengths above is taken from Leong’s original description; the largest specimen we have seen is a female with a length of 5.5 mm.
Anisops nasutus F IEBER Anisops nasuta Fieber, 1851: 60. Anisops nasuta; Brooks 1951: 416-418. Anisops nasutus; Nieser 2004: 87-88.
Distribution. - Widespread from India through the Malay Peninsula to southern China and through Indonesia to Philippines (Mindanao), Australia and Pacific islands (Nieser 2004). Not recorded from Thailand. Diagnosis. - Macropterous form. A medium sized to large pale yellow to greyish species. Length, male 6.0-7.8, female 6.0-7.0; width, male 1.3-18, female 1.4-1.9. Width of head slightly more than four times the anterior width of vertex. Labrum with scattered quite long bristles, more numerous in males than in females. Posterior margin of hind femur with 12-20 spines in dorsal and 40-60 spines in ventral row. Male. In dorsal view the head has a cephalic projection with a broadly indented apex, in lateral view the cephalic projection extends less than half the total length of the frons in front of the eyes. In frontal view the tylus has a wide median depression, appearing almost flat; the frons is narrowed in the middle and has dorsally a median oval depression bordered by a carina (Fig. 19). Synthlipsis about 0.25 times the anterior
262 N. Nieser, P.-p. Chen & P. Leksawasdi
width of vertex. Rostral prong slightly shorter than third rostral segment, originating near its proximal margin. Fore femur apically narrowed. Stridulatory comb on fore tibia with about 14 teeth, which are longest in the middle. Female. In dorsal view the head is rounded anteriorly; the synthlipsis 0.25-0.33 times the anterior width of vertex. Tylus flat. Brachypterous form unknown. Comparative notes. - Males are easily recognized by the structure of the cephalic projection and the frons. Females are very similar to those of A. bouvieri and A. kuroiwae but usually larger (length of females of A. nasutus 6.0-7.0, of A. bouvieri and A. kuroiwae 5.4-6.3). In addition there are 12-18 spines in the dorsal and 47-60 spines in the ventral row on posterior margin of hind femur. In A. bouvieri the dorsal row has 23-28 spines and in A. kuroiwae the ventral row has 40-50 spines (Figs 21, 22).
Anisops nigrolineatus LUNDBLAD Anisops nigrolineata Lundblad, 1933: 160-163. Anisops nigrolineata; Brooks 1951: 409-411 (redescription). Anisops nigrolineatus; Nieser 2004: 88. Material examined. - Thailand: CHIANG MAI Prov., Fang Dist., Doi Ang Khang, Office of forestry department, large pond in stream, stagnant, 30.X.2004, leg. PC, NN & RT, 1♂; W of Mae Rim, Mae Sa waterfall, 30-31.X.1995, leg. H. Zettel, 2♂ 4♀; Muang Dist., Doi Suthep NP, Montatharn Falls, 750-800m asl., 2.XI.1995, leg. H. Zettel, 2♂ 2♀ (NHMW); stream at entrance of Chiang Mai Zoo, near dam at bridge to C MU, virtually stagnant, bottom a 0.050.1m thick layer of black mud on sand, 11.XII.2004, leg. PC & NN., CN0499, 1♂; at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Anisops in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 1♀; Chom Thong Dist., Doi Inthanon NP, 70 km SW CMC, Wang Khwaay waterfall, pools in streambed downstream of waterfall, 31.I.2002, leg. NN, CD & AT, C0223, 1♀; Samoeng Dist., Doi Laung, Mae Tong Ting Village, 800m asl., artificial pond near the station of Chiang Mai regional Forest Division, 29.III.2001, leg. PC, B. Kavinsekan & S. Praiwan, C0146, 4♂ 2♀ 5Lv.; MAE HONG SON Prov., Phaa Boong, ca. 20 Km S of Mae Hong Son City, pool at base of waterfall, sampled in edge with Spirogyra-like Chlorophyta, floating plant debris and plant roots washed out in the water, stagnant, area 3x1, depth 1m, 12.XI.1995, leg. NN 6♂ 2♀; Pang Mapha Dist., Ban Taam Than Lod, 650m asl., 20.XII.1996, leg. P. Schwendinger, 1♀ (NHMW); PHRAE Prov., Baan Huay Kaet (along highway 101 to Nan), virtually stagnant pool in mountain stream, 5-10m wide, depth up to 0.7m, mostly 0.2m, water hyaline, clear, 17.XI.1995, leg. NN, 1♂; PHITSANULOK Prov., Phu Hin Rong Kla NP, near Huai Khamunnoi waterfall, 1100m asl., 6.III.2001, leg. PC, SP & WW, 2♂ 1♀; same stream near bridge of Mung hill tribe village, 1100m asl., 6.III.2001, leg. PC, SP & WW, 14♂ 2♀; PETCHABUN Prov., Nam Nao NP, Huai Ye Krua, near headquarters, 24.XI.1995, leg. H. Zettel, 1♂ (NHMW); SAKON NAKHON Prov., Phu Pan NP, 490-520m asl., 7-8.XII.1995, leg. P. Schwendinger, 1♂ 1♀ (NHMW); CHAIYAPHUN Prov., Phi Kalo NP, tributary to Nam Prom, 13.III.1994, leg. W.D. Shepard (1038), 2♂ (NHMW); UBON RATCHATHANI Prov., Kong Chiam, 16.III.1996, leg. Y. Hanboonsong,
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1♂ 1♀ (NHMW); MUKDAHAN Prov., Phu Pha Toep NP, 31.XII.1999, leg. P. Mazzoldi, 1♂ (NHMW); KANCHANABURI Prov., Thong Pha Phum NP, Somnoi, 9.II.2002, 907m asl., leg. S. Mongchoichana, 7♂ 4♀. Laos: VIANGCHAN Prov., Phou Kao Khouay NP, Nam Leuk, near Tad Leik waterfall, 200m asl., at light, 1-8.VI.1996, leg. H. Schillhammer, 1♀ (NHMW). First record for Laos. Vietnam: Dalat City, 21-27.IV.1994, leg. P. Patchlátko & L. Dembicky, 4♀ 2♀; Nam Cat Tien NP, 1-15.V.1994, leg. P. Patchlátko & L. Dembicky 9♂ 3♀ (NHMW). First record for Vietnam.
Distribution and habitat. - From India through SE Asia to Java and the Philippines. First recorded from Thailand by Hanboonson et al. (1996) from Ubon Ratchathani. This species prefers virtually stagnant parts of streams and pools and ponds associated with streams. Diagnosis. - Macropterous form (Fig. 34). A medium sized, rather slender species with little difference in size between sexes, length male and female 5.4-6.6; width, male and female 1.5-1.8. Fresh specimens are usually blackish with a greyish shimmer due to the translucent hemielytra, the black colour keeps at least some time when specimens are stored in alcohol. Width of head four to five times the anterior width of vertex; synthlipsis wide, about 0.5 times the anterior width of vertex. Tylus flat, and somewhat expanded laterally; labrum smooth without bristles. Pronotum comparatively long, its humeral width about 1.5 times its median length, which is over twice the median length of head. Male. The vertex protrudes in front of the eyes both in dorsal (Fig. 34) and lateral view. Rostral prong longer than third rostral segment, originating proximally on third rostral segment. Fore femur strongly narrowed distally. Stridulatory comb with about 22 teeth of which the apical 14 are long and slender, more than twice the length of the basal 8 which are oriented in a different direction (Fig. 30). Female. Vertex not protruding anteriorly of eyes. Brachypterous form not known, however, according to Brooks (1951) there is a pale form, which we have not seen; it may be either the brachypterous form or specimens stored for long time in alcohol. Comparative notes. - Males can be recognized by the unique structure of the stridulatory comb. Females differ from other Thai species by the ratio synthlipsis/anterior width of vertex in combination with Fig. 34: Anisops nigrolineatus, male, length 6.3 mm. the long pronotum.
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Anisops niveus (FABRICIUS), first record Notonecta nivea Fabricius, 1775: 690. Anisops nivea; Brooks 1951: 373-376. Anisops niveus; Nieser 2004: 88-89. Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Boonrod Beverage Company, Mount Phangan, stream, 15.I.1995, leg. PC & Anonymous, C9504, 4♂; NAN Prov., Pua Dist., Ban Mon village, 5 KM NE of Pua town, lotus pond at bus stop, 19.XI.1994, leg. PC & SP, 1♂; MUKDAHAN Prov., Phu Pha Toep NP. 31.XII.1999, leg. P. Mazzoldi, 1♂ (NHMW); KRABI Prov., Ao Luk Dist., P.N. Mountain resort, tiled basin with fountain in front of resort building, on Nymphaea-like plant, 20.XI.2004, leg. PC, NN & SP, CN0489, 2♂ 2♀; SONGKHLA Prov., fishing village near The Pha, small trench draining coconut orchards and meadows, water clear, light brown, hardly brackish although only 200-300 m from the seafront, 28.XI.1995, leg. NN, N9535, 1♂. First record for Thailand. Vietnam: 40 Km NW of An Khe Buon Loi, 14°10’N, 108°30’E, 620-750m asl., 28.III-12.IV.1995, leg. P. Pacholátko & L. Dembicky, 1♂ (NHMW). Indonesia: SUMATRA, Danau Toba, pond at Arjuna, 21.VII.1994, leg. D.R. Thomas, 1♂ 1♀; Huya Padang, Sungai Sila, 200m asl., 2°47’N, 99°14’E, 13.IV.1997, F. Malicky, 2♂ 2♀ (NCTN).
Distribution. - India through SE Asia to Sumatra. Diagnosis. - Macropterous form. A sordid white to dark grey species, length, male, 4.8-5.1, female 4.8-5.4; width, male 1.2-1.4, female1.3-1.8. In dorsal view the head is obtusely rounded to nearly truncate; width of head four to five times the anterior width of vertex; median length of head subequal to medium length of pronotum. Posterior margin of hind femur with 10-15 spines in dorsal and 35-40 spines in ventral row. Male. Synthlipsis 0.3-0.4 times the anterior width of vertex. In frontal view tylus and frons widely and shallowly excavate with a single lateral carina on each side, the median groove runs up to ventral quarter between the eyes without the carinae meeting (Fig. 17). Rostral prong shorter than third rostral segment, originating proximally on third rostral segment. Labrum with rather long hairs proximally, the lateral ones forming a pair of tufts which curve upwards along the carinae of the excavation of the tylus; the middle ones combined to an erect tuft. Fore femur apically somewhat narrowed. Fore tibia stout, stridulatory comb with about 15 teeth of subequal length. Female. Synthlipsis is 0.35-0.5 times the anterior width of vertex. Tylus flat, not swollen or excavate. Brachypterous form not known. Comparative notes. - Males are immediately recognized by the shape of the frons (Fig. 17). Females by the low number (10-15) of spines in the postero-dorsal row of the hind femur combined by the fact median length of head and pronotum are subequal.
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Anisops occipitalis BREDDIN Anisops occipitalis Breddin, 1905: 152. Anisops occipitalis; Brooks 1951: 344-347 (redescription). Anisops occipitalis; Nieser 2004: 89.
Distribution and habitat. - From West Malaysia through Indonesia to Australia and New Caledonia. Not recorded from Thailand, however, this species has been found in several localities throughout W. Malaysia, and is to be looked for in SE Thailand. This species is reported to colonize isolated and temporary ponds (Fernando & Leong 1976). Diagnosis. - Macropterous form. A large, pale yellowish to grey species. Length, male 6.6-8.1, female 6.6-8.3; width, male 1.6-2.2, female 1.6-2.4. In dorsal view, head anteriorly truncate to slightly rounded, vertex level with anterior margin of eyes; width of head 5-6.5 times the anterior width of vertex. Male. Synthlipsis wide, 0.4-0.5 times the anterior width of vertex. Tylus slightly swollen, labrum covered with short hairs. Rostral prong longer than third rostral segment, originating near its distal margin. Fore femur broadly rounded apically, its dorsal margin with a wide shallow indentation in apical third (Fig. 28). Stridulatory comb with 20-25 teeth decreasing in width from base to apex. Female. Synthlipsis 0.5-0.7 times the anterior width of vertex. Tylus flat, labrum bare. Brachypterous form, not recorded from the area under consideration. However, Lansbury (1965a) suggested that A. leucotheca Esaki (1928), occurring from the Solomon Islands to Samoa, may represent the brachypterous form of this species. Nieser & Chen (2005) found that indirect flight muscles in A. leucotheca from Vanuatu are not developed whereas indirect flight muscles in A. occipitalis from West Malaysia are fully developed, thus supporting Lansbury’s hypothesis. Comparative notes. - Usually recognizable by its length, intermediate between A. barbatus, which is larger, and the other species occurring in the area under consideration, which are smaller. Small females of A. barbatus may be of the same length as large females of A. occipitalis but they are broader, width 2.8 or more in A. barbatus whereas A. occipitalis females have a width up to 2.4. Females of the smaller species occurring in Thailand have a narrower synthlipsis.
Anisops ogasawarensis MATSUMURA Anisops scutellaris var. ogasawarensis Matsumura, 1915: 109. Anisops genji Hutchinson, 1927: 377-378. Anisops ogasawarensis; Esaki 1930: 214. Anisops genji; Brooks 1951: 371-373 (redescription). Anisops nivea Lansbury, 1964a: 216 (non Fabricius, 1775, misidentification). Anisops ogasawarensis; Miyamoto 1976: 197-198 (synonymizes A. genji with A. ogasawarensis).
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Distribution. - A. ogasawarensis is an eastern species mainly known from China and Japan. It has not been recorded from Thailand but as Lansbury (1964a) (under the name A. nivea) records a large sample (nearly 200 specimens) from N Vietnam it might occur in mountainous areas in N Thailand. Diagnosis. - Macropterous form. A medium sized pale yellow to dark grey species. Length, male 5.8-6.7, female 6.1-7.0; width both sexes 1.6-1.9. Width of head four to six times the anterior width of vertex. Posterior margin of hind femur with 12-15 spines in dorsal and 44-51 in ventral row. Male. In dorsal view the head has a v-shaped indentation. In some specimens the carinae project somewhat anteriorly of the eyes, suggesting a very short cephalic projection, see below under comparative notes. Synthlipsis ±1/3 anterior width of vertex. In frontal view tylus and frons are deeply excavate with two lateral carinae on each side, the median groove runs up to halfway between the eyes with the outer carinae meeting in an acute apex (Fig. 18). Rostral prong shorter than third rostral segment, originating proximally on third rostral segment. Labrum with rather long bristles, laterally united in a pair of tufts, which curve upwards, the middle one often united to a small tuft which points downwards. Fore femur apically somewhat narrowed. Fore tibia stout, stridulatory comb with about 13 teeth of equal length. Female. In dorsal view the head is nearly truncate anteriorly with the vertex slightly extending in front of the eyes; the synthlipsis is 0.35-0.50 times the anterior width of vertex. Tylus flat. Comparative notes. - Similar to A. niveus. Males are easily separated by the excavation of tylus and frons which is much stronger developed in A. ogasawarensis. Females can be separated by size, up to 5.5 mm in A. niveus, over 6.0 mm in A. ogasawarensis. If one considers the slight projection of the frontal carinae in front of the eyes as a cephalic projection, males would run to A. kuroiwae in the key. The difference is that apart from the cephalic projection in A. kuroiwae being much more strongly developed, in lateral view its ventral margin runs horizontally (Fig. 13), whereas in A. ogasawarensis the margin of the frons goes upwards along the margin of the eye.
Anisops tahitiensis LUNDBLAD, first record Anisops tahitiensis Lundblad, 1934: 121-123. Anisops tahitiensis; Brooks 1951: 376-378 (redescription). Anisops tahitiensis; Nieser 2004: 90-91. Material examined. - Thailand:KANCHANABURI Prov., Thong Pha Phum NP, Somnoi, 9.II.2002, 907m asl., leg. S. Mongchoichana, 1♂ ; PRACHUAP KHIRI KHAN Prov., Thap Sakae Dist., Huay Yang waterfall, downstream of waterfall at headquarters and other buildings of park; sampled at edge of stream, 22.XI.2004, lef PC, NN & SP, CN0495, 1♀; NAKHON SI THAMMARAT Prov., Pak Panang Dist., Baan Laem Talumphuk, stagnant fresh water pool near the sea, behind the strand wall, shaded, water light brown, clear, overgrown with Poaceae,
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14.XI.2004, leg. PC, NN & SP, CN0470, 13♂ 17♀; KRABI Prov., Ao Luk Dist., P.N. Mountain resort, tiled basin with fountain in front of resort building, on Nymphaea-like plant, 20.XI.2004, leg. PC, NN & SP, CN0489, 2♂ 5♀; TRANG Prov., Kantang Dist., Ban Nam Rab, Hat Yao fishing harbour, (rain) water standing in a boat on the shore, leg. PC, NN, SP, CN0483, 3♀. First record for Thailand.
Distribution. - From Andaman Islands, Thailand and Vietnam through Malesia to Australia, Tahiti, Guadalcanal and Okinawa. This species inhabits various types of stagnant waters, rarely found in running waters. Although it is often found near the sea and it may have tolerance for brackish water, our samples were all from fresh water habitats. Diagnosis. - Macropterous form. A medium sized brown to dark brown species, pronotum and scutellum sordid white, apex of abdomen blackish. Length, male 5.1-6.0, female 5.1-6.3; width male 1.3-1.5, female 1.3-1.7. Width of head five to seven times the anterior width of vertex. Posterior margin of hind femur with 20-30 spines in dorsal and 30-45 spins in ventral row. Male. In dorsal view the head is anteriorly truncate. Synthlipsis about 0.3 times the anterior width of vertex. In frontal view the tylus is swollen and laterally compressed forming a low, rather blunt, carina , which runs to basal half to two thirds of the frons, on the tylus the carina, has a short median slit. Rostral prong longer than third rostral segment, originating near its distal end (Fig. 12). Labrum without specialized hairs. Fore femur distally truncate. Stridulatory comb on fore tibia with 23-24 teeth decreasing in length towards apex (Fig. 31). Female. In dorsal view the head is virtually truncate anteriorly. Width of head 5.5-6 times the anterior width of vertex; synthlipsis about 0.5 times the anterior width of vertex. Tylus slightly swollen, frons without carina. Brachypterous form unknown. Comparative notes. - Due to the brown to dark brown colour, fresh specimens are similar to A. nigrolineatus which is, however, recognized by its short head, median length less than half the median length of pronotum. Brooks (1951) gives as body length for both sexes 5.1-5.5, although Lundblad (1934) stated that the length of a male is about 6.0 and of a female 6.7. Specimens from West Malaysia and Thailand have a length of male 5.3-6.0 and female 5.5-6.4 (personal observation). The figure of the stridulatory comb given by Lansbury (1964a) gives the impression that the length of the teeth increases from base to apex. Lundblad (1934) states the opposite, confirmed by our observations (Fig. 31).
Paranisops HALE, 1924 Until recently the genus Paranisops was considered to be an Australian endemic containing only two species (Lansbury 1964b). Nieser & Zettel (2001) reported the first species, P. leucopardalos, outside Australia, from Thailand. Chen et al. (2002) added a second species from Thailand, P. sawangi. All Paranisops species occur probably in a flightless,
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brachypterous and a flying, macropterous form. As far as is known, the brachypterous form is whitish to pale yellowish brown and the macropterous form blackish. As P. leucopardalos is only known from macropterous males and P. sawangi only from brachypterous specimens at present only males can be identified on base of their parameres.
KEY TO MALES OF PARANISOPS SPECIES OCCURRING IN THAILAND 1. -.
Parameres nearly symmetrical, left paramere with a slightly longer tip than right one (Figs 38, 39) ............................................................................. P. leukopardalos Parameres distinctly asymmetrical, left paramere boat-shaped (Figs 36, 37) ......... .............................................................................................................. P. sawangi
Paranisops leukopardalos NIESER & ZETTEL Paranisops leukopardalos Nieser & Zettel, 2001: 244-247. Material examined. — Thailand: MUKDAHAN Prov., Phu Pha Thup NP, 31.XII.1999, leg. P. Mazzoldi, 1♂ macr., paratype.
Distribution. - Only known from the type series, Phu Pha Thup NP, Mukdahan, Thailand. Diagnosis. - Macropterous male. Head pronotum and legs brownish yellow, eyes light castaneous, scutellum and hemielytra blackish brown to velvety black, corium with a large pale mark. Length 6.4- 6.8, width 1.9-2.0. In dorsal view the head is anteriorly truncate with vertex extending slightly in front of eyes. Width of head about six times the anterior width of vertex. Synthlipsis about 0.6 times the anterior width of vertex. In frontal view tylus slightly swollen, frons with a broad median carina which splits on the vertex in two, posteriorly diverging, carinae. Median length of head two thirds the median length of pronotum. Parameres only slightly asymmetrical (Figs 38, 39). Brachypterous form unknown. Comparative notes, see key.
Paranisops sawangi CHEN et al. Paranisops sawangi Chen, Nieser & Wattanachaiyingcharoen, 2002: 200-201. Material examined. - Thailand: PHITSANULOK Prov., Nakorn Thai Dist., Phu Hin Rong Kla NP, near Huai Khammunnoi waterfall, 1100m, 6.III.2001, leg. PC, SP & W. W, C0109, 3♂ 1♀ paratypes; Phu Hin Ron Kla NP, stream near bridge to Mung hill tribe village, 1100m,
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6.III.2001, leg. PC, SP & W. W, C0110, 7♀ paratypes; Phu Hin Rong Kla NP, downstream of Water wheel waterfall, low current velocity, bottom sand with some small pebbles, 7.IX.2004, leg. PC, NN, KB & TJ, CN0401, 8♂ 14♀; Phu Hin Rong Kla NP, road 2331, km 30, pond caused by the road blocking a small stream, open water with thick marginal vegetation, 7.IX.2004, leg. PC, NN, KB & TJ, CN0402, 1♂ 1♀. All brachypterous.
Distribution. - So far only known from Phu Hin Rong Kla NP, Phitsanulok, Thailand, where it is apparently not rare. Diagnosis. - Brachypterous form (Fig. 35). In dorsal view a sordid white to yellowish species with dark castaneous to blackish eyes and a brown to blackish apex of abdomen. Width of head about six times the anterior width of vertex. Synthlipsis 0.6-0.7 times the anterior width of vertex. In frontal view, tylus slightly swollen, frons with a broad median carina which splits on the vertex into two, posteriorly diverging carinae. Median length of head about 0.8 times the median length of pronotum. Male. Parameres asymmetrical (Figs 36, 37), apically with small teeth on the ventral (when folded over the genital capsule in rest) side. Macropterous form unknown.
36
37
38
39
35 Fig. 35: Paranisops sawangi, brachypterous male, length 6.7 mm (from Chen et al. 2002). Figs 36-39: Parameres of Paranisops spp., lateral view: 36 = P. sawangi right paramere, 37 = P. sawangi left paramere, 38 = P. leucopardalos right paramere, 39 = P. leucopardalos left paramere (from Chen et al. 2002).
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SUBFAMILY NOTONECTINAE Aphelonecta LANSBURY, 1965 This genus is apparently endemic in Continental SE Asia and Sundaland. A key to all six described species was provided by Zettel (1995). There is only one species recorded from Thailand and adjacent areas.
Aphelonecta gavini LANSBURY Aphelonecta gavini Lansbury, 1966: 629-630. Aphelonecta gavini; Nieser 2004: 91. Material examined. - Thailand: CHIANG MAI Prov., Chiang Dao Dist., stream just outside Doi Chian Dao Wildlife Sanctuary, stagnant bay, 7.XI.1995, leg. NN, N9506, 1♀; Muang Dist., campus of CMU, dept. of Physics, muddy shallow pond surrounded by palm trees, 13.II.2002, leg. PC & NN, C0234, 2♀; Mae Wang Dist., Doi Hui Muang, Ban Huay Kho stream, near Ban Hui Thong village, narrow stream along roadside in secondary forest, 460m asl., leg. PC, AT & CD, C0206, 1♀ Huay Kho stream, mostly dried up mountainous stream, 12.II.2005, leg. PC, NN & PL, CN0520, 1♀; PHITSANULOK Prov., Thung Saleang Luang NP, quiet bay of stream with dead leaves on bottom, shaded, depth 0.1-0.2m, 435m asl., 6.III.2002, leg. PC, NN & D. Wattanaiyingcharoen, C0252, 2♂ 3♀ 8 Lv.; TRANG Prov., Wang Wiset Dist., Loi Chan Phan Wang waterfall, many cascades on limestone, sample from stagnant pool with muddy bottom, in evergreen rainforest, no aquatic or marginal vegetation, 18.XI.2004, leg. PC, NN & SP, CN0485, 2♂ 1 Lv.
Distribution and habitat. - Malay Peninsula, Thailand and Vietnam (Zettel 1995; Nieser 2004). Original description based on a single male from Chiang Mai Prov., Chiang Dao. This species has a strong preference for virtually stagnant parts of streams. Diagnosis. - Macropterous form (Fig. 40). Length male 6.2-6.9, female 6.3-6.9; width both sexes 2.1-2.3. Colour in dorsal view, eyes reddish brown often flecked with black; vertex, anterior half of pronotum and most of scutellum and legs pale yellowish to light brown. Posterior half of pronotum translucent, appearing black due to underlying part. Hind wings black, shining through the translucent hemielytra. In dorsal view, vertex slightly protruding in front of eyes; greatest width of head about three times the anterior width of vertex; synthlipsis about half the anterior width of vertex. Dorsal margin of pronotal fovea directed straight caudad before turning laterally. Nodal furrow straight, directed dorsad, less than its own length removed from membranal suture. Brachypterous form unknown. Comparative notes. - About the same size and somewhat similar to Enithares mandalayensis, but usually darker and more flattened dorsoventrally. The lack of a subapical pointed protuberance on middle femur distinguishes this genus from Enithares.
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Enithares SPINOLA, 1837 The Asian and Australian species of Enithares have been revised by Lansbury (1968, 1973), who provides keys to males and females. Since then several other species have been described but these are nearly all restricted to Insular Asia, notably the Island of Sulawesi (Indonesia) (Nieser & Chen 1996; Chen et al. 2005). Synonymy is abbreviated, synonyms mentioned by Lansbury (1968) are not repeated. Nine species are known from continental SE Asia, of which, including one newly recorded in this paper, five occur in Thailand. Most live in virtually stagnant bays of streams or pools associated with streams. An exception is E. mandalayensis, which can be found in ponds and marshes away from streams. In addition, E. ciliata, although predominantly found in habitats associated with streams, has been found several times in water tanks. Like many Notonectinae, Enithares species usually float against the underside of the surface film when resting or awaiting prey. Most species are easily disturbed and then hide under water. Within a species there is in several cases a dark and a pale form. The hemielytra and hind wings are usually translucent and the colour of posterior part of thorax and abdominal dorsum shining through. Species of this genus are only known in the macropterous form. The development of the indirect flight muscles in correlation with the dark and light form has not yet been studied in this genus. Both males and females can be identified although males show more characters and their identification is consequently more reliable. Males have usually modified (curved and/or with indentations or expansions) fore tibiae and tarsi beset with dense pilosity on the inner side. This helps to keep a hold on females during
Fig. 40: Aphelonecta gavini, female, length 6.7 mm. Fig. 41: Enithares ciliata, male, length 9.2 mm.
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copulation. The caudal abdominal sternites are also different between males and females, but unfortunately the characteristic sternite VII, which in males is more elongate than in females (Figs 50, 51), is usually hidden inside the apex of abdomen. If the shape of the fore legs is not conclusive the easiest way to separate males and females is to lift the caudal sternites of the abdomen to reveal the male genital capsule (Figs 53, 54) or the paired, short and blunt ovipositors (first gonapophyses, Fig. 52) in the female. The genital capsule offers usually several characteristics for specific identification. Important are the general shape of the capsule, the shape of the symmetrical parameres and the shape of the lateral arms of the basal plate through which the aedeagus is protruded (Fig. 54). The genital capsule consists of abdominal segment IX (pygophore) which is dorsally mostly open with its tergite usually forming a bridge. Segment X or proctiger lies as a lid on part of segment IX. Inside lies the aedeagus or phallus and attached to it laterally are the parameres (Fig. 54). In the genus Enithares the parameres lie in a cleft, the part behind the cleft is called posterior lobe which is dorsally open and encloses the aedeagus. The genital capsule can be removed from soft specimens by prying with a bent insect needle around the capsule to cut the attaching tissues and then softly pulling the capsule out. Dry specimens should be softened first. In order to expose the lateral arms of the basal plate the genital capsule can be softened by soaking it in 10% potassium or sodium hydroxide for a few hours at room temperature, or boil for a few minutes. When the capsule is soft, gently insert a fine insect needle into the capsule at the cleft of a paramere and flip out the aedeagus and lateral arms of basal plate by moving the tip of the needle upwards. A character used specifically in identification of Enithares is the distance of the nodal furrow to the membranal suture, introduced by Lansbury (1968). To measure this distance one has to look at the specimen in lateral view and tilt it so that nodal furrow and membranal suture are in a horizontal plane. The distance between the nodal furrow and the membranal suture is measured from the inner (dorsal) tip of the furrow straight back to the suture in a line parallel to the lateral margin of the hemielytron (Figs 43, 44).
KEY TO ENITHARES SPECIES OF CONTINENTAL SE ASIA (female of E. intha unknown) 1. -. 2. -. 3. -. 4. -.
Males....................................................................................................................2 Females ..............................................................................................................10 Large species, length over 10 mm ........................................................................3 Smaller species, length up to 9.5 mm ...................................................................6 Caudal part of clavus raised and densely pitted with fine punctures (Fig. 47); basal part of embolium in ventral view greatly expanded (Fig. 46)...............E. stridulata Caudal part of clavus not raised or pitted; embolium not expanded ....................4 Lateral margins of ventral connexiva I-III and part of IV transversely ridged (Fig. 45) ........................................................................................................................5 Lateral margins of ventral connexiva I-IV not transversely ridged.......E. metallica
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 273
5. -. 6. -. 7. -.
Nodal furrow short and straight, directed dorsad, more than its own length removed from the membranal suture (Fig. 44) ................................................... E. chinensis Nodal furrow curved cephalad, less than its own length removed from the membranal suture (Fig. 43) .............................................................................E. tibialis Small species, length up to 7 mm; outer claw of mid tarsus conspicuously thickened (Fig. 48) ........................................................................................E. mandalayensis Larger species, length 8 mm or more ...................................................................7 Trochanter of middle leg forming a distinct spur at inner ventral margin (Fig. 48) ...8 Trochanter of middle leg rounded or bluntly angulate ...........................................9
e cl
co
t
48 s f t m f
42 45
47 46 p
49 43
co
f
x
c
t
m
44 Fig. 42: Enithares, right hemielytron: cl = clavus, co corium, f = nodal furrow, m = membrane. Figs 43-44: Enithares, distal part of hemielytron in lateral view, stippled line represents distance of nodal furrow to membranal suture: 43 nodal furrow less than its own distance removed from membranal suture, 44 nodal furrow more than its own distance removed from membranal suture, co = corium, f = nodal furrow, m = membrane. Fig. 45: Enithares, left connexival segments I-IV and proximal part of hemielytron in ventral view: e = embolium not dilated proximally, f = nodal furrow, t = transversely ridged segments I-IV of connexivum. Fig. 46: Enithares, proximal part of left hemielytron with proximally dilated embolium (e) in ventral view. Fig. 47: Enithares stridulata, male, right clavus in dorsal view. Fig. 48: Enithares malayensis, male, trochanter and femur of middle leg: s = spur at inner ventral margin of trochanter (from Nieser 2004). Fig. 49: Enithares, metasternum, ventral view: c = hind coxa, p = coxal plate, t = hind trochanter, x = metaternal xiphus (dotted, from Nieser 2004).
274 N. Nieser, P.-p. Chen & P. Leksawasdi
8.
Fore tibia distinctly curved, distally only slightly and smoothly expanded (Fig. 58) .. .......................................................................................................... E. malayensis -. Fore tibia straight; distally expanded into a blunt spur-like projection (Fig. 57) ... .................................................................................................................. E. intha 9. Second segment of middle tarsus widened, about as wide as long (Fig. 56) (in lateral view the first segment is often largely hidden, so the second segment is apparently the first); hind femur without a large ventral projection ...........................E. ciliata -. Second segment of middle leg not appreciably widened; hind femur with a large ventral projection distally (Fig. 60) ............................................................E. sinica 10. Small species, length 7.0-8.0 mm; metaxiphus as (Fig. 55mn) .....E. mandalayensis -. Larger species, length over 8.5 mm ....................................................................11 11. Lateral margins of ventral connexiva I-III and part of IV distinctly transversely ridged (Fig. 45) ...................................................................................................12 -. Lateral margins of ventral connexiva I-IV not distinctly transversely ridged .....13 12. Nodal furrow short and straight, directed dorsad, more than its own length removed from the membranal suture (Fig. 44) ................................................... E. chinensis
in st
si
50
me
53 l
52
ml
p
mn ci
51
54
55
ch
Figs 50- 51: Apex of abdomen of Enithares, ventral view: 50 male, 51 female (from Nieser 2004). Fig. 52: Enithares, apices of female ovipositor, ventral view (from Nieser 2004). Figs 53-54: Outline of genital capsule of Enithares spp.: 53 = E. chinensis, 54 = E. metallica. l = lateral arm of basal plate, p = paramere. Fig. 55: Outlines of metasternal xiphi of Enithares: ch = E. chinensis, ci = E. ciliata in = E. intha (redrawn after Lansbury 1974), me = E. metallica, ml = E. malayensis, mn = E. mandalayensis, si = E. sinica, st = E. stridulata.
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 275
-. 13. -. 14. -. 15. -.
16. -.
Nodal furrow curved cephalad, less than its own length removed from the membranal suture (Fig. 43) .............................................................................E. tibialis Length less than 10 mm; metaxiphus with a short and blunt apex (Fig. 55: ci, ml, si) .. .......................................................................................................................... 14 Length over 10.5 mm; metaxiphus with a long and narrow apex (Fig. 55: me, st) .... 16 Metaxiphus with smoothly convex lateral margins (Fig. 55: ml) ....... E. malayensis Metaxiphus with small blunt subapical lateral projections (Fig. 55: ci) ..............15 Metaxiphus with lateral margins slightly converging caudally; subapical lateral projections small (Fig. 55: ci)....................................................................E. ciliata Metaxiphus with lateral margins parallel to slightly diverging caudally; subapical lateral projections prominent (Fig. 55: si) .............................................................. .......................E. sinica (E. intha may key out with E. sinica, see Lansbury 1973) Metaxiphus with convex lateral margins; nodal furrow more than its own length removed from claval suture (Fig. 44) ...................................................E. stridulata Metaxiphus with concave lateral margins; nodal furrow less than its own length removed from claval suture (Fig. 43) ....................................................E. metallica
56
59
57
60
h f
58
62 61
63
Fig. 56: Enithares ciliata, male, tarsus of middle leg (from Nieser 2004). Figs 57-58: Enithares spp., male fore tibia: 57 = E. intha, 58 = E. malayensis (57 redrawn from Lansbury 1974). Fig. 59: Enithares mandalayensis, male middle tarsus, outer claw stippled (from Nieser 2004). Fig. 60: Enithares sinica, male, distal part of hind femur. Figs 61-62: Enithares tibialis, 61 = genital capsule, 62 = metaxiphus (redrawn after Liu & Zheng 1991). Fig. 63: Nychia sappho, brachypterous male, length 4.0 mm: f = pronotal fovea, h = eyes holoptic posteriorly (adapted from Chen et al. 2005).
276 N. Nieser, P.-p. Chen & P. Leksawasdi
Enithares chinensis BROOKS Enithares chinensis Brooks, 1948: 43. Enithares chinensis; Lansbury 1968: 389-390 (redescription). Material examined. - China: GUANGDONG Prov., Ruyang County, Ba-bao Shan NP, Lao Peng Keng stream, 1100m, 13-14.VIII.1990, leg. PC & L. Xie, C9011/12, 1♂ 1♀; Ba-bao Shan NP, Dong Gu Keng stream, 1100m, 15.VIII.1990, leg. PC & L. Xie, C9013, 1♂ 1♀. Laos: Laos, leg. R.V. de Salvaza, 1♀ (RMNH).
Distribution. — SE China and Laos. Not recorded from Thailand. Diagnosis (partly based on Lansbury 1968). - Length ♂ 10.8-11.0, ♀ 11.211.5; width ♂ 4.0-4.5, ♀ 4.1-4.5. Pale form in dorsal view, eyes castaneous with indistinct black spots, remainder yellowish with a brownish tinge, hemielytra and hind wings translucent, dorsal third of to entire pronotal fovea brownish. Dark form in dorsal view, eyes castaneous to dark brown, vertex yellowish to yellowish brown with a median brown spot posteriorly, pronotum and scutellum yellowish to black, abdomen black, hemielytra translucent with variable infuscations, hind wings smoky brown. Head in dorsal view rounded anteriorly, its width 2.1-2.2 times the anterior width of vertex. Anterior width of vertex twice synthlipsis or slightly less (vertex/synthlipsis 1.93-2.04). Dorsal margin of pronotal fovea directed slightly obliquely caudad before turning laterad. Nodal furrow short and straight, directed dorsal, more than its own length removed from membranal suture. Outer margin of connexivum in ventral view with a row of small transverse ridges on segments I-III and anteriorly on segment IV. Mesotrochanter rounded. Metasternum xiphus subtriangular (Fig. 55: ch). Male. Fore tibia and tarsus not modified except for ventro-anterior pilosity on distal part of tibia and on tarsus much stronger developed than in female. Genital capsule as in Fig. 53, parameres tapering with some hairs on their apex. Female. Conform the general diagnosis. Comparative notes. - This species is similar to E. tibialis which has, however, the nodal furrow less than its own length removed from membranal suture. In addition the caudal projection of the lateral arms of the basal plate of the aedeagus is narrow in E. tibialis and broad in E. chinensis (Figs 53, 61). For females, see also comparative notes under E. stridulata and E. metallica.
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Enithares ciliata (FABRICIUS) Notonecta ciliata Fabricius, 1798: 524-525. Enithares ciliata; Lansbury 1968: 413-416 (redescription, synonymy). Enithares ciliata; Nieser 2004: 93 Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Baan Khun Khon, stream 5 km downstream of Khun Khon waterfall, just outside NP, 33 km SW Chiang Rai city, 23.II.2002, leg. PC, NN, AT & CD, C0242, 1♀; Mae Fah Kuang Dist., Hilltribe village 15 km N of Maesalong Village, small muddy rain puddle along road, 29.X.2004, leg. PC, NN & RT, CN0449, 2♀; CHIANG MAI Prov., Mae Rim Dist., Mae Sa NP, 20 km N CMC, near waterfall, lentic part at cave, 27.XI.1994, leg. PC & SP, C9414, 1♂ 2♀; Queen Sirikit Botanical garden, Huay Mae Sa Noi, mountain stream, bottom sand and rock, water turbid grey due to construction work upstream, 2.II.2005, leg. PC & NN, CN0518, 1♂ 2♀; Huay Mae Sa Luang, 2.II.2005, larger mountain stream at entrance of QSBG, bottom sand and rock, water turbid grey, 2. II.2005, leg. PC & NN, CN0519, 1♀; Fang Dist., Doi Ang Khang, stream pools at Ang Khang Royal Horticulture Station, 1400m asl., 14.I.1995, leg. PC & K. Suriya, C9503, 5 Lv.; same, stream pond at forestry department station office, 31.X.2004, leg. PC, NN,& RT, CN0455, 6♂ 7♀; Chom Thong Dist., Doi Inthanon NP, 70 km SW CMC, Wang Khwaay waterfall, pools in streambed downstream of waterfall, 31.I.2002, leg. NN, CD & AT, C0223, 3♂ 2♀; Doi Inthanon NP, watertanks at guesthouse of Mae Chon Luang Agricultural Station, round tanks of fiber plastic, diameter ca 10m, depth 1m, no vegetation but may flowers of Prunus sp., floating on the water, 14.I.2005, leg. PC & NN, CH0503, 1♂ 3♀; pond in Mae Chon Luang Agricultural station, 18°35’54.8”N, 98°28’40.9”E, stagnant pond with dense vegtation of Hydrilla, partly covered with orange-brown iron ooze, sample from partly shaded, partly sunny edge with emerging Poaceae, 1306m asl., temp. 19°C, pH 6.7, 15.I.2005, leg. PC & NN, CN0506, 1♀; Doi Inthanon NP, Khun Huay Haeney, 18°34’23.5”n, 98°31’04.9”E, permanent mountain pond along road, heavyly shaded, in deciduous forest, no aquatic vegetation, pH 7.0, 15.I.2005, leg. PC & NN., CN0508, 1♂ 2♀; Doi Inthanon NP, Huay Mae Ab at Mae Aeb Nai Village, 18°30’N, 98°35’E, 860m asl., rockpools along narrow mountain stream, downstream of bridge, 23-24.I.2005, leg. PC, NN & PL, CN0515, 3♂ 3♀; same, 9.II.2005, leg. W. Srisuka, CN0521, 1♂ 1♀; Muang Dist., campus of CMU, narrow stream beside road near Dept. of Physics, stony bottom, water shallow, somewhat polluted, 13.II.2002. leg. PC & NN, C0235, 1♂ 1♀; stream at entrance of Chiang Mai Zoo, near dam at bridge to CMU, virtually stagnant, bottom a 0.05-0.1m thick layer of black mud on sand, 11.XII.2004, leg. PC & NN., CN0499, 1♂ 1♀; at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, Enithares in open vegetation free area up to 0.5m deep, 14.XII.2004, leg. PC & NN, CN04100, 9♂ 14♀ 3Lv.; cattle drinking tank in meadows, close to CN04100, concrete tank, water turbid green due to algal bloom, 14.XII, 2004, leg. PC & NN, CN04101, 9♂ 15♀; Mae Wang Dist., Huay Kho stream, dried mountain stream, pools connected by stretches of shallow running water, 12.II.2005, leg. PC, NN & PL, CN0520, 1♂ 1♀; Samoeng Dist., Doi Laung, Mae Tong Ting Village, 800m asl., artificial pond near the station of Chiang Mai Regional Forest Division, 29.III.2001, leg. PC, B. Kavinsekan & S. Praiwan, C0146, 2♂ 2♀ 3Lv.; MAE HONG SON Prov., Phaa Boong, ca. 20 km S of Mae Hong Son City, small pool at base of waterfall, sampled in edge with Spirogyra-like Chlorophyta, floating plant debris and plant roots washed out in water, stagnant, 3x1m, 1m deep, 12.XI.1995,
278 N. Nieser, P.-p. Chen & P. Leksawasdi
leg. NN, N9511, 1♀; Mae Sa Riang Dist., Baan Pak Tham, Mae Sa Riang River, 1 km E of Mae Sa Riang Town, 1100+ m asl. broad mountan stream, 19.II.2002, leg. PC, NN, CD & AT, C0239, 3♂ 2♀; UTTARADIT Prov., Lab Lae Dist., Baan Mae Phuul, Mae Phuul waterfall, 24 km NW Uttaradit city, 670m asl., 9.II.2002, leg. PC, NN, AT, CD & WJ, C0228, 2♀; PHITSANULOK Prov., Nakorn Thai Dist., Tung Saleang Luang NP, Khek River, 101 km E of Phitsanulok City, 500m N of headquarters, 17.XI.1994, leg. PC & SP, C9404, 1♀; Phu Hin Rong Kla NP, near Huai Khammunnoi Waterfall, 1100m asl., 6.III.2001, leg. PC, SP & WW, C0109, 1♂; same, Waterwheel waterfall, 8.IX.2004, leg. PC, NN, KB, & TJ, CN0401, 2♂ 1♀; same, road 2331, km 30, pond caused by the road blocking a small stream, open water with thick marginal vegetation, 7.IX.2004, leg. PC, NN, KB & TJ, CN0402, 1♀; CHON BURI Prov., Khao Khaew Open Zoo, ponds, 6.IV.2001, leg. PC, BK & S. Leepitakrat, C0158, 2♀ 6Lv.; PRACHUAP KHIRI KHAN Prov., Bang Saphan Dist., Saikhu waterfall, mountain stream downstream of waterfall, 22.XI.2004, leg. PC, NN, SP, CN0494, 1♂; NAKHON SI THAMMARAT Prov., Prom Khiri Dist., Khao Luang NP, Ai Khieo waterfall, rockpools along mountain stream downstream of waterfall, 12.XI.2004, leg. PC, NN & SP, CN0464, 1♀.
Distribution. - Widespread, Mauritius, Buthan, India, Sri Lanka, continental SE Asia, SE China and Indonesia: Sumatra (Lansbury 1968; Polhemus 1995). Although predominantly found in virtually stagnant bays and pools associated with streams this species occurs regularly in isolated pools and ponds including drinking tanks for cattle (CN04101) and rice-fields (Kirkaldy 1904 under the name E. abbreviata Kirby). The first record for Thailand is apparently by Lansbury (1968). It is by far the commonest species in Thailand, and probably the species studied by Chowanadisai (1986), under the name of E. templetoni Kirby (1891) which is a synonym of E. simplex Kirby (1891) and endemic in Sri Lanka, so clearly a misidentification. Diagnosis. - Length male 8.5-9.6, female 9.0-9.6; width, both sexes 3.0-3.6. Pale form can be entirely pale yellowish brown but usually has variable dark markings, it was encountered in one sample from Thailand only (CN04100). Dark form blackish in dorsal view, eyes dark brown, vertex and anterior half to two thirds of pronotum yellowish. Lateral margins of scutellum in living specimens with a greenish fluorescent stripe, which becomes yellow in dead specimens. Head in dorsal view rounded anteriorly, its width about 2.75 times the anterior width of vertex. Anterior width of vertex 2.1-2.2 times the synthlipsis. Dorsal margin of pronotal fovea directed straight caudad before turning laterad. Nodal furrow curved cephalad, less than its own distance removed from membranal suture. Mesotrochanter rounded. Lateral margins of metasternal xiphus caudally convergent and with blunt, not prominent, subapical projections (Fig. 55: ci). Male. Fore tibia on its inner side with a wide indentation halfway its length, distally of this indentation the tibia is widened into a triangular projection. Second segment of middle tarsus short and wide, about as wide as long (Fig. 56), the first segment is often largely hidden, so the second segment is apparently the first). Comparative notes. Males can be easily recognized by the middle tarsus, and also by the fore tibia. Females are recognized by the shape of the metasternal xiphus combined with their size.
The Notonectidae (Heteroptera, Nepomorpha) of Thailand and adjacent areas, with keys for identification... 279
Enithares intha PAIVA Enithares intha Paiva, 1918: 27. Enithares intha; Lansbury 1973: 228-229.
Distribution. - Only known by the type series (two males) from Inle Lake in Burma. As Inle Lake is only about 150 km as the crow flies removed from NW Thailand, this species might be found in NW Thailand. Diagnosis (adapted from Lansbury 1973). — Only males known. Length 8.08.5, width 3.0-3.5. General appearance in dorsal view shining reddish brown. Eyes with irregular black blotches. In dorsal view the head is anteriorly rounded, width of head 2.75 times the anterior width of vertex. Anterior width of vertex two times the synthlipsis. Dorsal margin of pronotal fovea directed obliquely laterad from the posterior margin of the eyes. Nodal furrow curved cephalad, less than its own length removed from membranal suture. Fore tibia distally expanded on inner margin (Fig. 57). Mesotrochanter with inner ventral angle produced into a spur-like projection (as in E. malayensis Fig. 48). Outer claw of middle tarsus thickened. Metaxiphus with a blunt apex and subapical infuscated expansions (Fig. 55: in). Comparative notes. The spur-like projection of the mesotrochanter keys males of this species out with E. malayensis which has the metaxiphus with smoothly converging, convex, lateral margins and an acute apex (Fig. 55: ml).
Enithares malayensis BROOKS, first record Enithares malayensis Brooks, 1948: 41. Enithares malayensis; Lansbury 1968: 427-428. Enithares malayensis; Nieser 2004: 93. Material examined. - Thailand: SATUN Prov., Wang Pa Chang Dist., Taleh Ban NP, pothole in narrow stream in primary evergreen forest, no aquatic vegetation, bottom sand with some plant debris, 16.XI.2004, leg. PC, NN & SP, CN0478, 1♂. First record for Thailand.
Distribution. - Indonesia (Riau Archipelago), Malay Peninsula and S Thailand. In the Malay Peninsula, where it seems to be the commonest species of Enithares, it is equally common in streams as in stagnant pools (Nieser 2004). Diagnosis (adapted from Nieser 2004). - Length, both sexes 9.0-9.4; width both sexes 3.2-3.6. Colour in dorsal view blackish. Eyes reddish brown to grey, vertex and anterior one third of pronotum brown. Middle third of pronotum yellowish and posterior third translucent but appearing blackish due to underlying scutellum shining through. Scutellum, apart from V-shaped yellowish area, and most of abdominal dorsum black, shining through translucent hemielytra.
280 N. Nieser, P.-p. Chen & P. Leksawasdi
Head in dorsal view rounded anteriorly, width of head three times or more the anteriorwidth of vertex. Anterior width of vertex twice the synthlipsis. Dorsal margin of pronotal fovea directed straight caudad before turning laterad. Nodal furrow curved cephalad, less than its own length removed from membranal suture. Metasternal xiphus with convex lateral margins. Male. Fore tibia curved forward, concave on anterior surface and convex on posterior surface. Inner ventral angle of mesotrochanter with a spur-like projection (Fig. 48). Middle tibia weakly convex along dorsal side and expanded distally. Comparative notes. - Males have the spur-like inner ventral angle of mesotrochanter in common with E. intha. The metaxiphi of E. intha and E. malayensis are, however, very different (Fig. 55: in, ml). Females should also be recognizable by the shape of the metaxiphus in combination with size.
Enithares mandalayensis DISTANT Enithares mandalayensis Distant, 1911: 331-332. Enithares mandalayensis; Lansbury 1968: 380-381 (redescription). Enithares mandalayensis; Nieser 2004: 93-94. Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Doi Ngyme, Campus of MFLU, ponds, 13-26.X.2004, leg. PC & NN, CN0440, CN0446, CN0448, CN0453 2♂ 7♀ 1Lv; CHIANG MAI Prov., Mae Dang Dist., Road 107 along Mae Ping River, at Royal Ping Garden & resort, 28.III.2001, leg. PC et al., C0144, 1LvV; Mae Wang Dist., Huay Kho stream, mostly dried up mountainous stream, 12.II.2005, leg. PC, NN & PL, CN0520, 1♀; Chom Thong Dist., Baan Mae Jam Fang Khaw, Ping River, under bridge, 82 km SW CMC, 5.II.2002, leg. PC, NN, AT & CD, C0227, 2♂; LAMPANG Prov., Hang Chat Dist., Mount Lung Khuntan, pond in Elephant Training Centre, 11.II.2002, leg. PC, NN, AT, CD & WJ, C0233, 1♀; PHITSANULOK Prov., Nakorn Thai Dist., 100 km NE Phitsanulok City, pond in garden, 6.III.2001, leg. PC, SP, WW, C0112, 1LvV; Muang Dist., outskirts of Phitsanulok City, Biological Control Centre, frog rearing pond, 7.III.2001, leg. PC, SP & WW, C0114a, 1♀; Mount Lung Khuntan, pond in Elephant Training Centre, 27.III.2001, leg. PC, SW & B. Kavinseksan, C0140, 1LvV; KANCHANABURI Prov., Thongphaphom Dist., Ban Passaduuklang, Kayen stream, 23.II.2002, leg. N. Sangpradub, 1♀; SATUN Prov., Wang Pa Chang Distr., Taleh Ban NP, pothole in narrow stream in primary evergreen forest, no aquatic vegetation, bottom sand with some plant debris, 16.XI.2004, leg. PC, NN & SP, CN0478, 1♂.
Distribution. - Continental SE Asia. In Thailand widespread but scarce, in isolated ponds and ponds associated with streams, first recorded from Thailand by Lansbury (1968) from Chiang Mai. Diagnosis. - Length male, 6.6-6.9, female 7.0-8.0; width both sexes 2.2-2.8. Usually a pale yellowish species, eyes reddish, vertex and anterior part of pronotum pale yellowish to light brown, posterior part of pronotum and hemielytra translucent with scutellum and dorsum of abdomen shining through; scutellum brownish
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anteriorly darker, dorsum of abdomen brown to grey. Head in dorsal view rounded anteriorly, vertex slightly protruding; width of head about 2.5-2.8 times the anterior width of vertex. Anterior width of vertex 2.3-2.5 times the synthlipsis. Dorsal margin of pronotal fovea directed straight caudad before turning laterad. Nodal furrow curved cephalad, less than its own length removed from membranal suture. Mesotrochanter rounded. Metasternal xiphus with lateral margins converging caudally with subapical indentations (Fig. 55: mn). Male. Fore leg, tibia distally and first tarsal segment with shallow indentation. Outer claw of middle tarsus curved and widened. Comparative notes. - The head ratios are very variable in this species, the values mentioned above are based on W Malaysian specimens (Lansbury 1968; Nieser 2004). Easily distinguishable from other Enithares occurring in Continental SE Asia by its small size combined with the shape of the metasternal xiphus. E. madalayensis and Aphelonecta gavini are of about the same size but Aphelonecta is lacking the subdistal spur on mesofemur. In Java and Sumatra occurs a very similar species, E. uncata Lundblad (1933), which might be a subspecies of E. mandalayensis (Lansbury 1968).
Enithares metallica BROOKS Enithares metallica Brooks, 1948: 38-40. Enithares metallica; Lansbury 1968: 420-421 (redescription). Enithares metallica; Nieser 2004: 94. Material examined. - Thailand: SAKON NAKHON Prov., Phu Pan NP, 480-520m asl., 7-8. XII.1995, leg. P. Schwendinger 2♂ 1♀ (NHMW).
Distribution. - Thailand, Vietnam and W Malaysia (Lansbury 1968; Nieser 2004). Type series from “Siam” (Brooks 1948). Diagnosis (adapted from Lansbury 1968 and Nieser 2004). - Length, both sexes 10.2-12.0; width, both sexes 3.9-5.5. Eyes castaneous. Pale form, vertex, pronotum and scutellum yellowish brown. Clavus and corium translucent, appearing grey to black due to underlying pigmentation. Opaque zone of membrane greyish yellow, remainder of membrane smoky brown. Dark form, vertex and anterior part of pronotum yellowish with foveae black, posterior part of pronotum translucent, appearing dark brown due to underlying pigmentation. Scutellum black with variable yellowish markings along lateral margins. Clavus, corium and membrane largely blackish with a metallic sheen, basal parts of clavus and corium and apical part of membrane translucent. Head in dorsal view rounded anteriorly, width of head about 2.5-2.8 times the anterior width of vertex. Anterior width of vertex 1.7-1.9 times the synthlipsis. Dorsal margin of pronotal fovea directed laterocaudad before turning laterad. Nodal furrow curved cephalad, less than its own length removed from membranal suture. Mesotrochanter rounded. Metaxiphus with concave lateral margins, resulting in a long acute tip.
282 N. Nieser, P.-p. Chen & P. Leksawasdi
Male. Mesofemur ventrally with a large shallow depression covered with long hairs. Comparative notes. - The dark form is recognizable by its metallic sheen.The light form by its size combined with the shape of the metaxiphus. E. chinensis, which is of approximately the same size, has the nodal furrow more than its own length removed from membranal suture, the lateral margins of connexival segments I-IV ridged, and the lateral arms of basal lobe broader distally (Figs 53, 54). E. stridulata, which is also of approximately the same size and has a somewhat similar metaxiphus, is not known to have a light form; males of this species are immediately recognizable by the raised and pitted caudal apex of the clavus.
Enithares sinica (STÅL) Notonecta sinica Stål, 1854: 241. Enithares sinica; Lansbury 1968: 378-380 (redescription). Material examined. - Laos: CHAMPASAK Prov., Bolavens plateau, Pak Song, 1250m asl., 27.XII.1996, leg. P. Schwendinger, 1♂ 1♀ (NHMW). First record for Laos. China: HENAN Prov., Xichuan, 4.VIII.1988, leg. Y.I. Zheng, 1♀; HUBEI Prov., Chang yang Co., Jin Yang Kou, 200m asl., 24.VII.1990, leg. L.C. Ren, 1♀; GUANGDONG Prov., Guangzhou Botanical garden, pond, 3.VII.1990, leg. PC, 1♂ 1♀; SICHUAN Prov., Leshan, canalized river at Wogouba, 29.VII.1989, leg. PC & NN, N8958, 1♀; Leshan, small pond on banks of river Wuyiuba, 30.VII.1989, leg. PC & NN, N8959, 1♀; YUNNAN Prov., Dali, trench and ponds on banks of small stream, 13.VIII.1989, leg. PC & NN, N8965, 1♀.
Distribution. - Japan, China, Philippines, Vietnam, Laos (Polhemus 1995). Not recorded from Thailand. Diagnosis. - Length, male 8.5-9.5, female 8.2-9.5; width both sexes 2.9-3.5. Eyes usually reddish brown. Pale form, yellowish to yellowish brown, posterior half of dorsum of abdomen often darker. Dark form, vertex and anterior half of pronotum yellow to yellowish brown; posterior half of pronotum translucent, appearing blackish due to underlying structures. Scutellum yellowish to yellowish brown with a triangular marking basally (according to Lansbury 1968 the black marking may be absent). Hemielytra and hind wings translucent with the grey to blackish dorsum of abdomen shining through. Head in dorsal view rounded anteriorly with vertex slightly protruding, width of head 2.3-2.6 times the anterior width of vertex. Anterior width of vertex 1.9-2.3 times the synthlipsis. Dorsal margin of pronotal fovea directed straight caudad before turning laterad. Nodal furrow straight, directed cephalad, less than its own length removed from membranal suture. Mesotrochanter rounded. Metasternal xiphus with subapical extensions laterally (Fig. 55: si). Male. Fore leg, first tarsal segment on its posterior side with a distinct patch of short black spines. Middle leg, trochanter short and angularly rounded, outer claw curved and widened. Hind femur with a large subdistal spur (Fig. 60). Female. Conform the general diagnosis.
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Comparative notes. - Males can be easily recognized by the large subdistal spur on hind femur. Females are recognized by the metaxiphus in combination with their size. However, females of E. intha may be very similar.
Enithares stridulata Brooks Enithares stridulata Brooks, 1948: 37-38. Enithares stridulata; Lansbury 1968: 366-368 (redescription). Material examined. - Thailand: CHIANG RAI Prov., Muang Dist., Pong Pra Bat NP, mountain stream, flooded two days ago due to heavy rain, sample from pond on banks, 13.X.2004, leg. PC, NN & RT, CN0441, 2♀ CHIANG MAI Prov., Doi Suthep, beside parking lot of hill tribe village, narrow stream shaded by bamboo, 4.XII.1994, leg. PC, C9421, 1♂ 1♀; NAN Prov., Pua Dist., Doi Phuka NP, Huai Nam Daen, mountain stream with torrents and boulders, sample from quiet pools at adge, 22.IX.2004, leg. PC, NN & SP, CN0426, 3♂ 1♀; Fang Dist., Doi Ang Khang, stream pond at forestry department station office, 31.X.2004, leg. PC, NN,& RT, CN0455, 1♂ 2Lv.; Chom Thong Dist., Doi Inthanon NP, Huay Rong Rian at Adminstration Building of Mae Chon Luang Agricultural Station, mountain stream in open area, limestone, bottom sand with pebbles and boulders, water colourless, clear, velocity of current variable, mostly high, no aquatic vegetation, 18°38’55.6” N, 98°28’46.0”E, 1345m asl., pH 6.0, temperature 19°C, 14.I.2005, leg. PC & NN, CN0502, 1♂ 1♀; Doi Inthanon NP, Huay Ton Phueng, narrow muddy stream in mixed pine forest, sample from virtually stagnant shallow pools with bottom of thin black mud, water colourless, clear, 18°39’11.7”N, 98°27’29.7”E, 1357m asl., pH 6.5, temperature 17°C, 15.I.2005, leg. PC & NN, CN0507, 2♂ 2♀; PHITSANULOK Prov., Nakorn Thai Dist., Phu Hin Rong Kla NP, Huai Khamunnoi waterfall, 1100m asl., 6.III.2001, leg. PC, SP & WW, C0109, 1LvIV, 1LvV; Phu Hin Rong Kla NP, Waterwheel waterfall, 8.IX.2004, leg. PC, NN, KB & TJ, CN0401, 1♂ 1♀; same, open wetland, 8.IX.2004, leg. PC, NN, KB & TJ, CN0402, 1♀; same, stream beside the temple, 8.IX.2004, leg. PC, NN, KB & TJ, CN0403, 1♂, 1Lv; same, roadside stream near main gate, 8.IX.2004, leg. PC, NN, KB & TJ, CN0404, 2♂ 1♀.
Distribution and habitat. - N Thailand, Laos, N Vietnam and Yunnan (Nieser et al. 2005). First recorded from Thailand: Chiang Mai by Lansbury (1968). Apparently restricted to the mountainous areas in the north of Continental SE Asia and Yunnan. Collected mostly in ponds associated with streams and virtually stagnant parts of streams. Diagnosis. - Length, male 12.2-13.0, female 11.7-12.3; width, male 5.0-5.3, female 4.5-5.0. Colour, in dorsal view, eyes brown to blackish; vertex and anterior part of pronotum yellowish to light brown, posterior part of pronotum translucent, usually appearing dark due to underlying structures shining through. Scutellum yellowish with an irregularly triangular dark brown spot of variable size at base. Clavus dark brown to blackish with outer margin greyish to entire clavus greyish and somewhat translucent; corium greyish translucent, usually with a dark band along costal margin and a variable dark marking along inner margin; basal apart of membrane usually dark brown to blackish, apical part translucent. Head, in dorsal view, anteriorly virtually straight, width of head 2.5-2.7 times the anterior width of vertex. Anterior width of vertex 1.7-1.9 times the synthlipsis. Dorsal
284 N. Nieser, P.-p. Chen & P. Leksawasdi
margin of pronotal fovea directed obliquely caudad before turning laterad. Nodal furrow straight to curved cephalad, more than its own length removed from membranal suture. Nodal furrow directed dorsad, tip inclined cephalad, more than its own length removed from membranal suture. Mesotrochanter rounded. Metasternal xiphus with a narrow point (Fig. 55: st). Outer margin of connexivum in ventral view with a row of small transverse ridges on segments I=III and anterior part of segment IV. Male. Basal half of embolium in ventral view distinctly expanded (Fig. 46); caudal apices of clavi raised and pitted (Fig. 47). Female. Conform the general diagnosis. Comparative notes. - Males of this species are easily recognized by the raised and pitted apices of clavi and the distinctly expanded basal part of the embolium. Females, which lack these distinctive characteristics, are very similar to females of E. chinensis. Lansbury (1968) uses in his key to females the ridged lateral margins of ventral connexival segments I-IV to separate E. chinensis from other females with nodal furrow clearly more than its own length removed from membranal suture. He overlooked, however, that E. stridulata also has lateral margins ridges on these connexival segments, though less distinctly than in E. chinensis. The size in nearly overlapping, length females of E. chinensis 11.0-11.5; of E. stridulata 11.7-12.5. The only reliable difference is in the metaxiphus, which in E. stridulata has a distinctly longer point than in E. chinensis (Fig. 55: ch, st). See also under E. tibialis.
Enithares tibialis Liu & Zheng Enithares tibialis Liu & Zheng, 1991: 66-69.
Distribution. — SE China (Hainan Island), Laos (Polhemus 1995). Not recorded from Thailand. Diagnosis (based on the original description). - Length, both sexes 12.3-14.5; width, both sexes 4.5-5.5. Colour, in dorsal view, yellowish, scutellum basally and medially black. head in dorsal view anteriorly rounded, width of head just over twice the anterior width of vertex, judging from the figure of the original description, head 2.3 times as wide as anterior width of vertex. Anterior width of vertex in male 1.5 times, in female 1.7 times the synthlipsis. Dorsal margin of pronotal fovea directed obliquely caudad before turning laterad. Nodal furrow curved cephalad, less than its own length removed from membranal suture. Mesotrochanter rounded to obtusely angular. Metasternal xiphus with a long narrow point (Fig. 62). Outer margin of connexival segments I-III and base of IV in ventral view minutely transversely ridged. Male. Genital capsule with lateral arms of basal plate caudally with a narrow projection (Fig. 61). Female. Conform the general diagnosis. Comparative notes. - This species is similar to E. chinensis and E. metallica. Males of both these species have the lateral arms of the basal plate of aedeagus stouter (Figs
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53, 54, 61). In addition E. metallica lacks the minute transverse ridges ventrally on connexival segments I-III. E. chinensis has the metaxiphus with a shorter and stouter apex and the nodal furrow more than its own length removed from the membranal suture. Females of E. stridulata, which are also quite similar, have the nodal furrow more than its own length removed from the membranal suture. Remark. - The ratio vertex: synthlipsis given by Liu & Zheng (1991) in their text and repeated above do not agree with their figure in which the vertex is twice the synthlipsis. Possibly they took the distance between eyes along posterior margin of head as synthlipsis instead of the smallest distance between eyes posteriorly, which in this species lies somewhat anteriorly of the posterior margin of the head.
Notonecta LINNAEUS, 1758 The genus Notonecta is common in temperate areas and, especially in the Old World, replaced by Enithares in tropical areas. It does not occur in Thailand but two species: N. montandoni Kirkaldy (1897) and N. violacea Kirkaldy (1897), which both have their main distribution in China, reach Burma (Polhemus 1995).
Nychia STÅL, 1860 The genus Nychia was revised by Lansbury (1985). It is a small genus with two to four species widespread in the Old World tropics. One species, N. marshalli (Scott 1872) is widespread throughout Africa and the Mediterranean. The identity of the type species, N. limpida Stål (1860) from Guandong Prov. in China is uncertain as there is only the head and pronotum left of the holotype and the species has never been collected again in China. The identity of N. infuscata Paiva (1918) described from Inle Lake in Burma is also uncertain. No specimens from the type area have been studied since its description, which was essentially on colour characters. Some specimens of N. sappho from NW Thailand, which is quite close to Inle Lake, show the same colour patterns as described by Paiva (1918) so probably N. infuscata is a synonym of N. sappho. In continental SE Asia only one species.
Nychia sappho KIRKALDY Nychia marshalli var. sappho Kirkaldy, 1901: 809-810. Nychia sappho; Lansbury 1985: 4-8 (redescription, synonymy). Nychia sappho; Nieser 2004: 94-95. Material examined. — Thailand: CHIANG RAI Prov., Chiang Saen Dist., Baan Sop Ruank, pond in Thai side of Golden Triangle, 50 km NE Chiang Rai city, 26.I.2002, leg. PC, AT, CD
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& WJ, C0214, 6♂ 12♀ brach., 6Lv.; Mae Sai Dist., Baan Sri Pa Dang, polluted pond, 72 km N of Chiang Rai city, 26.I.2002, leg. PC, AT, CD & WJ, C0215, 33exx.; Muang Dist., Baan Khun Khon, 33 km SW Chiang Rai city, 5 km downstream of Khun Khon waterfall, ponds behind hill tribe house, 23.II.2002, leg. PC, NN, AT &CD, C0243, 1♂ 1♀ brach.; campus MFLU, pond in front of president’s villa, water reservoir, water coming from Doi Ngam, 22.X.2004, leg. PC & NN, CN0446, 5♂ 2♀ brach.; campus MFLU, pond behind A1-2 residential building, in the water many Lentibulariaceae mixed with Poaceae, Nychia from shallow more open edge, 26.X.2005, leg. PC & NN, CN0448, 2♂ 2♀ brach.; Wang Chai Dist., Don Sila Subdist., stagnant ditch along the road, 1-6m wide, no aquatic vegetation but overgrown at palces by land plants, 3.XI.2004, leg. PC, NN & RT, CN0458, 3♂ 5♀ brach., 1♀ macr.; Wang Chia Dist., Nong Luang Lake, sampled at edge up to 0.5m deep, vegetation Eichornia and Mimosa, water clear to slightly opaque, bottom muddy with plant debris, 3.XI.2004, leg. PC, NN & RT, CN0459, 1♂ 5♀ brach.; CHIANG MAI Prov., Chiango Dao dist., stream pond near entrance of Chiang Dao Cave, 26.III.2001, leg. PC, BK & SW, C0136, 2♂ 1♀ brach., 1♀ macr., 1LvV.; Samoeng Dist., Doi Luang, Mae Sarb River, irrigation dam in mountain stream, 850m asl., 29.III.2001, leg. PC, BK & S. Praiwan, C0148, 2♀ brach.; Mae Wang Dist., Doi Hui Muang, Ban Hui Thong, Mae Khan River, 510m asl., 18.I.2002, leg. PC, AT & CD, C0207, 2♂ 3♀ macr.; Muang Dist., campus of CMU, dept. of Physics, muddy shallow pond surrounded by palm trees, 13.II.2002, leg. PC & NN, C0234, 1♂ 2♀ brach.; at foot of Doi Suthep, drying up pond in Agro-forestry Research Station, Faculty of Agriculture, CMU, 14.XII.2004, leg. PC & NN, CN04100, 1♂ 3♀ macr.; cattle drinking tank in meadows, close to CN04100, concrete tank, water turbid green due to algal bloom, 14.XII, 2004, leg. PC & NN, CN04101, 1♂ brach., 1♀ macr., 1Lv.; San Kamphaeng Dist., Ban Hui Phai, Mae On stream, 14 km SE CMC, 22.I.2002, leg. PC, AT & CD, C0208, 4♂ brach., 4♂ 3♀ macr., 3Lv.; Doi Saket Dist., Hui Hong Krai, Royal development Study Centre, large and deep open lake for rearing catfish, 1.XII.1994, leg. PC &SW, C9418, 1♂ brach., 2Lv.; Chom Thong Dist., Doi Inthanon NP, 68 km SW CMC, Mae Klang waterfall, 300m downstream, 31.I.2002, leg. NN, AT &CD, C0221, 1♂ macr.; Doi Inthanon NP, Mae Kleng waterfall, 4.XI.1995, leg. H. Zettel, 2♂ 2♀ brach.(NHMW); Chom Thong Dist., Baan mai San Tung, pond near the road at edge of longan orchard, 31.I.2002, leg. NN, AT & CD, C0225, 1♀ macr.; Chom Thong Dist., Ping River under bridge at Baan Mae Jam Fang Khaw, 82 km SW CMC, 5.II.2002, leg. PC, NN, AT & CD, C0227, 2♂ 4♀ brach., 6♂ 10♀ macr.; Mae Wang Dist., Huay Kho stream, dried mountain stream, pools connected by stretches of shallow running water, 12.II.2005, leg. PC, NN & PL, CN0520, 1♂ macr.; MAE HONG SON Prov., Sop Moei Dist., Baan Mae Khatuan, 5 km N of Sop Moei Town, Mae Yuam River, at bridge, broad mountain stream, 19.II.2002, leg. PC, NN, AT & CD, C0240, 3♀ macr.; PHRAE Prov., roadside pond somewhat downstream of Mae Kam waterfall, 20x10m, water turbid light brown due to suspended silt, bottom very soft mud, depth unknown, sample from edge with shade, some plant debris and some marginal plant material hanging in the water, 16.XI.1995, leg. NN, N9516, 11♂ 29♀ brach., 5♂ 9♀ macr.; LAMPANG Prov., Hang Chat Dist., Mount Lung Khuntan, pond in Elephant Training Centre, 27.III.2001, leg. PC, SW & BK, C0140, 1♀ brach., 1♂ macr., 11Lv.; LAMPHUN Prov., Muang Dist., Baan Pa Hei, 8 km NE of Lamphun city, Pig Hang canal (connected with Mae Ping River), 14.II.2002, leg. PC, NN, AT & CD, C0237, 1♀ macr.; UTTARADIT Prov., Muang Dist., Nan river under bridge near Tha Sao market, 3 Km N of Uttaradit city, 64m asl., ponds on banks, 9.II.2002, Leg. PC, NN, AT, CD & WJ, C0229, 9♂ 5♀ brach., 3♂ 2♀ macr., 28Lv.; Ban Khok Dist., Baan Muangchedon, Naam Pat barrage lake, 10 km W Ban Khok town, small ponds downstream of lake, 10.II.2002, leg. PC, NN, AT, CD & WJ, 1♀; PHITSANULOK Prov., Noenmaprang
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Dist., Tham Pha Tha Phon non hunting area, 80 km SE Phitsanulok city, Hua Khao village, lime stone mountain, Klong Wang Kang river outside Lod Cave, 5.III.2001, leg. PC, SP & WW, C0108, 1♀ brach.; Nakorn Thai Dist., Phu Hin Rong Kla NP, near Huai Khammunnoi waterfall, 1100m, 6.III.2001, leg. PC, SP & WW, C0109, 1♂ macr.; Thung Saleang NP, stream, 400m asl., 6.III.2002, leg. PC & NN., C0252, 2♂ 1♀ brach., 1♂ macr., 5Lv.; Muang Dist., outskirts of Phitsanulok City, Biological Control Centre, frog rearing pond, 7.III.2001, leg. PC, SP & WW, C0114a, 6♂ 6♀ brach., 4Lv.; outskirts of Phitsanulok City, fish pond near agricultural station of Naresuan University, 7.III.2001, leg. PC, SP & WW, C0115, 2♀ brach., 7Lv.; PHICHIT Prov., Nachilaparami Dist., 18 Km S of Pichit City, abandoned rice fields and road ditch, 15.IX.2004, leg. PC, NN, KB & TJ, C0414, 3♂ brach., 7Lv.; NAKHON SAWAN Prov., Muang Dist., Beung Boraphet NP, Beung Boraphet Lake, resort area, at edge of lake between emerging and floating vegetation including terrestrial plants growing into the water, 15.IX.2004, leg. PC, NN, KB & TJ, CN0410, 1♂ 2♀ brach.; Beung Boraphet NP, lotus pond at the gate to Freshwater Fisheries Museum, 15.IX.2004, leg. PC, NN, KB & TJ, CN0411, 1♀ brach.; Beung Boraphet Lake, canal in marsh near office station of National Park, 15.IX.2004, leg. PC, NN, KB & TJ, CN0412, 5♂ brach., 1♀ macr.; Muang Dist., Sawan park in the centre of Nakhon Sawan City, small lake with edge of Poaceae, which are trimmed and kept neat, 15.IX.2004, leg. PC, NN, KB & TJ, C0413, 1♂ macr.; SAKON NAKHON Prov., constructed wetland for water treatment near Nong Haan, overgrown marshy pond, Eichornia, Hydrilla, Ipomoea, Lotus, 23.XI.1995, leg. NN, N9526, 4♂ 6♀ brach., 1♂ 1♀ macr.; KHON KAEN Prov., Nam Phong Dist., 50 km N of Khon Kaen city, Nam Phong river, 27.XII.1994, leg PC & TH, C9426, 4♂ 6♀ brach., 1♂ 1♀ macr., 1Lv.;Phe Pen Kham NP, Baan Noon Hua Chang, Huay Sem Caen, 21.XI.1995, leg. H. Zettel, 1♂ 1♀ brach., 1♀ macr. (NHMW); Baan Noon Hua Chang, in reserved area upstream of Naam Phong dam, large pool upstream of cascade, over 1m deep, partly shaded by trees, some plants growing in the water, 21.XI.1995, leg. NN & TH, N9521, 4♂ 1♀ brach.; 10 min walk upstream of Naam Phong dam, open area, roockpools, small potholes, 21.XI.1995, leg. NN & TH, N9523, 2♂ 3♀ brach.; Muang Dist., Namphon River, down from Wat Tha Song Korn, 12 km E of Khon Kaen, polluted fishery pond, 28.XII.1994, leg. PC & JH, C9430, 10♂ 11♀ brach.; CHAYAPHUN Prov., Tat Ton NP, Tat Ton, 27.XI.1995, leg. H. Zettel, 1♂ brach. (NHMW); UBON RATCHATHANI Prov., Warin Chamrap Dist., pond alongside Moon River, 25.XII.1994, leg. PC & JH, C9423, 2♂ 3♀ brach.; SURIN Prov., Surin, 150m asl., 5-10.XII.1995, leg. P. Schwendinger, 1♂ macr. (NHMW); CHON BURI Prov., Khao Khaew Open Zoo, ponds, 6.IV.2001, leg. PC, BK & S. Leepitakrat, C0158, 1♂ 2♀ brach., 7Lv.; KANCHANABURI Prov., Thonphaphom Dist., Ban Taotan, Kayen stream, 25.II.2002, leg. N. Sangpradub, S0205 1♀ brach.; NAKHON SI THAMMARAAT Prov., Ron Phi Bun Dist., Mae Sethi watrefall, stream with lowland stream aspect downstream of waterfall, 15.XI.2005, leg. PC, NN & SP, CN0473, 1♀ brach., 1♀ macr.; SATUN Prov., Taleh Ban NP, small lake in front of office, depth over 1m, sampled at edge with marsh plants and Hydrilla, 26.XI.1995, leg. NN, N9530, 10♂ 21♀ brach.
Distribution. - Widespread from Burma through continental SE Asia, Philippines, and Indonesia to New Guinea and N Australia and to SW China (Nieser et al. 2005). In Thailand common throughout the country, found mostly in stagnant parts of streams and ponds not connected with streams. Diagnosis. Brachypterous form (Fig. 63). Length, male 3.6-3.9, female 4.2-4.8; width male 1.0-1.3, female 1.2-1.4. Eyes reddish, pronotum and scutellum whitish,
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hemielytra largely translucent with the whitish dorsum of abdomen shining through. Hemielytra along costal margin with a pruinose band which is variably infuscated, base of hemielytra also pruinose, in females extending caudally to a white to brownish pruinose band running to about one third the length of hemielytron, midway between costal and inner margins; some females with two variable brown spots cross hemielytral commissure. In dorsal view, vertex somewhat protruding in front of eyes; eyes holoptic posteriorly. Width of pronotum, male 0.9-1.0, female 1.0-1.1; pronotum laterally with a pair of foveae. Basal width of scutellum, both sexes 0.4-0.7; median length of scutellum 0.5-0.8. Hemielytra without claval suture, membrane reduced, hind wings absent. Male, fore tarsi two-segmented. Female, fore tarsi, one-segmented. Macropterous form. length, male 3.9-4.6, female 4.4-4.7; width, male 1.2-1.4, female 1.3-1.4. Colour as in brachypterous form except for hemielytra: clavus largely translucent; corium and mebrane pruinose; corium with a narrow longitudinal median translucent stripe; membrane whitish. Hind wings translucent with a vague whitish hue. Structurally essentially as brachypterous form except for: scutellum more strongly developed, basal width 0.9-1.2, median length 0.9-0.11. Hemielytra with claval suture present and membrane fully developed. Hind wings fully developed. Comparative notes. Immediately recognizable by the posteriorly holoptic eyes in combination with the lateral pronotal foveae.
РЕЗЮМЕ Това е едно изследване на Notonectidae от Тайланд с ключ за определяне на мъжките и женските, с диагнози за всички видове от континентална югоизточна Азия и данни за разпространението им в Тайланд. Пет вида са съобщени за първи път за Тайланд: Anisops exiguus HORVÁTH, A. lansburyi LEONG, A. niveus (FABRICIUS), A. tahitiensis LUNDBLAD и Enithares malayensis BROOKS. Освен това за първи път са съобщени Anisops exiguus за Индонезия и Лаос; A. nigrolineatus LUNDBLAD за Лаос и Виетнам; и Enithares sinica (STÅL) за Лаос.
REFERENCES Bare C.O. (1928). Haemoglobin cells and other studies of the genus Buenoa (Hemiptera Notonectidae). — The University of Kansas Science Bulletin, 18: 265-349. Blaustein L., M. Kiflawi, A. Eitam, M. Mangel & J.E. Cohen (2004). Oviposition habitat selection in response to risk of predation in temporary pools: mode of detection and consistency across experimental venue. — Oecologia, 138: 300-305. Brooks G.T. (1948). New species of Enithares (Hemiptera, Notonectidae). — Journal of the Kansas Entomological Society, 21: 37-54. Brooks G.T. (1951). A revision of the genus Anisops (Notonectidae, Hemiptera). — University of Kansas Science Bulletin, 34: 301-519.
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Chen P.-p., N. Nieser & W. Wattanachaiyingcharoen (2002). A new genus, Pleciogonus and four new species of semiaquatic and aquatic bugs from Thailand (Heteroptera: Gerromorpha, Nepomorpha), Tijdschrift voor Entomologie, 145: 193-212. Chen P.-p., N. Nieser & H. Zettel (2005). The aquatic and semiaquatic bugs (Heteroptera: Nepomorpha & Gerromorpha) of Malesia. — Fauna Malesiana Handbook 5. Brill, Leiden & Boston: i-x + 1-546. Chesson J. (1984). Effect of notonectids (Hemiptera: Notonectidae) on mosquitoes (Diptera: Culicidae): predation or selective oviposition ? — Environmental Entomology, 13: 531-538. Chowanadisai L. (1986). Giant back swimmer (Enithares templetoni) a possible bio-control agent for Anopheline larvae. — Mosquito Borne Diseases Bulletin, 3: 34-36. Distant L. (1911). Rhynchota 5. — The fauna of British India, including Ceylon and Burma. Taylor & Francis London: i-xii + 1-362. Eitam A., L. Blaustein & M. Mangel (2002). Effects of Anisops sardea (Hemiptera: Notonectidae) on oviposition habitat slection by mosquitoes and other dipterans and on community structure in artificial pools. — Hydrobiologia, 485: 183-189. Esaki T. (1928). Aquatic and semiaquatic Heteroptera. — Insects of Samoa, 2: 47-80. Esaki T. (1930). Übersicht über die Insektenfauna der Bonin (Ogasawara) Inseln, unter besonderer Berücksichtigung der zoogeographischen Faunencharaktere. — Bulletin of the Biogeographical Society of Japan, 1: 205-226. Fabricius J.C. (1775). Systema entomologiae, sistens insectorum clases, ordines, genera, species, adjectis synonymis, locis, descriptionibus, observationibus. Kortii Flensburgi & Lipsiae: i-xxx, 1-832. Fernando C.H. & C.Y. Leong (1976). A taxonomic study of the Malayan Notonectidae (Hemiptera: Heteroptera). — Indian Journal of Zoology, 4: 15-34. Fieber F.X. (1851). Rhynchotographieen. — Calvae Pragae: 1-64. [preprint of Abhandlungen der Böhmischen Gesellschaft der Wissenschaften, 1852; 7: 469-486.] Gorai A.K. & D.N.R. Chaudhuri (1962). Food and feeding habits of Anisops bouvieri Kirk. (Heteroptera: Notonectidae). — Journal of the Asiatic Society, 4: 134-139. Hanboonsong Y., P. Mungkandee & H. Zettel (1996). A preliminary list of aquatic and semiaquatic Heteroptera (Nepomorpha, Gerromorpha) from the Maekhong-Chi-Mung rivers area, Thailand. — Amemboa, 1: 19-24. Herrich-Schäffer G.A.W. (1850). Die Wanzenartige Insekten, 9. — Lotzbeck Nürnberg: 1-44. Hungerford H.B. (1922). Oxyhaemoglobin present in back swimmer, Buenoa margaritacea Bueno. — Canadian Entomologist, 54: 262-265. Hutchinson G.E. (1927). New or little known Notonectidae (Hemiptera-Heteroptera). — Annals and Magazine of Natural History (9), 19: 375-379. Kirkaldy G.W. (1901). On some Rhynchota, principally from New Guinea. (Amphibicorisae and Notonectidae). — Annali del Museo Civico di Storia Naturale Giacomo Doria, 20: 804-810. Kirkaldy G.W. (1904). Über Notonectiden (Hemiptera) I, II. — Wiener Entomologische Zeitung, 6: 93-135. Lansbury I. (1961). Notes on the genus Anisops (Hemiptera, Notonectidae) in the Hungarian National Museum. — Annales Historico-Naturales Musei Nationalis Hungarici, 53: 505-506.
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Lansbury I. (1962). Notes on the genus Anisops in Bishop Museum. — Pacific Insects, 4: 141-151. Lansbury I. (1964a). Some observations on the Notonectidae (Hemiptera-Heteroptera) of Viêt-Nam and adjacent regions. — Annales Zoologici Warszawa, 22: 203-219. Lansbury I. (1964b). A revision of the genus Paranisops Hale (Heteroptera: Notonectidae). — Proceedings of the Royal Entomological Society of London (B), 33: 181-188. Lansbury I. (1965a). Notes on the species of Anisops Spin. (Hemiptera-Heteroptera, Notonectidae) of Java. — Annales Zoologici Warszawa, 23: 57-68. Lansbury I. (1965b). A new tribe and genus of Notonectinae (Heteroptera: Notonectidae) from Borneo. — Pacific Insects, 7: 327-332. [not cited in the text] Lansbury I. (1966). Notes on the genus Aphelonecta (Hemiptera-Heteroptera: Notonectidae). — Pacific Insects, 8: 629-632. Lansbury I. (1968). The Enithares (Hemiptera-Heteroptera: Notonectidae) of the Oriental Region. — Pacific Insects, 10: 353-442. Lansbury I. (1973). Notes on the genus Enithares (Hem., Notonectidae). — The Entomologist’s Monthly Magazine, 109: 226-231. Lansbury I. (1984). Some Nepomorpha (Corixidae, Notonectidae and Nepidae) (HemipteraHeteroptera) of North-west Australia. — Transactions of the Royal Society of South Australia, 108: 35-49. Lansbury I. (1985). Notes on the identity of Nychia Stål (Hemiptera-Heteroptera: Notonectidae) in Australia. — The Beagle, Occasional papers of the Northern territory Museum of Arts and Sciences, 2: 1-9. Leong C.Y. (1962). The life-history of Anisops breddini Kirkaldy (Hemiptera, Notonectidae) — Annals and Magazine of Natural History (ser.13), 5: 377-383. Leong C.Y. (1963). Anisops lansburyi sp. n. (Hemiptera: Notonectidae) from Malaya. —Proceedings of the Royal Entomological Society of London (B), 32: 101-102. Liu G. & L. Zheng (1991). A new species of the genus Enithares from China (Hemiptera, Notonectidae). — Acta Scientiarum Naturalium Universitatis Nan Kaiensis : 66-68 (in Chinese with description of E. tibialis also in English). Lundblad O. (1933a). Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von Sumatra, Java und Bali. — Archiv für Hydrobiologie, Supplement Band 12: 1-195, 263-488, pls. 1-21. Lundblad O. (1933b). Some new or little known Rhynchota from China. — Annals and Magazine of Natural History (10), 12: 449-464. Lundblad O. (1934). Aquatic and semiaquatic Heteroptera of Tahiti. — Bernice P. Bishop Museum Bulletin, 113: 121-126. Matsumura R. (1915). Uebersicht der Wasser-Hemipteren von Japan und Formosa. — Entomological Magazine, Kyoto, 1: 103-119. Miller P.L. (1964). The possible role of haemoglobin in Anisops and Buenoa (Hemiptera: Notonectidae). — Proceedings of the Entomological Society of London (A), 39: 166-175. Miura T., R.M. Takahashi & R.J. Stewart (1979). Habitat and food selction by the mosquitofish (Gambusia affinis). — Proceedings of the California Mosquito Vector Control Association, 55: 45-49. Miura T., R.M. Takahashi & W.H. Wilder (1984). Impact of the mosquitofish (Gambusia affinis) on a rice field ecosystem when used as a mosquito control agent. — Mosquito News, 44: 510-516.
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Miyamoto S. (1976). On the scientific names concerning Japanese Heteroptera. 1. — Rostria, 26: 197-198. Nieser N. (1996). An illustrated key to the families of Nepomorpha in Thailand. — Amemboa, 1: 4-9. Nieser N. (1998). Introduction to the Notonectidae (Nepomorpha) of Thailand. — Amemboa, 2: 10-14. Nieser N. (2004). Guide to Aquatic Heteroptera of Singapore and Peninsular Malaysia III. Pleidae and Notonectidae. — The Raffles Bulletin of Zoology, 52(1): 79-96. Nieser N. & P.-p. Chen (1991). Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). — Tijdschrift voor Entomologie, 134: 47-67. Nieser N. & P.-p. Chen (1996). Six new taxa of Nepomorpha from Sulawesi and Mindanao. — Tijdschrift voor Entomologie, 139: 161-174. Nieser N. & P.-p. Chen (2005). The water bugs (Hemiptera: Nepomorpha and Gerromorpha) of Vanuatu — Tijdschrift voor Entomologie, 148: 307-327. Nieser N., P.-p. Chen, P. Leksawasdi, A. Thanyakam & C. Duangsupa (2004). Five new species of Nepomorpha (Heteroptera) from Southeast Asia. — Tijdschrift voor Entomologie, 147: 29-40. Nieser N., P.-p. Chen & C.M. Yang (2005). A new subgenus and six new species of Nepomorpha (Insecta: Heteroptera) form Yunnan, China. — The Raffles Bulletin of Zoology, 53: 189-209. Nieser N. & H. Zettel (2001). First record of Paranisops Hale, 1924 (Insecta: Heteroptera: Notonectidae) from Southeast Asia, with description of P. leucopardalos sp. n. — Annalen des Naturhistorischen Museums in Wien, 103B: 243-247. Paiva C.A. (1918). Aquatic Rhynchota from the Southern Shan States. — Records of the Indian Museum, 14: 19-32. Papáek M. (2000). Chapter 24. Small aquatic bugs with slight or underestimated economic importance. — In: Schaefer, C.W. & A.R. Panizzi (Eds.): Heteroptera of Economic Importance, CRC Press Boca Raton: 591-595. Papáek M. (2001). Small aquatic and ripicolous bugs (Heteroptera: Nepomorpha) as predators and prey: the question of economic importance. — European Journal of Entomology, 98: 1-12. Poisson R.A. (1926). L’ Anisops producta Fieb. (Hemiptera Notonectidae) observations sur son anatomie et sa biologie. — Archives de Zoologie Experimentale et Generale, 65: 181-208. Polhemus J.T. (1995). Family Notonectidae Latreille, 1802 – backswimmers. — In: Aukema, B. & C. Rieger (Eds.): Catalogue of the Heteroptera of the Palaearctic Region 1. The Netherlands Entomological Society, Amsterdam: 63-73. (ISBN 90-71912-12-4). Schuh R.T. & J.A. Slater (1995). True Bugs of the World (Hemiptera: Heteroptera) classification and natural history. — Cornell University Press, Ithaca & London: i-xii, 1-336 Stål C. (1854). Nya Hemiptera. — Öfversigt af Kungliga Vetenskapsakademiens Förhandlingar, 11: 231-235. Stål C. (1860). Hemiptera. Species nova descripsit. — Konglia Svenska Fregattens Eugenies Resa Omkring Jorden 1851-1853, 2: 219-298. Štys P. & A. Jansson (1988). Check-list of recent family-group and genus-group names of Nepomorpha (Heteroptera) of the world. — Acta Entomologica Fennica, 50: 1-44.
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Tawfik M.F.S., M.M. El-Husseini & H. Abou Bakr (1986). The biology of the notonectid Anisops sardea H.-S., an active mosquito predator in Egypt. — Bulletin de la Societé Entomologique d’Egypte, 66: 127-144. Truxal F.S. (1953). A revision of the genus Buenoa (Hemiptera Notonectidae). — The University of Kansas Sience Bulletin, 35: 1351-1523. Wells R.M.G., M.J. Hudson & T. Brittain (1981). Function of the hemoglobin and the gas bubble in the backswimmer Anisops assimilis (Hemiptera: Notonectidae). — Journal of Comparative Physiology, 142: 515-522. Zettel H. (1995). Neubeschreibungen von zwei Arten der Gattung Aphelonecta Lansbury aus Indonesien und den Philippinen sowie ein Bestimmungsschlüssel zu den Arten (Heteroptera: Notonectidae). — Linzer Biologische Beiträge, 27: 107-113.
S. Grozeva & Austral N. Simov (Eds) 2008Polynesia (Heteroptera, Veliidae) 293 A new genus of Microveliinae from the Islands, French ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 293-302. © Pensoft Publishers Sofia–Moscow
A new genus of Microveliinae from the Austral Islands, French Polynesia (Heteroptera, Veliidae) J.T. Polhemus1 & D.A. Polhemus2 1
Colorado Entomological Institute, 3115 South York Street, Englewood, Colorado, USA 80013 2 Dept. of Natural Sciences, Bishop Museum, 1525 Bernice St., Honolulu, HI 96817
ABSTRACT A new genus of Microveliinae, Tubuaivelia nov. gen. with type species T. michaili nov. sp., is described from Tubuai Island in the Austral Islands of French Polynesia. This represents the first endemic genus of Microveliinae known in the Pacific region east of the Tonga Trench. Comparisons are provided to other microveliine genera in the Pacific region, accompanied by illustrations of the female dorsal habitus, male venter, male genitalia, and tarsal structures. A distribution map and photograph of the type locality habitat are also provided, along with a discussion of the zoogeographic questions raised by the discovery of this new genus. Keywords: Heteroptera, Veliidae, new genus, new species, Austral Islands, Tubuai
INTRODUCTION In recent years, a number of new genera in the subfamily Microveliinae have been described from the Pacific region (Polhemus & Polhemus 1994, 2006; Polhemus & Polhemus 2000; Andersen & Weir 2001). To the east of the Tonga Trench, however, only members of the cosmopolitan genus Microvelia have heretofore been recorded. In this paper, we report the existence of a new, endemic genus of Microveliinae on the remote and relatively old island of Tubuai in the Austral Islands of French Polynesia, a discovery which substantially changes our biogeographic perception of this subfamily in the insular Pacific.
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The relationships of the Australasian microveliinae genera were reviewed by Andersen & Weir (2001); we consider this analysis correct, and a supplement to it is beyond the scope of the present paper. We would note, however, that two additional genera of Microveliinae have been subsequently discovered in the Pacific: Fijivelia on the Fijian islands of Kadavu and Viti Levu (Polhemus & Polhemus 2006), and the new genus from the Austral Islands, described herein as Tubuaivelia. These newly described genera necessitate a revision of the phylogenetic analysis of Andersen & Weir (2001), a project that when completed will provide significant new insights regarding the dispersal and diversification of this group within the Pacific region over time.
MATERIAL AND METHODS The collection of the new genus described herein occurred during a program of intensive aquatic biodiversity surveys on the islands of French Polynesia. Collections at a given site were made by visual searching, hand netting, and localized pyrethrin fogging of riparian and hygropetric habitats. Specimens were preserved in 75% ethanol, then transported to the Bishop Museum in Honolulu, Hawaii and the Colorado Entomological Institute in Englewood, Colorado for detailed analysis and identification. The holotype and allotype are deposited in the former institution, while paratypes are deposited in the latter. All measurements are given in millimeters. The CL number in the material examined section refers to a collection locality number series used by the authors to cross reference specimens with photographs, field notes, and other metadata.
Tubuaivelia nov. gen. (Figs 1–7)
Description. Size.- Apterous form, length of males 2.38–2.84 mm., females 2.48–2.94 mm; general body characteristics and size not sexually dimorphic, males and females similar. Colour.- Ground color brown, with extensive yellowish brown markings; head mostly yellowish brown dorsally, with a weakly developed median dark stripe, posteriorly meeting a well developed U-shaped brown marking reaching along inner eye margins, following ecdysial line; pronotum anteriorly with a yellowish transverse stripe behind vertex of head, not interrupted medially; head, pronotum, parts of abdomen laterally, acetabulae bluish pruinose; pronotal lobe mostly yellowish brown; mesonotum yellowish medially; connexiva mostly yellowish, segement VI bluish pruinose, dark at intersegmental sutures; female abdominal tergites I – VII yellowish medailly, II–III mostly bluish pruinose; venter mostly yellowish. Structural characters.- Only apterous forms known, shape narrow-elongate (Figs 1, 2). Head with eyes globose, exserted but not beyond anterolateral pronotal angles, separated by about three times the width of an eye, appressed against anterior pronotal
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Fig. 1: Tubuaivelia michaili, nov. sp., female, dorsal habitus.
Fig. 2: Tubuaivelia michaili, nov. sp., male, ventral view of body, rostrum omitted to show rostal groove.
margin, with tiny ocular setae. Head sloping anteriorly, posterior margin sloping caudodorsally, extending posteriorly almost one eye length, with usual three pairs of facial trichobothria plus numerous short setae; vertex with black glabrous pseudocellar spots, without evident median sulcus; gular region moderately long, plainly visible, rostral cavity not closed posteriorly, but open and V-shaped (Fig. 2). Rostrum long, reaching to posterior margin of mesosternum, segment I short, distally visible, segment II even shorter, segment IV about 6 times the length of II, segment III more than 13 times as long as II. Antennae slender, segments III and IV long, total length slightly more than 3/5 of body length. Thorax with pronotum not raised, posterior lobe slightly depressed, without median carina; collar absent; anterior and posterior lobes of about equal length, set off by a transverse row of a few obscure foveae, evanescent laterally; entire dorsum set with short decumbent dark and golden setae; mesonotum on midline slightly more than half of pronotal length, raised posteriorly, without wing pads; humeri not prominent, exposing metanotum laterally. Prothoracic acetabulae separated by the width of the rostrum, mesosternum depressed medially, forming an obvious rostral groove. Thoracic venter not diagnostic, with weakly formed tubercles on either side of mesosternal midline on posterior margin opposing an unmodified metasternum. Metasternal scent gland opening (omphalium) small but visible, marked by a small tubercle; scent channels prominent, curving slightly anterad to base of metacetabulae. Wing pads not present. Abdomen broad, with two (1 + 1) foveae on either side of midline between tergites I and II appearing silvery; without longitudinal carinae on tergites; tergites II – VI sub-
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equal in length, I shorter, VII longer. Abdominal sternites not set off from laterosternites; male ventrites not modified. Legs slender, long, hind legs longer than others; anterior femur set beneath with short light setae, unmodified in males; anterior tibia of male with a comb of minute black setae occupying about 1/7 of tibial length; middle tibia slightly curved; middle and hind femora set ventrally with long slender setae; all tarsi of moderate length (Fig. 1), claws long, slender; both up- and down curving arolia of moderate length, slender. Male genitalia with ventrite VII not modified; pygophore simple, unmodified; parameres small, rounded, symmetrical (Fig. 4); proctiger strongly modified, with prominent, wing-like lateral projections (Fig. 3); tergite IX subtriangular, rounded at apex, not protruding. Female with tergite VIII on same plane as VII, slightly sloping posteriorly; proctiger forming an anal plate or lid, bent ventrad, concealing gonocoxae and genital opening. Type-species. – Tubuaivelia michaili nov. sp. Etymology. – The generic name Tubuaivelia is derived from Tubuai, the island of discovery, and Velia, the nominate genus of the family. Gender feminine. Biological notes. – The only known species of Tubuaivelia was collected at the base of a mossy, vertical, seeping rheocrene formed by the forewall of a water diversion dam on the upper midreach of a tributary to the Vaiapu River in central Tubuai (Figs 6, 7). The insects were taken by applying a light pyrethrin fog to this wet face, and collected when they fell into the pool below. Although we suspect this genus may be hygropetric, it was not possible to establish at the time of collection whether the insects were inhabiting the vertical wet wall, or sheltering in dark pockets at its base. Comparative notes. - Tubuaivelia differs from all other microveliine genera by the rostral cavity open posteriorly and posterior margin V-shaped (Fig. 2); by possessing a very long rostrum that reaches past the posterior margin of the mesosternum; by possessing wing-like processes laterally on the male proctiger; and by lacking a median sulcus on the head, with only a faint dark line present instead. Tubuaivelia also has the head posteriorly produced, extending well behind the hind margin of the eyes (Fig. 1).
4
3
5
Figs 3–5: Tubuaivelia michaili, nov. sp., structural details. Fig. 3. Male proctiger. Fig. 4. Male paramere. Fig. 5. Male terminal tarsal segment, hind leg.
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This latter character state is shared with species of Aegilipsicola, Tanyvelia (Polhemus & Polhemus 1994), Fijivelia (Polhemus & Polhemus 2006), Nesidovelia and Microvelopsis (Andersen & Weir 2001). Distribution. – Based on current knowledge the genus Tubuaivelia is monotypic, and is apparently endemic to the island of Tubuai in the Austral Islands of French Polynesia (Fig. 6).
Tubuaivelia michaili nov. sp. (Figs 1–7)
Type material: Holotype, apterous male: FRENCH POLYNESIA, Austral Islands, Tubuai Island, headwater reach of Vaiapu River at diversion weir above end of service road, 90 m. [295 ft.], 23°21’58”S, 148°28’33”W, 9 Sept. 2007, CL 5092, J. T. & D. A. Polhemus (BPBM). - Paratypes: FRENCH POLYNESIA, Austral Islands, Tubuai Island: 3 apterous males, 4 apterous females, same data as holotype, J. T. & D. A. Polhemus (BPBM, JTPC). Description. Size.- Apterous male, length 2.38–2.84 mm (= 2.54, n = 4); width 0.79–0.80 mm (= 0.79, n = 4). Apterous female, length 2.48–2.95 mm (= 2.67, n = 4); width 0.83–0.92 mm, (= 0.88, n = 4). Colour.- Apterous male: see generic description.
5
Fig. 6: Satellite view of Tubuai island showing location of type locality for Tubuaivelia michaili, nov. sp. (dark square) in the upper Vaiapu River catchment.
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Structural characters.- Apterous male: see generic description, with following additions: Head of moderate length, sloping anteriorly; length 0.51, width 0.53; width of eye/ interocular space, 0.09/0.35. Pronotum of moderate length, posterior margin straight medially length (midline): width, 0.23 : 0.70; mesonotum length, 0.12; humeri obscure. Abdomen with tergites with I–VI subequal, tergite VII longest (0.16, 0.19, 0.16, 0.14, 0.16, 0.16, 0.28 respectively). Abdominal venter bearing short, fine, pale setae; ventrites IV–VI depressed medially; first genital segment (segment VIII) with a small semi-circular lip medially directed ventrad; genital capsule small, not protruding.
7 Fig. 7: Habitat of Tubuaivelia michaili, nov. sp., on the upper Vaiapu River in central Tubuai. Specimens were taken by applying a light pyrethrin fog to the wet lower face of the cement wall along the downstream side of the diversion structure.
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Legs clothed with numerous short, appressed, pale setae, intermixed on ventral surface of fore femur with longer, erect, pale, pilose setae and a few longer dark setae; all legs unarmed, lacking setal tufts or other modifications. Antennae thickly clothed with short to long setae, segments I and IV also bearing a few long, spine-like black setae. Antennae slender, formula I : II : III : IV; 0.35 : 0.28 : 0.40 : 0.47. Legs with proportions as follows: Femur, tibia, tarsal 1, tarsal 2 of fore leg, 0.65 : 0.47 : 0.23 : 0.0; of middle leg, 0.84 : 0.70 : 0.09 : 0.19; of hind leg, 0.88 : 1.02 : 0.14 : 0.23. Male genitalia with parameres small, rounded, symmetrical (Fig. 4); proctiger with prominent tapering, symmetrical, wing-like lateral processes (Fig. 3). Apterous female: Similar to male in general structure and coloration, but slightly larger. Connexiva not raised, converging posteriorly but not touching, evenly tapering toward tip of abdomen, posterolateral right angles slightly infolded, thickly set with inwardly directed long black setae; abdominal venter unmodified. Etymology. - The name michaili honors Michail Josifov for his many contributions to the study of Heteroptera.
DISCUSSION Tubuaivelia is similar in size and general appearance to a suite of microveliine genera occurring further to the west in Melanesia and Australia. In particular, the posterior projection of the head vertex behind the eyes is a character state shared within the Microveliinae with the genera Aegilipsicola and Tanyvelia from New Guinea, Fijivelia from Fiji, and Nesidovelia and Microvelopsis from Australia. This would suggest that the ancestor of Tubuaivelia originated in the Melanesian region and reached the Austral Islands from the west. The alternative hypothesis would be that Tubuaivelia is a local derivitive of the Microvelia prompta lineage, members of which are widespread on the archipelagoes of French Polynesia (including the Austral Islands, where undescribed species occur), but this would require a significant amount of convergent evolution in relation to the Melanesian genera mentioned above. The prominent lateral processes on the proctiger of Tubuaivelia are a unique character state within the Microveliinae, although many other microveliines (and many other Veliidae) do possess much smaller, angulate lateral processes on the basal portion of the proctiger. Similarly, the posteriorly open rostral cavity with its V-shaped posterior termination, the long rostrum, and the prominent medial rostral groove on the ventral thorax are all apomorphic states within the Microveliinae. In addition, the rostrum reaching between the hind coxae and the open V-shaped posterior margin of the rostral cavity are also unique character states for Tubuaivelia when compared to all other described genera of Microveliinae, as well as a number of undescribed genera in the J. T. Polhemus Collection. Tubuaivelia thus stands as highly modified taxon that has apparently evolved unique character states in extreme isolation.
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ZOOGEOGRAPHIC CONSIDERATIONS The island of Tubuai lies in the south-central Pacific Ocean, approximately 640 km. south of Tahiti at latitude 23°23’S and longitude 149°27’W. With a total land area of 45 km2, Tubuai is the largest of the Austral Islands (also known as the Tubuai Islands), a hotspot chain extending for a distance of 1500 km in an ESE-WNW direction, subparallel to the Society Islands (which include Tahiti) to the north. The Australs are the geologically oldest hotspot-derived high islands currently extant on the Pacific Plate (Clouard & Bonneville 2001), with the high islands in their northwestern sector (Rimatara, Rurutu, and Tubuai) retaining significant elevation even at geological ages that equate to an atoll-forming stage in other hotspot chains. This anomalous elevation in relation to age appears to be linked to the composition of the islands’ basalts, which consist of phonolites that possess isotopic signatures indicating a mixing of upper mantle melts with previously subducted oceanic crust, presumably as a result of the Australs hotspot crossing an ancient subduction zone (Dupuy et al. 1988). This has produced hard basalts with exceptional resistance to erosion. Tubuai contains two distinct eruptive epicenters, with ages ranging from 12.4-9.5 Ma (Liotard & Barsczus 1989; Maury et al. 1994). The Hanareho volcano in the west is a breached cone with an age of 9.5 Ma, while the Herani volcano in the east is older, composite structure, with ages of 12.4 Ma at Mt. Panee, and 9.9 Ma at Mt. Taitaa. The latter peak is also the highest point on Tubuai, with an elevation of 422 m. The island is also surrounded by an extensive barrier reef and lagoon, with the extent of the enclosed platform indicating that the island was once over twice its current size (Fig. 7). Of the two volcanoes, only the older Herani volcano, on which Mt. Taitaa lies, still retains perennial streams. Tubuai and the older Austral islands to the northwest, Rimatara and Rurutu, represent some of the most persistent high islands presently found in the insular Pacific. Although both Rimatara (28.6–14.4 Ma)* and Rurutu (12.0–8.4 Ma) are still high islands with ages older than Tubuai (Clouard & Bonneville 2001), both the size and elevation of Rimatara are now relatively minimal, the island having a total land area of only 8 km2 and a maximum elevation of 83 m at Mt. Uahu, while Rurutu has been secondarily elevated following a period of erosion in which its area and elevation were significantly reduced from its current extent of 32 km2 with a maximum elevation of 389 m at Mt. Taatioe. By contrast, the island of Raivavae (6.6 Ma), the youngest high island considered unequivocally part of the Austral chain (Rapa may be of separate origin), has a maximum elevation of 437 m. at Mt. Hiro, and is therefore barely higher than Tubuai despite being only half its age. Therefore within the old hotspot archipelago of the Australs, it is Tubuai that still retains a biologically significant combination of relatively large size, elevation and age. The above factors are reflected in the composition of the Heteroptera biota occurring on Tubuai. The island supports no less than 3 endemic monotypic genera of emesine *
Although Rimatara is clearly older than Rurutu, the geological ages cited here, although the best available, were considered suspect by Clouard & Bonneville (2001).
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Reduviidae (Wygodzinsky 1966): Tubuaitaita, Tubuaia, and Pelmatomesa (although new species of the latter are known to us from both Rurutu and Raivavae). To this assemblage may now be added the endemic genus of Veliidae, Tubuaivelia, described herein. Based on recent collections, the island also supports an undescribed endemic species of Microvelia, allied to M. prompta from Tahiti, and an undescribed endemic genus of Miridae. This unusual assemblage of endemic elements makes Tubuai a land mass of particular zoogeographic significance in the south-central Pacific. The evolution of Tubuaivelia may therefore be the product of relatively long isolation. In the sector of the Pacific in which the Austral Islands lie, all surrounding island groups except the Tuamotu Archipelago to the northeast (47.4–25.1 Ma, and now worn down to atolls) and the Cook Islands to the northwest (18.8–1.5 Ma, with only a few atolls and small high islands remaining) are of significantly younger age (Clouard & Bonneville 2001). If the older age determinations on Rimatara are accepted, then the Austral Archipelago may have occupied an isolated position as the only major land masses in the south-central Pacific from the late Oligocene into the early Miocene, a period which also saw significant arc-related island building to the west in the Melanesian region (Polhemus 2007). Although the precise phylogenetic relationships of Tubuaivelia are as yet unknown, we consider it possible that they will be traced back to the southern sector of the Oligocene arcs that formed in the southwest Pacific, perhaps to Fiji, where another locally endemic microveliinae, Fijivelia, has recently been described (Polhemus & Polhemus 2006).
ACKNOWLEDGMENTS The authors wish to extend their profound gratitude to David Hembry and Brad Balukjian of the University of California, Berkeley, who provided unfailingly cheerful and competent companionship in the field during surveys in the Austral Islands. Special thanks are also due to the staff of the Gump Biological Station of the University of California, on Moorea, particularly Dr. Elin Claridge, for coordinating the logistics of our Australs survey. This research was supported by grant DEB-0451971 from the U. S. National Science Foundation, Washington, DC, and thanks are also due to this agency for its continued support of research on the taxonomy and biogeograpy of Indo-Pacific Heteroptera. In addition, JTP completed this research as an adjunct faculty member in the Bioagricultural Sciences Department at Colorado State University. This is contribution 2008-008 to the Pacific Biological Survey.
РЕЗЮМЕ Описан е нов род Tubuaivelia nov. gen. (Microveliinae) с типов вид T. michaili nov. sp. от остров Тубуай, Южните острови на Френска Полинезия. Това е първият ендемичен род от подсем. Microveliinae, известен за района на Тихия океан, източно от падината
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Тонга. Направено е сравнение с други родове Мicroveliinаe от Тихоокеанския район, c илюстрации на дорзалния хабитус на женските, коремната страна на мъжките, мъжките гениталии, и структурите на тарзуса. Представени са също карта на разпространение и снимка на типовото находище, с дискусия по зоогеографски въпроси, възникнали с откриването на този род.
REFERENCES Andersen N.M. & T.A. Weir (2001). New genera of Veliidae (Hemiptera: Heteroptera) from Australia, with notes on the generic classification of the subfamily Microveliinae. — Invertebrate Taxonomy, 15: 217–258. Clouard V. & A. Bonneville (2001). Age of seamounts, islands and plateaus on the Pacific Plate. — Université de Polynésie française, Faaa: 1-42. Dupuy C., H.G. Barsczus, J.M. Liotard & J. Dostal (1988). Trace element evidence for the origin of oceanic island basalts: an example from the Austral Islands (French Polynesia). — Minerology and Petrology, 98: 293–302. Liotard J.-M. & H.G. Barsczus (1989). Gènése des foïdites phonolitiques de Tubuai (Archipel des Australes, Océan Pacifique Sud): Intervention d’un magma d’affinité carbonatique. — C. R. Acad. Sci. Paris, 308, Ser. II: 1261–1266. Maury R.C., M. El Azzouzi, H. Bellon, J.-M. Liotard, G. Guille, H.G. Barsczus, C. Chauvel, C. Diraison, C. Dupuy, P. Vidal & R. Brousse (1994). Géologie et pétrologie de lîle de Tubuai (Australes, Polynésie français). C. R. Acad. Sci. Paris, 308, Ser. II: 1341–1347. Polhemus D.A. (2007). Tectonic geology of New Guinea, pp. 137–164. — In: A.J. Marshall & B.M. Beehler (Eds.): The Ecology of Papua, 1 Periplus Editions Ltd., Singapore: xxxi + 1-749. Polhemus D.A. & J.T. Polhemus (2000). Additional new genera and species of Microveliinae (Heteroptera, Veliidae) from New Guinea. — Tijdschrift voor Entomologie, 143: 91–123. Polhemus J.T. & D.A. Polhemus (1994). Four new genera of Microveliinae (Heteroptera, Veliidae) from New Guinea. — Tijdschrift voor Entomologie, 137: 57–74. Polhemus J.T. & D.A. Polhemus (2006) A new genus and two new species of Microveliinae from Fiji. — Russian Entomological Journal, 15 (2): 73–77. Wygodzinsky P.W. 1966. A monograph of the Emesinae (Reduviidae: Heteroptera). — Bulletin of the American Museum of Natural History, 133: 1–614.
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ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 303-307. © Pensoft Publishers Sofia–Moscow
Contribution to the knowledge of the Isometopinae (Heteroptera, Miridae) of the Balkan Peninsula Lj. Protić Natural History Museum Belgrade, Serbia. E-mail:
[email protected]
ABSTRACT Isometopus mirificus Mulsant & Rey was first recorded in Serbia 100 years ago in Ruma (Horváth 1907). We found a second specimen of this species in Serbia on June 29th, 2006 in the vicinity of Belgrade (locality Vrčin) on Pirus sp. We present also data on the Isometopinae for the area of ex Yugoslavia, as they were accidentally omitted in the Catalogue (Proti 1998). Keywords: Balkan Peninsula, Distribution, Isometopinae, Serbia
INTRODUCTION It is a great honor and responsibility to write a contributing article for this „Festschrift“, published for the celebration of 80 years of life of honored professor Michail Josifov. Professor Josifov is one of the few remaining heteropterists with the broad knowledge and ability to study the entire suborder Heteroptera. Recent times are marked with a decreasing number of heteropterists who are able to study the entire Heteroptera as does Professor Josifov. As I wished to contribute to a better knowledge of the Heteroptera fauna in the Balkan Peninsula, which was studied by the Professor throughout his life, I decided to write this modest contribution to the knowledge of Isometopinae, a subfamily well represented in the rich scientific opus of Professor Josifov. The greatest centers of distribution and diversity of the subfamily Isometopinae are the tropics. Altogether 31 genera and 170 species have been described worldwide (Akingbohungbe 1996).
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In the European fauna, Isometopinae are rare, represented by two genera: Isometopus and Myiomma.The genus Isometopus is represented by five species in Europe, three of them in the Balkan Peninsula: I. intrusus, I. longirostris and I. mirificus; and two species, I. nigritulus and I. peregrinus, are on Cyprus. The genus Myiomma is represented in the Balkan Peninsula by only one species, M. fieberi. Only single records of these species are so far known both for Europe and Balkan Peninsula (Fieber 1861; Puton 1873; Josifov 1964, 1974, 1990, 1993; Gogala & Gogala 1986, 1989; Rietschel 2000; Gogala 2006). In the Catalogue of Heteroptera of Yugoslav countries (Proti 1998) Isometopinae were accidentally omitted. This error is now corrected, as here we bring together all the data on this subfamily for the area of ex Yugoslavia. The second purpose for this contribution is the discovery of new specimens of Isometopinae collected in Serbia after more than 100 years.
LIST OF SPECIES The abbreviations of the states are according to the Catalogue of Heteroptera of the Palearctic Region (Kerzhner & Josifov 1999). In the Catalogue abbreviation YU included both Serbia and Montenegro, which became separate countries in 2006, and their new abbreviations are now RS for Serbia and ME for Montenegro. Isometopus intrusus (Herrich-Schaeffer, 1835) BU Asenovgrad on Populus ( Josifov 1964); Black Sea coast on Salix ( Josifov 1974); CR Košute-Sinj 5.6.1933. on Verbascum nigrum L. (Novak & Wagner 1955); RS Vrdnik (Horváth 1900); SL „Krain“ = Kranjska Carniola (Fieber 1861); Ljubljansko Barje: Log, Lukovica 17.7.1984, 26.8.1984, 24.7.1988, 30.6.1992, 29.6.2001. on Malus leg. A. & M. Gogala (Gogala & Gogala, 1986, 1989); Istra: Koštabona, Škrline, 21.6.2001, leg. I. Sivec (Gogala 2006). Catalogue of Heteroptera of the Palearctic Region / Distribution on Balkan Peninsula: BU CR GR RO SL YU Isometopus longirostris Josifov, 1993 BU Kressnensko-Hance, Kressna-r. 16.8.1984. 4 ♂, 14.8.1986. 2♂, 7.8.1991. 3♂. Female unknown ( Josifov 1993). Isometopus mirificus Mulsant & Rey, 1879 RS Ruma (Horváth 1907); Grocka: Vrčin 29.06.2006. 2♀, 1 larva on Pirus sp. leg. A. Stojanović NHMBeo (Fig. 1).
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Catalogue of Heteroptera of the Palearctic Region / Distribution on Balkan Peninsula: BU GR RO YU Isometopus nigritulus Akingbohungbe, 1996 CY Akingbohungbe 1996. Isometopus peregrinus Akingbohungbe, 1996 CY Akingbohungbe 1996. Myiomma fieberi Puton, 1872 BH Domanovići leg. Dr. Hensch (Horváth 1889). Thanks to the kindness of Dr B. Aukema, we cite the paper by Horváth (1889). Catalogue of Heteroptera of the Palearctic Region / Distribution on Balkan Peninsula: BH.
CONCLUSIONS We conclude that Isometopinae are rare in the Balkan Peninsula, as they have been recorded from only a handful of localities. Two species are represented in Serbia: I. intrusus and I. mirificus. So far, the literature data for these two species are for only one locality,
Fig. 1: Isometopus mirificus Mulsant & Rey, the specimen from Serbia. Drawing: ing. Aleksandar Stojanović, entomologist from Natural History Museum in Belgrade.
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in Srem. To the Heteroptera collection of the Natural History Museum in Belgrade was added in 2006 an additional specimen of I. mirificus from the broader vicinity of Belgrade, at the locality of Vrčin, from the trunk of an old pear tree. In this paper we would also like to substitute the citation of YU in the Catalogue of Heteroptera of the Palearctic Region for species Isometopus intrusus and I. mirificus in RS (Serbia).
РЕЗЮМЕ Подсемейство Isometopinae е представено на Балканския полуостров от четири вида. Само два вида Isometopus intrusus HERRICH-SCHAEFFER и Isometopus mirificus MULSANT & REY са съобщени за Сърбия преди 100 и повече години (HORVÁTH 1900, 1907). На 29 юни 2006 нов екземпляр от I. mirificus е намерен в околностите на Белград на Pirus sp. В Каталога по Heteroptera на Югославските републики (PROTIĆ 1998), Isometopinae е случайно пропуснат. Сега тази грешка е корегирана и са представени всички данни за това подсемейство в бивша Югославия.
REFERENCES Akingbohungbe A.E. (1996). The Isometopinae (Heteroptera: Miridae) of Africa, Europe, and the Middle East. — Delar Tertiary Publishers, Ibadan, Nigeria: 1-170. Fieber F. X. (1861). Die europäischen Hemipteren. Halbflügler. (Rhynchota Heteroptera). — Gerold’s Sohn, Wien:113-444. Gogala A. (2006). Heteroptera of Slovenia, III: Miridae. — Annales, Annals for Istrian and Mediterranean Studies, Series historia naturalis, 16(1): 77-112. Gogala A. & M. Gogala (1986). Seznam vrst stenic ugotovljenih v Sloveniji (Insecta: Heteroptera) . — Biološki Vestnik, 34: 21-52. [In Slovenian]. Gogala, A. & M. Gogala (1989). True Bugs of Slovenia (Insecta: Heteroptera). — Biološki Vestnik, 37: 11-44. Horváth G. (1889). Notes synonymiques et geographiques sur les hemipteres palearctiques. — Revue d’Entomologie, 8: 325-331. Horváth G. (1900). Fauna regni Hungariae. Animalium Hungariaehucusque cogitorum enumeratio systematica 111. Arthropoda Ordo: Hemiptera. Budapest. Horváth G. (1907). Supplementum ad Faunam Hemipterorum regni Hungariae. — Annales Musei Nationalis Hungarici, 5: 500-506. Josifov M. (1964). Artbestand und Verbreitung der Insekten von der Ordnung Heteroptera in Bulgaria, Teil II. — Bulletin de l’Institut et Musée de Zoologie, 16: 83-150 [In Bulgarian] Josifov M. (1974). Die Heteropteren der Bulgarischen Schwarzmeerkuste. — Bulletin de l’Institut de Zoologie et Musée, 39:5-27. [In Bulgarian]. Josifov M. (1990). Über die Verbreitung mancher Heteropterenarten auf der Balkanhalbinsel. III (Insecta) . — Acta Zoologica bulgarica, 40: 3-11.
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Josifov M. (1993). Eine neue mediterrane Isometopus-Art (Insecta: Heteroptera: Miridae: Isometopinae). — Reichenbachia Staatliches Museum für Tierkunde Dresden, 30(3): 7-8. Kerzhner I. M. & M. Josifov (1999). Family Miridae Hahn, 1833 – In: Aukema, B. & C. Rieger (Eds) Catalogue of the Heteroptera of the Palaearctic Region, The Netherlands Entomological Society, Amsterdam, Vol. 3: 1-576. Novak P. & E. Wagner (1955). II Prilog poznavanju faune Hemiptera Dalmacije (Hemiptera-Heteroptera). — Zaštita bilja 30, Dodatak Prilozi entomofauni Jugoslavije: 1-10. Proti Lj. (1998). Catalogue of the Heteroptera fauna of Yugoslav countries. Part one. — Natural History Museum, Belgrade. Special issue 38: 1-215 Puton A. (1873). Notes pour Servir à l’étude des Hémiptéres. — Annales de la Société entomologique de France, 3: 11-26. Rietschel S. (2000). Funde von Isometopus mirificus Mulsant & Rey, 1878 und Cardiastethus fasciiventris (Garbiglietti, 1869) im Oberrhein-Gebiet (Heteroptera, Isometopidae und Anthocoridae). — Heteropteron, 9: 9-11.
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ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 309-326. © Pensoft Publishers Sofia–Moscow
The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria * W. Rabitsch Federal Environment Agency, Spittelauer Lände 5, A-1090 Wien, Austria. E-mail:
[email protected] Department of Evolutionary Biology, University of Vienna, Althanstraße 14, A-1090 Wien, Austria. E-mail:
[email protected]
ABSTRACT During the past approximate 25 years 84 Heteroptera species were published for the first time for Austria. These species account for 9.4 % of the 896 Heteroptera species currently known to occur in Austria and means that species were added at a rate of more than 3 per year, including previously overlooked species. Two thirds of the species (68 %) were overlooked for taxonomic reasons or because of their cryptic appearance or biology; the remaining species (32 %) are considered new arrivals. Excluding single records, 18 species (21 %) are considered established new arrivals. This is a rate of 0.72 species establishing per year. Most of these species are generally expanding their distribution and increasing in abundance in Austria and elsewhere in Europe and most species reached Austria probably due to current favourable climatic conditions (14 species), followed by species introduced with ornamental plants or transport (4 species). The dominance of arboricolous, phytophagous species with a high degree of host plant specialization corroborates previous findings. The native region of 14 of these 18 species is the Mediterranean, resulting in a current “Mediterranization” of the Heteroptera fauna of Austria. Such a movement probably has happened before (1932–1940), but most of the species failed to establish in Central Europe. Most species arrive in the north-east of Austria, which may be a preferred invasion gateway for southeast-mediterranean species, due to the barrier effect of the Alps. Beside climatic factors, habitat change – particularly *
This paper is dedicated to Michail Josifov at his 80th birthday and his stimulating biogeographical and taxonomic contributions to the Heteroptera of Europe.
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availability of ornamental plants in urban settings – is considered an important driving force of recent additions to the Heteroptera inventory of Austria. Further monitoring of Heteroptera occurrence in Austria is needed to detect and verify this pattern. Keywords: Austria, climatic change, non-native species, Heteroptera
INTRODUCTION The accelerating pace of human impact on the environment in the last decades has caused a multitude of alterations, such as changes of element cycling and of climate as well as a dramatic decline of biodiversity (Sala et al. 2000; Chapin et al. 2001). However, at the local scale, biodiversity in terms of species numbers can rise due to man-made introductions of species beyond natural boundaries. These non-native (alien) species may harm native species or have economic cost; non-natives are considered second only to land use change as driving force for loss of global biodiversity (Vitousek et al. 1997; Wilcove et al. 1998) and are responsible for an economic loss equal to about 5 % of the world GNP (Pimentel 2002). Several driving forces are responsible for changes in the fauna of a given area. Beside anthropogenic forces (intentional release, unintentional introduction with plant material or as stowaways with goods or stored products), there are natural biotic and abiotic forces (the edges of species ranges are not stable and natural progression and regression of ranges do occur) and – of course – an interaction of both (e.g. Dukes & Mooney 1999; Gaston 2003). It is, therefore, difficult to uncouple the driving forces of range changes, but climatic change is clearly one of the most important (e.g. Parmesan 1996, 2006; Cannon 1998; Parmesan et al. 1999; Burton 2001; McCarty 2001; Hill et al. 2002; Walther et al. 2002, 2005; Root et al. 2003). In this paper, new additions to the Heteroptera fauna of Austria over the past approximate 25 years are summarized and discussed in the light of their possible driving forces of change. A descriptive, not statistically based, prediction for a future scenario is made.
METHODS The available literature on the Heteroptera of Austria (see Rabitsch 2006) was checked for new species records since and including the seminal paper of Melber et al. (1991), considering records back to 1982. The present study therefore deals with new additions to the species inventory for approximately the last 25 years. While collating data for a critical check-list of Austrian species (Rabitsch 2004a, 2005b) and a red list of endangered species of Lower Austria (Rabitsch 2007), voucher specimens were verified in several museum and private collections (Burgenländisches Landesmuseum, Eisenstadt; Kärntner Landesmuseum, Klagenfurt; Muséum d`Histoire Naturelle, Geneva; Naturhistorisches Museum, Wien;
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Niederösterreichisches Landesmuseum, St. Pölten; Oberösterreichisches Landesmuseum / Biologiezentrum, Linz; Zoologisches Museum, Universität Hamburg; Zoologische Staatssammlung, München). Altogether, approximately 40,000 data entries were analysed. Species were assigned to different categories as to the reason for their addition to the national list (new arrival or overlooked) as well as the likely driving force for arrival where appropriate: (i) species overlooked due to taxonomic changes, (ii) species overlooked due to cryptic biology (including rare species), (iii) new arrivals, probably due to climatic change, (iv) new arrivals, probably due to unintentional introduction (either with ornamental plant trade or with transport of goods). Further, the status of the species in Austria was evaluated from available data, and biological features of the species were compiled from the literature.
RESULTS During the past approximate 25 years 84 Heteroptera species were published for the first time (or confirmed as being present) for Austria (Tab. 1). This is 9.4 % of the 896 Heteroptera species currently known to occur in Austria (Rabitsch 2004a, 2005b; Rabitsch et al. 2007; Rabitsch & Heiss 2008) and means that species were added at a rate of more than 3 per year, including those previously overlooked. Thirty-seven species were recorded based on historic material (collected before 1980) and 12 of these were not found again. Fifty-seven species are considered to have been overlooked and 27 species are considered as new arrivals. Within the overlooked species, 16 species were overlooked probably due to taxonomic changes, 25 species were overlooked probably due to their cryptic habit or biology and 16 species were overlooked for both reasons. Of the 27 new arrivals, the establishment of 9 species is doubtful or unknown because they are known only from a single record. This leads to 18 established new arrivals; a rate of 0.72 species added per year. The following comparisons are made between the complete data set (n=84) and the new arrival data set excluding doubtful records (n=18). Climatic conditions are considered to be the main driving force (14 species) for the 18 new arrivals, followed by introduction with ornamental plants or transport (4 species). Almost all of these species (15 of 18) are considered to be expanding their range and currently are increasing in abundance within in Austria and beyond. The native region of 14 of these 18 species is the Mediterranean (7 Ponto-Mediterranean, 4 Holo-Mediterranean, 2 Atlanto-Mediterranean, 1 North-Mediterranean), two species are of Eastern Palaearctic origin, one species is of Eurosiberian origin and one species (Leptoglossus occidentalis Heidemann, 1910) is of Nearctic origin. Most species of the complete data set belong to Miridae (30 species), followed by Lygaeidae (15 species) and Anthocoridae (8 spp.). The same families dominate within the newly arrived species: Miridae (5 species), Anthocoridae and Lygaeidae (3 species each). Most of the species occur in the Pannonic region of Austria (31 % in Lower Austria, 24 % in Burgenland, 15 % in Vienna). Considering stratum preference, arboricolous spe-
312 W. Rabitsch
cies, living on trees and shrubs, dominate (45 %), followed by herbicolous and epigaeic species (30 % and 21 %, respectively) in the new arrival species and with similar proportions in the overlooked species. As expected, the proportion of epigaeic species with a more concealed way of life is rather high within the overlooked-cryptic species (46 %). The average body size of the Heteroptera species in Table 1 is 5.3 mm. Almost all of the species develop a single generation per year. Whereas no preponderance of a particular mode of hibernation was found in the complete data set, hibernation as adults clearly prevails among the new arrival species (70 % vs 20 % as eggs and 10 % as larvae). Phytophagous species dominate both data sets (46–52 %), followed by zoophagous species (25–27 %) and zoophytophagous species (18–21 %). There is a high degree of food plant or prey specialization in both data sets. Monophagous, oligophagous and polyphagous species (30–40 %) were almost equal. Almost all new arrivals are macropterous and capable of flying. Notable exceptions are the regularly apterous (rarely winged) Metapterus caspicus (Reduviidae) and the sexually dimorphic Mimocoris rugicollis (Miridae), the latter with winged males only. Habitat selection of new arrivals revealed a preference in decreasing order of open-land, forest-land and synanthropic habitats with not a single aquatic or semiaquatic species.
DISCUSSION In Europe, during the last decades, an increasing number of insect species have shifted their range northwards from the Mediterranean (Reemer et al. 2003; Hickling et al. 2005; Parmesan 2006), including Heteroptera (e.g. Netherlands, Aukema 2003; UK, Judd & Hodkinson 1998; Kirby et al. 2001; Hickling et al. 2006; see review by Musolin & Fujisaki 2006; Musolin 2007). However, as recently argued by Gaston (2003) climate (or weather) is not the sole ultimate factor governing or driving range expansions. In particular, habitat change is of special relevance for the successful establishment of many species in the new area. It is also clear that there are additional forces responsible, such as unintentional human-induced transportation by vehicles and goods, the intentional release for biological control and natural dispersal. This is confirmed by the present results, which suggest climatic conditions are the main driving force for new arrivals of Heteroptera to Austria, including some doubtful cases, and supplemented by unintentional introductions with ornamental plants and goods. Most of the new arrivals are currently increasing their range, so it is expected that their abundance and distribution will continue to increase in the future in Austria and beyond. The Mediterranean region is the major source of new arrivals of Heteroptera in Austria for the last decades with Ponto-Mediterranean species dominating. The intracontinental species exchange within Europe clearly exceeded the transcontinental introductions of non-native Heteroptera to Europe (Rabitsch, in prep.). The arrival of additional species in Austria is considered a matter of time. Among these are the Nearctic Corythucha arcuata (Say, 1832) (Tingidae), the Eastern-Asian Stephanitis takeyai Drake
Berytidae
Lygaeidae
Miridae Brachynotocoris puncticornis Reuter
Arocatus longiceps Stål Berytinus consimilis (Horváth)
Family
new arrival new arrival
overlooked new arrival
new arrival new arrival new arrival overlooked
Invasion process
2002 North-Medi- new terranean arrival
1995 Ponto-Mediterranean 2003 Ponto-Mediterranean
Year Native of first region known record in Austria Acetropis longirostris Miridae 1983 Ponto-MediPuton terranean Agramma atricapillum Tingidae 1982 Holo-Medi(Spinola) terranean Amphiareus obscuriceps Anthocoridae 1998 East-Palae(Poppius) arctic Anthocoris sarothamni Anthocoridae 1921 AtlantoDouglas & Scott Mediterranean Aradus krueperi Aradidae 1974 North-MediReuter terranean Aradus kuthyi Aradidae 1991 Ponto-MediHorvath terranean
Species
local
climatic?
single record, establishment probable
introduced- expansive ornamental climatic? local single record, establishment probable climatic expansive? single record, establishment probable
expansive?
cryptic
introduced- extinct ornamental
climatic?
expansive endangered, halophilous expansive
climatic
Comment
expansive
Status in Austria
climatic
Driving force
Rabitsch 2003b
Adlbauer & Friess 1996 Rabitsch 2003c
Melber et al. 1991, Rabitsch unpubl. Adlbauer 1992
Rabitsch 2004a
Friess 2000
Melber et al. 1991
Melber et al. 1991
Reference
4.5 W
6.0 B, W, N, O, St, K, T 6.6 N
5.5 St
8.8 B, W, N
3.4 N
2.7 B, W, N, K
3.0 B, N
7.5 B, N
Mean Known body distribution size in Austrian provinces
i
az i, l m o m w
az i, l m o m w
z o m sy
z p m w
p o m o
ar
un e zp m m w
ar un- i p m m sy bi epi un i zp m m o
ar
ar
epi, un? i he ar un i
he un
p m m o
V H T F W P
he un e
S
Table 1: Heteroptera species reported as occurring in Austria for the first time since (and including) Melber et al. (1991). If two references are given, the first mentioned the species for the first time for Austria and the second reports an earlier finding. Mean body sizes are given in mm. Abbreviations for Austrian provinces: B – Burgenland, W – Vienna, N – Lower Austria, O – Upper Austria, S – Salzburg, St – Styria, K – Carinthia, T – Tyrol, V – Vorarlberg. Further abbreviations: S = Stratum: ar – arboricolous, end – endogaeic, epi – epigaeic, he – herbicolous, para – ectoparasitic, ep – epineustic, aq – aquatic; V = Voltinism: un – univoltine, bi – bivoltine, un-bi – univoltine or bivoltine, az – asynchronous; H = Hibernation: e – egg, l – larvae, i – imago; T = Trophism: p – phytophagous (including seed-feeding), m – mycetophagous, z – zoophagous, zp – zoophytophagous, h – haematophagous; F = food preferences: o – oligophagous, p – polyphagous; W = wing development: m – macropterous, b – brachypterous, a – apterous; P = habitat preference: o – open-land (wet and dry meadows, rural sites), w – wood-land (forests, forest edges, woodland), aq – aquatic and semiaquatic, sy – synanthropic, c - caves.
The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria
313
Criocoris nigricornis Reuter Deraeocoris flavilinea (A. Costa) Dichrooscytus gustavi Josifov Dicranocephalus setulosus (Ferrari) Dybowskyia reticulata (Dallas)
Brachysteles parvicornis (A. Costa) Campylomma annulicorne (Signoret) Campylosteira bosnica Horváth Capsus pilifer (Remane) Capsus wagneri (Remane) Cardiastethus fasciiventris (Garbiglietti) Chorosoma gracile Josifov Cimex dissimilis (Horváth) Compsidolon pumilum ( Jakovlev) Conostethus roseus (Fallén)
Species
Stenocephalidae Pentatomidae
Miridae
Miridae
Miridae
Miridae
Miridae
Cimicidae
Rhopalidae
Anthocoridae
Miridae
Miridae
Tingidae
overlooked 2002 North-Medi- new terranean arrival 2003 European overlooked 1982 Holo-Medi- new terranean arrival 1971 East-Palae- new arctic arrival?
1950 European climatic? introduced? introducedornamental introducedtransport cryptic
cryptic
cryptic
taxonomy, cryptic taxonomy
cryptic
climatic?
taxonomy
taxonomy, cryptic taxonomy
Friess et al. 2005
Melber et al. 1991, Rabitsch 1999b Rabitsch 2001b
Rabitsch 1999b
Melber et al. 1991, Ressl 1995
Niederer 2003
Reference
critically endangered, psammophilous
13.0 S 4.0 N
Rabitsch 2000
4.0 W, N, K, V
6.7 W, N
3.5 B, N
3.0 N
3.3 B
5.0 N
11.2 B
2.5 St
5.8 N, O
5.4 B, N
1.8 W
2.8 B, N, V
2.0 V
Mean Known body distribution size in Austrian provinces
Moulet 1995
Rabitsch 2004b
Rabitsch 2002b
Rabitsch 1999b
Melber et al. 1991, Rabitsch unpubl. Rabitsch 1999b
single record, Moulet 1995 status unknown? Rabitsch 1999b
Comment
unknown single record, not established local single record, establishment probable
expansive
expansive
local
local
local
local
extinct?
expansive
local
local
extinct
local
taxonomy, cryptic
overlooked
Status in Austria
local
Driving force
overcryptic looked?
Invasion process
1956 Ponto-Medi- overterranean looked 1983 Eurosiberian overlooked 1934 Eurosiberian overlooked 2000 Atlantonew Mediterra- arrival nean 1960 Eurosiberian overlooked 1997 Eurosiberian overlooked 1965 Holarctic overlooked 1999 Eurosiberian overlooked?
Year Native of first region known record in Austria Anthocoridae 2000 AtlantoMediterranean Miridae 1971 Eurosiberian
Family
un
i
z o m w
p m m o
un e
he un
i
bi e (p) he un i
ar
ar
he un e
he un e
p o m o
p o m o
p o m sy
z p m sy
p o m o
p o m o
he un e zp m m o
he un- e p o m o bi para az i, l h o a c
ar
he un e
p p b, o (m) p m m o
un e zp m m w
epi, un i end he un e
ar
z o m, w b
V H T F W P
epi, un? i he
S
314 W. Rabitsch
Family
Ischnocoris punctulatus Fieber Kalama aethiops (Horváth) Kleidocerys privignus (Horváth) Lasiacantha hermani Vásárhelyi
Geocoris erythrocephalus (Lepeletier & Serville) Geotomus brunnipennis Wagner Halticus major Wagner Haploprocta sulcicornis (Fabricius) Heterotoma planicornis (Pallas) Himacerus boops (Schiodte) Himacerus major A. Costa Hypseloecus visci (Puton) Icus angularis Fieber
1970 Holo-Mediterranean 1988 Ponto-Mediterranean 1932 North-Mediterranean 1961 Holo-Mediterranean 2000 Holo-Mediterranean 1988 European?
Nabidae
Tingidae
Lygaeidae
Tingidae
Lygaeidae
Lygaeidae
Miridae
1950 Eurosiberian
1982 Eurosiberian
Nabidae
Miridae
Coreidae
Miridae
1935 North-Mediterranean 1944 South-European 1940 Holo-Mediterranean 1941 European?
Cydnidae
overlooked overlooked overlooked overlooked overlooked overlooked? overlooked overlooked overlooked new arrival overlooked overlooked
1954 Holo-Medi- new terranean arrival
1976 Ponto-Medi- overterranean looked
taxonomy, cryptic
taxonomy
climatic
cryptic
cryptic
cryptic
cryptic
cryptic
taxonomy
cryptic
taxonomy, cryptic taxonomy
climatic
cryptic
Year Native InvaDriving of first region sion force known process record in Austria 1998 Eurosiberian overcryptic looked?
Lygaeidae
Fulvius oxycarenoides Miridae (Reuter)
Emblethis brachynotus Lygaeidae Horváth
Species Comment
Reference
local
local
local
local
extinct
local
local
local
common
extinct
local
extinct
Melber et al. 1991, Rabitsch unpubl.
Melber et al. 1991
Friess et al. 2005
Péricart 1999
Rabitsch 1999b
Melber et al. 1991, Rabitsch unpubl. critically endan- Melber et al. 1991 gered vulnerable Heiss et al. 1991, Rabitsch 1999b Ressl 1995
Heiss et al. 1991, Rabitsch unpubl. Moulet 1995
Rabitsch 2001a
critically Rabitsch 1999b endangered, psammophilous local single record, Melber et al. 1991 establishment probable expansive Adlbauer 1995, Rabitsch unpubl.
local
Status in Austria
2.9 B, N
5.2 B, W, N
2.6 St
2.7 N, T
3.9 N
3.3 N
8.3 B, W, N
6.7 B, W, N
5.0 W, N, O
11.2 N
2.7 B, N
4.5 B
3.8 B, N, St
3.5 B
4.9 N
Mean Known body distribution size in Austrian provinces
i
p o m o
i
i
b
o
p p m o
p m m w
z p m o
z p
z p m w
p m m o
p m m o
p m m o
z p m o
he un- i bi
p o b, o (m)
epi un- i p p m o bi epi, un- i? p p m, o he bi? (b) ar un i p m m w
epi un
epi, un e he ar un e
he, un e ar epi un e
he un
end, un i epi he un e
i
un i? z p m w
bi
V H T F W P
epi, un he
ar
epi
S
The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria
315
Micronecta scholtzi (Fieber) Micracanthia fennica (Reuter) Microvelia buenoi Drake Microvelia pygmaea (Dufour) Mimocoris rugicollis (A. Costa) Nagusta goedelii (Kolenati) Neolygus zebei (Günther) Notonecta meridionalis Poisson Nysius graminicola (Kolenati) Oncotylus viridiflavus (Goeze) Orius agilis (Flor)
Metacanthus annulosus (Fieber) Metapterus caspicus (Dohrn)
2003 Ponto-Mediterranean 1988 Ponto-Mediterranean
Berytidae
1982 North-Mediterranean 1971 Boreo-montane 1993 Holarctic
Anthocoridae
1985 East-Palaearctic
1920 North-Mediterranean Miridae 1987 Ponto-Mediterranean Reduviidae <1999 Ponto-Mediterranean Miridae 2000 Ponto-Mediterranean? Notonectidae <1975 Ponto-Mediterranean Lygaeidae 1983 Trans-Palaearctic Miridae <1930 Eurosiberian
Veliidae
Veliidae
Saldidae
Corixidae
Reduviidae
Invasion process
overlooked overlooked overlooked overlooked new arrival new arrival overlooked overlooked overlooked? overlooked new arrival?
new arrival new arrival
new arrival overlooked 2001 Ponto-Medi- new terranean arrival
Year Native of first region known record in Austria 2005 North America 1896 Eurosiberian
Family
Leptoglossus occidentalis Coreidae Heidemann Lygaeus simulans Lygaeidae Deckert Megalocoleus naso Miridae Reuter
Species Status in Austria
expansive common local local local
climatic taxonomy taxonomy cryptic cryptic
local
local
climatic
climatic?
extinct
common
local
common
expansive
expansive
expansive
taxonomy
taxonomy
cryptic
taxonomy
climatic
climatic
climatic
introduced- expansive ornamental taxonomy common
Driving force
only males, established?
Comment
Heiss 1996, Rabitsch 1999a Melber et al. 1991
Rabitsch & Zettel 2000 Melber et al. 1991
Rabitsch 2001b
Rabitsch & Heiss 2008 Rabitsch 2001a
Günther & Strauss 2006 Rabitsch & Zettel 2000 Rabitsch 1999a
Melber et al. 1991
Melber et al. 1991
Rabitsch & Heiss 2005 Melber et al. 1991, Rabitsch unpubl. Rabitsch 2001b, Rabitsch & Matocq 2005 Rabitsch 2004b
Reference S
1.8 B, N
7.1 N, V
4.3 B
15.2 N
5.8 B, W, N
14.3 St
4.0 K
1.9 V
1.9 B, W, N
2.3 St
2.3 B, W, N, V
10.1 B, N
5.0 B, W
11.7 B, W, N, St, K, T 3.4 B, N
un
un
i
un e un
p
? m w
z p m aq
p m m w
z p a, o b, (m) l o p b, aq m e, z p m o i? i z p b, aq (m) i z p b, aq (m) e zp p b, w (m) i z p m w
i
i
p o m o
p o m o
p o m sy
he un
i
z p b, o m
epi un- i? p p m o bi? he un e p m m o
aq
ar
ep unbi ep unbi ar, un epi ar un
aq unbi epi un
epi un
ar
he un e
i
i
V H T F W P
he un
18.0 W, S, St, K, T ar
Mean Known body distribution size in Austrian provinces
316 W. Rabitsch
Psallus anaemicus Seidenstücker Psallus assimilis Stichel Psallus confusus Rieger Psallus cruentatus (Mulsant & Rey) Psallus lapponicus Reuter Psallus lentigo Seidenstücker
Miridae
Miridae
Miridae
Miridae
Miridae
Miridae
< 1900 Ponto-Medi- overterranean looked 1946 European? overlooked 1923 Eurosiberian overlooked 1983 South-Euro- new pean? arrival? 1957 Boreo-mon- overtane looked 2003 Ponto-Medi- new terranean? arrival?
overlooked overlooked overlooked overlooked overlooked
Peritrechus Lygaeidae <1950 Holo-Medimeridionalis Puton terranean Phoenicocoris dissimilis Miridae 1933 Ponto-Medi(Reuter) terranean? Picromerus conformis Pentatomidae 1960 Ponto-Medi(Herrich-Schäffer) terranean Pinalitus coccineus Miridae 1984 Ponto-Medi(Horváth) terranean Pionosomus opacellus Lygaeidae 1933 Eurosiberian Horváth
Oxycarenus lavaterae Lygaeidae (Fabricius)
Lygaeidae
overlooked new arrival new arrival
Orsillus depressus (Mulsant & Rey) Orsillus reyi Puton
Lygaeidae
climatic
cryptic
climatic
taxonomy
taxonomy
local
extinct?
local
local
common
common
local
cryptic
taxonomy
3.9 B
local
cryptic
single record, Rabitsch 2004a status unknown single record, Rabitsch 2003c establishment probable
Melber et al. 1991, Rabitsch 2003c Melber et al. 1991, Rabitsch 2005a Melber et al. 1991
critically Melber et al. 1991, endangered, Rabitsch unpubl. psammophilous Rabitsch 2003b
Melber et al. 1991
ar
4.4 N
4.2 T
3.4 B, W
3.4 B, O
3.6 B, W, N
3.5 W, N
2.7 B, N
ar
ar
ar
ar
ar
ar
p p b, o m
un e zp m m w
un e zp o m w
un e zp m m w
un e zp m m w
un e zp m m w
un e zp m m w
i
un e? p m m w epi, un end
ar
p o m w
p p m o
un e? z o m w
un e
i
un i, p o m sy (l) un i, p o m sy (l) az i, p o m w (l)
12.5 N
ar
ar
ar
i zp o m o
V H T F W P
he un
S
ar
5.4 B, W, N, St
7.4 B, W, N, O, St, K, T 7.3 St
2.0 W
Mean Known body distribution size in Austrian provinces
3.4 V
Rabitsch 2001b
Reference
epi un
single record, establishment probable
Comment
4.7 B, N
local
Status in Austria
introduced- expansive Melber et al. 1991, ornamental Rabitsch unpubl. introduced- local single record, Adlbauer 1999 transport not established climatic? expansive Rabitsch & introduced? Adlbauer 2001, Rabitsch 1999b cryptic local Melber et al. 1991, Rabitsch 2001b cryptic extinct single record, Rabitsch 1999a status unknown cryptic local Rabitsch 2001b
Year Native InvaDriving of first region sion force known process record in Austria Anthocoridae 2000 Holo-Medi- new climatic terranean arrival
Family
1948 Holo-Mediterranean 1998 Holo-Mediterranean 1966 Holo-Mediterranean
Orius pallidicornis (Reuter)
Species
The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria
317
Family
Tritomegas rotundipennis (Dohrn) Tuponia elegans ( Jakovlev) Velia currens (Fabricius) Xylocoris obliquus A. Costa
Psallus wagneri Ossiannilsson Rhynocoris rubricus (Germar) Spathocera dalmanii (Schilling) Stygnocoris cimbricus (Gredler) Stygnocoris similis Wagner Temnostethus dacicus (Puton) Thyreocoris fulvipennis (Dallas)
1920 Eurosiberian?
1984 North-Mediterranean 1988 Ponto-Mediterranean 2001 AtlantoMediterranean 1940 North-Medi- overterranean looked
Lygaeidae
Lygaeidae
2002 Ponto-Mediterranean Veliidae <1925 South-European Anthocoridae 2006 Holo-Mediterranean
Miridae
Cydnidae
Cydnidae
Anthocoridae
new arrival overlooked overlooked?
<1900 Ponto-Mediterranean 1937 Eurosiberian
overlooked? overlooked overlooked overlooked overlooked overlooked? new arrival
Reduviidae
Coreidae
Driving force
cryptic
local
introduced- expansive ornamental taxonomy local
extinct
expansive
climatic
taxonomy
local
local
common
extinct
extinct
regional
local
single record, not established
single record, establishment probable
6.6 K
Friess 2001a, Rabitsch 2003a Rabitsch et al. 2007
2.1 B
3.2 W, N
6.5 K
3.7 N
3.0 B
3.1 B
2.3 N, O, K, T
6.0 B, N
15.6 N
3.5 B
5.1 W, N, O, St, K, T, V 3.8 N
4.6 B, N
Mean Known body distribution size in Austrian provinces
Rabitsch 2002a
Rabitsch 2001a
Rabitsch 2001b
Melber et al. 1991
Melber et al. 1991, Rabitsch 2001b Heiss 1997b, Rabitsch 2003a Melber et al. 1991
Rabitsch 2004a
Melber et al. 1991
Rieger & Rabitsch 2006 Rabitsch 2003c
common
Reference
Rabitsch 2003c
Comment
common
Status in Austria
cryptic
cryptic
taxonomy
cryptic
taxonomy
taxonomy
overtaxonomy looked overtaxonomy looked overtaxonomy looked?
Invasion process
1983 European?
Year Native of first region known record in Austria 1952 North-Mediterranean 1919 West-Palaearctic 2003 European?
Miridae
Psallus lucanicus Miridae Wagner Psallus montanus Miridae Josifov Psallus pseudoplatani Miridae Reichling
Species
p m m o
z p m o
un
un e
i
i
i
p m m sy
p m m w
z p b, w m p m m o
ep un- i, z p a, aq bi (l) (m) epi un i z p m, o b
ar
he un
epi, un end
ar
epi un i, p o m o (e) epi un? i? p p m o
i
l
un e zp m m w
un e zp m m w
un e zp o m w
un e zp m m w
V H T F W P
he, un epi epi un
ar
ar
ar
ar
S
318 W. Rabitsch
The Times They Are A-Changin’: Driving forces of recent additions to the Heteroptera fauna of Austria
319
& Maa, 1955 (Tingidae) and some Mediterranean species, e.g. Tuponia spp. (Miridae) and Nezara viridula (Linnaeus, 1758) (Pentatomidae). It is not possible to predict the “next” species to arrive in Austria, but it is very likely that it will be a mirid or tingid from the Mediterranean region. In other words, central Europe is becoming more Mediterranean-like in temperature and precipitation and in species diversity. Regional scenarios predict an increase in winter and summer temperatures of 3°C and of 1.5-2.5°C in spring and autumn for Austria until 2048 (Matulla et al. 2004). A northward shift of current species distributions is to be expected. The idea of a Mediterranization of Austria’s heteroptera fauna is not new. Hölzel (1954) wrote: “We are witnessing an immigration of [Heteroptera] species from the south and see a northwards migration in formerly owned areas” (my translation). Unfortunately, most of his new records did not withstand critical taxonomic verification (Rabitsch 2003a). Mediterranean species have shifted their range northwards in the past as well. Current data indicate a nested occurrence of these species between 1932–1940, which were the climatically most favourable years of the 20th century before the exceptional increase around 1990 (Auer et al. 2001). Coincidentally, identical and additional Mediterranean species were also reported at this time from Slovakia and the Czech Republic (e.g. Icus angularis Fieber, 1861 (Lygaeidae), Monosteira unicostata (Mulsant & Rey, 1852) (Tingidae)) (Stehlík & Vavrinová 1999, Stehlík 2002), but not thereafter. Whereas this historic shift may be considered merely temperature-driven, the recent shift is promoted by a combination of temperature increases and changes in habitat characteristics. The increasing cultivation of native and non-native plants for horticultural and ornamental reasons, the utilization of anthropogenically changed habitats, the distribution patterns and dispersal routes are superimposed by climatic change. In conclusion, several driving forces work in concert, so that ultimate explanations of recent range expansions will remain controversial. But habitat change is at least as important as current temperature increase for Heteroptera. This trend continues, and continuous monitoring is necessary to detect these changes. Many of the Mediterranean species given in Tab. 1 live on cultivated ornamental food plants. It is therefore difficult to elucidate the mechanism which triggered their expansion and to separate climatic change from habitat change or if both driving forces act together in concert. New arrivals were also found to be associated with introduced ornamental plants in the U.K. by Kirby et al. (2001). However, as Kirby et al. (2001) noted, it is a surprisingly low number of species compared with the rather high number of introduced ornamental plants. Most species also show some degree of food plant or prey specialization. A similar pattern was found for the non-native Heteroptera species introduced to or within Europe (Rabitsch, in prep.). Being polyphagous and nonselective seems not to be of particular advantage for Heteroptera. Most species (70 %) occur in the dry, Pannonic, north-east of Austria, which may indicate an invasion gateway for Ponto-Mediterranean species. The Alps are an effective barrier for Mediterranean species, which have to move west- or eastwards and only rarely penetrate into the Alps. Projecting the estimate of 0.76 new arrival species per
320 W. Rabitsch
year to a decade (7.6) exceeds the estimated rate of new arrivals per decade for Europe (Rabitsch, in prep.). This is likely to be due to the geographical location of the country, and changes of the Heteroptera fauna of Austria may be indicative for tomorrow’s Heteroptera fauna of Europe. The average body size of the Heteroptera species (5.3 mm) is similar to the average of the Central European Heteroptera species (5.6 mm, Rabitsch et al., in prep.). This contradicts the assumption that small species are more easily overlooked and indicates that body size is not of particular relevance for predicting additions of overlooked and new arrivals in a given territory. Body size also was not significantly different between newly arrived and overlooked species (5.8 vs 5.0 mm, respectively; p>0.05, t-test), although a cryptic way of life (epigaeic species) prevails among the overlooked species. Hibernation in the adult stage prevails in the new arrival species, which may be an effect of the greater robustness and defence capabilities of adults, compared to nymphs and eggs. To successfully conquer new territories, it is obviously advantageous to be able to fly, which is confirmed by the macroptery of almost all new arrivals. It is puzzling, however, to imagine how a regularly apterous species such as Metapterus caspicus or a sexually wing-dimorphic species such as Mimocoris rugicollis expands its range in a relatively short time. Beside possible anthropogenic translocation, passive wind dispersal of flight-unable individuals and larvae has to be taken into account (Rietschel 2007). New arrivals do not seem to prefer a particular type of habitat since they can be found in a wide variety of biotopes, but a slight preference for synanthropic sites in the most recent arrivals (since 2000) was found (Deraeocoris flavilinea, Dichrooscytus gustavi, Leptoglossus occidentalis, Tuponia elegans), indicating the increasing relevance of introductions with the transport of goods. Many species have been overlooked in the past and were discovered only recently. This is understandable in case of taxonomic changes, but unfortunate for rare species in cryptic habitats because it results in uneven data quality in time and space for these insects. Museum specimens not only serve as taxonomic vouchers but also document past faunas. Heteroptera in general is a thermophilous taxon (Honek & Kocourek 1990); abundance and diversity decrease with increasing altitude and latitude. Therefore the Heteroptera might profit from rising temperatures more than other insect groups. Considering that correlations between temperature and range changes often will remain tentative, phenological and elevation shifts of resident species seem to be much more conclusive indicators of climatic change than multi-factor driven interpretations of range changes (e.g. Ellis et al. 1997; Konvicka et al. 2003; Roy & Sparks 2000; Stefanescu et al. 2003). Data on changes in elevation range of Austrian Heteroptera are not readily available, but phenological shifts were already observed and will be presented elsewhere (Rabitsch, in prep.). In conclusion, Heteroptera are not only suitable indicators of diversity (e.g. Duelli & Obrist 1998; Virolainen et al. 2000), but also are valuable indicators of global change and deserve further study.
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ACKNOWLEDGEMENTS I thank Nikolay Simov for the invitation to contribute to this Festschrift. Berend Aukema and Petr Kment are thanked for helpful comments on the manuscript, David Roy and Al Wheeler are especially thanked for gentle language editing. Parts of the work were written within the EU-FP6-STREP DAISIE (http://www.europe-aliens.org).
РЕЗЮМЕ През последните приблизително 25 години 84 вида Heteroptera бяха публикувани за първи път в Австрия. Тези видове представляват 9.4 % от 896 вида Heteroptera, известни в днешно време за Австрия, и това означава, че са били добавяни със скорост нe по-малко от 3 на година, включително пропуснати преди това пропуснати видове. Две трети от видовете (68%) са били пропуснати поради таксономични причини или заради техния прикрит начин на живот или биологични особености, останалите видове (32%) се разглеждат като „пришълци”. С изключение на един единствен вид, 18 вида (21%) са доказани „пришълци”, установявани със скорост от 0.72 вида за година. Повечето от тези видове общо разширяват ареала си и увеличават обилието си в Австрия и другаде в Европа. По-голямата част от тези видове достигат Австрия вероятно благодарение на благоприятни климатични условия (14 вида), следвани от видовете, пренесени чрез декоративни растения или транспорта (4 вида). Превъзходството на дендробионтните фитофагите, с висока степен на специализация към хранителното растение е в съгласие с предишни открития. Родина за 14 от 18-те видове е Средиземноморието, което води до „медитеранизация” на хетероптерната фауна в Австрия. Такова движение вероятно е имало и преди (1932-1940), но много от видовете не са успели да се приспособят в Централна Европа. Много видове се появяват в Североизточна Австрия, което може би е предпочитан път за проникване на югоизточни средиземноморски видове, избягвайки по този начин бариерния ефект на Алпите. В сравнение с климатичните фактори, промяната на хабитата – особено поява на декоративни растения в урбанизираните райони – се разглежда като важен движещ фактор за съвременни допълнения към хетероптерната фауна на Австрия. За да бъде установен и проверен този модел е необходимо провеждане на мониторинг на Heteroptera в Австрия.
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S. Grozeva & N. 2008 First record of Pinochius Carayon, 1949 from theSimov Oriental(Eds) Region, with description of a new species... 327 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 327-337. © Pensoft Publishers Sofia–Moscow
First record of Pinochius CARAYON, 1949 from the Oriental Region, with description of a new species from Vietnam (Heteroptera: Schizopteridae) D. Rédei Department of Zoology, Hungarian Natural History Museum, H-1088 Budapest, Baross u. 13, Hungary. E-mail:
[email protected]
ABSTRACT The family Schizopteridae (Heteroptera) is reported for the first time from Vietnam. Pinochius Carayon, 1949, known from Africa and Australia so far, is reported first from the Oriental Region by the description of Pinochius josifovi nov. sp. from North Vietnam. The new species is unique within its congeners described so far by the presence of a posteriorly directed right-side process on tergite VIII. Keywords: Heteroptera, Dipsocoromorpha, Schizopteridae, Pinochius, new species, Vietnam, morphology, fore wing venation.
INTRODUCTION Carayon (1949) described a new genus and species of Schizopteridae (regarded as a subfamily within Cryptostemmatidae sensu lato, a family identical with the current Dipsocoromorpha, at that time), Pinochius africanus, from Cameroon. Simultaneously, Wygodzinsky (1950a) established the new genus Seabranannus for the accommodation of four new species, anterii, dundundo, imitator, and similis, from Angola; later in that year he synonymized Seabranannus with Pinochius (Wygodzinsky 1950b). One further species, Pinochius nitidicollis was later described from Egypt by Linnavuori (1966). Quite unexpectedly, Hill (1985) discovered the presence of Pinochius in Australia and described a new species, P. australiensis from Queensland. Hill noted that some
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differences of the Austrialian species seem to be of generic importance, and therefore probably a new genus should be erected for its accommodation, however, this question is still open. Pinochius is treated here as a single genus, currently comprising six African and one Australian species. Little is known about the biology of the species. Most specimens of the previously described species were collected in woodland habitats, frequently in riverine forests, in most cases in humid places. Specimens were usually obtained from vegetal debris at the ground, rarely captured by sweep-netting. The single known species of P. nitidicollis was accidentally collected in a hotel room in Giza, Egypt (Štys 1990); this might be regarded as indirect information that at least some species of Pinochius are attracted to light. The dipsocoromorphan fauna of Vietnam is poorly investigated. Only members of two families, Ceratocombidae (4 species of Kvamula Štys, 1982) and Hypsipterygidae (1 species of Hypsipteryx Drake, 1961) have been reported from Vietnam so far (Štys 1982, Rédei 2007). Several new records and new species are expected from the country. Among unidentified material deposited at the Hungarian Natural History Museum, Budapest, specimens of Pinochius from North Vietnam have been found. Their examination concluded that they represent an undescribed species which is described as new in this paper.
MATERIAL AND METHODS Structures were examined from uncleared glycerine-alcohol mounts by using an XSZN107 optical microscope supplied with a camera lucida. Measurements were taken using a micrometer eyepiece. The abdomen was examined after a very short and careful clearing in 10% KOH solution.
TAXONOMY Genus Pinochius Carayon, 1949 Pinochius Carayon, 1949: 239. Type species by original designation: Pinochius africanus Carayon, 1949. Seabranannus Wygodzinsky, 1950a: 34 (synonymized by Wygodzinsky 1950b: 391). Type species by original designation: Seabranannus anterii Wygodzinsky, 1950. Wygodzinsky 1950a (morphology, larval morphology, key to African species); Emsley 1969 (morphology); Štys 1970 (morphology); Hill 1985 (redescription, morphology, diagnostic characters).
Diagnosis. Easily recognized among Schizopteridae: Schizopterinae, also among all other Dipsocoromorpha, by its unique wing venation, formed by a strongly elevated, carina-like longitudinal vein and three oblique veins connecting this vein with those form-
First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species... 329
ing the leading edge of the remigium (for an overview of the fore wing venation of most genera of Schizopteridae, see Emsley 1969). A detailed redescription, discussion of the morphology and diagnostic characters of the genus were presented by Hill (1985). Revised diagnosis. In order to accommodate the new species described below in Pinochius, the definition of the genus as recognized by Wygodzinsky (1950a, as Seabranannus), Štys (1970) and Hill (1985) should be modified as follows: tergite VIII without processes or with a single right-side process.
Pinochius josifovi, nov. sp. (Figs 1–10, 13–20)
Type material. Holotype (♂): “VIETNAM prov. \ Nam há, Tuong \ linh, nr. Phu ly,” [printed]; “No. 565. \ 25. V. 1966. \ leg. Topál” [printed]”; deposited in the Hungarian Natural History Museum, Budapest. The specimen is mounted on card, the abdomen is preserved in intact condition in a plastic microvial with glycerol and is pinned with the specimen. Other specimens examined. “VIETNAM, Tanh liet \ SE of Hanoi \ 23. IV. 1966, No. 140 \ leg. Gy. Topál” [printed] (1♀, mounted on card); “VIETNAM, Lao Cai prov., \ Sa Pa distr. Sin Chai, \ “Legendary place”, \ 22°20’93”N 103°46’06”E” [printed]; “10. XI. 2003, leg. M. Földvári, \ L. Peregovits & Á. Kőrösi” [printed] (1♀, dissected, body parts are mounted on slide in Euparal 3C-239); deposited in the Hungarian Natural History Museum, Budapest. — Because none of the females was captured together with the male, it was decided to exclude these specimens from the type series because their conspecificity is not undoubtful. Diagnosis. Readily differs from all the other known congeners by the antler-line appendage of the proctiger composed of a curved body and two branches, and the eighth tergite bearing a right-hand process directed caudad. Description. Macropterous male. General facies as in Fig. 1, body wide ovoid, very small. Colour: rather uniformly dark brown dorsally; fore wings widely paler distally; extremities stramineous. Body surface and pilosity: Head and pronotum with dense, relatively long, adpressed pubescence. Antennal segments I and II with short, adpressed pubescence; segments III and IV with sparse, diagonal, fine pilosity, the hairs are about 20 times as long as diameter of segment III. Mesoscutellum with long hairs (as on pronotum) at its basal half and with very short, strong, spine-like hairs each originating from a distinct tubercle (Fig. 2). Legs with short, rather dense, semierect pilosity. Fore wings with extremely short pilosity, hardly visible in 400× magnification; clavus, remigium between the carina-like longitudinal vein and the vein running along claval furrow, and anterior marginal cells of remigium with several minute, wart-like tubercles of different size, usually forming small groups, each bearing a short hair (Fig. 2); tubercles neighbouring claval furrow on clavus and remigium larger and darker; veins with long hairs usually forming two rows on
330 D. Rédei
the carina-like central longitudinal vein and the three oblique veins. Abdominal tergites bare, sternites and pygophore with rather dense and long pilosity. Head (Figs 3, 4) strongly declivent, almost vertical; head across eyes about 1.9 times as wide as interocular distance; with indistinct carinate rim mediad to antennal insertion; bucculae well developed, surrounding labial segment I. Labium (Fig. 4) 3-segmented,
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Figs 1–7: Pinochius josifovi nov. sp., male. 1: habitus; 2: clavus and mesoscutellum of left fore wing; 3: head, pronotum and mesoscutellum, dorsal view; 4: head and prothorax, ventral view; 5–7: tarsi of fore (5), mid (6) and hind (7) legs. Measurements in mm. Lettering: acv = anterior claval vein; cf = claval furrow; pcv = posterior claval vein; scu = mesoscutellum; wc = wing coupling apparatus.
First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species... 331
slightly surpassing apex of prothoracic xyphus; segment I wide, segments II and III narrow, segment III distinctly longer than segment II. Antenna: segment I only slightly longer than its diameter; segment II pyriform, about 1.6 times as long as segment I; segments III and IV gracile, segment III distinctly thickened at its base; segment IV gradually thickening towards apex, abruptly narrowing apically; greatest diameter of segment IV (close to its apex) about 2 times greater than diameter of segment III. Thorax. Pronotum (Figs 3, 4) short, trapezoid in dorsal view; about 2.0 times as wide across humeral angles as total length along meson; about 1.3 times as wide as head; with a distinct collar anteriorly; lateral margins slightly concave. Mesoscutellum (Fig. 2) short, triangular; about 1.5 times as wide at its base as its length along meson; apex very shortly mucronate, globular, with several minute tubercles (Fig. 2). Propleura, pteropleura and pterosterna studied only in female; see below. Legs. Hind coxa with well developed adhesive pad apically (Fig. 9); fore, mid and hind (Fig. 9) trochanters distinctly subdivided into two pseudosegments; fore, mid and hind femora subequal in length, without modifications; hind tibia distinctly longer than fore and mid tibiae; fore (Fig. 5) and mid (Fig. 6) tarsi 3-segmented, relatively short, distinctly widened at joint of segments II and III; hind tarsus (Fig. 7) 2-segmented, with a very short segment I and a long, narrow segment II; fore, mid, and hind tarsi each with a pair of claws identical in size and shape, each bearing a distinct basal tooth; pulvilli capitate apically. Fore wing (Fig. 10) surpassing apex of abdomen; basal half sclerotized, not transparent, becoming more transparent towards apex; central longitudinal vein strongly elevated, carina-like; anterior marginal vein wide, sharply delimited; veins between central longitudinal vein and posterior margin of remigium (marked with dotted line in Fig. 8) delimiting the small central and posterior cells obsolete, can be traced mostly by their pilosity. Presumable homologies of venation (Figs 11, 12) are treated below under Discussion. Pregenital abdomen (Figs 13, 19, 20) with slight dextral asymmetry; no orifices of dorsal glands. Tergites III–VII (Figs 13, 20: t3–7) very weakly sclerotized, difficult to observe; tergites III and IV slightly, V–VII strongly shortened at right side. Sternites III–VII (Figs 13, 19, 20: s3–7) distinctly sclerotized; sternites III–VI nearly symmetrical; sternite VII strongly asymmetrical, deeply excavated posteriorly at the right side, its posterior margin widely covering base of ventral surface of pygophore. Sternites VI and VII each with a pair of distinct spiracles. Terminalia (Figs 13–20). Tergite VIII (Figs 13, 19, 20: t8) distinctly sclerotized; strongly asymmetrical; having a horizontal part at level of tergites III–VII and a long vertical part projecting to pygophore at the right side (Figs 19, 20: t8*); with a distinct, stout, apically narrowing process (Figs 13, 19, 20: pt8; Fig. 12) at its right side, projecting above base of appendage of the proctiger; with a distinct spiracle at a short projection at its left margin, and an indistinct spiracle, difficult to observe, at the overlap of the right side of the tergite and the pygophore. Sternite VIII could not be recognized, membraneous or maybe missing. Pygophore (= segment IX) (Figs 13, 19, 20: py) with dextral asymmetry, laterotergites absent. Proctiger with a large, antler-shaped (Figs 13, 19, 20: apg1; Fig. 15) and a short, apically widened (Figs 13, 19: apg2) appendage. Large appendage
332 D. Rédei
consisting of a long, curved body directed anteromediad and two branches projecting mediad, apically each of them with minute peg-like projections; body strongly widened in the horizontal plane at its middle; the widened part bearing a small, flattened, triangular process ventromedially. Left paramere (as incompletely seen in situ, Figs 13, 18: lp) large, approximately triangular. Right paramere (as incompletely seen in situ, Figs 13, 18–20: rp, Figs 16, 17) elongate, apex projecting above left paramere; apical part narrow, curved,
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Fig. 8: Pinochius josifovi nov. sp., female, pterothorax, ventral aspect. Figs 9–10: Pinochius josifovi nov. sp., male. 9: right hind coxa and trochanter, anterior surface; 10: fore wing (as seen in situ, not flattened). Measurements in mm. Lettering: ad = adhesive pad; cx = coxa; tr1 and tr2: proximal and distal pseudosegments of trochanter. Figs 11–12: Two alternative hypotheses of the fore wing venation homologies of macropterous Pinochius. 11: as interpreted by Emsley (1969); 12: as interpreted by the author (dc = discal cell).
First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species... 333
gradually narrowing towards apex; bearing a small, flattened, rounded process subapically; basal part distinctly widened, stout. Phallus (as incompletely seen in situ; Fig. 13) with a long, narrow vesica (Figs 13, 19, 20: v) forming about 2.5 coils, bearing a long, apically abruptly narrowing process (Fig. 13: pv) at about 2/5 of its total length. Measurements (in mm). Holotype male. Body length from apex of head to apex of fore wing 1.45. Width of head across eyes 0.504, interocular distance 0.264. Length
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Figs 13–18: Pinochius josifovi nov. sp., male. 13: abdomen from dorsal view; 14: process of tergite VIII; 15: large appendage of proctiger; 16, 17: right paramere, two different aspects; 18: left paramere and apex of right paramere, dorsal view, in situ. Measurements in mm. Lettering: apg1 = large appendage of proctiger; apg2 = small appendage of proctiger; lp = left paramere; pt8 = process of tergite VIII; py = pygophore; rp = right paramere; s3–s7 = sternites III–VII; t3–t8 = tergites III–VIII; v = vesica; pv = process of vesica.
334 D. Rédei
of antennal segments I : II : III : IV as 0.064 : 0.104 : 0.360 : 0.256. Length of pronotum along meson (including collar) 0.328, length of collar along meson 0.050, humeral width 0.664, width of collar 0.344. Length of mesoscutellum 0.156, basal width 0.240. Length of fore wing 1.16. Lengths of femur, tibia, and tarsus (tarsal segments I : II : III) of fore leg 0.368, 0.328, and 0.140 (0.044 : 0.084 : 0.080); of mid leg 0.352, 0.312, and 0.130 (0.040 : 0.078 : 0.060); lengths of femur, tibia, and tarsus (tarsal segments I : II) of of hind leg 0.360, 0.520, and 0.176 (0.048 : 0.158). Females (specimens from Tanh Liet / Sin Chai). Body length from apex of head to apex of fore wing 1.65 / 1.75. Width of head across eyes 0.518 / 0.546, interocular distance 0.280 / 0.301. Humeral width of pronotum 0.777 / 0.784. Macropterous female. Similar to male but slightly larger and darker; head across eyes about 1.80–1.85 times as wide as interocular distance; pronotum about 1.45–1.5 times as wide as head. Propleura separated from pronotum by a distinct horizontal carina; proepimeron with a distinct oblique carina projecting from acetabulum to small projection on posterior margin; proepimeral carina distinctly more conspicuous than carina separating pronotum and propleura. Pteropleura and -sterna (studied only in female). Mesosternum (Fig. 8) with a small median incision anteriorly, without posterior protuberance between mid coxae; metasternum projecting posteriorly between hind coxae, projecting part distinctly widened apically; pterothoracic sternum with an elongate median protuberance anteriad to hind acetabuli, apparently of metasternal origin. Etymology. It is a great pleasure to dedicate this interesting new species to Prof. Michail Josifov on the occasion of his 80th birthday.
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Figs 19–20: Pinochius josifovi nov. sp., male. 17: abdomen from ventral view; 18: abdomen from lateral view. Measurement in mm. Lettering as in Fig. 13; in addition: t8* = right, vertical part of tergite VIII.
First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species... 335
Habitat and biology. The male holotype was collected at light. One of the females (from Tanh Liet) was beaten from trees (Gy. Topál’s unpublished collecting diary), the other one (from Sin Chai) was collected by sweep-netting from herbaceous riparian vegetation in a riverine forest (L. Peregovits, personal communication).
DISCUSSION Zoogeography. Pinochius josifovi nov. sp. from North Vietnam extends considerably the known range of the genus, reported from Africa and Australia so far. However, similar discoveries might be expected among Dipsocoromorpha, since they are usually rare in collections due to their very small size and cryptic habits. A most probably undescribed species of Pinochius, represented only by a single female collected in India, is deposited at the Hungarian Natural History Museum. Therefore, occurrence of Pinochius is expected from other parts from the Oriental Region too. The only other genus of Schizopteridae which is known to occur in more than one of the great zoogeographical realms is Hypselosoma Reuter, 1891, reported from the Afrotropical (Madagascar), Oriental, Australian and Pacific Regions as well as from marginal areas of the Palaearctic Region ( Japan) so far. The discovery of Pinochius josifovi nov. sp. also means the first report of the family Schizopteridae from Vietnam. Fore wing venation. Little is known about the homologies of wing venation of Dipsocoromorpha. Emsley (1969: 126, fig. 32, redrawn in Fig. 11) presented a hypothesys for the fore wing venation in Pinochius. However, by the microscopic examination of the fore wing of P. josifovi nov. sp., it is without doubt that the wide vein forming the leading edge of the remigium includes two, perpendicularly running tracheae in most of its length (their run is clearly traceable distad to the second marginal cell), therefore, it might be originated from two veins as also noted by Hill (1985: 276). The author’s interpretation of the fore wing venation of Pinochius is illustrated in Fig. 12. It seems probable that the wide marginal vein forming the leading edge of the remigium is originated from the fusion of C and Sc. Most probably the strongly elevated, carina-like central longitudinal vein is at least in its basal part the fused R+M (its presence is a common feature in all Heteroptera), and the longitudinal vein of the remigium running along the claval furrow is the Cu (as in most other Heteroptera) as suggested by Emsley. In this case, the small, triangular or more or less tetragonal cell in the middle of the wing posterior to the carina-like central longitudinal vein is delimited proximally by m-cu, posteriorly by Cu, therefore, it is homologous with the cubital (= discal) cell commonly found in Enicocephalomorpha and many derived Heteroptera. The posterior vein of the remigium is, like in most other Heteroptera, most probably the continuation of the anterior claval vein (its origin is not clear in Heteroptera, tentatively labelled as 1A in this study) and not that of the posterior vein as suggested by Emsley. The origin of the second and third oblique veins are not clear; the second is almost certainly originated from M (probably M1), the third is maybe the second branch of M (= M2).
336 D. Rédei
Relationships within Pinochius. Hill (1985) considered in details the possibility of P. australiensis being generically distinct of the African species. The presence of a posteriorly directed right-side process on tergite VIII is unique within the species of Pinochius known so far. Because of this character, P. josifovi nov. sp. might differ from the other species at subgeneric or even generic level. However, among the African species of Pinochius, only the male genitalia of P. anterii and P. dundundo are known (the other four species, unfortunately also the type species of the genus, were described based only on a single female each), furthermore, the new species strongly agrees in several respects to all the other species. Because of these reasons, no changes are introduced in the generic level taxonomy of this group in the present paper, but it is noted that Pinochius seems to be in need of redefinition. Pinochius josifovi nov. sp. shares the following characters with P. australiensis (conditions in the African species P. anterii and P. dundundo are given in parentheses) (all data are taken from the original descriptions): pterothoracic sternum with an elongate median protuberance anteriad to hind acetabuli, apparently of metasternal origin (with well developed median protuberance between mid coxae, apparently of mesosternal origin); claws with distinct basal tooth (without tooth); right paramere with a small subapical process (without subapical process). Other characters shared with P. anterii and P. dundundo but not with P. australiensis are as follows: lateral margin of pronotum slightly concave (convex); sternite VII of the male is strongly asymmetrical (nearly symmetrical); vesica is long, narrow, bearing a process (short, lacking process). Consequently, the phylogenetic relationships of the above species are not evident. Hill (1985) noted important differences of the prothoracic carinae between the African species and P. australiensis. In this respect, P. josifovi is similar to P. australiensis, having a proepimeral carina which is distinctly more conspicuous than the carina separating pronotum and propleura; however, the proepimeral carina is not distinctly angulate but nearly straight, oblique. The re-examination of this character in the African species is necessary.
ACKNOWLEDGEMENTS I am greatly indebted to Lionel Hill (Department of Primary Industries, Water and Environment, Devonport) for his careful reading, valuable comments and suggestions to the manuscript. Thanks are due to András Kun and László Peregovits (Hungarian Natural History Museum, Budapest) for information about the locality of one of the specimens.
РЕЗЮМЕ Семейство Schizopteridae (Heteroptera) се съобщава за първи път за Виетнам. Родът Pinochius CARAYON, 1949 е известен от Африка и Австралия досега и с описанието на
First record of Pinochius Carayon, 1949 from the Oriental Region, with description of a new species... 337
Pinochius josifovi nov. sp. от Северен Виетнам се съобщава за първи път в Ориенталската област. Новият вид е уникален сред описаните досега видове от този род с наличието на насочен назад израстък на осмия тергит.
REFERENCES Carayon J. 1949. Description d’un Schizopterinae africain, Pinochius africanus n. g., n. sp., et remarques sur les Hémiptères Cryptostemmatidae. — Bulletin du Muséum d’Histoire Naturelle, 2e série, 21(2): 239–242. Emsley M.G. 1969. The Schizopteridae (Hemiptera: Heteroptera) with the description of new species from Trinidad. — Memoirs of the American Entomological Society, 25: i + 1–154. Hill L. 1985. A new species of Pinochius Carayon (Hemiptera: Schizopteridae), first record from Australia. — Journal of the Australian Entomological Society, 24: 275–279. Linnavuori R. 1966. Two new Heteroptera species from Egypt. — Annales Entomologici Fennici, 32(3): 263–264. Rédei D. 2007. A new species of the family Hypsipterygidae from Vietnam, with notes on the hypsipterygid fore wing venation (Heteroptera: Dipsocoromorpha). — Deutsche Entomologische Zeitschrift, 54(1): 43–50. Štys P. 1970. On the morphology and classification of the family Dipsocoridae s. lat., with particular reference to the genus Hypsipteryx Drake (Heteroptera). — Acta entomologica bohemoslovaca, 67: 21–46. Štys P. 1982. A new Oriental genus of Ceratocombidae and higher classification of the family (Heteroptera). — Acta entomologica bohemoslovaca, 79: 354–376. Štys P. 1990. Enicocephalomorphan and dipsocoromorphan fauna of W. Palaearctic (Heteroptera): composition, distribution and biology. — Scopolia, Suppl. 1: 3–15. Wygodzinsky P.W. 1950a. Schizopterinae from Angola (Cryptostemmatidae, Hemiptera). — Publicações Culturais da Companhia de Diamantes de Angola, 7: 9–48. Wygodzinsky P.W. 1950b. Contribution towards the knowledge of the family “Cryptostemmatidae” (Hemiptera). — Revista Brasileira de Biologia, 10(4): 377–392.
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S. Grozeva & N. Simov (Eds)Chorosomatini) 2008 The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: from the New World... 339 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 339-348. © Pensoft Publishers Sofia–Moscow
The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World: Chorosoma josifovi nov. sp. M.D. Schwartz1, C.W. Schaefer2 & J.D. Lattin3 1
Agriculture & Agri-Food Canada, Research Centre, 960 Carling Avenue, Ottawa ON K1A OC6, Canada. E-mail:
[email protected] 2 Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs CT 06269-3043, U.S.A. E-mail:
[email protected] 3 Department of Botany and Plant Pathology, Oregon State University, Corvallis OR 97331-2902, U.S.A.
ABSTRACT Except for one Ethiopian species, Chorosoma until now has been an exclusively Palearctic genus. Fifty years ago, a new species was discovered in the northwestern United States; we name this species Chorosoma josifovi in honor of the 80th birthday of Michail Josifov, who described C. gracile among his many other contributions to our knowledge of Heteroptera. The new species resembles the widespread C. schillingii in some features, and resembles other species in some other features.
INTRODUCTION The six species in Chorosoma are all Palearctic (Göllner-Scheiding 1983; Dolling 2006) except C. xenocles Linnavuori, which occurs in Ethiopia (Linnavuori 1976). In 1956, several specimens of the genus were collected in the state of Idaho, U.S.A.; and since then others have been found in Oregon and Nevada, all adjacent states in the far northwest of the United States (Fig. 1). So far, there have been no collections of the species from Utah or from north of Oregon—that is, from central Washington state north through the dry land areas of British Columbia and the Yukon
340 M.D. Schwartz, C.W. Schaefer & J.D. Lattin
Fig. 1: Map of the distribution of Chorosoma josifovi nov. sp. (Pacific Ocean at lower left)
Territory of western Canada into Alaska (Henry 1988; Scudder 1997; Maw et al. 2000; Paiero et al. 2003). Comparison of determined specimens, illustrations, and descriptions of the six described Chorosoma species with the North American specimens, makes clear that these collections represent a new species. We describe this new species here, and name it for Dr. Michail Josifov, in honor of his many fine contributions to the study of Heteroptera (including the description of Chorosoma gracile) and in happy celebration of his 80th birthday. Because most other species of Chorosoma occur in the Palearctic, we assume that the ancestor of this new species has immigrated to the New World from the Old. Although members of the genus occur widely throughout eastern Russia and the former eastern Soviet Republics, and occur also in Mongolia, no species of Chorosoma appears to have been recorded so far from far-northeastern Russia, near Beringia (see Martynova 1975; Göllner-Scheiding 1983; Vinokurov 1988; Dolling 2006), where the two continents were most recently connected during the Pleistocene, although Vinokurov & Kanyukova (1995) record it from Siberia. Indeed, the preference of Chorosoma for very dry grasslands suggests a movement into the New World long ago, before the growth of the boreal forests. We are now studying the systematic and biogeographic relationships and history of this new species, and we shall discuss these elsewhere (Schaefer, Schwartz & Lattin, in preparation).
The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 341
TAXONOMY Chorosoma josifovi nov. sp. (Figs 2-5) Description. Male: Gracile, abdomen extending beyond membrane of hemelytron for distance subequal to length of membrane; total length 10.14-11.60 mm (Fig. 2). COLORATION: Entirely pale yellowish brown to sandy, except for reddish brown speckling on head, antennal segments I and II, and dorsum and sublaterally on venter of abdomen; hemelytron and membrane hyaline, embolium and veins concolorous with rest of body, except sometimes faintly brown on radial vein; reddish brown lateral stripe between antennal insertion and eye and faint lateral stripe on abdominal venter submedial to connexivum; brown marks at base of suture between clypeus and mandibular plate, tip of labium, variable curved mark on medial periphery of callus, and short apical stripe dorsal to procoxal cleft and ventral to lateral carinate margin of pronotum; abdominal dorsum with prominent, wide, irregular, dark brown or black line just medial to connexivum extending from tergum 1 (1T) (beside scutellum) to or slightly beyond 4T or 5T, lines occasionally shorter or longer; some specimens with line anteriorly, between lateral lines; trichobothrial bases and abdominal spiracles not contrastingly colored; hind
Fig. 2: Chorosoma josifovi nov. sp., dorsal habitus.
342 M.D. Schwartz, C.W. Schaefer & J.D. Lattin
leg with dark brown on apicomedial surface of tibia, apical half of tarsomere I; tarsomere III and pretarsus dark brown; tips of parameres dark brown to black. SURFACE and VESTITURE: Discretely punctate, except callar region smooth and slightly rugose; moderately densely covered with very short, appressed pale setae; setal bases of postcallar pronotum often dark; antennal segment IV also with sparse, slightly longer, suberect setae; apical portion of tibia, and tarsomeres with longer setae; medial surface of metatibia with erect, pale, bristlelike spines with length subequal to diameter of tibia. STRUCTURE: Lateral margins of head subparallel, except for bulge of eye; base of head slightly raised, ridgelike, raised area extending forward sublaterally to enclose ocellar tubercle; tubercle low. Eyes rather prominent, distance across them (lateral margin to lateral margin) approximately 80% of maximum width of pronotum. Antenna relatively long, antennal segment I thicker than others, subequal in length to head (clypeus to ocellar line), parallel-sided laterally, convex medially; II-IV slender, II slightly thicker proximally, IV slightly clavate, II≈III>I=IV (see Measurements). Labium reaching mesocoxa, segment I>II>IV>III. Pronotum narrow, broadened slightly caudally, lateral margins very slightly sinuate, posterior margin straight, callus broad, disc densely punctate, median carina faint, especially apically; humeral area produced laterally into broad rounded tubercle. Scutellum narrow apically, slightly concave medially with faint median carina, apex rounded, distinctly concave, slightly raised. Apex of hemelytron membrane reaching apex of abdominal segment V. Pretarsus with strongly curved claw, reaching beyond apex of broad and apically attenuate pulvillus. GENITALIA: Pygophore with rather widely spaced lateral and paralateral lobes; both lobes longer than medial lobe (Figs 3, 4). Parameres distinctly narrow apically, tip strongly sclerotized, curved laterally (Fig. 5). Aedeagus with tightly coiled vesical base; apical region with sclerotized and membranous lobes. Female: Same as male, but larger (Table 1); apex of hemelytral membrane reaching middle of 5T. medial lobe
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The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 343
0.10 mm
Fig. 5: Chorosoma josifovi nov. sp., right paramere, lateral view; scale = 0.10 mm.
TERMINALIA: Proctiger prominently exposed ventrally, covered dorsally by exposed 8T and 9T; ovipositor concealed by seventh sternum. MEASUREMENTS: Table 1. ABBREVIATIONS: CNC, Canadian National Collection, Ottawa; CWS, collection of Carl W. Schaefer, Storrs; OSUC, Oregon State University, Corvallis; UIM, University of Idaho, Moscow; UNV, University of Nevada, Reno; USNM, United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. HOLOTYPE: ♂. [U.S.A.] 1) “Blow Sand Mtns./ Churchill Co. NV/ T16N, R30E [39.204442, -18.591547, 1317m]/ viii-2-1979. 2) R. C. Bechtel/ R. W. Rust/ Collectors. 3) Oryzopsis hymenoides. 4 [red] HOLOTYPE/ Chorosoma josifovi/ Schwartz, Schaefer, & Lattin.” Deposited in USNM. PARATYPES [22 specimens with blue PARATYPE labels; no specimen is in perfect condition]: U.S.A.: Idaho: Fremont Co.: Sand Dunes, Saint Anthony [43.962428, -111.854627, 1488m], 16 July 1956, W. L. Barr, 1♂, 1♀ (CNC), 3♂ (UIM). Nevada: Churchill Co.: same labels as holotype, 1♂, 2♀ (CWS), 1♀ (OSUC), 3♂, 2♀ (USNM); Blow Sand Mt., 28 mi SSE Fallon, June 1979, R. W. Rust, 1♀ (OSUC, UNV), 2♀ (USNM). Oregon: Harney Co.: Alvord Desert Sand Dunes, 18 June-1 July 1986, N. Cobb, ex: Oryzopsis hymenoides, Sporobolus airoides, 1♂ (OSUC); T36S R35E S8 SW [42.428350, -118.420327, 1281m], 20 July 1979, Lightfoot and Cobb, 1♀ (OSUC); sample 44 [in pencil], Neil Cobb, 2♂ (OSUC) [we strongly suspect that these specimens were collected at one of the previous localities in the Alvord Desert]. ADDITIONAL SPECIMENS [sex undetermined because of very poor condition]: Idaho: Fremont Co.: 6 mi NW of Saint Anthony, sand dunes, 16 July 1956, W. F. Barr,
344 M.D. Schwartz, C.W. Schaefer & J.D. Lattin
Table 1. Measurements (mm.) of Chorosoma josifovi nov. sp. Specimens from Idaho (U.S.A) differ somewhat from those from Oregon and Nevada, and are listed separately. Male
Total length Length: head to apex hemelytron Head lengtha
Idaho Oregon/Nevada (N=4) (N=4) 11.31 (10.75-11.60) 10.29 (10.14-10.53)
Female Idaho Oregon/Nevada (N=1) (N=2) 14.80 13.33 (12.74-13.91)
8.25 (7.80-8.80)
7.45 (7.28-7.54) (N=3)
9.70
8.84 (8.19-9.49)
1.33 (1.25-1.40)
1.29 (1.29) (N=2)
1.50
1.53 (N=1)
1.13 (1.09-1.16) (N=2)
1.33
1.23 (1.19-1.32)
0.70 (0.66-0.73) (N=2)
0.83
0.89 (0.86-0.91)
1.17 (1.15-1.18) Ocular distance (from outer edge of each eye) Interocular width 0.69 (0.66-0.70) Antennal segments: I
1.52 (1.48-1.60)
1.41 (1.39-1.41) (N=2)
1.68
1.75 (N=1)
II
2.32 (2.25-2.45)
2.05 (1.98-2.11) (N=2)
2.48
2.38 (N=1)
III
2.17 (2.05-2.30)
1.90 (1.82-1.98) (N=2)
2.15
2.05 (N=1)
IV
1.58 (N=1)
1.64 (1.52-1.75) (N=2)
----
1.45 (N=1)
Pronotal length
1.38 (1.30-1.38)
1.19 (1.19) (N=2)
1.66
1.39 (1.32-1.45)
Pronotal width
1.43 (1.41-1.45)
1.35 (1.32-1.39) (N=2)
1.70
1.64 (1.52-1.75)
Rostrum length
3.11 (3.00-3.20)
2.78 (2.54-2.94) (N=3)
3.10
2.96 (2.81-3.10)
Scutellum length
0.81 (0.75-0.86)
0.71 (0.60-0.73) (N=3)
0.95
0.85 (0.79-0.92) (N=3)
Scutellum width
0.49 (0.44-0.54)
0.56 (0.53-0.59) (N=3)
0.55
0.62 (0.61-0.63 (N=3))
a
from tip of clypeus to anterior edge of ocellar line
1 spec. consisting of head and thorax only (CWS). Nevada: Churchill Co.: same labels as holotype, 4 spec. consisting of head and thorax only (all in CWS). VARIATION: There is some variation in coloring and patterning, most noticeably between the specimens from Idaho and those from Oregon and Nevada (see below). The Oregon specimens also show other variation. In particular, the two sublateral dark lines on the abdominal dorsum may extend only as far as the third tergite (3T), or as far as 7T. The latter occurs in the male holotype (Oregon); the former in a female collected on the same day at the same place. In other specimens also collected with these (same place and time), the dark lines end at 4T or 5T; this occurs in the majority of specimens. When the dark lines extend the full length of the abdomen, the insect appears dark; nevertheless, the ground color is pale. Minor variation also occurs in the degree of overall paleness, and in the degree of darkness, and extent, of the various markings given in the Description. None of this variation is consistent by sex or population.
The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 345
DIFFERENCES BETWEEN UNITED STATES POPULATIONS: There are some differences between the specimens collected in 1956 in Idaho (at 1488 m), and those collected later (1979, 1986) in Oregon (at 1281 m) and Nevada (at 1317 m), although structurally they are the same. These populations are separated by habitat inhospitable to Chorosoma and are not close to one another (the closest localities are 322 km apart, and the farthest are 724 km apart). The specimens from Idaho are in general darker than the Oregon-Nevada specimens. They also differ with respect to several measurements (Table 1). The Idaho specimens are somewhat, but consistently, larger. Perhaps of greater significance, would be differences in absolute and relative lengths of head and antennal segments. These appear to be important in distinguishing Chorosoma species (see key in Linnavuori 1976). We have calculated several ratios to aid discussion (Table 2). The head length/ antennal-segment I ratios do not differ greatly, either by sex, United States state, or date of collection (the last two of course are a single variable). This ratio indicates that the first antennal segment is longer, but not greatly longer, than the head (measured from clypeal tip to an imaginary line drawn across anterior edge of ocelli). Similarly, the first antennal segment is about two-thirds the length of the second, and there is little variation among the North American specimens. None of these measurements (Table 1) or ratios (Table 2) suggests that these different North American populations are not conspecific. There may be significant interspecific differences in the genus, however, which we shall consider in the future (in preparation). HABITAT AND FOOD PLANTS: All specimens of Chorosoma josifovi were collected on dune grasses in or on sand dunes, from ~1280-1490 m elevation. The recorded food plants are the native grasses Oryzopsis hymenoides (Roemar & Schultes) Ricker (“ricegrass”) (now Achnatherum hymenoides (Roemer and Schultes) Barkworm) and Sporobolus airoides (Torr.) Torr. (“finetop saltgrass”). Both grasses grow on sandy, or very sandy, soil (Fernald 1950; Gleason & Cronquist 1991); S. airoides in alkaline meadows (Gleason & Cronquist 1991) and saline flats (Fernald 1950). With respect to habitat and food plant, Chorosoma josifovi is similar to other species of Chorosoma, at least to the European ones. C. xenocles was collected in (near?) Addis Ababa, Ethiopia (Linnavuori 1976), whose elevation is, for the most part, less than 3000 m. COMPARISON WITH OTHER SPECIES: The following discussion is based on identified specimens of Chorosoma brevicolle Hsiao, C. gracile Josifov, C. longicolle Reuter, and C. schillingii (Schilling); in addition, for these species and for C. xenocles LinnaTable 2. Ratios of several measurements of United States Chorosoma josifovi nov. sp. (data from Table 1). Idaho specimens
Oregon & Nevada specimens
Male
Female
Male
Female
Head length/antennal I
0.86
0.89
0.91
0.86
Antennal I/II
0.66
0.68
0.69
0.74
Head to hemelytron apex/body length
0.73
0.66
0.72
0.66
346 M.D. Schwartz, C.W. Schaefer & J.D. Lattin
vuori, we have trusted descriptions, illustrations, and characters in Linnavuori (1976). Distinguishing among these species is not easy, because in all species there is considerable intraspecific variation in color and in structure. For example, the most widespread species, Chorosoma schillingii, varies in size: we have identified specimens from “England” (no further data) that are considerably larger than other specimens from Kazakhstan (England male 14.43 mm, Kazakhstan male 12.35; England female 17.42, Kazakhstan female 14.43). And, judging from the illustrations of parameres and genital capsules in Josifov (1968, Figs 7-9, 13-17), Linnavuori (1976, Figs 12c, d), and Pehlivan (1981, Fig. 54), these structures also vary considerably. We shall discuss these differences, similarities, and intraspecific variations elsewhere (Schaefer, Schwartz, & Lattin, in preparation). Chorosoma josifovi differs from other Chorosoma species chiefly and most consistently in being considerably paler (especially the Oregon-Nevada specimens) and more uniformly colored. In particular, the dorsum of the abdomen in the other species we have seen is darker than in C. josifovi. Because these other species occur in the same sort of dry habitat as C. josifovi, the paleness of the latter cannot be attributed entirely to its habitat; therefore, this color difference appears to be significant. Chorosoma josifovi is of moderate size (Table 1), not much different in length from the other species, except C. brevicolle, which is larger (17 mm; Hsiao 1964; the “1.7” in the figure caption is a misprint); some C. schillingii are large, too: a female from “England” is 17.42 mm long, but other specimens are of average length (11-14 mm), like other species. Although except for color, distinguishing C. josifovi from other species is not easy, there are more subtle consistent differences, especially in genitalia, that convince us the New World specimens represent a new species. These differences too will be discussed elsewhere. In Chorosoma schillingii, the two sublateral dark lines on the abdomen’s dorsum extend to the tip of the abdomen or onto the sixth tergite; in other species they usually end no more posteriorly than on the fourth or fifth tergite. However, the variation noted above renders this character less reliable than may have been thought. There are several keys to the species of Chorosoma: Martynova (1975), Linnavuori (1976), and Putshkov (1986). Species are often separated in these keys not by distinct discrete differences but by relative differences such as antennal ratios. But here too confusion may occur: for example, antennal IV of C. longicolle is “fast so lang wie III” ( Josifov 1968, p. 256); or equal in males but III shorter in females (Martynova 1975, Putshkov 1986); or, the ratio of the two is 48: 58 (III: IV) (Linnavuori 1976, key, couplet 3). Also used to distinguish species is the relative length of the antenna, usually measured by the antennal I: head length ratio. In this regard the comparative and total lengths of the antennal segments of C. josifovi are very close to those of C. longicolle (compare our Fig. 3 with Josifov 1968, Fig. 1). The paramere of Chorosoma josifovi (Fig. 5) is longer and more slender than that of C. longicolle, and bent laterally at the tip more than is the paramere of C. longicolle, whose paramere tip is bent almost dorsally, along the main plane of the paramere’s body. In these respects the C. josifovi paramere more closely resembles those of C. gracile, C. schillingii ( Josifov 1968, Figs 13-22; Linnavuori 1976, Figs 12c, d), and C. xenocles
The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 347
(Linnavuori 1976, Fig. 12b). In all species there is ventrally a median extension of the male’s genital capsule (actually, the fused cuplike sclerite and median projection: see Schaefer 1977, 1978). Like the paramere, this is more slender and longer in C. josifovi (Figs 3, 4) than in C. longicolle; in the former (and in C. schillingii [ Josifov 1968, Fig. 7]) it is about three-fourths of the paramere’s length, but slightly more than half that length in C. gracile ( Josifov 1968, Fig. 10) and in C. longicolle. The paralateral lobe (Figs 3, 4) is less prominent than in C. gracile ( Josifov 1968, Figs 10-12) and C. schillingii ( Josifov 1968, Figs 7-9; Schaefer 1978, Fig. 3). However, it is more prominent than that of C. longicolle (Martynova 1975, Fig. 9; CWS, personal observation). Finally, Martynova (1975) suggests that C. brevicolle is actually a composite of C. macilentum and one or more other described species. The close similarity of the species and the great intraspecific variation, point clearly to a need for a generic revision of Chorosoma, which must include an analysis of populations over the entire range of the genus. We shall discuss this need in greater detail elsewhere. Note: Part of this paper was presented at the Fourth European Hemiptera Congress (Ivrea, Italy, September 2007), by CWS.
ACKNOWLEDGEMENTS We thank W. L. Barr, University of Idaho (UIM) and R. W. Rust, University of Nevada, Reno for originally bringing the specimens from Idaho and Nevada respectively to our attention. We also appreciate the assistance extended to us by R. G. Foottit for access to the collection and facilities of the Canadian National Collection. We are grateful also to V. Kask and J. O’Donnell (University of Connecticut) and B. B. Hall (Oregon State University) for the dorsal habitus illustrations; to Fedor Konstantinov, Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZISP), for translating several Russian-language papers; to D. Lightfoot and N. Cobb (while at OSU) for their collecting at the Alvord Desert site in Oregon; and to Dr. I. M. Kerzhner (ZISP) for gently revealing to us several major and minor errors in this paper. And we thank T. J. Henry (U.S. National Museum, Smithsonian Institution) for the loan of specimens. Finally, CWS is grateful to the sponsors and organizers (and especially to Dr. Peter Mazzoglio) of the Fourth European Hemiptera Congress, for support which allowed him to present part of this paper at the Congress.
РЕЗЮМЕ По екземпляри, намерени преди 50 години в северозападната част на САЩ, се описва нов вид Chorosoma, наименована C. josifovi, в чест на 80 годишнина на M. Josifov като признание като признание за приноса му към изучаването на Hetoroptera. Новият вид наподобява по някои белези широкоразпространения C. schillingii. Това е втория представител на рода извън границите на Палеарктика.
348 M.D. Schwartz, C.W. Schaefer & J.D. Lattin
REFERENCES Dolling W.R. (2006). Family Rhopalidae Amyot & Serville, 1843. — In: B. Aukema & C. Rieger (Eds.): Catalogue of the Heteroptera of the Palaearctic region, vol. 5 Pentatomomorpha II. The Netherlands Entomological Society, Amsterdam: 8-27. Fernald M.L. (1950). Gray’s manual of botany. 8th edition. Dioscrides Press, — Portland, Oregon: 1-1632. Gleason H.A. & A.C. Cronquist (1991). Manual of vascular plants of northeastern United States and adjacent Canada. 2nd edition. — The New York Botanical Garden, New York, New York: 1-910. Göllner-Scheiding U. (1983). General-Katalog der Familie Rhopalidae (Heteroptera). — Mitteilungen des Zoologischen Museums Berlin, 59: 37-189. Henry T.J. (1988). Family Rhopalidae Amyot & Serville, 1843 (=Corizidae Douglas & Scott, 1865). The scentless plant bugs. — In: Henry T.J. & R.C. Froeschner (Eds.): Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E.J. Brill, Leiden: 625-644. Hsiao T.-Y. (1964). New Coreidae from China (Hemiptera, Heteroptera) III. — Acta Zoologica Sinica, 16: 251-262. Josifov M. (1968). Eine neue Chorosoma-Art aus Bulgarien (Heteroptera, Rhopalidae). — Reichenbachia, 10: 255-258. Linnavuori R. (1976). Taxonomical studies on African Coreoidea (Heteroptera). —Notulae Entomologicae, 56: 89-96. Martynova G.P. (1975). Species of the genus Chorosoma Curtis (Heteroptera, Rhopalidae) of the U.S.S.R and Mongolia.— Nasekomye Mongoli,i 3: 79-85. (in Russian). Maw H.E.L., R.G. Foottit, K.G.A. Hamilton & G.G.E. Scudder (2000). Checklist of the Hemiptera of Canada and Alaska. — Ottawa, NRC Research Press: viii, 1-220. Paiero S.M., S.A. Marshall & K.G.A. Hamilton (2003). New records of Hemiptera from Canada and Ontario. — Journal of the Entomological Society of Ontario, 134: 115-129. Pehlivan E. (1981). Türkiye Stenocephalidae, Rhopalidae ve Alydidae (Heteroptera: Coreoidea) Faunasi, Üzerinde Sistematik Araştirmalar. — E.Ü. Ziraat Fakültesi, Izmir, Turkey: 1-189. (in Turkish) Putshkov V.G. (1986). Bugs of the family Rhopalidae (Heteroptera) of the fauna of the USSR. — Opredeliteli po Faune SSSR 146: 1-132. (in Russian) Schaefer C.W. (1977). The genital capsule of the trichophoran male (Hemiptera: Heteroptera: Geocorisae). International Journal of Insect Morphology and Embryology, 6: 277-301. Schaefer C.W. (1978). The genital capsule of the Rhopalidae (Hemiptera: Heteroptera: Coreoidea). — Annals of the Entomological Society of America, 71: 659-666. Scudder G.G.E. (1997). True bugs (Heteroptera) of the Yukon. — In: Danks, H.V. & J.A. Downes (Eds.): Insects of the Yukon. Biological Survey of Canada, Ottawa: 241-336. Vinokurov N.N. (1988). Heteroptera of Yakutia.—Amerind Publishing Co. Pvt. Ltd., New Delhi: i-xi + 1-328. (English translation of Russian original) Vinokurov N.N. & E.V. Kanyukova (1995). Heteroptera of Siberia. Nauka, Novosibrisk (in Russian).
S. Grozeva & N. Simov 2008 (Heteroptera, Microphysidae) 349 A new Loricula species (Eds) from Bulgaria ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 349-356. © Pensoft Publishers Sofia–Moscow
A new Loricula species from Bulgaria (Heteroptera, Microphysidae) N. Simov National Museum of Natural History, 1 Tzar Osvoboditel Blvd., 1000 Sofia, Bulgaria. E-mail:
[email protected]
ABSTRACT Loricula (Myrmedobia) josifovi nov. sp. from Strandzha Mts., Bulgaria is described. Loricula (Myrmedobia) coleoptrata (Fallén, 1807) and Loricula (Mymericula) bedeli (Montandon, 1887) are reported as new to Bulgarian fauna. Keywords: Loricula (Myrmedobia) josifovi nov. sp., Microphysidae, Bulgaria, new records.
INTRODUCTION The examination of material of Microphysidae from collection of Dr. Michail Josifov and material collected by author in various regions in Bulgaria led to identify a new Loricula species which is described in this work with new faunistic records for Bulgarian fauna. Microphysids were collected by hand, pitfall traps and MSS (“milieu souterrain superficiel”) traps. MSS traps were made from PVC pipe with diameter of the whole 8 cm and length of 60 or 80 cm and were set vertically in limestone at 60 and 80 cm depth. Plastic cup, tied to polythene rope, and filled with solution of ethilenglycol with few drops of formalin was put at the bottom of the pipe. Solid plastic covers in order to avoid penetration of superficial fauna and infiltration of water during heavy rains covered traps. In the present paper the systematics of family Microphysidae, proposed by Popov (2004) is used.
350 N. Simov
RESULTS So far 9 microphysid species are known from the territory of Bulgaria (Tab. 1), including new species and new records reported below (Péricart 1996; Simov & Josifov 2004; present paper).
Loricula (Myrmedobia) josifovi nov. sp. Myrmedobia distinguenda (non Reuter 1884): Josifov 1964: 88.
Holotype: 1 male, labeled [Bulgaria, Ropotamo, 8.6.1959, M. Josifov leg]. Paratypes: 7 males and 4 females, as holotype; 1 female, labeled [Bulgaria, Ropotamo, 10.07.1967, M. Josifov leg]. Holotype and paratypes stored in the collection of Dr. Michail Josifov in Institute of Zoology, Sofia. Diagnosis: General habitus like representatives of subgenus Myrmedobia Bärensprung 1857, with thick rostrum, reaching anterior coxae, second rostral segment reach only the middle of eyes, relatively short preocular part of head. The new species belongs Table 1: List of Bulgarian Microphysidae. BG1 - zone of Quercus-forests south of Balkan Mts., except mountain regions in Southwest Bulgaria; BG2 - zone of Quercus-forests north of Balkan Mts., BG3 - mountainous regions above upper limit of Quercus (above 800–1000m). Species
BG1
BG2
+
+
BG3
Genus Loricula Curtis, 1883 Subgenus Loricula Curtis, 1883 Loricula (s.str.) bipunctata (Perris, 1857) Loricula (s.str.) pselaphiphormis Curtis, 1833
+
Loricula (s.str.) ruficeps (Reuter, 1884)
+
+
Subgenus Myrmedobia Bärensprung, 1857 Loricula (Myrmedobia) coleoptrata (Fallén, 1807)
+
Loricula (Myrmedobia) distinguenda Reuter, 1884
+
Loricula (Myrmedobia) exilis (Fallén, 1807)
+
Loricula (Myrmedobia) jakovlevi Péricart, 1969
+
Loricula (Myrmedobia) josifovi nov. sp.
+
Subgenus Mymericula Popov, 2004 Loricula (Mymericula) bedeli (Montandon, 1887)
+
+
A new Loricula species from Bulgaria (Heteroptera, Microphysidae)
351
to a species group with flattened lateral edging of the pronotum and with collar placed between anterior rounded pronotal angles. In well developed flattened edging (mostly in anterior half ) of pronotum, shape of pronotum, hemelytra and body the new species is similar to Loricula (Myrmedobia) exilis (Fallén, 1807), but easy recognized from the latter by smaller size, mat pronotum, shorter head and bigger value of ratio between II antennal article and distance between eyes. Description: Males (Fig. 1). Body length from head to apices of hemelytra 1.8-1.95 mm. Generally oblong, 2.2-2.4 as long as wide. General coloration brownish; head and pronotum dark brown; eyes light brilliant with reddish spots, hemelytra pale brown, almost transparent, except reddish-brown cuneus, membrane transparent, light smoky. Ventral side of the body reddish-brown. Dorsal surface of head covered with long pale hairs (Fig. 5). Dorsal surface of pronotum and hemelytra (except membrane) with very short, dense adpressed pale hairs and long, raised, pale hairs (Fig. 5, Fig. 8). Membrane with very short dense adpressed pale hairs. Lateral parts of pronotum, callosities and external margin of exocorium with bulb-like cuticular projections (Fig. 5, Fig. 7 and Fig. 8). Head short (Fig. 3, Fig. 5), 1.15x-1.25x as wide as long, with one pair of setae; preocular part about 0.5x as long as ocular + postocular part; head 0.9-1.1x as long as pronotum; frons 2.17-2.36x wider than diameter of eyes; rostrum robust, reaching anterior coxae, rostral segment II reaching only middle of eyes, II and IV are of equal lenght; antennae slender and pilose (Fig. 9), pale brown, all segments are equally thickened, segments II-IV covered with long pale hairs, 2x longer than diameter of antennal segments, antennal segment II 1.46x as long as distance between eyes and 0.76-0.78x as long as the diatone; proportions of antennal segments I-IV: 3.5-9.5-7-10. Pronotum mat, except callosities, transverse 1.85-2.1x as wide as long with flattened lateral edging
1
2
Fig. 1: Habitus SEM photograph of Loricula (Myrmedobia) josifovi nov. sp., male, dorsal view (scale bare 1 mm). Fig. 2: Habitus SEM photograph of Loricula (Myrmedobia) josifovi nov. sp., female, dorsal view (scale bare 1 mm).
352 N. Simov
3
4
5
6
7
8
9
10 Fig. 3: Loricula (Myrmedobia) josifovi nov. sp. - head and pronotum, male, dorsal view (scale bare 0.1 mm). Fig. 4: Loricula (Myrmedobia) josifovi nov. sp. - head and pronotum, female, dorsal view (scale bare 0.1 mm). Fig. 5: Loricula (Myrmedobia) josifovi nov. sp. - head and pronotum, male, lateral view (scale bare 0.1 mm). Fig. 6: Loricula (Myrmedobia) josifovi nov. sp. - head and pronotum, female, lateral view (scale bare 0.1 mm). Fig. 7: Loricula (Myrmedobia) josifovi nov. sp. - lateral part of pronotum of females – vestiture and bulb-like cuticular projections, dorsal view (scale bare 0.01 mm). Fig. 8: Loricula (Myrmedobia) josifovi nov. sp. - hemelytral vestiture, male, dorsal view (scale bare 0.1 mm). Fig. 9: Loricula (Myrmedobia) josifovi nov. sp. - antennae, male, dorsal view (scale bare 0.1 mm). Fig. 10: Loricula (Myrmedobia) josifovi nov. sp. - hemelytra, male (scale bare 1 mm).
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(mostly in anterior half ) (Fig. 3, Fig. 5), lateral margins converging anteriorly; posterior margin 1.5-1.7x as wide as anterior one; collar distinct, placed between rounded pronotal angles; anterior part of pronotum with pair of low callosities and distinct transversal impression. Mesoscutum 0.8x as long as the scutellum, the latter mat. Lateral margins of hemelytra slightly convex and parallel, external margin of exocorium distinctly thickened along its length up to the cuneal fracture, apex of cuneus subacute; proportion of length of hemelytron, corium and cuneus about 10-5-2; length of cuneus about 0.2x (0.2-0.21) as long as hemelytra and about 0.4x (0.39-0.43) as long as the corium; membrane with one cell and two veins, one originating from the cell and branching and other short vein branching from corium; membrane transparent, slightly smoky, with dark spot along the apical part of the cell (Fig. 10). Legs generally brown, in pale specimens yellowishbrown, apical and basal part of tibiae and apical part of femora slightly paler than the rest part; tarsi brownish-yellow; tibiae and femora slender and pilose, metatibiae weakly curved with long erected pale hairs on inner side, these hairs 3x as long as the diameter of metatibiae, their tarsi long, about 0.36x as long as tibiae. Measurements (in mm): Body length (including hemelytra) 1.8-1.95, width 0.770.87; head length 0.29-0.31, width (diatone) 0.34-0.36, distance between eyes 0.19, length of preocular part 0.1-0.11, length ocular + postocular part 0.19-0.21, eye width 0.08, antennal segments I: 0.1, II: 0.27, III: 0.2, IV: 0.27-0.29: rostral segments I: 0.06, II: 0.11, III: 0.13, IV: 0,11; pronotum: length 0.29-0.34, width 0.36-0.37 (ant.) and 0.56-0.61 (post.); length of open part of mesoscutum 0.1-0.11; length of scutellum 0.14-0.157; length of hemelytron, corium and cuneus (outer side): 1.47-1.57, 0.73-0.77, 0.29-0.33; length of posterior legs: femora 0.4, tibiae 0.61-0.62, tarsi 0.13 (0.03: 0.1), length of long erected pale hairs on inner side of tibiae 0.09-0.1; posterior tibiae width 0.03. Females (Fig. 2). Body length from head to apices of abdomen 1.45-1.5 mm. Generally rounded, 1.4-1.5 as long as wide. Micropterous. General coloration dark brown; head reddish brown, eyes light brilliant, anterior pronotal angles, lateral part of hemelytra, connexivum and apical part of abdomen brown to dark brown, in pale specimens lateral part of hemelytra pale brown. Ventral side of the body generally reddish-brown; mesosternum and metasternum reddish. Dorsal surface of head covered with long pale hairs (Fig. 6). Dorsal surface of pronotum and hemelytra with very short, dense adpressed pale hairs and long, raised, pale hairs (Fig. 6). Dorsal side of abdomen with long, raised, pale hairs (Fig. 2, Fig. 6). Head, lateral parts of pronotum, callosities, hemelytra and dorsal surface of abdomen with bulb-like cuticular projections (Fig. 4, Fig. 6, Fig. 7). Head short (Fig. 4, Fig. 6), 1.14x-1.19x as wide as long, with one pair of setae; preocular part about 0.75x as long as ocular + postocular part; head 1.05-1.1x as long as pronotum; frons 3-3.3x wider than diameter of eyes; rostrum robust, reaching base of anterior coxae, rostral segment II reaching only middle of eyes; antennae slender and pilose, pale brown, all segments are equally thickened, segments II-IV covered with long pale hairs, 2x longer than diameter of antennal segments, antennal segment II 1x as long as distance between eyes and 0.6-0.62x as long as the diatone; proportions of antennal segments I-IV: 3-7.5-6-8.5. Pronotum mat, except callosities, transverse 1.8-1.87x as
354 N. Simov
wide as long with flattened lateral edging (mostly in anterior half ) (Fig. 4, Fig. 6), lateral margins parallel; collar not distinct, placed between rounded pronotal angles; anterior part of pronotum with pair of low callosities and distinct transversal impression. Scutellum mat. Hemelytra reduced to minute leathery pads covering only the base of abdomen. Legs generally pale brownish-yellow; tibiae and femora slender and pilose, metatibiae with long erected pale hairs on inner side, these hairs 3x as long as the diameter of metatibiae, their tarsi long, about 0.43x as long as tibiae. Measurements (in mm): Body length 1.45-1.5, width 0.95-1.1; head length 0.3, width (diatone) 0.34-0.36, distance between eyes 0.21, length of preocular part 0.13, length ocular + postocular part 0.17, eye width 0.07, antennal segments I: 0.09, II: 0.21, III: 0.17, IV: 0.24: rostral segments I: 0.06, II: 0.13, III: 0.16, IV: 0,1; pronotum: length 0.27-0.29, width 0.5-0.53; length of scutellum 0.11; length of posterior legs: femora 0.42, tibiae 0.54-0.55, tarsi 0.13 (0.03: 0.1), length of long erected pale hairs on inner side of tibiae 0.09-0.1; posterior tibiae width 0.03. Etymology: The new species is named after my colleague, friend and teacher in heteropterology Dr. Michail Josifov in occasion of his 80th anniversary. Distribution: Up to now the new species was known only by the type locality in Bulgaria (Fig. 11). Locality is situated in Strandzha Mts., 5-10 m a.s.l., Ropotamo Reserve, about 5 km from outflow of Ropotamo River in Black Sea, near the old road from Primorsko Village to Sozopol Village, Bugras District ( Josifov in litt.). Ecological remarks: The species inhabit wet grassland communities in oak forest near Ropotamo river ( Josifov 1964). Males and females were caught on grasses with mosses among them ( Josifov in litt.).
New records of microphysids for the Bulgarian fauna Loricula (Myrmedobia) coleoptrata (FALLÉN, 1807) 1♂, Bulgaria, N 41º 33’ 48”, E 023º 37’ 17”, Southern Pirin Mts., 1970 m a.s.l., 1 km south of Orelek Peak, subalpine zone, 10 pitfall traps, 02.07.-31.07.2007, leg. N. Simov & M. Langourov. A West Palaearctic species. Until now it has been announced for the territory of the Balkan Peninsula from Croatia (Horváth 1897; Péricart 1972, 1996; Gogala 2004) (Fig. 11).
Loricula (Mymericula) bedeli (MONTANDON, 1887) 3♀♀, Bulgaria, N 41º 27.612, E 023º 29.448, Southern Pirin Mts., 494m a.s.l., SE slope of Sveti Iliya Hill, near Kalimantsi Village, communities of Quercus coccifera, MSS trap at 80 cm depth, 08.06.-27.06.2006, leg. S. Lazarov.
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Fig. 11: Map of distribution on Balkan Peninsula of: - Loricula (Myrmedobia) josifovi nov. sp.; - Loricula (Myrmedobia) coleoptrata (Fallén, 1807); - Loricula (Mymericula) bedeli (Montandon, 1887).
1♂, Bulgaria, N 41º 24’ 33”, E 023º 36’ 06”, Slavyanka/ Alibotoush Mts., 1250 m a.s.l., Alibotoush Biosphere Reserve; Pinus nigra wood with Ostrya carpinifolia, Quercus sp., Sorbus torminalis, Fagus sylvatica, Juniperus oxycedrus, Luzula sp.; 5 pitfall traps; 01.08.-29.08.2007, leg. M. Langourov & N. Simov A West Mediterranean species. Until now it has been announced for the territory of the Balkan Peninsula from Greece and Albania (Horváth 1916; Péricart 1972, 1996) (Fig. 11).
ACKNOWLEDGEMENTS The study was partly supported by the grants B-1523/05 and B-1525/05 of the Bulgarian Ministry of Education and Science, National Science Fund. I also thank to Dr. Yuri Popov for kindly provided papers.
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РЕЗЮМЕ Описва се видът Loricula (Myrmedobia) josifovi nov. sp. от България, Странджа планина. Видът принадлежи към групата видове със сплеснат латерален ръб на пронотума и е най-близък до Loricula (Myrmedobia) exilis (FALLÉN, 1807). Новият вид лесно се отличава от последния по по-малките размери, по-късата глава, матовия пронотум и по-големите стойности на съотношението между второто членче на антените и разстоянието между очите. Видовете Loricula (Myrmedobia) coleoptrata (FALLÉN, 1807) and Loricula (Mymericula) bedeli (MONTANDON, 1887) се съобщават за първи път за територията на България.
REFERENCES Gogala A. (2004). Heteroptera of Slovenia, II: Cimicomorpha I. — Annales, Annals for Istrian and Mediterranean Studies, Series historia naturalis, 14 (2): 237–258. Horváth G. (1897). Ordo Hemiptera. — In: Fauna Regni Hungariae III. Arthropoda (Insecta Hemiptera). Regia Societas Scientiarium Naturalium Hungarica, Budapest: 5–64 (in Hungarian and Latin) Horváth G. (1916). Albania Hemiptera-Faunaja. (Fauna Hemipterorum Albaniae). — Annales Musei Nationalis Hungarici, 14: 1–16. Josifov M. (1969). Artenzusammensetzung und Verbreitung der Insekten von der Ordnung Heteroptera in Bulgarien, Teil II. — Bulletin de’l Institut de Zoologie et Musée, 16: 83–150. (In Bulgarian, German Summary) Péricart J. (1972). Hémipterès Anthocoridae, Cimicidae, Microphysidae de l’Ouest – Paléarctique. — Faune de l’Europe et du Bassin Méditerranéen, 7. Masson & Cie éditeurs, Paris, 1–402. Péricart J. (1996). Family Microphysidae Dohrn, 1859 – little pirate bugs, minute bugs. — In: Aukema B. & C. Rieger (Eds) Catalogue of the Heteroptera of the Palaearctic Region. The Netherlands Entomological Society, Amsterdam, Vol. 2: 79–84. Popov Y.A. (2004). New microphysids (Heteroptera: Cimicomorpha, Microphysidae) from Baltic amber and taxonomy of this family. — Prace Muzeum Ziemi, 47: 97–107. Simov N. & M. Josifov (2004). Contribution to the Bulgarian fauna of Heteroptera. — Historia naturalis bulgarica, 16: 89–94.
Grozeva & N. Simov (Eds) 2008 Xenicocephalus - an enigmaticS.genus of American Enicocephalidae (Heteroptera): a new male-based... 357 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 357-376. © Pensoft Publishers Sofia–Moscow
Xenicocephalus - an enigmatic genus of American Enicocephalidae (Heteroptera): a new male-based species from Suriname* P. Štys1 & P. Baňař2 1
Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44 Praha 2, Czech Republic. E-mail:
[email protected] 2 Moravian Museum, Department of Entomology, Hviezdoslavova 29a, Brno, CZ-627 00 and Forestry and Game Management Research Institute, Department of Forest Protection, Jíloviště – Strnady, Praha 5 – Zbraslav, CZ-156 04, Czech Republic. E-mail:
[email protected]
ABSTRACT Wygodzinsky & Schmidt (1991) established Xenicocephalus giganticus, nov. gen., nov. sp. (Hemiptera: Heteroptera: Enicocephalomorpha: Enicocephalidae: Enicocephalinae) from Colombia - the species was based on a single, incomplete female; for description of the genus also the larvae were employed. We describe Xenicocephalus josifovi nov. sp. based on a male from Suriname, provide new diagnosis of the genus and discuss the relationship and autapomorphies of the genus. The description of X. josifovi covers also male genitalia and internal reproductive organs, and the ducts and reservoirs of persisting dorsoabdominal gland. The classification of Xenicocephalus within the Enicocephalinae is confirmed. A new type of raptorial forelegs, unique among the Enicocephalomorpha and Euheteroptera and suggesting oligophagy on a special prey, is characteristic of Xenicocephalus. Also the construction of reservoirs of a dorsoabdominal gland is unique – the ducts leading from a single orifice on mediotergite 4 are double and enter two large, differently shaped and exceedingly sclerotized reservoirs within the abdominal segment 6. Keywords: Hemiptera: Heteroptera: Enicocephalidae, taxonomy, Xenicocephalus - redescription, X. josifovi nov. sp. (Suriname), morphology: forelegs, male genitalia, dorsoabdominal gland. * This paper is dedicated to the 80th birthday of Misha Josifov, an eminent student of many groups of Heteroptera, their cataloguer and keen collector, but first of all good and generous man and forever remaining friend.
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INTRODUCTION Wygodzinsky & Schmidt (1991) = W&S described a new genus and species of Enicocephalinae: Enicocephalini, Xenicocephalus giganticus Wygodzinsky & Schmidt 1991, from Colombia (“Magdalena”*: Sevilla; the holotype in AMNH, New York, examined by P.Š. in 1990), on basis of an incomplete female adult (head and prothorax with their appendages missing).The larvae of Xenicocephalus were available from several C and S American countries (Colombia: Santa Marta and Cundinamarca; Panama; Costa Rica; Guayana: MazaruniPotaro District); their conspecificity with X. giganticus could not be demonstrated, and hence the larvae were excluded by W&S from the type series. However, some characters of the female L5 from Santa Marta have been used for completion of the generic diagnosis because the last instar larvae and adults of the enicocephalids share usually (always?) characters of the head and the armature of forelegs (W&S; Baa, Štys, pers. observ.). W&S described and illustrated the above larva, and provided figures of some characters also for an early larval instar from Panama (probably a female owing to its strikingly incrassate forelegs - PŠ). There has been no additional information on Xenicocephalus since its original description. Štys (2002) could easily and correctly fit the genus in the key of the Enicocephalomorpha of the World but some doubts about its taxonomic position have persisted because of our ignorance of its critical features (architecture of head, pronotum, forelegs and their armature, male terminalia). Moreover, the general facies of Xenicocephalus resembles (“superficially” - W&S) that of the little known monotypic Oriental genus Megenicocephalus Usinger, 1945 (Enicocephalidae: Megenicocephalinae). A new species of Xenicocephalus from Suriname, based on a complete adult male, is described here - its description is focused on those features that could not have been studied by W&S. An extended diagnosis of the genus is provided, and its morphology and relationships are discussed.
MATERIAL AND METHODS Only a card-mounted holotype was available for study. Glue used for its mounting could not have been dissolved in water (even after boiling), and after relaxing the specimen and dissecting parts of it, the glue has covered the objects by a continuous, transparent film obscuring the details. Consequently, we could not study those body parts, which were most heavily affected (the pterothorax except for mesoscutellum; midlegs, hindwings, and the basis of abdomen). The arrangement of some internal organs is well visible through the semitransparent abdominal cuticle of the holotype, dry-mounted some 40 years ago, and only relaxed and *
The Colombian town Sevilla (4.16 N, 75.58 W) is actually situated in the Department Valle del Cauca.
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observed in alcohol. We have used the opportunity for brief descriptions of the reservoirs of the persisting dorsoabdominal larval gland, and male gonads. Comparison of Xenicocephalus nov. sp. to X. giganticus is tentative since part of the differences may be species-specific or stage-specific, others may reflect sexual dimorphism, and some may be idiosyncratic and of no diagnostic value. To facilitate the comparison, we have used the same nomenclature of forewing veins and cells as W&S, except for that on clavus. All measurements are given in millimetres. The following abbreviation are used: FW - forewing(s), L - length, max - maximum, min - minimum, W - width; Cx coxa, F - femur, Ta - tarsus, Ti - tibia, Tr - trochanter, subscripts 1,2,3 - concerning first, second, third pair of legs, respectively (e.g., Tr1 - trochanter of the foreleg). Genus Xenicocephalus WYGODZINSKY & SCHMIDT, 1991 Xenicocephalus Wygodzinsky & Schmidt, 1991: 37 (keyed), 200-205 (O.D.) Xenicocephalus: Štys (2002): 350 (listed), 354 (keyed)
Type species: Xenicocephalus giganticus Wygodzinsky & Schmidt, 1991 by original designation Distribution: Southern Central America and northern South America (W&S) Diagnosis: Adults large, broad and flat, at least 7.9 mm long. Most of the cuticle covered by black granules; no true setigerous tubercles, trichobothria-like setae and strong bristles. Head very short (shorter than pronotum), the preocular part short, antennifers close to eyes, transverse constriction well marked, posterior lobe transverse. Labium thick and short. Antennal segments 2 and 3 terete, 4 subfusiform. Pronotum of three distinct lobes, much wider than long; midlobe with a Y-shaped median not interrupting its posterior margin but with a complex, depressed basis, and with 2+2 lateral depressions; posterior margin ample, strikingly broad. Foretrochanter with a ventral crest. Ventral faces of incrassate forefemur and foretibia concave, sharply delimited and lacking any vestiture;. apicitibial armature of 7 spiniform setae (♂; ♀ ?) directed outwards. Foretarsal armature of 1+1 proximal spiniform setae and 1 distal spiniform seta + 1 distal simple seta; anterior claw stout, posterior reduced in a stump. Mid- and hindtibiae proximally bent, with 2 apical combs associated with a diverse number of spiniform setae each; tarsi two-segmented with two claws each. Macropterous; venation of forewing complete, both basal and discal cells present and closed, the former shorter only by one fifth. Abdomen membranous (exc. distal segments and terminalia), both dorsal and ventral laterotergites individualized; connexival edge sharp. Male opening of dorsal gland associated with two large and strongly sclerotized reservoirs situated within the abdominal segment 6. Pygophore small, cup-shaped, subtriangular, parameres bi-partite, guide elongate, subtriangular, pointed; no remnants of other genital elements present; tergum 10 fused with but distinct from the pygophoral bridge. Female ventrite 8 produced in a short tab. Forewing lobes of larva 5 contiguous.
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Differential diagnosis: The peculiar construction of forefemur and foretibia with ventral faces sharply delimited, concave and devoid of vestiture as well as presence of black granules all over body are autapomorphies of Xenicocephalus; they are not shared with any known genus of the Enicocephalomorpha.
Xenicocephalus josifovi, nov. sp. (Figs 1-33)
Etymology: josifovi - a patronym derived from the surname of the Bulgarian heteropterist, Michail Josifov. Type material.Holotype: ♂, “Suriname Onverdacht \ Para Distr. 07.07.1962.\ 5°36’N-55°09’W. \ on light, mercury \ PHvDoesburg ♂2000” [printed] “RMNH \ 12774” [printed]; card mounted, left foreleg and right hindleg mounted on separate card. Measurements. Total body L – 7.9, W (max.) – 2.36. Head. Total L (without neck) - 1.08 Anterior lobe L – 0.68; distance of eye to apex of antennifer – 0.22; distance of eye to basis of antennifer - 0.04; eye L – 0.28; preocular W (min) - 0.42; diatone (max W across eyes) – 0.66; min interocular distance, dorsal – 0.35; min interocular distance, ventral – 0.15. Posterior lobe: L – 0.40, W – 0.63. Labium. Total L – 0.79; segment 1 - 0.09; segment 2 L – 0.20; segment 3 – 0.31; segment 4 - 0.19. Antenna. Segment 1 L – 0.35; segment 2 L – 0.71; segment 3 L – 0.66; segment 4 L – 0.64. Pronotum. Total L (max) – 1.44; collum: L (median) – 0.22, W (max) – 0.75; midlobe: L (max) – 0.55, W (max) – 1.33; hindlobe: L (max) – 0.64, L (median) – 0.36, W (max) – 2.22. Forewing. L – 5.35, W (max) – 1.86. Foreleg. F1: L – 1.33, W (max) – 0.44; Ti1: L – 1.15, W (max) – 0.26; Ta1: L – 0.28, W (max) – 0.15; anterior claw L (basis – apex) – 0.19; posterior claw L (basis – apex) – reduced. Midleg. F2: L – 1.26, W (max) – 0.24; Ti2: L – 1.15, W (max) – 0.15; Ta2: L (without claw) – 0.35, W (max) – 0.13. Hindleg. F3: L – 1.55, W (max) – 0.29; Ti3: L – 1.37, W (max) – 0.15; Ta3: L (without claw) – 0.51, W (max) – 0.20. General shape. Macropterous, strikingly large and broad, flat, nearly parallel-sided. Head unusually small, contrasting with ample pronotum, and long and broad forewings and abdomen. Body 3.35 times as longer as wide; proportions of body parts, head L (without neck) : pronotum L (max) : FW = 1 : 1.3 : 5. Coloration. Nearly monochromatic, non-contrastingly coloured, without distinct pattern. Groundcolour pale yellow-brown; FW olive-brown to blackish brown; labium yellow; venter yellow to orange; legs stramineous. Mesoscutum laterally to parapsidal sulci contrastingly whitish (in the area normally covered by pronotum in the encicocephalines). Texture. Cuticle basically matt, somewhat lustrous on the pronotum, mesoscutellum and FW; usually densely covered by small, wart-like, subsphaerical black granules (as described and illustrated for X. giganticus by W&S); the granules scarce on FW and abdomen, missing on antennae, labium, whitish parts of mesothorax (present in the area between the parapsidal sulci), some parts of forelegs, and all the mid- and hindlegs. Most of the cuticle of
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legs not smooth but covered by “sand-grain granules” visible only under high magnification and resembling rows of loaf-shaped tiles. Some areas (e.g. F1, Ti1, F3) also with small and thin but rather conspicuous conical tubercles. Setigerous tubercles absent. Vestiture. The whole surface of body and its appendages covered by “soft”, golden, usually straight, semierect to erect pile of thin golden macrotrichia, never arising from setigerous tubercles, never containing spiniform, trichobothrium-like, thickened or otherwise outstanding setae* (excepting specialized setae on legs). Body, antennae and legs very densely covered by “soft”, rather long, thin, setae. Antennae, head and pronotum with rather long, semierect macrotrichia forming a radiating fringe along lateral margins of the posterior lobe of head and collum, and a unidirectional fringe along lateral and posterolateral margins of pronotum (dorsal view); the excised medial sector of posterior margin of pronotum bare. Venter of head with uniformly long, erect, curved, nearly unidirectional hairs, twice that long as the pile on the dorsal surface of head. Dorsal side of labium with short macrotrichia, ventral sides with mixed short and long, and straight and curved setae, some of them on segments 3 and 4 outstanding. Eyes densely covered by short macrotrichia. Macrotrichia on lateral margins of mesoscutellum longer than elsewhere, curved and ruffled. Lateral and ventral sides of thorax with rather long, sparse, curved, uniform pubescence. Ventral side and margins of the abdomen with uniform, dense, rather short pile, with no outstanding long setae (not even on margins of posterior segments); only those on sides of the pygophore longer and denser. Dorsum of abdomen with very short, straight, rather sparse filiform setae. Forewings with four types of macrotrichia: (a) Long, straight, erect to semierect setae distributed densely all over the veins and on those sclerotized parts formed originally by veins (in proximal two thirds of FW). (b) Short, erect, black (sic!) setae distributed among the long ones. (c) Short, curved macrotrichia, usually in two rows, on or along margins of distal sectors of veins, semierect to appressed. (d) Short, multiple-rowed, curved macrotrichia along the anterior and apical margins of FW and on hypocostal lamina; these setae very dense basally; the same type of macrotrichia occurring also on the proximal part of the anterior margin of hindwing, both on its dorsal and ventral side. Legs. (For the specialized rows of tubercles and spines along F1 and Ti1 and the apicitibial and tarsal armatures see under Fore leg.) Coxae and trochanters without particulars; femora with very dense pile of diagonal macrotrichia all over (excepting ventral face of forefemora), densest and longest on F1; long macrotrichia mixed with uniformly long short ones**. Ti1 with diagonal to erect (particularly distally) macrotrichia all over, except the bare area in the proximal part of depression of its posterior face, and specialized, silvery appressed setae in the distal part of this depression, and except bare area on its ventral (adfemoral) face. Ti2 and Ti3 with sparse, erect macrotrichia on dorsal face, denser and diagonal on ventral face. *
W&S obviously had in mind that strong or otherwise outstanding setae are missing while stating that the “macrotrichia” in Xenicocephalus are absent. ** Short macrotrichia were called “microtrichia” by W&S.
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Abdomen. Dorsum bare; venter with uniform, short, appressed hairs, slightly longer and suberect on its medial, convex part. Connexival edge with a dense pilosity, very short on proximal segments, gradually longer towards the apex of abdomen, with no outstanding setae. Head (Figs 1-2, 10) strikingly short. Preocular part abbreviated owing to extremely short genae; the latter slightly diverging in dorsal view; antennifers conspicuous, strongly diverging, reaching nearly the level of the outermost margin of eyes. Dorsum of preocular lobe convex, but provided with an inconspicuous, flat, wedge-shaped elevation, tapering and disappearing between the eyes. Eyes large, facets separately convex, dorsal ocular index 4.26, ventral ocular index 2.59; in lateral view, the eyes by far not reaching the level of dorsum of head, but strongly exceeding its ventral outline. Postocular constriction sharp, long and deep. Postocular lobe strikingly transverse, lateral margins regularly rounded; its anterodorsal surface convex but mediodorsally broadly and shallowly concave (no median); its posterodorsal surface nearly flat, gradually declivous towards the neck; ocelli large, set widely apart, interocellar distance much larger than distance eye-ocellus. Antennae (Fig. 1). Segments 1-3 terete, their widths gradually decreasing distad, 4 subfusiform. Antennal formula (the longest segment first) 2,3,4,1. Labium (Fig. 2) short and thick, nearly reaching the posterior margin of eyes. Segment 1 broad and abbreviated, directed anteroventrad; 2 short, widening distad, directed ventrad, its upper margin straight; 3 rather long, its dorsal margin straight, ventral moderately convex; 4 short, conical. Labial formula 3, 2 = 4, 1. Pronotum tripartite (Figs 1, 10), wider than long (ratio L, max : W, max 0.65), its lobes of a strikingly different width, constrictions between lobes linear and deep. Collum with a distinct precollum; the collar median marked by a shallow groove; lateral sides of collum with a transverse sulcus delimiting a subventral, posteriorly directed, broad, hook-shaped tubercle. Midlobe lateral margins simply rounded, posterior margin convex, not interrupted. The midlobe median proceeding from the anterior midlobe margin as an inverted Y-shaped impression, with the stem (in first third of midlobe) linear, sharply delimited, and the branches thick and vaguely delimited, embracing a large, shallow, medial, subcircular depression (in second third of midlobe). Proximal part of this depression with an anchor like, dark (hypersclerotized?) anchor-shaped structure provided with a short, thick, tongue-shaped stem. Posterior third of midlobe with no median structure but with 1+1 submedial, longitudinal, shallow depressions arising from the anchor-shaped structure. Lateral areas of dorsum with 1+1 extensive, indistinctly delimited depressions distant from lateral margins*; the deepest anterolateral part of the depression provided with a whitish macula devoid of black tubercles. Similar, less well delimited whitish maculae (formed largely by arrangement of black tubercles and lack of them) occurring also mesad of and posterad to the depressions. Hindlobe strikingly ample, lateral and posterolateral margins broadly and continuously rounded; posterola*
These isolated and strictly dorsal depressions are unlike sharply delimited 1+1 Y-shaped impressions or 1+1 pits often connected with lateral margins in many other enicocephalines.
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Fig. 1: Xenicocephalus josifovi nov. sp., male holotype, head and pronotum, dorsal view. Fig. 2: Xenicocephalus josifovi nov. sp., male holotype, head, lateral view. Fig. 3: Xenicocephalus josifovi nov. sp., male holotype, forewings. Fig. 4: Xenicocephalus josifovi nov. sp., male holotype, left foreleg, posterior view. Fig. 5: Xenicocephalus josifovi nov. sp., male holotype, left foretrochanter, anterior view.
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Fig. 6: Xenicocephalus josifovi nov. sp., male holotype, right hindleg, anterior view. Fig. 7: Xenicocephalus josifovi nov. sp., male holotype, abdomen, dorsal view. Fig. 8: Xenicocephalus josifovi nov. sp., male holotype, abdomen, ventral view. Fig. 9: Xenicocephalus josifovi nov. sp., male holotype, abdomen, dorsal view, detail of dorsoabdominal gland.
teral angle indistinct, broadly obtuse; posteromedial margin broadly and deeply excised, convex, only the very midpoint of the excision concave. Median marked by a linear keel extending to the posterior margin, and forming there a concave triangle with a concave basis. “Proepimeral lobe” broad, short, visible in lateral view only; the upper part of its posterior margin concave, the lower one (ventral) broadly rounded, the lobe(s) fusing with the posterior prosupracoxale and externally delimited from the latter by a shallow depression only. Mesoscutellum ample, triangular, mucronate, the mucro long, broad, parallel-sided, apically rounded. Lateral parts of prothorax (as seen in lateral view). Collum subdivided by a horizontal sulcus (situated at the level of notopleural impression - see below) in dorsal and lateral parts; the dorsal one formed by a ventrally narrowing, elongate, apically rounded elevation (provided with black granules) surrounded by flat precollar and post-elevation parts (both lacking the granules); ventral part of collum split by a sulcus in equally long precollar and collar parts. Sulcus between collum and midlobe considerably widening ventrad. Lateral parts of pronotal midlobe widely reflected ventrad, and the true propleuron
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restricted to a broad and deep notopleural impression delimiting the supracoxalia from the notum. Anterior part of notopleural sulcus with a deep propleural pit resembling a spiracle. Anterior supracoxale moderately large, subrectangular, delimited from the posterior supracoxale by pleural sulcus not extending onto the notopleural impression; posterior supracoxale ample, widening posteroventrad and fused with the “proepimeral lobe”. Proacetabula free posteriorly, not enclosed by the “proepimeral lobes”. Prosternum: eusternum simple, triangular; sternellum extensive, forming mesal parts of proacetabula, apex subtriangular, produced but apically rounded, its median with a broad, shallow groove. Foreleg (Figs 4-5, 11-13, 16-20). Coxa. All the faces covered with sand-grain granules. Trochanter (Figs 4-6, 16-18) bearing on distal two thirds of its posteroventral edge a prominent ridge (Figs 5, 20), resembling a rough mountain crest increasing distad, apically rounded (in anterior view, of two equally long lobes) and much exceeding the basis of femur both dorsad and distad. In strictly ventral view, two strikingly distinct, chocolate-coloured sinuate lines (sharply contrasting with the stramineous background) are delimiting the ridge and terminate on a pair of equally coloured distalmost tubercles each. Femur (Figs 4, 11, 19, 23-24) incrassate, nearly uniformly thick in all its length, ratio L : max W 3.02; distinctly curved, moderately C-shaped; all the ventral face* concave, with absolutely no vestiture, parallel-sided and sharply delimited at both anterior and posterior edges by a row of macrotrichia and irregularly distributed black granules intermixed with a row of conspicuous, high, non-setigerous conical tubercles (Fig. 24). Basal third of anterior face of forefemur nearly bare, with a few short setae, without sculpture. All the other faces (exc. the ventral concavity) covered with both black and sand-grain granules. The concave face densely tiled by small, broad, transverse, scale-shaped structures (Fig. 23). Neopatella** formed by two distinct slerites: the proximal one, crescent-shaped, articulating with the distidorsal concave posterior margin of femur and with the distal sclerite, the latter shaped as a broadly open and rounded V, and providing contact with the small, basidorsal process of tibia. Tibia (Figs 4, 11). Nearly straight, cylindrical, of uniform width, only the dorsal outline slightly curved; ratio L to max W 4.4. All the ventral face moderately concave, with no vestiture, the edges of the tibial concavity less sharply delimited than those of the femoral one. Anterior edge with 14, posterior edge with numerous conical tubercles of the same shape as on the femur. Anterior face approximately with 50 black granules, posterior face with several hundred of them. Sculpture of ventral concavity as in the femur. All the faces of foretibia covered with sand-grain granules. Cleaning comb very long, consisting of 4042 setae. Apicitibial process moderately long, only slightly protruding, with no pointed projection. Apicitibial armature (Fig. 12) of 7 straight, spiniform setae: four in ventral *
The architecture of the ventral side of forefemur described in the following text is unique among all the Enicocephalomorpha. See Discussion. ** For definition and discussion of neopatella (forming a “knee” between femur and tibia) see ŠTYS & BAŇAŘ (2007).
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Fig. 10: Xenicocephalus josifovi nov. sp., male holotype, head and pronotum, dorsal view. Fig. 11: Xenicocephalus josifovi nov. sp., male holotype, left foreleg, anterior view. Fig. 12: Xenicocephalus josifovi nov. sp., male holotype, left foreleg, apicitibial armature, posterior view. Fig. 13: Xenicocephalus josifovi nov. sp., male holotype, left foreleg, tarsal armature, posterior view. Fig. 14: Xenicocephalus josifovi nov. sp., male holotype, right hindtibia, anterior apicitibial comb. Fig. 15: Xenicocephalus josifovi nov. sp., male holotype, right hindtibia, posterior apicitibial comb.
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row, three in dorsal row. Spiniform setae situated (and well visible) exclusively on posterior face and directed outwards (instead of inwards as in the other enicocephalines). Tarsus one-segmented, cylindrical, narrowing distad. Tarsal armature (Fig. 13) of 1+1 proximal thin, spiniform setae, 1 short and thin distal seta (anterior) and 1 distal spiniform seta (posterior). Claws: the anterior one stout and regularly curved, the posterior one reduced in an inconspicuous tubercle. Specialized sense organs on foreleg. Basal rim of forecoxa anteromesally with coxal rim organ, consisting of cluster of 7-8 badly visible, diversely directed, straight setae. Condylar trochanteral organ of six short setae (Fig. 16). Anterior trochanteral organ (Fig. 17) of 4+2 campaniform sensilla (4 in a row and 2 isolated); posterior trochanteral organ (Fig. 18 ) of 5+1 campaniform sensilla (5 in straight row and 1 isolated). Anterior femoral organ (Fig. 19) of 5 nearly basal campaniform sensilla (4 in straight row and 1 isolated ); posterior femoral organ absent. Midlegs short and slender, not studied in detail. Femur with conical tubercles on both dorsal and ventral faces, femur and tibia covered by sand-grain granules; no black granules. Tibia strongly curved in basal third, sulcate; apex with two combs. Tarsus 2-segmented, basitarsus very short, nearly without dorsal surface; claws short and thin, moderately curved, the posterior one stouter. Hindlegs (Figs 6, 14-15, 21-22) relatively long and slender. Anterior face of hindtrochanter with trochanteral organ, consisting of 5+1 campaniform sensilla (Fig. 21), one large isolated campaniform sensillum and two strong, prominent setae. Posterior face of hindtrochanter with trochanteral organ, consisting of 6+1 campaniform sensilla (Fig. 22). Femur with conical tubercles on the dorsal and ventral faces. All faces of femora and tibiae covered with small sand-grain granules. Tibia curved in basal third (Fig. 6), sulcate. Anterior apicitibial comb on hindtibia formed by 19 setae on ventral face and two strong, longer spiniform setae (Fig. 14), posterior comb formed by 15 setae on posterior face and 4 strong, spiniform ventral setae (one short, three longer, subequal in length) (Fig. 15); the spiniform setae not incorporated in combs but situated behind them and more ventrad. Tarsus and claws as on the midleg. Forewings (Figs 3, 25-26) 2.87 times as long as wide (max), reaching the apex of abdomen. Anterobasal transverse anchor-like vein well developed; anteradial furrow distinct. Details of venation and shape as described and illustrated for X. giganticus by W&S, with the following exceptions (situation in X. giganticus in parantheses): - right FW in upper position (x left FW); - costal margin much more concave in basal third; - diffusion of the sclerotized material from proximal sectors of remigial veins, veins along costal margin, and particularly the veins on clavus, onto surrounding areas of the wing membrane, the meeting point of AA1+2 and AA3+4 thus forming nearly a plate (x not such a diffusion); - proportions of veins delimiting the basis of basal cell or associated with the latter different (cf. Figs 25-26 x W&S, Fig. 148A): - r-m a distinct crossvein (x nearly a point);
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Fig. 16: Xenicocephalus josifovi nov. sp., male holotype, foretrochanter, condylar trochanteral organ. Fig. 17: Xenicocephalus josifovi nov. sp., male holotype, foretrochanter, anterior trochanteral organ. Fig. 18: Xenicocephalus josifovi nov. sp., male holotype, foretrochanter, posterior trochanteral organ. Fig. 19: Xenicocephalus josifovi nov. sp., male holotype, forefemur, anterior femoral organ. Fig. 20: Xenicocephalus josifovi nov. sp., male holotype, foretrochanter, detail of trochanteral process. Fig. 21: Xenicocephalus josifovi nov. sp., male holotype, hindtrochanter, anterior trochanteral organ. Fig. 22: Xenicocephalus josifovi nov. sp., male holotype, hindtrochanter, posterior trochanteral organ. Fig. 23: Xenicocephalus josifovi nov. sp., male holotype, forefemur, detail of femoral concavity. Fig. 24: Xenicocephalus josifovi nov. sp., male holotype, forefemur, detail of anteroventral face.
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- claval AP branching from AA3+4 distinct (AP not illustrated by W&S); - vein connecting discal cell with wing margin extremely short; - FW covering the abdomen, the anterior margins of wings and lateral margins of abdomen coinciding (x abdomen broader and longer than FW - a female character ?). Hindwings not examined. Abdomen (Figs 7-9, 27-33) oval, its lateral margins diverging up to intersegmental line 5-6, then converging distad. All the eight pregenital segments and pygophore exposed, little sclerotized (except for parts of the terminalia). Pregenital dorsum. Mediotergites 1 and 2 fused, delimited by a transverse apodeme, lateral parts amalgamated. Dorsal laterotergite (1&2) triangular, delimited from mediotergite1&2 by anterior longitudinal incisions coinciding with more posterior longitudinal sulci. Mediotergites 3-8 nearly fused with corresponding dorsal laterotergites, no distinct intratergal sulci present, but the original areas clearly separated by longitudinal impressions becoming more pronounced distad and separating dorsal laterotergites (provided with sclerites) from mediotergites (provided with lateral, paired, sclerotized maculae marking the insertions of dorsoventral muscles). Sclerites of dorsal laterotergites 3-5 vaguely delimited, those of 6 and 7 strongly sclerotized and spreading across laterotergal boundaries onto lateral mediotergites; dorsum 8 fully sclerotized; the sclerotized areas always bearing the black cuticular granules. Posterior margin of the more anterior segment (from 1&2 up to 7) always slightly overlapping the anterior margin of the following segment. Posterior margin of dorsum 8 broadly and deeply subrectangularly excised, posterolateral angles of the segment prominent, rounded, subrectangular. Pygophore situated within the excision of segment 8, not at all telescoped. Anterior part of mediotergite 4 with a simple, minute, ellipsoid, crevice-like, intrasegmental opening of a dorsoabdominal gland (Figs 27-28, 30); cuticle surrounding the opening unmodified. Duct of the gland (Fig. 28) thin and non-sclerotized in segment 4; double and wide in segment 5, becoming fully sclerotized in distal half of this segment. In segment 6, the thick left duct (Fig. 28) entering a strongly sclerotized proximal part of a horizontally situated left bulbous reservoir (Fig. 28) whose distal flat, elongate, diagonal, flask-shaped part, longer than segment 6, is occupying the whole median region of the latter. A short, most proximal part of the left bulbus differentiated from the distal one. The narrow right duct (Fig 28) percurrent along mediotergite 6, then forming a loop, turning anterad, and at the anterior margin of mediotergite 6 entering a differentiated proximal part of vertically situated, elongate, right plate-shaped reservoir (Figs 30-31), visible in ventral and lateral views only. In ventral view, the right reservoir stretching from distal fifth of segment 5 up to proximal fifth of segment 7. Connexival edge sharp, strikingly distinct on segments 1-8. Abdomen - pregenital venter. (Ventrite 1 with sharply delimited and sclerotized triangular ventral laterotergites, and a sclerotized medial part; the latter damaged and, consequently, presence or absence of opening of basiabdominal scent glands not ascertained. All further observations concern an untreated, not cleared specimen.) Each of the ventrites 2-7 subdivided in a medial convex region and lateral flat parts; the latter
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Fig. 25: Xenicocephalus giganticus, female, venation associated with the discal cell of forewing (modified from Wygodzinsky & Schmidt, 1991: Fig. 148A), highly schematised. BC - basal cell, DC - discal cell. Fig. 26: Xenicocephalus josifovi nov. sp., male holotype, venation associated with the discal cell of forewing, highly schematised. BC - basal cell, DC - discal cell. Figs 27-29: Xenicocephalus josifovi nov. sp., male holotype, distal part of abdomen and internal organs (dorso-abdominal gland and its reservoirs and reproductive organs as seen through the abdominal cuticle) in dorsal view; coiling of ducts of the resrvoir is not illustrated and laterotergal sclerites are omitted. The right reservoir is visible as a shade only. 27. Total view. 28. Left reservoir; scheme. 29. Terminalia and reproductive organs. Lettering: py - pygophore, t - testis, vd - vas deferens. Fig. 30: Xenicocephalus josifovi nov. sp., male holotype, distal part of abdomen with the right reservoir of dorso-abdominal gland in ventral view (the left reservoir not visible), laterotergal sclerites omitted. Lettering: py - pygophore (part of ).
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subdivided again in lateral, sclerite-bearing regions and non-sclerotized submedial regions. The sclerite-bearing regions corresponding to ventral laterotergites and provided with black granules; the sclerites indistinctly delimited on segments 2-5, strongly sclerotized and sharply delimited on 6-7. However, no intrasegmental sulci present, except for those delimiting medial part of ventrite 2. Posterior margins of segments 1-7 slightly overlapping anterior margins of the followings segments. Segment 8 fully sclerotized, subdivided only in lateral flat (laterotergal) and medial convex parts; posterior margin bisinuate, formed by three moderately convex lobes, the medial one (functioning as a subgenital plate) about as long but twice as wide as the lateral ones. Abdominal spiracles 2-8 minute, situated just mesad to laterotergal sclerites, situated in about first fifths of segment width. (Spiracle 1 not found.) Terminalia (Figs 29, 32-33). Pygophore fully exposed on dorsum (only basal tenth covered by segment 8), ventrally covered by subgenital plate 8, apparently largely immobile. The visible part (dorsal view) cup-shaped, subtriangular, apically rounded, dorsal bridge (= tergum 9) complete, posterior foramen situated at the dorsal and posterodorsal side. Dorsal area of the foramen with a proximal, arcuate sclerotized strip turning lateroventrad and disappearing there within the cavity of the pygophore; the strip representing tergum 10, distinct from but probably immovably fused with anterior margin of dorsal foramen. A membraneous segment 11 filling up the mediodorsal part of the foramen, while its mediolateral parts occupied by paired, bipartite, immobile and non-articulating, two-dimensional parameral sclerites formed by proximal plate (covered by tergum10 just under its lateral apodemes and easily mistaken for them) and thinner distal rod. In dorsal view, only the thin sclerotized frame of the distal acutangular, apically rounded part of the guide is visible. The guide fully visible in posterior view: shaped as a narrow, slightly convex, well sclerotized triangular frame, with apex pointing dorso-anteriorly; the solid base hardly sclerotized, fused with posteroventral wall of the pygophore, its basilateral margins produced into apodemes within the wall.The space between the guide and the membrane containing the parameres filled up by everted, coiled, thin intromittent organ. Paired, spherical testes (Fig. 29) situated in distal 3/5 of segment 7; thick vasa defferentia retaining paired condition till the proximal margin of the dorsal foramen of pygophore, being dilated in segment 8 in seminal vesicles. Differential diagnosis. Xenicocephalus josifovi nov. sp. (based on a male) and X. giganticus (based on an incomplete female) can be distinguished by some features in the venation of forewing, which are not sex-linked and can hardly be explained by individual variation (schemes in Figs 25-26). The most striking are the following (condition in X. giganticus in parentheses - see W&S, Fig. 148A): - r-m normal cross-vein, as long as cu-an and longer than Cu1a, emanating from apex of discal cell (r-m extremely short, forming a contact point of discal cell with R only, shorter than any other cross-vein and than Cu1a); - apex of discal cell nearly reaching the margin of wing, Cu1a very short and rather indistinct (apex of discal cell distant from wing margin, Cu1a moderately long and distinct);
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- proportions of sectors of longitudinal veins forming the proximal part of discal cell and the associated crossveins strikingly different in both species. Only the examination of further material of both sexes might test our above characters and reveal further potential differences. The type localities of both species are nearly 2200 km apart and are situated in biotically different Neotropical provinces (Andean and Venezuelan, respectively) - this supports our conviction that two species are involved.
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Fig. 31: Xenicocephalus josifovi nov. sp., male holotype, right reservoir in dextral, ventro-lateral view; scheme. Figs 32-33: Xenicocephalus josifovi nov. sp., male holotype. 32. Pygophore as observed in situ in the posterior foramen of abdominal segment 8, dorsal view. Only the distinct structures are shown (membranous and sclerotized structures of pygophore not clearly distinguishable as well as margins of its posterior foramen). Shapes of minute parameres and guide distorted owing to angle of observation. 33. Genitalia in situ; basically dorsal view (slightly more posterior than Fig. 32 - compare the different shape of guide and distance guide - paramere); tentative scheme, open to reinterpretation. The guide, in truly posterior view, has the same shape as in Fig. 33, only the sclerotized basal part between its arms is distinct. Lettering: 8LT - dorsal laterotergite; 8MT - mediotergite; 8V - dorsal surface of produced part of ventrite 8; 10T - tergum 10; 11 - membraneous, seemingly amorphous valves of segment 11; a - two sublinear apodemes (strikingly black) of unknown homology, precise position and function, situated within the lumen of pygophore; g - guide (in nearly dorsal view); or - orifice representing probably a secondary gonopore through which the coiled ejaculatory duct (?), situated within the pygophore and providing for a tertiary gonopore, may be everted; pb - basal part of a paramere (covered by 10T, and coinciding with its lateral apodeme); pd - distal part of a paramere; py - pygophore (segment 9).
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DISCUSSION 1. Discrepancies between original description of Xenicocephalus and X. giganticus by WYGODZINSKY & SCHMIDT (1991) and our findings in X. josifovi The important discrepancies concern (a) the construction of forefemur and foretibia (studied by W & S in ♀ larva only), (b) number of spines in fore apicitibial armature (ditto) - 3-5 in X. giganticus, 7 in X. josifovi, (c) composition of the armature of foretarsus (ditto), (d) presence of apicitibial projection in X. giganticus (ditto) and its absence in X. josifovi, (e) length and width of abdomen relative to forewings (adult ♀ studied by W&S) - broader and longer in X. giganticus, equally long in X. josifovi, (f ) pattern formed by sclerites on dorsum and venter of abdomen (ditto), (g) architecture of apicitibial comb on hindleg (ditto). Taking into account the unusual accuracy of both Wygodzinsky´s and Schmidt´s studies and particularly their illustrations, we suggest the following tentative assessment of the above discrepancies. The characters (a), (b), (c) and (d) are possibly larval characters different from adult situation; in this case the statement by W&S “the basic features of the raptorial forelegs in enicocephalids are identical in larva 5 and adults” (modified by PŠ) have no universal validity. Alternatively, these characters may be female characters different from those of adult male. Minor cases of sexual dimorphism in these characters were recorded by W&S in several genera, the only striking case being the Enicocephalinae: Alienatinae, an extremely derived group with winged males and apterous females. However, such variation in number of fore apicitibial spines as recored in Xenicocephalus (3-7) is unique and open to doubt. The characters (c) and (d) may, of course, be also species-specific. There is no doubt that the characters (e) and (f ) are sex-linked. We suspect that the ambiguous statement by W&S (p. 201) on architecture of the hind apicitibial combs (not accompanied by an illustration) - “Spines even with arc (i.e. neither within arc, nor outside of it) formed by bristle combs” - might have been caused by erroneous observation: a sex-linked variation of this character is unknown in enicocephalids. Description of “scent gland auricle” associated with mesothoracic FW grooves of X. giganticus as provided by W&S (p. 201) refers actually to a wing-holding device characteristic of the Enicocephalomorpha (Štys 1998).
2. Relationship of Xenicocephalus The general facies of Xenicocephalus - flat, broad, robust, short-headed - is unique among the American genera of the Enicocephalomorpha and among Enicocephalidae of the World. Only the Oriental genus Megenicocephalus Usinger, 1945 (Megenicocephalinae) with so far single species M. chinai Usinger, 1945 from the continental Malaysia is superficially similar by its robust stature, incrassate and curved forefemora and foretibia,
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and crested foretrochanter. However, all the elements of the similarity are homoplasic, and Megenicocephalus differs strikingly by its universal aposematism, ambulatory forelegs, the three lobes of pronotum indistinctly delimited, presence of incipient costal fracture on forewings, etc. These genera do not share any synapomorphies and cannot belong to the same clade. On the other hand, despite the below reviewed strange autapomorphies, Xenicocephalus clearly belong to the Enicocephalidae: Enicocephalini as assumed by W& S, and as suggested by architecture of pronotum, forewing structure and venation, character of fore apicitibal armature, male pygophore provided with guide and immobile parameres but no other major genital structures, and distinct abdominal tergum 10.
3. Autapomorphies of Xenicocephalus Only a few characters are briefly mentioned since most of them will be studied in future in a broader comparative context. 3.1. Black granules These cuticular, uniformly black structures are probably unique in the Enicocephalidae. They are definitely not melanization centres, and we must admit a complete ignorance on their biochemical composition and potential function. It seems significant that their unique presence seems to be associated with an equally unique absence of differentiation among the macrotrichia, except for those forming specialized apicitibial and tarsal organs. 3.2. Crested fore trochanter The crested or tuberculate or otherwise modified fore trochanter (and/or forecoxa) is diagnostic for the Enicocephalidae: Phallopiratinae and Megenicocephalinae, and it occurs mosaic-like also in the Enicocephalinae (e.g., Xenicocephalus, Systelloderes loebli Štys & Baa, 2007 from New Caledonia). We have already briefly discussed its potential function (Štys & Baa 2007). 3.3. Apicitibial armature of foreleg The spiniform or spatulate setae (“platellets”) of the apicitibial armature are normally directed inwards, towards the foretarsus with its own armature and towards the claw(s) curved towards the tibial process. When stretched, the foreleg armature is well suited for boring into and thus grasping a soft prey (e.g. chrysomelid larvae; P.Š., pers. observ. on an Australian Oncylocotis sp.) or holding a small prey (e.g., a collembolan, P.B., pers.observ. on a Madagascan Euchelichir sp.). In Xenicocephalus, the spines are directed outwards, can hardly cooperate with the tarsal armature and claws, and a different mode of holding the prey is suggested. As far as we know the situation is unique among the Enicocephalidae.
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3.4. Bare and concave ventral faces on forefemur and foretibia The architecture of forefemora and foretibiae in Xenicocephalus has been sufficiently explained and illustrated in the descriptive part; it is unique in all the Heteroptera. Femur and tibia seems to form a raptorial organ (see sub 3.3) serving for holding a rounded, potentially strongly sclerotized prey. We can only suggest that Xenicocephalus must be trophically specialized and catches the prey by a unique method. 3.5. Apicitibial armature of hindleg In most enicocephalids, the apicitibial combs on mid-and hindtibiae are formed by thin macrotrichia (a notable ecception being Brevidorsus arizonensis Kritsky, 1977), and the strong, spiniform setae participate in formation of the comb are diversely situated, but nearly always potentially functioning as comb guarding setae. The notable exceptions are Xenicocephalus and Hymenocoris Uhler, 1892, possibly also Urnacephala Wygodzinsky & Schmidt, 1991, in which the setae are situated between the comb and the basitarsus, resembling thus rather a situation on forelegs. Also the high number of spiniform setae in Xenicocephalus is unusual. For a review of American genera see W&S: Fig. 14 and SEM photographs within the monograph; there is no review available for the Old World genera. 3.6. Male terminalia The occurrence of immobile structures representing the parameres which lost the articulation and were transformed in generally flat sclerites is a synapomorphy (?) of most Enicocephalinae (probably paraphyletic) and all the Alienatinae and Megenicocephalinae. The bipartite condition found in Xenicocephalus josifovi is highly unusual and requires a new morphological interpretaion. 3.7. Male dorsoabdominal gland All the enicocephalomorphans possess a single opening of the persisting dorsoabdominal gland onto the abdominal mediotergite 4. Very few data on the gland itself and potential sexual differences in size and degree of persistence are available to allow any generalizations. However, in all the heteropterans examined, in which the gland persists (including a few genera of the Enicocephalinae), the gland(s) is situated direct beneath the opening(s). The described situation in Xenicocephalus josifovi, with two large, sclerotized and asymmetrical reservoirs situated far more caudad than the external orifice, is unique. One of us (PŠ) can only recall an existence of a similar, large, plate-shaped and strongly sclerotized reservoir in Maoristolus sp. (Aenictopecheidae: Maoristolinae; New Zealand), found during dissection but not sufficiently appreciated and not studied in detail. We included this situation among autapomorphies of Xenicocephalus, but our opinion is open to reinterpretation.
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ACKNOWLEDGEMENTS The project was funded by the Grant Agency of the Academy of Sciences of Czech Republic (IAA 60110706), which also supported senior author´s trip to Leiden (The Netherlands) where the holotype of the new species was discovered in the collections of the Naturalis Museum. For a substantial assistance while working there I am grateful to my long-standing friend Pieter van Doesburg (Leiden) and to some suggestions to Dominique Pluot-Sigwalt (Paris).
РЕЗЮМЕ WYGODZINSKY & SCHMIDT (1991) описват Xenicocephalus giganticus, nov. gen., nov. sp. (Hemiptera: Heteroptera: Enicocephalomorpha: Enicocephalidae: Enicocephalinae) от Колумбия – видът е описан на базата на една, не запазена изцяло женска; за описанието на рода са използвани и ларви. Ние описваме Xenicocephalus josifovi nov. sp. на базата на мъжки екземпляр от Суринам, с нова диагноза на рода и обсъждане на родствените връзки и апоморфиите на рода. Описанието на X. josifovi включва също мъжките гениталии и вътрешните репродуктивни органи, дуктуси и резервоари на дорзоабдоминалната жлеза. Потвърдена е класификацията на Xenicocephalus в подсем. Enicocephalinae. Нов, уникален сред Enicocephalomorpha и Euheteroptera, тип хватателни предни крака е характерен за род Xenicocephalus, който предполага олигофагия със специфична жертва. Конструкцията на резервоарите на дорзоабдоминалната жлеза е уникална - дуктусите започват от един единствен отвор на четвърти медиотергит, двойни са и се отварят в два големи, различни по форма, и изключително склеротизирали резервоари в шести коремен сегмент.
REFERENCES Štys P. 1998. Evolutionary origin of squamiform microsculpture on the forewing-holding devices (frenae) in Heteroptera. — European Journal of Entomology, 95:307-310. Štys P. 2002. Key to the genus-group taxa of the extant Enicocephalomorpha of the World, their list, and taxonomic changes (Heteroptera). — Acta Universitatis Carolinae Biologica, 45 (2001): 339-368. Štys P. & P. Baa 2007. First species of Systelloderes (Hemiptera: Heteroptera: Enicocephalidae) from New Caledonia. — Acta Entomologica Musei Nationalis Pragae, 47: 3-15. Usinger R.L. 1945. Classification of the Enicocephalidae (Hemiptera, Reduvioidea). — Annals of the Entomological Society of America, 38: 321-342. Wygodzinsky P.W. & K. Schmidt 1991. Revision of the New World Enicocephalomorpha (Heteroptera). — Bulletin of the American Museum of Natural History, 200: 1-265.
S. Grozeva & N.1895) Simov (Eds) 2008 The Recognition of “Botocudo” ornatulus (Bergroth, of Australia, the Designation of the Lectotype... 377 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 377-383. © Pensoft Publishers Sofia–Moscow
The Recognition of “Botocudo” ornatulus (BERGROTH, 1895) (Hemiptera, Prosorrhyncha, Rhyparochromidae, Antillocorini) of Australia, the Designation of the Lectotype, and a Redescription of the Type Specimen, with a Discussion of the Nomenclature of Botocudo KIRKALDY * M.H. Sweet Department of Biology, Texas A&M University, College Station, Texas, 77807, USA
ABSTRACT The type specimen of Cligenes ornatulus Bergroth was discovered in the Victoria Museum, Melbourne, Australia. The specimen is designated as the Lectotype, and is redescribed to allow recognition of the species. Its nomenclatural history as Botocudo ornatulus (Bergroth) is discussed, and evidence given that ornatulus is not a member of the Neotropical genus Botocudo Kirkaldy and is only nomenclaturally placed in Botocudo pending further studies on the Old World species assigned to Botocudo. Keywords: Australia, Botocudo ornatulus, Hemiptera, Lectotype, Lygaeoidea, Nomenclature
INTRODUCTION The quotation marks around the generic name Botocudo need explanation. Bergroth (1895) in his review of the genus Cligenes Distant 1893, placed 13 species in the genus, *
I am pleased to dedicate this paper to Dr. Michail Josifov in honor of his research on the Heteroptera.
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describing four new species: Cligenes minutus, C. noualhieri, C. picticollis, and Cligenes ornatulus, the last being the only species described from Australia. Bergroth (1895) noted that the genus Salacia Stål, 1874 was preoccupied and proposed that Distant’s genus Cligenes receive the species described in Salacia, that is, Salacia diluticornis (Stål, 1858), Salacia delineata Distant, 1893, Salacia picturata Distant, 1893, Salacia pilosulus Stål, 1874, Salacia punctatus Distant, 1893, and Salacia pusio (Stål, 1858). He also placed Salacia sericeus Lethierry, 1881 and Tropistethus flavicornis Signoret, 1880, in Cligenes Distant. Kirkaldy (1904) also discovered that Salacia was preoccupied and proposed the new name Botocudo for the species described in Salacia that Bergroth had placed in Cligenes Distant. An exception was Salacia signandus Distant, 1903 that Distant (1904) moved to Cligenes. Both Distant (1910) and Bergroth (1913) listed Botocudo as a synonym of Cligenes Distant, although Bergroth (1913) considered Stål’s (1874) division a. of Salacia to be Cligenes Distant s. str. and Salacia division aa. to be Antillocoris Kirkaldy, 1904, itself a replacement name for Pygaeus Uhler, 1894, which in turn was listed as a synonym of Cligenes (Slater, 1964). Bergroth (1913) considered Botocudo Kirkaldy, 1904 to be a synonym of Antillocoris Kirkaldy, 1904, in which he placed most species of Cligenes s. l. of the Old World, probably because of line priority in Kirkaldy’s (1904) listing of replacement names for preoccupied genera. Other species were described later in Cligenes (see Slater, 1964). Several species were later moved by Barber (1952) from Cligenes to Antillocoris (see Slater, 1964), and Barber thus established the present usage of Antillocoris. Scudder (1962) fixed Pygaeus pallidus Uhler as the type species of Pygaeus Uhler, thus of Antillocoris Kirkaldy. Scudder (1967) rightly restricted Cligenes to only Cligenes distinctus Distant, 1893, which species he fixed (Scudder, 1962) as the type species of Cligenes Distant. Distant (1893) was dubious about the species other than distinctus that he placed in Cligenes as he put marks by these placements in Cligenes. Scudder (1967) next moved to the synonym Botocudo, as the next available name, all 22 other species in Cligenes listed in Slater (1964). This greatly enlarged the concept of Botocudo to include virtually any small generalized antillocorine, thus the genus placement of ornatulus Bergroth is uncertain, especially as the type species of Salacia Stål, 1874 is Aphanus diluticornis Stål, 1858 from Rio de Janeiro, Brazil (Stål, 1858), which was fixed by Scudder (1962) as the type species of Botocudo. Slater and Brailovsky (1994) made much of the complex concept of Botocudo as a cosmopolitan genus, but actually, it is what was left after Cligenes (s. str.) was removed. While vigorous revisionary work has been done on the New World Antillocorini by Slater et al. (1977); Slater (1980); Baranowski & Slater (1987, 2005); Brambila (2000a, 2000b); (Slater & Brailovsky (1994, 2004); Baranowski & Brambila (2001); and O’Donnell (2004), little has been done on the Old World Botocudo, so I am reluctant to describe a new genus for ornatulus in isolation from the complex Old World Antillocorini fauna. Moreover, Brambila (in e-mail) is revising Botocudo sensu stricto as a New World genus and intends to work on the Old World fauna of “Botocudo.” Slater (1976) in his discussion of the lygaeid fauna of SW Australia, noted that while Bergroth’s description fits the specimens of the species he collected in SW Australia,
The Recognition of “Botocudo” ornatulus (Bergroth, 1895) of Australia, the Designation of the Lectotype... 379
the identity of Botocudo ornatulus was uncertain as Woodward (in litt.) had said to Slater that there are several species of Antillocorini in Australia, which I found also in my field work in Australia. The present contribution is therefore to establish the identity of the species Botocudo ornatulus (Bergroth), which hopefully will promote the badly needed revisional work on the Old World species presently remaining in Botocudo as well as the Antillocorini fauna in Australia and the Old World Tropics. In comparing the description of Botocudo diluticornis (Stål) by Slater & Brailovsky (1994) with Botocudo ornatulus, it is clear to me that Botocudo ornatulus is not congeneric with B. diluticornis. For example, B. ornatulus has four fore femoral spines and a straight apical corial margin while B. diluticornis has mutic femora and a concave apical corial margin. Nor is B. ornatulus congeneric with Cligenes distinctus of Distant as it lacks the deep prosternal groove. The description of a new genus here, however, would, I believe, be premature and should be made in context of a revision of the Antillocorini and “Botocudo” of the Old World and with additional correctly identified specimens of B. ornatulus to permit dissection of these tiny insects to ascertain their abdominal and genitalic structure. I therefore here leave ornatulus in “Botocudo” pending ongoing work on Botocudo s.l. being done by Brambila (in e-mail). To show the problem awaiting study, at the time of the Slater and O’Donnell catalogue (1995) 36 species were included in Botocudo, of which 29 were Old World in contrast to Bergroth’s concept of Cligenes when it was synonymized with Botocudo, in which 11 of the 13 species were New World. Brambila (2000b) added two more New World species to Botocudo s.str. Thus the quotation marks around the name Botocudo, to emphasize the need for work on the Old World “Botocudo” fauna. “Botocudo” was the only Antillocorini described from Australia until Slater (1983) transferred Lethaeaster from the Targaremini to the Antillocorini and listed specimens of L. anthocoroides Breddin, the type species of Lethaeaster from Northern Queensland. Also Tomocoris punctatus Woodward, which has many records from Queensland, was transferred by Slater & Woodward (1982) from the Targaremini to the Lilliputocorini as Lilliputocoris punctatus (Woodward). However, Tomocoris punctatus may turn out to be an Antillocorini when the nymphs are discovered, as it superficially resembles many Antillocorini.
Botocudo ornatulus (BERGROTH, 1895) Because Bergroth (1895) in his description of Cligenes ornatulus mentioned Mus. Melbourne, I, during fieldwork in Australia, visited the Museum Victoria in Melbourne where I was graciously allowed to search through their insect holdings. (I was also looking for Getes fusciceps Bergroth, 1916 from Tasmania, the description of which was published in the Proceedings of the Royal Society of Victoria, which was headquartered in Melbourne. I mention this in hope that someone may locate Bergroth’s type specimen of Getes fusciceps. I did not find it at the Museum Victoria). To my pleased surprise, I found in the Victoria Museum a single macropterous male specimen with a label in
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Bergroth’s distinctive handwriting, Cligenes ornatulus Bergr. Nat. Mus. Victoria. Also on the pin is a red circle with the numeral 124 and another label saying: From C. French, Jun 15.11.11. This corresponds precisely with the data in Bergroth’s paper where after his description of Cligenes ornatulus, he wrote: Australia (Queensland) –– Mus. Melbourne, Coll. French. Bergroth may have received the data about Queensland from elsewhere, perhaps from French himself. Probably based on this information, Cassis & Gross (1995) localized Botocudo ornatulus to the central coast of Queensland, Australia. However, I recently found in my field work that there were two evidently “Botocudo” species under figs in the city of Brisbane, Queensland, of which the larger species, I think, is Botocudo ornatulus itself, but careful comparison is needed. I also found two species of “Botocudo” in Adelaide, South Australia, one smaller, the other larger under figs. Because in his description of Cligenes ornatulus, Bergroth mentions a macropterous form and a brachypterous form, as well as a range of length (2.5-2.6 mill. [mm]) and there is no type label on the specimen, I therefore cannot unambiguously recognize this specimen as the Holotype; I therefore here designate this specimen as the Lectotype of Cligenes ornatulus Bergroth. The description by Bergroth (1895) is in Latin and brief. I here redescribe the Lectotype and include additional available morphological and mensural data to aid in the recognition of this species from the other Australian species of Antillocorini. As there is only one specimen, the Lectotype, available, it could not be dissected or cleared. I hope data can be added about the genitalia and abdominal characteristics in cleared specimens accurately identified as Botocudo ornatulus (Bergroth).
Redescription of Lectotype (all measurements in mm) Macropterous male. Oval in general form. Coloration: head and anterior pronotal lobe yellowish tan, paler yellow on anterior margin of pronotum and across the transverse depression of pronotum; longitudinal dark line in middle of pronotum; posterior lobe of pronotum, abdominal and thoracic sternum, metapleuron, and scutellum dark rusty brown, except for pale scutellar apex, pale posterior margin of mesopleuron and pale metapleural flange; corium and clavus whitish with three brown blotches on corium, one on posterior apex, on medial costal margin along lateral margin, and on interior angle; membrane smoky brown; antennae with antennal segments 1 and 4 light brown, 2 and 3 darker brown; legs light brown entirely. Investiture and surface texture: dorsum of head and thorax nude, glabrous; head trichobothria long, length 0.195; venter of abdomen finely sericeous with short adpressed hairs; posterior lobe of pronotum moderately punctate, slight row of punctures along midline, and along the anterior margin of pronotum; scutellum uniformly finely punctate; clavus with three rows of fine punctures, about 30 along claval suture; corium translucent with three weak longitudinal rows of fine punctures, each puncture darker
The Recognition of “Botocudo” ornatulus (Bergroth, 1895) of Australia, the Designation of the Lectotype... 381
than light corial surface; pleura nude, except for row of 9 punctures in groove delimiting metapleural flange; fine punctures in area above the scent gland, and along the propleuron above the acetabulum. Morphology: head subtriangular, transverse, moderately declivent, length 0.24, width across eyes 0.49; eyes large, interocular distance 0.29; ocelli large, ocellus width 0.049; bucculae low; no deep ventral rostral groove under head or pronotum. Pronotum not elevated, bluntly carinate laterally, at base a little sinuate laterally, and lightly depressed transversely to set off posterior lobe that is distinguished more by coloration and stronger punctation; pronotum length 0.54; width 1.49. Scutellum flat, with no carina, length 0.61, width 0.61. Metathoracic scent gland auricle sessile, narrow, curving moderately posteriorad; evaporatorium covers ventral one-third of metapleuron; metapleural height 0.37. Hemelytron exceeds apex of abdomen; hemelytral epipleura set off by low carina from hemelytron, distal margin of corium straight, not concave; veins weak in hemelytron, m-fracture long, curving mesad, more than half length of corium; commissure much shorter than scutellum; length 0.17; length of hemelytron 0.95; distance from apex of corium to apex of membrane 0.54. Abdominal sterna four and five fused, suture between sterna curving strongly anteriorad to nearly reach the lateral suture between sterna three and four. Spiracles and trichobothria invisible as specimen is glued on card. Antennae moderately thick, segment one a little clavate, segments 2 and 3 fusiform, antennal segment one exceeding apex of head by distance from antenniferous tubercles to apex of tylus; antennal segment lengths I: 0.29, II: 0.67, III: 0.20, IV missing. [Bergroth (1895) says the fourth segment is a little longer.] Fore femora incrassate, with four sharp spines in a row; labium and thoracic sterna invisible under specimen glued on card. First metatarsal segment as long as 2 and 3 together, lengths: I: 0.17, II: 0.05, III: 0.12. Total body length 2.56. Brachypter was not available to study. Bergroth (1895) said that in the brachypter the pronotum is a little narrower, the hemelytron attains the base of the fifth abdominal segment, and the membrane is very short.
ACKNOWLEDGEMENTS I thank the staff at the Museum Victoria for their gracious assistance in locating the specimen of Cligenes ornatulus Bergroth. I thank Ms Julieta Brambila and Dr. Jane O’Donnell for reviewing the manuscript and offering many valuable suggestions.
РЕЗЮМЕ Типовият екземпляр на Cligenes ornatulus BERGROTH е открит в Музея Виктория, Meлбърн, Aвстралия. Екземплярът е обозначен като лектотип, и е преописан, за да позволи признаването на вида. Номенклатурната история на пробема с Botocudo ornatulus
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(BERGROTH) е обсъдена и са представени доказателства, че видът ornatulus не се отнася към неотропичния род Botocudo KIRKALDY и само номенклатурно е поставен в този род, докато се водят изследвания на видовете от род Botocudo от Стария свят.
REFERENCES Barber H. G. 1952. The genus Antillocoris Kirkaldy in the United States (HemipteraLygaeidae). –– Bulletin of the Brooklyn Entomological Society, 47: 85–87. Baranowski R.M. & J. Brambila (2001). Caymanis, a new genus of Antillocorini from the Cayman Islands (Hemiptera: Lygaeidae). –– Florida Entomologist, 84: 117–322. Baranowski R.M. & J.A. Slater (1987). New genera and species of Antillocorini from Trinidad and Brazil (Hemiptera; Lygaeidae). –– Florida Entomologist, 70: 381–391. Baranowski R.M.& J.A. Slater (2005). The Lygaeidae of the West Indies. –– Florida Agricultural Experiment Station Bulletin 403, Gainesville, Florida: x + 1-266. Bergroth E. (1895). Le genre Cligenes Dist. (Lygaeidae). –– Revue d’Entomologie, 14: 142-145. Bergroth E. (1913). On some Heteroptera from the Philippine Islands. –– Annales de la Société Entomologique de Belgique, 57: 150–154. Brambila J. (2000a). A review of Cligenes with the description of a new genus, Valeris (Hemiptera: Rhyparochromidae: Antillocorini). –– Florida Entomologist, 83: (303–315. Brambila J. (2000b). Two new species of Botocudo from Trinidad, Tobago, and Grenada (Hemiptera: Rhyparochromidae: Antillocorini). –– Insecta Mundi, 14: 187–192. Cassis G. & G.F. Gross (1995). Australian Biological Resources Study. Zoological Catalogue of Australia. 27. 3B Hemiptera: Heteroptera (Pentatomomorpha). –– Commonwealth Scientific and Industrial Research Organisation (CSIRO) Publishing, Canberra, Australia: 1–524. Distant W.L. (1893). Biologia Centrali–Americana. Insecta. Rhynchota. Hemiptera–Heteroptera. Vol. I. Supplement. –– Taylor & Francis, London: 378–462. Distant W.L. (1904). The fauna of British India including Ceylon and Burma. Rhynchota. Volume II. (Heteroptera). –– Taylor & Francis London: 1–503. Distant W.L. (1910). The fauna of British India including Ceylon and Burma. Rhynchota. Volume V. Heteroptera: Appendix. –– Taylor & Francis, London: xii + 1–362. Kirkaldy G.W. (1904). Bibliographical and nomenclatorial notes on the Hemiptera. N. 3. — Entomologist, 37: 279–283. O’Donnell J. E. (2004). Notes on the phylogenetic relationships of Neotropical Antillocorini, with descriptions of three new species (Heteroptera: Rhyparochromidae: Rhyparochrominae). –– Journal of the New York Entomological Society, 108: 114–129. Scudder G.G.E. (1962). The World Rhyparochrominae (Hemiptera: Lygaeidae) 1.New synonymy and generic changes. –– Canadian Entomologist, 94: 764–773. Scudder G.G.E. (1967). Rhyparochrominae types in the British Museum (Natural History) (Hemiptera: Lygaeidae). –– Bulletin British Museum (Natural History) Entomology, 20: 253–285. Slater J.A. (1964). A catalogue of the Lygaeidae of the World. Volume 1 & 2. –– University of Connecticut, Storrs, Connecticut: 1–1668.
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Slater J.A. (1976). The biology, distribution and taxonomy of some Lygaeidae of Southwest Australia (Hemiptera: Heteroptera). –– Journal of the Australian Entomological Society, 15: 129–151. Slater J.A. (1980). Systematic relationships of the Antillocorini of the Western Hemisphere (Hemiptera: Lygaeidae). –– Systematic Entomology, 5: 199–226. Slater J.A. (1983). On the systematic position of Lethaeaster with descriptions of a new genus and four new species of Antillocorini (Hemiptera: Lygaeidae). –– International Journal of Entomology, 25: 285-296. Slater J.A. & H. Brailovsky (1994). The type species of the genus Botocudo with description of a new species from Mexico and second record of Paurocoris from the United States (Hemiptera: Lygaeidae). –– Journal of the Kansas Entomological Society, 67: 1016. Slater J. A. & H. Brailovsky (2004). The rectification of the type species of Acolhua Distant with the description of a new species (Heteroptera: Rhyparochromidae), –– Journal of the New York Entomological Society, 111: 207-210. Slater J. A. & J. E. O’Donnell (1995). A catalogue of the Lygaeidae of the World (1960–1994). –– The New York Entomological Society, New York, New York: xv + 1–410. Slater J.A., M.H. Sweet, and R. M. Baranowski (1977). The systematics and biology of the genus Bathydema Uhler (Hemiptera: Lygaeidae). –– Annals of the Entomological Society of America, 70: 343–358. Slater J.A. & T.E. Woodward (1982). Lilliputocorini, a new tribe with six new species of Lilliputocoris, and a cladistic analysis of the Rhyparochrominae (Hemiptera, Lygaeidae).–– American Museum Novitates, 2754: 1–24. Stål C. (1858). Bidrag till Rio Janeiro –Traktens Hemiptera–fauna. –– Kongliga Svenska Vetenskaps-Akademien Handlingar, 2(7): 1-84. Stål C. (1874). Enumeratio Hemipterorum, Part 4. –– Kongliga Svenska Vetenskaps-Akademien Handlingar, 12(1): 1–186.
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Grozeva Simov 2008 Saldidae) from Xizang, China Calacanthia josifovi nov. sp.,S. a new species&ofN. shore bugs(Eds) (Heteroptera,
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ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 385-389. © Pensoft Publishers Sofia–Moscow
Calacanthia josifovi nov. sp., a new species of shore bugs (Heteroptera, Saldidae) from Xizang, China N.N. Vinokurov Institute for Biological Problems of Cryolithozone, Siberian Branch of the Russian, Academy of Sciences, 677980 Yakutsk, Russia. E-mail:
[email protected]
ABSTRACT Calacanthia josifovi nov. sp. is described from Xizang, Tibet, China. Keywords: Heteroptera, Saldidae, shore bugs, China, Xizang, Tibet
INTRODUCTION Six species are known in the Holarctic genus Calacanthia Reuter, 1891. Of them, two very closely related species C. trybomi ( J. Sahlberg, 1878) and C. alpicola ( J. Sahlberg, 1880) are widespread along the coast of the Arctic Ocean, the latter is sometimes considered a variety (Kiritshenko 1951) or a synonym of C. trybomi (Kerzhner. & Jaczewski 1964). Other species (C. angulosa (Kiritshenko, 1912), C. grandis Cobben, 1985, C. sichuanica Chen & Zheng, 1987 and C. tibetana Drake, 1954) are characteristic only for the Chinese fauna (Lindskog 1995). An additional species collected in Xizang (Tibet) at the height of 4400 m is described below. The type material of the new species (including the holotype) is deposited in the collection of Professor E. Heiss (Innsbruck, Austria), and part of paratypes, in the collection of the Zoological Institute of the Russian Academy of Sciences (St. Petersburg, Russia).
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Calacanthia josifovi nov. sp. Body wide, obovate, twice as long as wide (Fig. 1). Head and hemelytra dull; pronotum and scutellum moderately shining. Pubescence of dorsum golden-brown, that of ventral part white and dense. Head black; vertex and frons flat, scabrous. Eyes large, brown, 1.3-1.6 times as long as width of frons (Figs 2, 3). Ocelli separated by 1/3 their diameters or sometimes almost contacting. Preocellar spots small, ovate or like skewed transverse stripe, connected to the wide light spot at the lower edge of eye through narrow light stripe. The latter spot is located in the area near the lower edge of eye in females (Fig. 2) and extended down onto mandibular plate in male (Fig. 3). There are three pairs of long trichobothria: at the base of postclypeus, at the middle of frons and on vertex. Mandibular plate, transversal crest and maxillary plate yellow or often (in female) reddish. Anteclypeus yellow; basal part sometimes with a dark spot; lateral lobes dark in female and usually light in male. Postclypeus black. Labrum yellow. Rostrum reaching middle coxae, shining, black with first segment yellow. Antennae long. Segment 1 black in outer and ventral side, rather thickened, with sparse erect black bristles; inner side flattened, in male yellow, in female often reddish. Segment 2 thin, with short erect black hairs, in the middle with 2-3 long erect black hairs; external latero-ventral side black, internal latero-dorsal side yellow-brown; inner side of segment flattened in male; in female, segment cylindrical. Segments 3 and 4 subequal in length, black, with short semi-erect black hairs. Pronotum trapezoid, in male 2.7-3 times, in female 2.3-3.4 times as wide as long. As in other Tibetan Calacanthia, anterolateral edges of pronotum provided with light calloused bulges bearing erect white hairs. Lateral margins straight or slightly convex, with one row of short erect black hairs, narrowly yellow in light specimens; the yellow edging frequently reduced to short light stripe near anterolateral edges of pronotum. Callus convex, wide, not reaching lateral margins of pronotum; median pit of callus transverse, deep. Thorax shining, black. Coxal covers of fore and middle legs and narrow light stripe on outer side of metathorax whitish yellow. Coxae black; trochanter yellow. Femora from Fig. 1: Calacanthia josifovi, nov.sp., female. below and on hind side black, from above
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yellow-brown, with small, sometimes merging black spots. Fore and middle femora with small, often fused black spots. Hind femora above with long narrow black stripe. Tibiae yellow-brown, with long black setae, near base with black spot, at apex narrowly black. Middle tibiae sometimes with fused black-brown spots in the middle. Hind tibiae with small dark spots at bases of setae. Tarsal segments 1 and 3 brown; segment 2 brownyellow with dark brown base. Scutellum flat, dimly shining on external edge and at base, mat and rough in the middle, with thin transverse wrinkles at apex. Semi-brachypterous specimens, hemelytra, hardly covering apex of abdomen, dull, coal-black (Figs 8-12). Hind wings rudimentary. Hypocostal plate well developed, without secondary ridge (Fig. 7). Clavus with a large yellow spot in apical part and 1 or
2
6 5 4
3
7
12 8
9
10
11
Figs 2-12: C. josifovi: 2,3 – frontal view of head (2 – female, 3 – male); 4-6 – antennae: 4 – C. angulosa, 5, 6 – C. sichuanica; 7-12 – left forewing (7 – underside with hypocostal lamina, 8-12 – pigment variation of wings).
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2 small yellow spots at base. Centre of clavus in light colored females sometimes with a minute yellow spot. Light area on hemelytra well developed in females (Figs 8-10), but in males hemelytra black with few light spots, mainly on exocorium and membrane (Figs 11, 12). Male genital structures as in Figs 13-15. Parandria as depicted (Fig. 13). Paramere with obtuse outgrowth laterally (Fig. 14); dorsum covered with long and dense dark hairs; processus hamatus thickened; processus sensualis weakly noticeable, with wisp of long hairs (Fig. 15). Subgenital plate of female on the outer side bordered with yellow stripe, which is narrow at apex and widened laterally (Fig. 16). Ovipositor often projecting from abdomen. Teeth of blade of gonapophysis I prominent, hook-like and sharp (Fig. 17). Apex of gonapophysis II as in Fig. 18; spermatheca almost 2.5 times as long as its base, slightly inclined and behind the middle narrowed (Fig. 19). Measurements (in mm) - ♂ (♀): body length 4.1-4.4 (5.1-5.5); body width 2.1-2.2 (2.5-2.7); head width 1.14-1.19 (1.29-1.34); vertex width across ocelli 0.41-0.46 (0.460.54); frons at narrowest width 0.29-0.47 (0.36-0.39); length of antennal segments 1-4: 0.40-0.47 (0.40-0.43), 0.71-0.80 (0.83-0.87), 0.43-0.57 (0.51-0.60), 0.50-0.57 (0.540.62); length of pronotum 0.53-0.64 (0.51-0.70). Material examined. China, Xizang (East Tibet). Holotype (♂): [“Chola Shan” pass, road Yanjing – Markam, 50 km S of Markam, ca 4400 m, 29º16´N 98º38´E, mixed forest,
16
19
18 14
17
15
13
Figs 13-18: C. josifovi: 13 – parandria, 14, 15 – paramere in various positions, 16 – subgenital plate, 17 – blade of gonapophysis I, 18 – gonapophysis II, 19 – spermateca.
Calacanthia josifovi nov. sp., a new species of shore bugs (Heteroptera, Saldidae) from Xizang, China
389
24–27.vi–2.vii.1997 (M. Trýzna et O. Šafránek lgt)]. Paratypes: [“Chola Shan” pass road Yanjing – Markam, 50 km S of Markam, ca 4400 m, 29º16´N 98º38´E, mixed forest, 24–27.vi–2.vii.1997 (M. Trýzna et O. Šafránek lgt)], 4 ♂, 2 ♀; [“Tuntala Shan” pass, road Markam – Zogang, ca 5000 m, 29º43´N 98º02´E, alpine meadow, 20.vi–2.vii.1997 (M. Trýzna et O. Šafránek lgt)], 1 ♂, 1 ♀; [“Tamala Shan” pass, road Qamdo – Toba, 20 km NE Qamdo, 31º16´N 97º18´E, 16.vii.1997 (M. Trýzna et O. Šafránek lgt)], 2 ♂, 3 ♀. C. josifovi nov. sp. possesses some features distinguishing it from other Tibetan representatives of the genus Calacanthia, first of all, the poorly expressed anterolateral edges of pronotum. It is similar to C. angulosa and C. tibetana in body length and short adpressed hairs of dorsal part, but the second antennal segment in the last two species is thick and flattened (Fig. 4). C. sichuanica also has a thickened second antennal segment (Figs 5, 6), and the head between eyes in addition to short hairs bears coarse, long, erect hairs. C. grandis has long thin antennae as well, but it is the largest species of the genus with a body length of 7.5 mm.
ACKNOWLEDGEMENTS The author is thankful to Prof. E. Heiss (Innsbruck, Austria) for kind providing his collection of Saldidae for study and to Prof. I.M. Kerzhner for advice and correction of the English text. The study was supported by grant from the Russian Foundation for Basic Research (No. 05-04-49917).
РЕЗЮМЕ Calacanthia josifovi nov. sp. описан от Сидзан, Тибет, Китай.
REFERENCES Chen P.P. & L. Zheng (1987). A new species and some new records of shore bug from China. — Acta Zootaxonomica Sinica, 12 (4): 393-397. Kerzhner I.M. & T.L. Jaczewski (1964). Order Hemiptera (Heteroptera). — In: G.Ya. Bei-Bienko (Ed.) Keys to the insects of the European part of the USSR. 1. Nauka, MoskwaLeningrad: 655-845. Kiritshenko A.N. (1951). True bugs of the European part of the USSR (Hemiptera): key and bibliography. Opredeliteli po faune SSSR. 42. — Izdatel’stvo AN SSSR. MoskwaLeningrad: 1-433. Cobben R.H. (1985). Additions to the Eurasian saldid fauna, with a description of fourteen new species (Heteroptera, Saldidae) — Tijdschrift voor Entomologie, 128: 215-270. Lindskog P. (1995). Family Saldidae. — In: Aukema, B. & Rieger, Chr. (Eds). Catalogue of the Heteroptera of the Palaearctic Region, 1. Netherlands Entomological Society: 116-137.
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S. of Grozeva & N. Simov (Eds) 2008 Emesinae) from Madagascar Mangabea barbiger, new species Collartidini (Heteroptera, Reduviidae,
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ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 391-402. © Pensoft Publishers Sofia–Moscow
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar* C. Weirauch Department of Entomology, University of California Riverside, Riverside CA 92521, USA. E-mail:
[email protected]
ABSTRACT A new species of the endemic Malagasy genus Mangabea Villiers (Emesinae-Collartidini), Mangabea barbiger, nov. sp., is described from Eastern-Central Madagascar. A revised diagnosis of Mangabea Villiers is provided. A distribution map for the two species of Mangabea and digital macrophotographs, scanning electron micrographs, and illustrations for the new species are provided that illustrate species and genus level characters. A table with diagnostic characters for the three described recent genera of Collartidini is given. Keywords: Collartidini, Emesinae, Madagascar, Reduviidae, systematics.
INTRODUCTION The fauna of Madagascar currently comprises more than 230 species of Reduviidae or assassin bugs (Maldonado 1990). Judging from currently available catalogs and revisions (Maldonado 1990; Putshkov & Putshkov 1985; Villiers 1968, 1979), the vast majority of these taxa are endemic to the fourth largest island in the world. About 150 of the 230 species were described by André Villiers in a series of papers (e.g., Villiers 1948, 1949a, 1952, 1957, 1971), with the remaining species mostly described in the late 19th and early 20th century. Since Villiers’ descriptive and revisionary activity roughly half a century ago, no new data have been published on the Malagasy reduviid fauna. *
This paper is dedicated to Michail Josifov on the occasion of his 80th birthday.
392 C. Weirauch
During an ongoing project based at the Californian Academy of Sciences, San Francisco, more than 2800 specimens of Reduviidae were collected and sorted to family level in that institution. While further sorting this material to species level in our lab, two male specimens of a Collartidini (Emesinae) were discovered that differ in several characters from the only described Malagasy species in this taxon, Mangabea orientalis Villiers and warrant the description of a new species. Wygodzinsky (1966) created the tribe Collartidini within Emesinae to accommodate the genera Collartida Villiers and Stenoramphus [sic] Elkins. With the description of the monotypic genus Mangabea from Madagascar (Villiers 1970) Collartidini currently comprise three described, extant genera. The description of one new genus from New Caledonia that will considerably extend the know distribution is under way (Villarosa Garcia and Wall, in prep.). Collartidini is currently considered to be the sistergroup of all remaining Emesinae due to several characters that may be plesiomorphic within the subfamily (Wygodzinsky 1966). Villiers (1949b) had initially described Collartida in the tribe Leistarchini (Emesinae), but later moved it to the emesine tribe Ploiariolini (Villiers 1961). Elkins (1962) described Stenoramphus within Saicinae, a taxon that may be the sistergroup of Emesinae (Wygodzinsky 1966). The position of Collartidini within the clade Emesinae-Saicinae is supported by characters such as the lack of ocelli in the adult (Usinger 1943), the absence of dorsal abdominal scent glands (Wygodzinsky 1966; Weirauch 2006), and the membranous hemelytron (Wygodzinsky 1966). Within that clade Collartidini share synapomorphic forward opening acetabula of the fore leg with the remaining Emesinae (Wygodzinsky 1966). The three described, recent genera are identified by a combination of the vestiture of the head, trochanter, and fore femur and fore wing venation (Elkins 1962; Wygodzinsky 1966; Villiers 1970). In addition to describing the second species of Mangabea this contribution aims at illustrating diagnostic features of the genus using digital micrographs and scanning electron photographs and on providing a synopsis of diagnostic characters within the tribe. In addition, this paper is intended as the first step into future revisionary studies on the reduviid fauna of Madagascar.
METHODS, TERMINOLOGY, AND ABBREVIATIONS Habitus photographs and photographs of details of head and thorax were taken on a Microptics-USA photographic system with an Infinity Photo-Optical K-2 lens system and a Canon EOS 1D digital camera. Five to ten images were taken at different focal planes and stacked with Helicon Focus 4.16 software. Scanning electron micrographs (SEMs) of the uncoated paratype were obtained using a Hitachi TM-1000 Tabletop Electron Microscope (UCR, University of California Riverside). Drawings were made with a Nikon SMZ 1500 stereo microscope with drawing tube. White arrow heads in the figures point to characters that are diagnostic on the species level, black arrow heads indicate genus level diagnostic features.
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar
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Map: Localities for M. orientalis (from Villiers 1970) were georeferenced using GEOLocate (Rios & Bart 2005). These coordinates and those provided on the labels of the recently collected material of the new species were mapped using DIVA-GIS software (Hijmans et al. 2007). The satellite map was obtained from www.maplibrary.org. Terminology: For the description of leg and wing structures, the appendages are referred to as being stretched out laterally, not in rest. The three stout setae on the fore coxa that point anteriad when the leg is in resting position are therefore described as being located on the outer surface of the coxa (Fig. 1d).
a
b
c
e
d
f
Fig. 1: Mangabea barbiger, nov. sp., habitus of holotype (a-d) and paratype (e, f ), digital microphotographs. a, lateral view; b, dorsal view; c, ventral view; d, head and thorax, lateral view; e, head and thorax, dorsal view; f, left hemelytron, dorsal view. Scale bars represent 1mm.
394 C. Weirauch
Abbreviations of terms used in the legends: bp, basal plates; end, endosoma of aedeagus; M, medial vein; M+Cu, joint vein representing medial and cubital veins; par, paramere; PCu, post cubital vein; pht, dorsal phallothecal sclerite; prpy, apical, medina process on pygophore; R, radial vein; rm, crossvein between R and M; rmcu, cross vein between R and M+Cu; sp8, spiracle of abdominal segment 8. Institutional abbreviation used: CAS, Californian Academy of Sciences, San Francisco.
TAXONOMY Revised diagnosis of Mangabea Villiers, 1970 Villiers, 1970: 809 [new genus] Type species: Mangabea orientalis Villiers, 1970 Revised diagnosis (Tab. 1): Recognized among extant genera of Collartidini by the moderate total size (7.5-9.1mm) (Fig. 1a), head elongate with postocular reduced to narrow strip behind eyes (Fig. 2b), vestiture on ventral surface of head consisting of one pair anterior and posterior to margin of eye and two pairs or one pair and a fascicle of stout setae ventrolaterally on gena (Fig. 2b, c); second labial segment (first visible) not reaching anterior margin of eye (Fig. 1d); ventral surface of second labial segment with dense vestiture in apical half (Fig. 1c); ventral surface of third segment with dense vestiture (Fig. 2d); posterior pronotal lobe covering metanotum, slightly longer or shorter than anterior lobe (Fig. 1e); fore coxa with three spinelike setae (Fig. 1d); spinelike setae on fore femur extending to apex (Fig. 3a); basal cell roughly pentagonal; discal cell long, more than 3/4 of length between rm and tip of wing; crossvein proximal to rm vein present (rmcu cross vein) (Figs 1f, 4b).
Mangabea barbiger, new species (Figs 1–6, table 2)
Holotype: Male: MADAGASCAR: Fianarantsoa: Parc National Ranomafana, radio tower at forest edge, 21.251°S 47.40716°E, 1130 m, 12 Jun 2005 - 27 Jun 2005, M. Irwin, R. Harin’Hala, 1♂ (UCR_ENT 00005202) (CAS). Diagnosis: Recognized within the genus Mangabea by the fascicle of stout setae on the anterior area of the gena (Figs 2a-c) [two pairs of single setae in M. orientalis], the anterior pronotal lobe not distinctly swollen (Figs 1a, b, d, e) [swollen in M. orientalis], the posterior pronotal lobe slightly longer than the anterior lobe (Figs 1a, d) [anterior lobe slightly longer than posterior in M. orientalis], the anteocular region anterior to the antennifer narrow (Fig. 1e) [broader in M. orientalis], the ventral row of stout setae in the apical half of the fore femur consisting of only few, short single spines (Fig. 3a) [alternating long setae with three or four setae interspersed in M. orientalis]. Also recognized by the slightly bigger body size.
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar
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Description: Male: medium sized (total length, holotype: 8.9 mm), head elongate, eyes globular and very large, thorax relatively short, abdomen elongate, legs slender, wings surpassing the apex of the abdomen; general dull brown colour without conspicuous markings. Vestiture: Body and appendages covered with evenly spaced, short setae with tuberculate bases (Figs 2a, e, f, 3a), longer on male segment 8 and pygophore (Fig. 3f ); ventral surface of head with three pairs of long, stout setae located posterior to antennifer, at anterior and at posterior margins of eye (Figs 2b, c), fascicle of more than ten stout setae on gena ventrad of apex of maxillary plate (Figs 2b, c); second labial segment (first visible) Table 1: Synopsis of diagnostic characters for the three recent genera of Collartidini. Character states are assembled from the literature (Elkins 1962; Villiers 1970) and original observations. Collartida VILLIERS
Mangabea VILLIERS
Stenoramphus ELKINS
Total size
5mm
7.5-9.1mm
8mm
Size of postocular region
always distinct, size virtually absent depending on wing morph; generally smaller in male than in female present present
very distinct (only female known)
present
present
present
one pair of setae
two pairs or setae or fascicle no
one pair of setae and fascicle no
Single pair of setae posterior to margin of eye Single pair of setae anterior to margin of eye Vestiture on anterior area of gena Second labial segment reaching anterior margin of eye Ventral surface of first visible labial segment Ventral surface of second visible labial segment Length of posterior lobe of pronotum
yes
single setae single setae not covering metanotum
present
apical half with dense row of stiff setae vestiture dense vestiture row of stiff setae covering metanotum, 1.5 times longer than anterior lobe not extending to apex
Spinelike setae of fore femur
extending to apex
covering metanotum, about as long as anterior lobe extending to apex
Spinelike setae on fore coxa
two
three
three
Shape of basal cell of hemelytron Length of discal cell of hemelytron
roughly rhombic
roughly pentagonal
roughly triangular
more than 3/4 of length between rm and tip of wing present
about 3/4 of length between rm and tip of wing absent
extending to at most half length between rm and tip of wing Crossvein proximal to rm vein absent
396 C. Weirauch
with fascicle of long and stout setae on ventral surface in apical half of segment (Fig. 2c), third labial segment (second visible) with long setae on entire ventral surface (Fig. 2d), fourth segment (third visible) with medium-length, stout setae on ventral surface in basal half (Fig. 2d); scapus of antenna with moderately dense, very long, slender setae in addition to short, regular setae (Fig. 2a); fore coxa , in addition to short vestiture, with three stout, long setae on outer surface (Figs 1d, 2e; most basal seta broken off, but represented by large socket), fore trochanter with three stout setae on anterior surface (Fig. 3b, two of the setae broken off, but represented by large sockets), ventral surface of fore femur with about 7 long, stout setae in basal 2/3, interspersed with 1-3 short setae, apical 1/3 with three short setae (Fig. 3a), fore tibia with relatively dense vestiture (Fig. 3c). Structure: Head (Figs 1d, e, 2a, b): elongate, anteocular portion long (Fig. 1d, e, 2a, b), postocular represented by narrow strip posterior to eye (Fig. 2b), about two times longer than width across eyes, pronotum about 1.5 times longer than head (Fig. 1d), apex of stout, bulging antennifer at mid point between anterior margin of eyes and apex of clypeus (Fig. 2b), head anterior to antennifer narrow in dorsal view (Fig. 1e), maxillary plate very small, consisting of slightly bulging lobe (Fig. 2b), mandibular plate very small and flat (Fig. 2b); gena with pronounced, elongate anterior portion (Fig. 2b), gula gently curving, clypeus slender, not produced, labrum small, elongate (Fig. 1e, 2b). Eyes: eyes globulose in dorsal perspective, almost touching at midline (Fig. 1e), reniform in lateral view, reaching dorsal surface of head, almost reaching ventral surface of head (Fig. 2b); consisting of relatively few, large ommatidia (Figs 2a, b). Antenna: extremely long, slender; scapus (directed posteriad) reaching metanotal spine (Figs 1a,b), pedicellus about 3/4 length of scapus, basiflagellomere about half the length of pedicellus, distiflagellomere of similar length as Table 2: Measurements of Mangabea barbiger, nov. sp. in mm
Length total
head
Holotype
8.90
1.20
Paratype
9.10
1.40
anteocular ant. pron. post. pron. scutellum lobe lobe 0.73 0.75 0.98 0.28 0.80
0.90
1.00
Labium
0.35
wing 6.00 6.10
Antenna
lab. 2
lab. 3
lab. 4
seg. 1
seg. 2
seg. 3
seg. 4
Holotype
0.68
0.55
0.35
3.60
2.90
1.38
ca. 2.5
Paratype
0.65
0.63
0.45
3.50
na
na
na
Width head Holotype
0.78
Paratype
0.82
ant. pron. post. pron. abdomen lobe lobe 0.95 1.35 1.66 0.98
1.45
1.90
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar
397
pedicellus (curved in the holotype, therefore measurement of 2.5 mm rough approximation). Labium (Figs 1d, 2b-d): second (first visible) labial segment slender, elongate, not reaching anterior margin of eye, third (second visible) labial segment slender and elongate, of about same length as second segment, fourth (third visible) segment slender, about 2/3 length of preceding segments, tapering towards apex. Thorax (Figs 1a, b, d, d, 2e, f ): pronotum longer than wide, unarmed, with anterior area corresponding to proepisternum distinctly lower than disc of anterior lobe, anterior lobe subquadrate with slightly bulging disc divided by distinct median, longitudinal groove, very deep and wide in posterior a
c
e
b
d
f
Fig. 2: Scanning electron micrographs of Mangabea barbiger, nov. sp., paratype. a, head and thorax, lateral view; b, head, lateral view; c, ventral surface of head showing fascicle and second labial segment, lateral view; d, third labial segment, lateral view; e, prothoraxic acetabulum and base of coxa, lateral view; f, meso- and metathorax, lateral view. Scales in micrometers unless indicated otherwise.
398 C. Weirauch
half of disc; anterior and posterior lobes separated by distinct furrow (Fig. 1e); posterior lobe about as long as wide, distinctly wider than anterior lobe, slightly depressed medially and with raised lateral areas in posterior half of lobe, posterior margin slightly concave (Fig. 1e). Scutellum semicircular with slightly pointed tip, raised rim of median spatulate area, and depressed median area (Fig. 1e). Proepisternum longer dorsally than ventrally, separated from proepimeron by distinct pleural suture; supracoxal cavity bulging laterally. Mesepisternum large, mesepimeron narrow. Metepisternum very large, anterior lobe forming part of mesocoxal cavity (Fig. 2f ); metepimeron obsolete (Fig. 2f ). Prosternum with a
b
c
d
e
f
Fig. 3: Scanning electron micrographs of Mangabea barbiger, nov. sp., paratype. a, fore leg, anterior surface, lateral view; b, fore trochanter, anterior surface; c, apex of fore tibia, anterior surface; d, fore tarsus, anterior surface; e, fore pretarsus, oblique, lateral view; f, abdominal segment 8 and pygophore, lateral view. Scales in micrometers unless indicated otherwise.
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar
399
prosternal process elevated, apex pointed (Fig. 2e); Stridulitrum with transverse cuticular ridges present. Meso- and metasternum of similar length, slightly longer than prosternum (Fig. 1c). Legs (Fig. 3): slender, fore leg distinctly stouter and shorter than mid and hind leg, hind leg longer than middle leg (Fig. 1a), tarsi with three, slender tarsomeres, first tarsomere very short, second and third tarsomere of equal length (Fig. 3d), claws symmetrical, very slender, gently curved, in oblique angle to distal tarsomere, base of claw not swollen, parempodia setiform, long, slender, and symmetrical (Fig. 3e); fore leg with coxa very long and slender (Figs 3a), trochanter as in Fig. 3b, femur straight, relatively slender, with one row of spines on medial surface (Figs 3a), tibia straight, of similar diameter over entire length (Fig. 3a); mid and hind legs with coxae ovoid, femora and tibiae very long and slender. Fossula spongiosa and metacoxal comb absent. Hemelytron (Figs 1b, f, 4b): elongate, corium beset with short setae restricted to narrow anterior area (Fig. 4b), basal area between R, M+Cu, Pcu, and posterior margin of wing slightly more heavily sclerotized than actual membrane, M and Cu fused, basal cell trapezoidal, discal cell very long and slender, additional cross vein apart from rm vein between R and M+Cu, here referred to as rmcu cross vein. Abdomen (Figs 1a, c, 3f ): elongate ovoid, lateral margins smooth, dorsal laterotergites indistinct, ventral laterotergites distinct (Fig. 3f ); second to seventh spiracle small, circular, on mediosternites (Figs 3f ), eight spiracle on dorsolateral surface of segment 8 (Figs 5b, c). GENITALIA (Fig. 5): segment 8 well developed, membranous on dorsal surface; pygophore elongate ovoid, with triangular process on posterior margin (Fig. 5b); parameres slender, curved, apex pointed (Figs 5a-c); aedeagus (Figs 5d, e) with basal plates stout and strongly curved, ponticulus basilaris slender, basal plate extension relatively short and stout, basal plate struts short, phallothecal sclerite wider than long, heavily sclerotized, endosoma with dorsal, lateral, slightly sclerotized lobes, one apical median and three pairs of lateral, membranous lobes beset with small spicules. b
a
c
M+Cu R
Pcu
basal cell
M+Cu
M+Cu
Pcu
R
R
rm
rmcu Pcu
rm distal cell
Cu
Pcu rm
Cu
Cu M
M M
Fig. 4: Comparison of venation of the hemelytron in the three recent genera of Collartidini; the anterior wing margin is on the left. a, Collartida sp. (after Wygodzinsky 1966); b, Mangabea barbiger, nov. sp. (original); c, Stenoramphus nubifera (Distant) (after Elkins 1962)
400 C. Weirauch
a
b
d
pht
end
1 mm
sp8
bp 1 mm
par
c
prpy
par
pht
e end bp
Fig. 5: Male genitalia of Mangabea barbiger, nov. sp. a, pygophore, ventral view; b, pygophore, dorsal view; c, pygophore, lateral view; d, semi-inflated aedeagus, dorsal view; e, semi-inflated aedeagus, lateral view.
Fig. 6: Collecting sites of Mangabea barbiger, nov. sp., ( squares) and Mangabea orientalis Villiers ( circles) in Madagascar. Records for the latter species are derived from Villiers (1970) and were georeferenced using GEOLocate software.
Mangabea barbiger, new species of Collartidini (Heteroptera, Reduviidae, Emesinae) from Madagascar
401
Coloration: general coloration dull brown, with distal parts of appendages and ventral surfaces of thorax abdomen somewhat paler (Figs 1a-c). Head: uniformly brown, labrum pale (Fig. 1e). Antenna: brown, flagellomeres light brown. Labium: brown, apical half of second segment, and third and fourth segment light brown. Thorax: uniformly brown, sterna somewhat paler. Legs: Coxae light brown, trochanters, femora, tibiae, and tarsi pale brown. Hemelytron: uniformly dull brown. Abdomen: tergites pale brown; mediosternites pale brown, laterosternites somewhat darker; pygophore pale brown. Female: unknown. Measurements (in mm): see Tab. 2. Etymology: from Latin adjective “barbiger” meaning “carrying a beard” for the diagnostic fascicle of setae on the ventral surface of the head. Distribution (Fig. 6): known from two localities at high altitudes (1100m) in Ranomafana National Park in the Eastern part of Madagascar. Biology: unknown; the two known specimens were collected in malaise traps, indicating that at least males fly. Both malaise traps were located in high altitudes, one in rainforest and one in mixed tropical forest. Discussion: the sparse distribution records of the two described species of Mangabea may indicate that M. orientalis occurs in low land rainforest in the North of the Island, whereas M. barbiger may be restricted to higher altitudes. Additional material examined: MADAGASCAR: Fianarantsoa: Parc National Ranomafana, Vohiparara, at broken bridge, 21.22616°S 47.36983°E, 1100 m, 19 Mar 2002, M. Irwin, R. Harin’Hala, 1♂ (UCR_ENT 00005201) (CAS).
ACKNOWLEDGEMENTS I thank Norman D. Penny, Californian Academy of Sciences (CAS), for sorting to family level, organizing, and loanining about 2800 specimens of Reduviidae from the CAS Madagascar Project. Collections from Madagascar were partially supported by grant DEB-0072713 from the National Science Foundation to B.L. Fisher and C.E. Griswold. Fieldwork that provided the basis for this work could not have been completed without the support of the Malagasy people and I would especially like to acknowledge the efforts of Rin’ha Harinhala. Thanks also to Wei Song Hwang and Guanyang Zhang (UCR) for sorting Malagasy Reduviidae to morpho species. I also thank Michael A. Wall for reviewing an earlier version of this manuscript.
РЕЗЮМЕ Описан е нов вид хетероптера Mangabea barbiger, nov. sp. от източната част на Централен Мадагаскар, представител на ендемичния мадагаскарски род Mangabea VILLIERS (Emesinae-Collartidini). Предложена е ревизирана диагноза за род Mangabea VILLIERS. Пред-
402 C. Weirauch
ставени са карта на разпространението на двата вида от род Mangabea, дигитални микрофотографии, микрофотографии под сканиращ микроскоп, и рисунки на новия вид, илюстриращи видовите и родови диагностични белези. Представени в Таблица са диагностичните белези на трите съвременни рода Collartidini.
REFERENCES Elkins J. C. (1962). Three new Saicine genera (Hemiptera, Reduviidae). — Journal of the Kansas Entomological Society, 35: 421-429. Hijmans R., L. Guarino, A. Jarvis & R. O’Brien (2007). DIVA-GIS. Maldonado J. (1990). Systematic catalogue of the Reduviidae of the World. — Caribbean Journal of Science, Special publication No. 1, University of Puerto Rico, Mayagüez, Puerto Rico: 1-694. Putshkov V.G. & P.V. Putshkov (1985). A catalogue of assassin-bug genera of the world (Heteroptera, Reduviidae). Published by the authors, Kiev: 1-137. Rios N.E. & H.L. Bart Jr. (2005). GEOLocate. Georeferencing software for Natural History Collections. Version 2.13. Usinger R.L. (1943). A revised classification of the Reduvioidea with a new subfamily from South America. — Annals of the Entomological Society of America, 36: 602-618. Villiers A. (1948). Les Reduviidae malgaches. III. Stenopoditae. Gen. Paraghesquierea nov., Antanambeus nov., Fitana nov. et Gallienius nov. — Revue de Zoologie et de Botanique Africaine, 40: 328-332. Villiers A. (1949a). Les Reduviidae Malgaches. VI. Piratinae. Formes nouvelles. — Bulletin du Muséum national d’histoire naturelle, 21: 707-709. Villiers A. (1949b). Revision des Emesides africains (Hemipteres, Reduviides). — Mémoires du Muséum national d’histoire naturelle, n. s., 23: 257-392. Villiers A. (1952). Les Heteropteres Malgaches du Museum National de Praha. VII (2eme note). — Acta Entomologica Musei Nationalis Pragae, 28: 431-433. Villiers A. (1957). Les Reduviides de Madagascar XI. Saicinae. — Bulletin du Museum National d’Histoire Naturelle, deuxieme serie, 29, 310-314. Villiers A. (1961). Localisation et descriptions d’Emesides africains. Revue de Zoologie et de Botanique Africaine., 63: 33-65. Villiers A. (1968). Insectes Hemipteres Reduviidae: 28. — In: Office de la Recherche Scientifique et Technique d’Outre-Mer, C. N. R. S.: Faune de Madagascar: 1-198. Villiers A. (1970). Nouveaux Hemipteres Reduviidae de Madagascar et des Comores. — Annales de la Société entomologique de France, 6: 809-824. Villiers A. (1971). Nouveaux Hemipteres Reduviidae Emesinae de la region Malgache. — Annales de la Société entomologique de France, 7: 849-883. Villiers A. (1979). Insectes Hemipteres Reduviidae (2eme partie): 49. — In: Office de la Recherche Scientifique et Technique d’Outre-Mer, C. N. R. S. Faune de Madagascar: 1-202. Weirauch C. (2006). Dorsal abdominal glands in adult Reduviidae (Cimicomorpha, Heteroptera). Deutsche entomologische Zeitschrift, 53: 91-102. Wygodzinsky P. (1966). A monograph of the Emesinae (Reduviidae, Hemiptera). — Bulletin of the American Museum of Natural History, 133: 1-614.
S.genus Grozeva & N. Simov (Eds)and 2008 New species of the mirine plant bug Castanopsides Yasunaga its assumed sister genus Mahania... 403 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80 th Anniversary of Michail Josifov, pp. 403-417. © Pensoft Publishers Sofia–Moscow
New species of the mirine plant bug genus Castanopsides YASUNAGA and its assumed sister genus Mahania POPPIUS from Nepal, with a new synonymy of the genus Liocapsus POPPIUS (Heteroptera, Miridae, Mirinae) T. Yasunaga1 & R. K. Duwal2 1
Research Associate, American Museum of Natural History, Central Park W at 79th Street, New York, NY 10024, c/o Nameshi 2-33-2, Nagasaki, 852-8061, Japan. E-mail:
[email protected] 2 Laboratory of Insect Biosystematics, Division of Entomology, College of Agriculture and Life Sciences, Seoul National University, San 56-1, Silim9-dong, Gwanak-gu, Seoul 151-742, Korea. E-mail:
[email protected]
ABSTRACT Three new species of the Asian mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania Poppius are described from Nepal. Dorsal habitus and genital structure for each species are provided as digital images. Phylogeny and zoogeography of Castanopsides and Mahania are discussed. Paracyphodema Lu & Zheng is proposed as a junior synonym of Liocapsus Poppius, thus a single representative known from continental China, P. inexpectatus (Zheng & Liu), is transferred to Liocapsus. Keywords: Castanopsides, Liocapsus, Mahania, new combination, new species, new synonymy, phylogeny, zoogeography.
INTRODUCTION Castanopsides was proposed by Yasunaga (1992) to accommodate a single species from southwestern Japan. Subsequently, a revision placed eight additional species in
404 T. Yasunaga & R.K. Duwal
the genus with discussion on the phylogeny and zoogeography (Yasunaga 1998). Any reliable sister genus was not determined in the revision but recent comprehensive work (Yasunaga & Schwartz 2007) suggests a close relationship between Castanopsides and Mahania Poppius. Continuing fieldworks carried out by T. Yasunaga (TY) in Nepal from 2005 to 2006 found three undescribed species of Castanopsides and Mahania, which are diagnosed and described with biological information. The phylogeny is also discussed. Further, during checking related papers, a Chinese genus Paracyphodema Lu & Zheng was found to be congeneric with Liocapsus Poppius. Therefore, Paracyphodema is synonymized with Liocapsus, and a single representative of the former genus is transferred to the latter. This paper is written to celebrate 80th birthday of Dr. Michail Josifov.
MATERIAL AND METHODS All type specimens designated in this study were collected by TY as part of a cooperative program between JICA Senior Volunteer and Natural History Museum, Tribhuvan University, Kathmandu, and will be deposited in the collections of American Museum of Natural History (AMNH); Natural History Museum, Tribhuvan University, Kathmandu (NMTU); and TY personal collection, Nagasaki (YCN). All measurements are given in millimeters. Digital images of live individuals were made by TY, using Canon EOS Kiss Digital camera body + Canon-Olympus mount adapter + Olympus Macrophoto System (Auto Extension Tube with 50 mm macrolens and T10 Ringflash). Terminology of the genitalia follows Yasunaga (1998) and Yasunaga & Schwartz (2007). The following abbreviations are used to describe and show the genitalic structures properly: DLP dorsal labiate plate (♀) DOS dorsal sac (♀) DS dorsal structure of posterior wall of bursa copulatrix or “posterior wall” (♀) FP fin-like process (of phallotheca) (♂) GP secondary gonopore (♂) HP hypophysis (♂) IRL interramal lobe of posterior wall (♀) IRS interramal sclerite of posterior wall (♀) LLB lateral lobe of posterior wall (♀) LS left lateral sclerite (♂)
MS OL PT RS SD SG SL SP SR VLP VLV
median sclerite (♂) oviductus lateralis (or lateral oviduct) (♀) phallotheca (♂) right lateral sclerite (♂) seminal depository (♀) spermathecal (or vermiform) gland (♀) sensory lobe (or shaft) (♂) spiculum (♂) sclerotized ring (♀) ventral labiate plate (♀) valvula (♀)
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 405
Checklist of Castanopsides and Mahania C. dasypterus (Reuter, 1906) — Continental China. C. falkovitshi (Kerzhner, 1979) — FE Russia (Primorskij Territory), N Japan, NE China. C. gotohi (Yasunaga, 1998) — Taiwan. C. hasegawai Yasunaga, 1992 — SW Japan. C. katsutai nov. sp. — Nepal. C. kerzhneri ( Josifov, 1985) — Continental China, FE Russia (Primorskij Territory), Japan, Korea. C. michaili nov. sp. — Nepal. C. taiwanus Yasunaga, 1998 — Taiwan. C. takaii Yasunaga, 1998 — Japanese Yaeyama Group (S Ryukyus), Taiwan. C. montanus Yasunaga, 1998 — Taiwan. C. potanini (Reuter, 1906) — Continental China, FE Russia, Japan, Korea. M. elongata Poppius, 1915 — Nepal, N India. M. ptilophalla nov. sp. — Nepal.
DESCRIPTIONS OF NEW SPECIES Genus Castanopsides YASUNAGA Castanopsides Yasunaga, 1992 (n. gen.): 45, type species: C. hasegawai Yasunaga, 1992: 46, original designation; 1998: 100 (diag., redesc., key, biol.); Kerzhner & Josifov 1999: 81 (cat.); Yasunaga 2001: 226 (diag.); Zheng et al. 2004: 216 (diag.).
Diagnosis. Recognized by the moderate to large size (5.5-8.5 mm), more or less shining dorsum provided with the sericeous pubescence and simple setae, narrow but distinct basal transverse carina of the vertex, short and filiform antennal segment IV that is shorter than the segment I, rather short labium that is not much exceeding apex of the mesocoxa, shiny and usually punctate pronotum, flattened inward surface of the right paramere, triangularly developed sensory lobe of the left paramere, and presence of an apical, fin-like process on the phallotheca and a vesical spiculum. Further diagnostic characters were provided by Yasunaga (1998). Discussion. This genus has hitherto been known by 9 eastern Palearctic species. Two additional species here described from Nepal represent the first records from the Oriental Region. However, occurrences of these Oriental members are apparently restricted to the temperate and subboreal Quercus forests. Therefore, the temperate forests on southern slopes of the Himalayas (1,500-3,500 m altitude) appear to provide the southernmost habitats for Castanopsides.
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Castanopsides was often confused with Arbolygus Kerzhner (a synonym of Philostephanus Distant), because of the similarity of overall appearance. The similarity apparently is simply superficial, and the male and female genitalic structures imply Mahania Poppius is a reliable sister genus of Castanopsides (Yasunaga & Schwartz 2007). Zheng et al. (2004) replaced Castanopsides falkovitshi in Arbolygus (= now Philostephanus). Having no synapomorphy of Philostephanus, however, falkovitshi is better to be regarded as a member of Castanopsides.
Castanopsides katsutai YASUNAGA & DUWAL nov. sp. Figs 1, 2, 7, 8, 13-15, 20, 22
Diagnosis. Recognized by the oval, broad and a little elongate body in both sexes, short antennal segment II that is much shorter than the metafemur, weak median sclerite (MS) and spiculum (SP) on the vesica, and narrow sclerotized rings (SR) that are separated from each other. Description. Body generally brown, more or less darker in ♂, oval, broad, a little elongate, not much sexually dimorphic; dorsal surface rather shining, with uniformly distributed, sericeous pubescence and simple brown setae. Head pale brown; frons usually with a dark, median stripe, with paired transverse sutures on each side of midline. Antenna pale brown; segment II not much incrassate toward apex, shorter than basal width of pronotum; apical 1/8-1/6 of segment II usually darkened; segments III and IV brown, with pale bases. Labium brown, slightly exceeding apex of mesocoxa; apical half of segment IV darkened. Pronotum brown, rather oily shiny, usually with darker posterior half that is provided with a mesal, narrow, pale stripe, coarsely but distinctly punctate; calli somewhat shagreened; collar pale brown, about as thick as apex of antennal segment II; scutellum with yellowish apex, rather arched, irregularly and transversely wrinkled; pleura more or less darkened, especially in ♂, with pale brown metathoracic scent efferent system. Hemelytron partly tinged with red; apex of corium pale red; membrane smoky brown, with pale red veins. Coxa and leg pale brown; each femur with 3-4 dark, apical rings that are sometimes reduced or obliterated in ♀; ♂ metafemur usually widely darkened; tibial spines pale brown; apical part of tarsomere III dark brown. Male genitalia: Parameres as in Figs 7 & 8; hypophysis (HP) of right paramere widened apically (7); left paramere with a sharply projected sensory lobe (8). Vesica (13) widely membranous; spiculum (SP) and median sclerite (MS) reduced, weak. Female genitalia: Sclerotized rings (SR) narrowed, elongate-oval, separated from each other (14); posterior wall of bursae with narrow lateral lobe (LLB, 15). Measurements (♂/ ♀): Body length 6.7-7.1/ 6.8-7.7; head width across eyes 1.051.11/ 1.12-1.16; vertex width 0.43-0.45/ 0.48-0.50; lengths of antennal segments I-IV 0.62-0.70, 1.92-2.16, 0.88-1.01, 0.48-0.50/ 0.64-0.70, 1.80-1.85, 0.93-0.96, 0.48-0.56; total labial length 2.40-2.48/ 2.59-2.64; mesal pronotal length including collar 1.12-1.20/ 1.24-1.32; basal pronotal width 1.68-2.26/ 2.40-2.50; width across hemelytron 2.56-
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 407
2.84/ 3.12-3.20; lengths of metafemur, tibia and tarsus 2.16-2.43, 3.16-3.48 0.72-0.77/ 2.44-2.64, 3.55-3.68, 0.74-0.80. Holotype ♂: NEPAL, Rasuwa District, Langtang Himal National Park, Lama Hotel ~ Rimche, 2,340~2,440 m alt., on flowers of Quercus, 8. vi. 2006, T. Yasunaga (AMNH). Paratypes: NEPAL. Rasuwa District, Langtang Himal National Park: same data as for holotype, 2♂7♀ (YCN); same data except for date, 3. vi. 2007, 10♂15♀ (NMTU, YCN); Gomnachok ~ Lama Hotel, 2,700~2,500 m alt., on flowers of Quercus, 7. vi. 2006, T. Yasunaga (YCN).
1
2
3
4
5
6
Fig 1: Castanopsides katsutai, ♂. Fig. 2: Ditto, ♀. Fig. 3: C. michaili, ♂. Fig. 4: Ditto, ♀. Figs 5-6: Mahania ptilophalla, holotype ♂.
408 T. Yasunaga & R.K. Duwal
Etymology. Named in honor of Mr. Nobuji Katsuta who helped TY’s fieldworks in Langtang Himal National Park; a noun in genitive case. Distribution. Nepal (subboreal forests grown with Quercus trees). Discussion. This new species is easily distinguished from other congeners by the broad body, short antennal segment II, reduced sclerotization of the vesica and narrowed sclerotized ring. TY collected many individuals by sweeping inflorescences of Quercus trees (mostly Q. semecarpifolia) that are regarded as the breeding host, during trekking subboreal forests of the Langtang Himal. HP
HP
HP
HP
7
9
SL SL
11 HP
HP
SL
SL
8
10
HP
12 HP
SL
katsutai
michaili
Mahania ptilophalla
Figs 7-12: Right (7, 9 & 11) and left (8, 10 & 12) parameres. Figs 7-8: Castanopsides katsutai. Figs 9-10: C. michaili. Figs 11-12: Mahania ptilophalla. Morphological abbreviations corresponding to those mentioned in the text.
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 409
Castanopsides michaili Yasunaga & Duwal nov. sp. Figs 3, 4, 9, 10, 16-18, 20, 22 Castanopsides sp.: Yasunaga 1998: 116 (diag.).
Diagnosis. Recognized by the elongate-oval body, dark general coloration, pale head and base of antennal segment III, pale cuneus with the darkened apical 1/4, short vesical spiculum that is fused with the median sclerite at apex, and enlarged sclerotized rings that are subcontiguous to each other. 13
14
SD
VLV II
15
MS
SR IRL
GP
DLP
SP
DS
LLB
PT FP VLP
SG MS
DOS
OL
IRS
SD
16
LS VLP
DLP SP
SR
GP
FP
FP
17 SG LLB
OL DS
IRL
SP PT RS?
19 IRS
GP
18
GP PT
Figs 13-19: Male phallic organ (13, 16 & 19) and female genitalia (14, 15, 17 & 18). Figs 1315: Castanopsides katsutai. Figs 16-18: C. michaili. Fig. 19: Mahania ptilophalla. Morphological abbreviations corresponding to those mentioned in the text.
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Description. Body generally dark brown, subparallel-sided (♂), chestnut brown, elongate-oval (♀); dorsal surface shining, with uniformly distributed, silky reclining pubescence and brown simple setae. Head brown to pale brown, oily shiny; frons usually with a yellow, mesal stripe; tylus infuscate. Antenna dark brown; segment I and basal 1/5-1/4 of II pale brown in ♀; ♂ segment II rather thickened; base of segment III pale brown. Labium pale brown, slightly exceeding apex of mesocoxa; apical half of segment IV darkened. Pronotum shiny chestnut brown, uniformly and finely punctate, with uniformly distributed, silky, semierect pubescence; calli dark brown, with a yellow mesal stripe; collar creamy yellow, contrasting to dark pronotal disk, about as thick as base of antennal segment II; scutellum fuscous, with yellow apex (♂), chestnut brown, darkened or somewhat striped medially (♀); pleura widely darkened; metathoracic scent efferent system darkened dorsally. Hemelytron dark chocolate brown (♂), chestnut brown (♀); cuneus yellow, with darkened apical 1/5-1/4 and reddish inner margin; membrane smoky brown, with more or less reddish veins. Coxa and leg pale brown; pro- and mesofemora with 2-3 obscure, pale red rings; apical half of metafemur and base of metatibia reddish brown; tibia with brown spines; apical part of tarsomere III dark brown. Male genitalia: Parameres as in Figs 9 & 10; right paramere rather shortened (9); left paramere with roundly projected sensory lobe (10). Vesical spinules on membranous areas distinct; spiculum (SP) short, fused with median sclerite (MS) at apex and forming a ring (16). Female genitalia: Sclerotized rings (SR) large, subcontiguous to each other (17); lateral lobe (LLB) narrow, indistinct (18). Measurements (♂/ ♀): Body length 5.9-6.4/ 6.5-7.4; head width across eyes 1.051.08/ 1.08-1.16; vertex width 0.40-0.41/ 0.45-0.48; lengths of antennal segments I-IV 0.69-0.72, 2.08-2.12, 0.96-0.99, 0.43-0.48/ 0.67-0.72, 2.01-2.12, 0.97-1.06, 0.45-0.51; total labial length 2.06-2.12/ 2.20-2.31; mesal pronotal length including collar 1.20-1.23/ 1.22-1.40; basal pronotal width 2.06-2.10/ 2.16-2.38; width across hemelytron 2.322.38/ 2.44-2.86; lengths of metafemur, tibia and tarsus 2.37-2.43, 3.19-3.36 0.74-0.77/ 2.40-2.75, 3.26-3.60, 0.72-0.81. Holotype ♂: NEPAL, Rasuwa District, Langtang Himal National Park, Dhunche (1,950 m alt.), at light [of a lodge balcony], 8-9. vi. 2006, T. Yasunaga (AMNH). Paratypes: NEPAL. Rasuwa District, Langtang Himal National Park: same data as for holotype, 2♂6♀ (NMTU, YCN); Lama Hotel ~ Rimche, 2,340 m~2,440 m alt., on flowers of Quercus, 8. vi. 2006, T. Yasunaga, 1♂ (YCN). Additional specimen: NEPAL. Kathmandu [Valley, Lalitpur], Godavari [= Godawari], 30. iv. 1983, Takakuwa, 1♀ (National Institute of Agro-Environmental Sciences, Tsukuba, Japan). Etymology. Named in honor of Dr. Michail Josifov who contributed much to clarification of Asian faunas of the Miridae; a noun in genitive case. Distribution. Nepal (temperate to subboreal forests grown with Quercus oaks). Discussion. This new species is similar in general appearance to the continental Chinese C. dasypterus (Reuter), from which it can be distinguished by the paler head, pale base of the antennal segment II, and structure of the vesical sclerites. Judging from
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 411
the form of the male and female genitalia, C. michaili is evidently allied to C. gotohi Yasunaga from Taiwan. The present new species corresponds to what was reported from Nepal as Castanopsis sp. (Yasunaga 1998). Only a teneral female was available at the moment. Zheng et al. (2004) redescribed C. dasypterus based on 27 specimens collected from 5 provinces of continental China. The illustrations of the vesica (Fig. 67) apparently do not fit that of the holotype of C. dasypterus (Kerzhner (1979) and Yasunaga (1998) illustrated it). The figured vesica is rather similar to that of C. michaili or C. gotohi as exhibiting the distinctly spinulate membranous areas and a shortened spiculum, or may represent an undescribed species as lacking the median sclerite. Further examination is desirable to correctly ascertain how many species are included in the 27 above-mentioned Chinese specimens.
Genus Mahania POPPIUS Mahania Poppius, 1915: 60 (n. gen.), type species: M. elongata Poppius, 1915: 61, original designation; Yasunaga & Schwartz 2007: 173 (diag., redesc.). Philostephanus Distant (part): Carvalho 1952: 92, 95 (n. syn., cat.); 1959: 187 (cat.); Schuh 1995: 862 (cat.).
Diagnosis. Recognized by the elongate body, shortened labium that is not exceeding the mesocoxa, flattened inward surface of the right paramere, strongly projected sensory lobe of the left paramere, a narrow keel along the phallotheca, and a single spiculum and developed right lateral sclerite on the vesica. Further diagnostic characters were provided by Yasunaga & Schwartz (2007). Discussion. Mahania has been represented by a single species, M. elongata, known from Nepal and north India, south slope of the Himalayas. The following species is the second member of this unique genus.
Mahania ptilophalla YASUNAGA & DUWAL nov. sp. Figs 5, 6, 11, 12, 19, 22
Diagnosis. Recognized by the small size, wholly dark antennal segment II that is much longer than basal width of the pronotum, and highly modified form of the vesica. Description. Male (female unknown): Body generally brownish, partly tinged with red, elongate, nearly parallel-sided; dorsal surface widely chocolate brown, weakly shining, with uniformly distributed, silky, reclining pubescence and brown, simple setae. Head shiny brown, almost smooth. Antenna dark brown; segment I about as broad as II; segment II much longer than basal width of pronotum, equal to metafemur; segment III and IV filiform, with whitish bases. Labium pale brown, short, reaching base
412 T. Yasunaga & R.K. Duwal
of mesocoxa; apical half of segment IV dark brown. Pronotum shining, widely darkened posterior to calli; collar creamy yellow, about as thick as base of antennal segment II; scutellum chocolate brown, rather flat, with pale extreme apex; pleura unicolorously pale brown. Hemelytron chocolate brown; embolium and lateral margin of cuneus reddish brown; membrane smoky brown, with pale veins. Coxa and leg pale brown; tibial spines pale brown; apical half of tarsomere III dark brown. Male genitalia: Parameres as in Figs 11, 12; right paramere slender, flattened inwardly (11); sensory lobe of left paramere with a flat, conspicuous protuberance (12). Vesica (19) highly modified and sclerotized, with feather-like, hairy appendages; spiculum flattened and broadened; phallotheca with a narrow keel. Measurements (♂): Body length 6.4; head width across eyes 1.15; vertex width 0.36; lengths of antennal segments I-IV 0.96, 2.71, 1.08, 0.53; total labial length 1.58; mesal pronotal length including collar 1.20; basal pronotal width 2.11; width across hemelytron 2.40; lengths of metafemur, tibia and tarsus 2.16, 3.48, 0.72. Holotype ♂: NEPAL, Kathmandu, Nagarjun (1,600~1,800 m alt.), on Quercus sp., 16. v. 2005, T. Yasunaga (AMNH). Etymology. From Greek, ptilon (soft feather, down) in combination with phallus (male phallic organ), referring to the unique feather-like pubescence on the vesica of this new species. Distribution. Nepal (temperate Quercus forest). Discussion. As the vesica has peculiar structure, this new species is at first sight difficult to be accommodated in any known genera. But the principal coloration and structure of the dorsal surface, short rostrum, inwardly flattened right paramere, developed sensory lobe of the left paramere and a narrow keel of the phallotheca apparently represent diagnostic characters of Mahania. We currently consider the highly modified form of the vesica as simply an autapomorphy.
PHYLOGENETIC DISCUSSION (Fig. 20)
Phylogeny of Castanopsides was inferred and discussed by Yasunaga (1998), based on an evaluation of 14 characters. Although the hypothesis appears to have well reconstructed the phylogenetic relationships in the genus, any reliable outgroup, or sister genus, was not confirmed at the moment. Since a recent revision (Yasunaga & Schwartz 2007) shows distinct monophyly of some related members and two new members are here included in Castanopsides, a more reliable analysis now can be provided. Among any known genera of the tribe Mirini, Mahania is assumed to be the closest outgroup, as discussed by Yasunaga & Schwartz (2007). A cladogram shown in Fig. 20 is based principally on a character set by Yasunaga (1998). Main additions and modifications (synapomorphies and autapomorphies) in the present analysis are summarized as follows:
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 413
1. Right paramere strongly flattened inwardly; vesica ventrally with a spiculum extending from subapical part of ejaculatory (seminal) duct. 2. Phallotheca apically with a fin-like process. 3. Labium shortened; sensory lobe of left paramere well developed, strongly projected dorsally; phallotheca with a narrow, keel-like projection or margin; vesical (right) lateral sclerite well sclerotized. 4. Vesica widely membranous, with weak spiculum and median sclerite. 5. Vesica covered with distinct spinules, with a shortened and broadened spiculum. 6. Vesical spiculum fused with median sclerite apically. Members of Castanopsides apparently prefer to inhabit the temperate to cold temperate climate zones. Most of them are considered to be associated with fagaceous oaks (Castanopsis, Quercus, etc.), and to have a univoltine life cycle. Both immatures and newly emerged adults are frequently found on inflorescences of these oaks that usually bloom from late spring to early summer. Habit and habitat of Mahania species are also similar to those of Castanopsides but the occurrences of Mahania members are presumably restricted to temperate forests of the Himalayas. Such biological and distributional information suggests that Castanopsides is likely to be originated from the Himalayas as supposed for many Asian organisms. Monophyly of Castanopsides and Mahania is supported by two synapomorphies (1). In addition to these apomorphies, many similarities are found in their general appearances and habitat preferences. Of all known members of Castanopsides (except for C. falkovitshi that is specialized in some structures and host preference), C. katsutai, a typical Himalayan Quercus inhabitant, is presumably the most basic member. On the other hand, C. michaili is considered sister to C. gotohi, known from Taiwanese mountains. Although distribu-
Fig. 20: Cladogram showing hypothesized phylogenetic relationships of Castanopsides species and its assumed sister genus Mahania, modified from Yasunaga (1998). See text for principal apomorphies (1-6).
414 T. Yasunaga & R.K. Duwal
tions of these two related species are isolated from each other, further investigations in continental China will discover additional sisters (see discussion on michaili). Mahania ptilophalla possesses a very specialized and highly modified vesica that is currently considered simply autapomorphic. Because this peculiar species is represented by the holotype alone, further efforts are required to reconfirm its generic placement by the female genitalic structure.
Fig. 21: Habitats of Castanopsides katsutai + C. michaili (upper, inflorescences of Quercus semecarpifolia in Langtang Himal National Park) and Mahania ptilophalla (temperate forest of Mt. Nagarjun, Kathmandu).
New species of the mirine plant bug genus Castanopsides Yasunaga and its assumed sister genus Mahania... 415
PROPOSAL OF NEW SYNONYMY AND NEW COMBINATION Genus Liocapsus POPPIUS Liocapsus Poppius, 1915: 15 (n. gen.), type species: Liocapsus brevirostris Poppius, 1915, original designation; Carvalho 1959: 109 (cat.); Schuh 1995: 788 (cat.); Yasunaga & Schwartz 2007: 165 (diag., redesc., key, biol.). Paracyphodema Lu & Zheng in Zheng et al., 2004: 462, 760, type species: Cyphodema inexpectata Zheng & Liu, 1992, original designation. nov. syn.
Discussion. During checking up on some related Asian taxa for this study, we found a mirine genus Paracyphodema recently proposed to accommodate a single Chinese species, Cyphodema inexpectata, by Lu & Zheng (in Zheng et al. 2004) is evidently congeneric with Liocapsus Poppius. The dorsal habitus image and genitalic illustrations suggest that inexpectata is very similar in overall appearance to L. brevirostris Poppius known from North India. Since we have not examined the genitalia of the Chinese species, however, we just propose the new synonymy of Liocapsus and the following new combination: Liocapsus inexpectatus (Zheng & Liu) nov. comb. Liocapsus at first sight resembles Castanopsides and Mahania, but the structures of the pronotum and the male and female genitalia are completely different from the latter genera. Liocapsus is now regarded as an evident monophyletic group and considered sister to Philostephanus Distant and Orientocapsus Yasunaga & Schwartz (Yasunaga & Schwartz 2007).
C. katsutai + C. michaili
C. michaili
M. ptilophalla
Fig. 22: Map showing distributions for new species of Castanopsides and Mahania.
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ACKNOWLEDGMENTS We express our gratitude to Dr. S. Miyamoto (Fukuoka, Japan) for continuous advice and to Dr K. Shrestha, Assoc. Prof. P. K. Shrestha and Mr. B. Kanal (NMTU) for encouraging our research activities in Nepal. We are also indebted to JICA ( Japan International Cooperation Agency) Nepal Office for funding cooperative program of TY and to Department of National Parks and Wildlife Conservation of Nepal, Kathmandu for issuing permissions to investigate and collect mirids in National Park areas. Mr. N. Katsuta (Kathmandu, Nepal) and his wife Ms. K. Ito ( JICA Expert) were trekking the Langtang Himal National Park with TY and helping his fieldwork much. We are grateful to Drs. S. Grozeva and N. Simov (Sofia, Bulgaria) who kindly invited us to this commemorative volume for Dr. Josifov. Thanks are extended to an anonymous reviewer for improving the manuscript with useful comments and suggestions.
РЕЗЮМЕ Описани са от Непал три нови вида хетероптери от родовете Castanopsides YASUNAGA и Mahania POPPIUS, представители на подсемейство Mirinae. Представени са дигитални фотографии на дорзалният набитус и гениталната структура. Обсъдени са филогенията и зоогеографията на Castanopsides и Mahania. Paracyphodema LU & ZHENG е синонимизиран с Liocapsus POPPIUS, и по този начин единственият представител на рода, известен от континентален Китай, P. inexpectatus (ZHENG & LIU), е преместен в род Liocapsus.
REFERENCES Carvalho J.C.M. (1952). On the major classification of the Miridae (Hemiptera) (With keys to subfamilies and tribes and a catalogue of the world genera). — Anais da Academia Brasileira de Ciencias, 24: 31-110. Carvalho J.C.M. (1959). A catalogue of the Miridae of the world, Part IV. — Arquivos do Museu Nacional, Rio de Janeiro, 48: 1-384. Kerzhner I.M. (1979). Novye poluzhstkokryrye (Heteroptera) s Dal’nego Vostoka SSSR [New Heteroptera from the Soviet Far East]. – Trudy zoologitsheskogo Instituta Akademija Nauk SSSR, Leningrad, 81: 14-65. (In Russian) Kerzhner I.M. & M. Josifov (1999). Miridae Hahn, 1833. — In: Aukema, B. & C. Rieger (eds.): Catalogue of the Heteroptera of the Palearctic Region, vol. 3, Cimicomorpha II. The Netherlands Entomological Society: 1-576. Poppius B. (1915). Zur Kenntnis der Indo-Australischen Capsarien I. – Annales historiconaturales Musei nationalis Hungarici, 13: 1-89.
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Schuh R.T. (1995). Plant bugs of the world (Insecta: Heteroptera: Miridae). Systematic catalog, distributions, host list and bibliography. — The New York Entomological Society: xii + 1329. Yasunaga T. (1992). New genera and species of the Miridae of Japan (Heteroptera). – Proceedings of the Japanese Society of systematic zoology, 47: 45-51. Yasunaga T. (1998). Revision of the mirine genus Castanopsides Yasunaga from the eastern Asia (Heteroptera: Miridae). — Entomologica Scandinavica, 29: 99-119. Yasunaga T. (2001). Family Miridae, plant bugs — In: Yasunaga T., M. Takai & T. Kawasawa (eds.): A Field Guide to Japanese Bugs II. Zenkoku Noson Kyoiku Kyokai Publ. Co. Ltd., Tokyo: 112-276, Figs 2-331. (In Japanese) Yasunaga T. & M.D. Schwartz (2007). Revision of the mirine plant bug genus Philostephanus Distant and allies (Heteroptera: Miridae: Mirinae: Mirini). — Tijdschrift voor Entomologie, 150: 101-180. Zheng L., N. Lu, G. Liu & B. Xu (2004). Hemiptera, Miridae, Mirinae. Fauna Sinica, Insecta Vol. 33. — Science Press, Beijing: xix + 797, 8 pls. (In Chinese, with English keys and descriptions of new taxa)
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S. genus Grozeva & N. Simov (eds.)and 2008. New species of the mirine plant bug Castanopsides Yasunaga its assumed sister genus Mahania... 419 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of 80th Anniversary of Michail Josifov, p. 419. © Pensoft Publishers Sofia–Moscow
Appendix: List of new taxa described in this volume Calacanthia josifovi nov. sp. Vinokurov, 2008 – Saldidae Castanopsides katsutai nov. sp. Yasunaga & Duwal, 2008 – Miridae Castanopsides michaili nov. sp. Yasunaga & Duwal, 2008 – Miridae Catoplatus josifovi nov. sp. Golub, 2008 – Tingidae Chorosoma josifovi nov. sp. Schwartz, Schaefer & De Lattin, 2008 – Rhopalidae Dictyonota michaili nov. sp. Günther, 2008 – Tingidae Euchilofulvius josifovi nov. sp. Gorczyca, 2008 – Miridae Josifovius nov. gen. Konstantinov, 2008 – Miridae Leprosoma tenuimarginatum nov. sp. Gapon, 2008 – Pentatomidae Loricula (Myrmedobia) josifovi nov. sp. Simov, 2008 – Microphysidae Mahania ptilophalla nov. sp. Yasunaga & Duwal, 2008 – Miridae Mangabea barbiger nov. sp. Weirauch, 2008 – Reduviidae Omocoris unicolor nov. sp. Konstantinov, 2008 – Miridae Pilophorus josifovi nov. sp. Kerzhner, 2008 – Miridae Pilophorus josifovianus nov. sp. Duwal & Yasunaga, 2008 – Miridae Pinochius josifovi nov. sp. Rédei, 2008 – Schizopteridae Raglius alboacuminatus josifovi nov. ssp. Linnavuori & Heiss, 2008 – Lygaeidae Sigara (Pseudovermicorixa) josifovi nov. sp. Chen & Nieser, 2008 – Corixidae Tubuaivelia nov. gen. Polhemus & Polhemus, 2008 – Veliidae Tubuaivelia michaili nov. sp. Polhemus & Polhemus, 2008 – Veliidae Xenicocephalus josifovi nov. sp. Štys & Baa, 2008 – Enicocephalidae
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