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L-Bulletin of the 'V Fisheries Research ; Board of Canada ^^ cC% ^^ L9 1 L-Bulletin of the 'v Fisheries Research ; Board of Canada DFO - Library MPO - Bibliothèque _-__a ^t Environnement * (F I VYiI@I^YII U01b9 12039458 isULLETiN OF TUF FIS?iF.R1FS RESEARCH 13IJARU t3F CANADA. Female genitalia in Chironomid ^ IM an other ^ ! NematoGer ' a-^1°I Y^ rtrf{faul ,.n^sty^: 1T/^^'s a...... phylogeniei keys ^^^.. OW^.n1, ^^^ IM-72, ^ Ole A. Saeth^ ^•• skwowao -. ® lb Fish. CanULLETIN 197 S-i ttawa 1977 223 "^ 1' 13 no. 197 JC op Fishenes an0 Enronmenl Pêches et Envirdmement C,-da Canada Fishene5 and ^aemca Op pklw Manne Sennce et de h ^ Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys Bulletins are designed to interpret current knowledge in scientific fields pertinent to Canadian fisheries and aquatic environments. Recent numbers in this series are listed at the back of this Bulletin. The Journal of the Fisheries Research Board of Canada is published in annual volumes of monthly issues and Miscellaneous Special Publications are issued periodically. These series are for sale by Supply and Services Canada, Printing and Publishing, Ottawa, KlA 0S9. Remittances must be in advance, payable in Canadian funds to the order of the Receiver General for Canada. Editor and Director of Scientific Information J. C. STEVENSON, PH.D. Deputy Editor J. WATSON, PH.D. JOHANNA M. REINHART, M.SC. Assistant Editors D. G. COOK, PH.D. J. CAMP Production-Documentation MICKEY LEWIS MONA SMITH Department of Fisheries and the Environment Fisheries and Marine Service Scientific Information and Publications Branch 116 Lisgar Street, Ottawa, Canada KlA OH3 BULLETIN 197 Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys 01e A. Sæther Department of Fisheries and the Environment Fisheries and Marine Service Freshwater Institute Winnipeg, Manitoba DEPARTMENT OF FISHERIES AND THE ENVIRONMENT FISHERIES AND MARINE SERVICE OTTAWA 1977 "2.;b 1\) o I C) Minister of Supply and Services Canada 1977 Available by mail from: Printing and Publishing Supply and Services Canada Ottawa, Canada KlA 0S9 or through your bookseller A deposit copy of this publication is also available for reference in public libraries across Canada Canada: $6.00 Catalog No. Fs 94-197 Other countries: $7.20 ISBN 0-660-00764-9 Price subject to change without notice •Coyer design by Christine Rusk• Contents \Ili/ viii ABSTRACT/ RÉsumÉ 1 INTRODUCTION 1 MATERIALS AND METHODS 2 ABBREVIATIONS 3 MORPHOLOGY OF FEMALE GENITALIA IN INSECTA AND DIPTERA 8 FEMALE GENITALIA OF DIPTERA OTHER THAN CHIRONOMIDAE 8 Infraorder Tipulomorpha 9 Infraorder Psychodomorpha 10 Infraorder Culicimorpha 12 Infraorder Bibionomorpha 15 Suborder Brachycera 15 Summary 25 FEMALE GENITALIA OF CHIRONOMIDAE 29 Phyletic relationships within Chironomidae 33 Key to females of subfamilies and tribes of Chironomidae 44 Subfamily Tanypodinae 44 Tribe Coelotanypodini 44 Key to females of Coelotanypodini 45 Tribe Tanypodini 45 Tribe Anatopyniini 45 Tribe Macropelopiini 46 Preliminary key to females of Macropelopiini 48 Key to some Nearctic females of Procladius Skuse 48 Tribe Pentaneurini 49 Phyletic relationships within Pentaneurini 50 Preliminary key to females of Pentaneurini 54 Subfamily Aphroteniinae 56 Subfamily Podonominae 56 Tribe Podonomini 57 Tribe Boreochlini Contents (concluded) 59 Subfamily Telmatogetoninae 60 Subfamily Diamesinae 63 Tribe Heptagyini 64 Tribe Lobodiamesini 64 Tribe Boreoheptagyiini 64 Tribe Harrisonini 64 Tribe Diamesini 65 Key to females of Diamesini 73 Tribe Protanypodini 73 Subfamily Prodiamesinae 74 Key to females of Prodiamesinae 76 Subfamily Orthocladiinae 76 Phyletic relationships within Orthocladiinae 87 Preliminary key to females of Orthocladiinae 130 Subfamily Chironominae 130 Phyletic relationships within Chironominae 138 Tribe Tanytarsini 140 Preliminary key to females of Tanytarsini 154 Tribe Pseudochironomini (new tribe) 154 Key to adults of Pseudochironomini 156 Tribe Chironomini 157 Preliminary key to females of Chironomini 169 Key to sufficiently known females of Glyptotendipes subgen. Phytotendipes Goetgh. 171 Key to examined females of Cryptotendipes 193 Summary 194 PROSPECTIVES 194 ACKNOWLEDGMENTS 195 REFERENCES 203 INDEX vi ABSTRACT SE-THEn, O. A. 1977. Female genitalia in Chironomidae and other Nematocera: morphol- ogy, phylogenies, keys. Bull, Fish. Res. Board Can. 197: 209 p. Elements of abdominal segments VIII-X or XI form the female genitalia in Diptera. Major genital elements include tergites (T) and sternites (S) VIII and IX, gonocoxites (Gc) VIII and IX (which may bear a gonostylus), gonapophyses (Gp) VIII and IX (which form a notum and rami), cerci, seminal capsules, and spermathecal ducts. Their development in various dipteran families (Nematocera and Brachycera) is discussed, the theoretical pro- genitor of the Diptera illustrated, and the female genitalia of 17 families of Diptera described (including 33 nonchironomid species representing 24 genera). Of the Chirono- midae, 189 species representing 123 genera have been illustrated and a few additional species examined. Within the Chironomidae, T VIII is reduced in the Telmatogetoninae only. Gc VIII is perhaps present in Afrochhis harrisoni Freem., but otherwise absent. Gonocoxapodemes (Gca) VIII are present in Diamesinae, Orthocladiinae, and Chironominae. Gp VIII is, in the basic design, simple and well developed; it is divided into a dorsomesal and a ventrolateral lobe in some Dilmesinae, Prodiamesinae, most Orthocladiinae, and most Chironominae; occasionally it bears an apodeme lobe in the same subfamilies. Gc IX carries a gonostylus in Telmatogetoninae only, and is fused with T IX forming gonotergite IX in Tanypodinae, Aphroteniinae, and Podonominae. T IX is partly or fully divided into two setigerous protrusions in some Diamesinae, Prodiamesinae, Orthocladiinae, and in Lnuterborniella (Chironominae). In Telmatogetoninae there are no seminal capsules, but there are two or three in the remaining subfamilies. Within the Chironomidae, the Telmatogetoninae forms the sister group of the other subfamilies which can be divided into two sister groups of subfamilies: Tanypodinae + Aphroteniinae + Podonominae, and Diamesinae + Prodiamesinae + Orthocladiinae + Chironominae. Keys and diagnoses are given to subfamilies, tribes, genera, and some species of some genera of female Chiron omidae. The phylogeny of each subfamily is discussed and synapomorphic diagrams presented for the Orthocladiinae and Chironominae using charac- teristics for males, pupae, and larvae, in addition to those of the females. In the Ortho- cladiinae, Diplocladins Kieff. probably forms the sister group of the remaining genera. In the Chironominae the tribes Tanytarsini and Pseudochironomini (new tribe) apparently form the sister group of the Chironomini. vii RÉSUMÉ SJETHER, O. A. 1977. Female genitalia in Chironomidae and other Nematocera: morphol- ogy, phylogenies, keys. Bull. Fish. Res. Board Can , 197: 209 p. Des éléments des segments abdominaux VIII—X ou XI forment les organes génitaux femelles des Diptera. Les principaux éléments génitaux comprennent les tergites (T) et les sternites (S) VIII et IX, les gonocoxites (Ge) VIII et IX (pouvant porter un gonostyle), les gonapophyses (Gp) VIII et IX (formant un notum et rami), les cerques, les capsules séminales et les conduits des spermatophores. L'auteur analyse leur développement chez diverses familles de Diptera (Nematocera et Brachycera), figure le progéniteur théorique des Diptera et décrit les organes génitaux femelles de 17 familles de Diptera (y compris 33 espèces autres que des Chironomidae, représentant 24 genres). Parmi les Chironomidae, 189 espèces représentant 123 genres sont figurées, et quelques autres espèces sont examinées. Chez les Chironomidae, T VIII n'est réduit que chez les Telmatogetoninae. Ge VIII est peut-être présent chez Afrochlus harrisoni Freem., mais est absent dans les autres cas. Les gonocoxapodèmes (Gca) VIII sont présents chez les Diamesinae, les Orthocladiinae et les Chironominae. Gp VIII, dans sa forme basale, est bien développé; il est divisé en lobes dorsomésal et ventrolatéral chez certains Diamesinae, Prodiamesinae, la plupart des Ortho- cladiinae et la plupart des Chironominae; dans ces mêmes sous-familles, il porte parfois un apodème. Ge IX ne porte un gonostyle que chez les Telmatogetoninae, et il se fusionne avec T IX pour former le gonotergite IX chez les Tanypodinae, les Aphroteniinae et les Podonominae. T IX est partiellement ou complètement divisé en deux excroissances sétigères chez certains Diamesinae, Prodiarnesinae, Orthocladiinae et chez Lauterhorniella (Chironominae). Chez les Telmatogetoninae il n'y a pas de capsules séminales, mais il y en a deux ou trois dans le reste des sous-familles. - Parmi le Chironomidae, les Telmatogetoninae constituent le groupe soeur des autres sous-familles, pouvant se diviser en deux groupes sœurs de sous-familles: Tanypodinae + Aphroteniinae + Podonominae et Diamesinae + Prodiamesinae + Orthocladiinae + Chironominae. L'auteur donne des clés et des diagnoses des sous-familles, tribus, genres et de quelques espèces de certains genres de Chironomidae femelles. Il discute de la phylogénie de chaque sous-famille et présente des diagrammes synapomorphes des Orthocladiinae et des Chironominae, se servant des caractéristiques des mâles, des pupes et des larves, en plus de celles des femelles. Chez' les Orthocladiinae, Diplocadius Kieff. constitue proba- blement le groupe soeur des autres genres. Chez
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