Diptera Chironomidae
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Bulletin of the Fisheries Research Board of Canada DF O 0]/ 'rissocladius, ^s aratrissocladius, a )ome related gener Fist Can 5H BULLETIN 195 2[3 Ottawa 1976 Bv?_13 Fisheries and Environment Péches et Environnement Canada Canada no. 195 Fisheries and Service des pecheS JC Marine Service et Oe la mer Reuision of ilydrobaenus rrissocladius Zalutschia Paratrissacladius and some related genera (Diptera: Chironomidae) 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 aré Fis/,crie: 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, K1A 0S9. Remittances must be in advance. payable in Canadian funds to the order of the Receiver General for Canada. Editor and Director STEVENSON, PH.D. of Scientific Information J. C. Deputy Editor J. WATSON, PH.D. JOHANNA M. REINHART, M.SC. Assistant Editors D. G. COOK, PH.D. J. CAMP Production-Documentation MONA SMITH MICKEY LEWIS Department of Fisheries and the Environment Fisheries and Marine Service Scientific Information and Publications Branch Ottawa, Canada KIA 0E6 BULLETIN 195 Reuision of }^t^drebaeyrus Zrissacladius ,Z"alutschta paratrtssacladtus and some related genera (Diptera: Chironomidae) Ole A. Saether Department of Fisheries and the Environment Fisheries and Marine Service Freshwater Institute Winnipeg, Man. R3T 2N6 DEPARTMENT OF FISHERIES AND THE ENVIRONMENT FISHERIES AND MARINE SERVICE Ottawa 1976 a. I N ■ 9 5 © Minister of Supply and Services Canada 1976 Available by mail from: Printing and Publishing Supply and Services Canada Ottawa, Canada K1A 0S9 or through your bookseller A deposit copy of this publication is also available for reference in public libraries across Canada Canada: $7.00 Catalogue No. Fs94-195 Other countries: $8.40 ISBN 0-660-00974-9 Price subject to change without notice Ottawa •Coyer design by Christine Ettisk• Contents 1 ABSTRACT/RÉSUMÉ 3 INTRODUCTION 4 METHODOLOGY 5 MORPHOLOGY 9 PHYLETIC RELATIONSHIPS 9 Relationships of the subfamily Orthocladiinae 13 Previous position of the treated genera 14 The anagenesis or evolutionary progress 23 Kladogenesis or phylogenetic branching 33 KEY TO SOME GENERA OF ORTHOCLADIINAE 33 Key to males of some genera of Orthocladiinae 35 Key to females of some genera of Orthocladiinae 37 Key to pupae of some genera of Orthocladiinae 39 Key to larvae of some genera of Orthocladiinae 41 Baeoctenus GEN.N. 48 Oliveria GEN.N. 54 Hydrobaenus FRIES 1830 80 H. conformis group 93 H. lapponicus group 104 H. lugubris group 114 H. pilipes group 114 H. pilipes subgroup 138 H. distylus subgroup 158 H. calvescens sp.n. 160 H. tumidistylus sp.n. 163 Trissocladius KIEFFER 1908 173 Zalutschia LIPINA 1939 194 Z. mucronata group 206 Z. tatrica group 219 Z. vockerothi sp.n. 222 Z. tornetraeskensis group 250 Freemaniella GEN.N. 253 Paratrissocladius ZAVIiEL 1937 v Contents (concluded) 266 SomE SPECIES PREVIOUSLY PLACED IN OR RESEMBLING TriSSOCiadiUS KIEFFER SENSU BRUNDIN 276 ACKNOWLEDGMENTS 277 REFERENCES 284 INDEX vi 1 Abstract SFETHER, O. A. 1976. Revision of Hydrobaenus, Trissocladius, Zalutschia, Paratrissocladius, and some related genera (Diptera: Chironomidae). Bull. Fish, Res. Board Can. 195: 287 p. Taxonomic revisions of the genera Hydrobaenus Fries, Trissocladius Kieff., Zalutschia Lipina, Paratrissocladius Zavel, and some of their closest relatives are ' presented, along with ecological comments and distribution records. Methodology and morphology are briefly discussed. The phyletic relationships of some subfamilies of Chironomidae are discussed, suggesting that Orthocladiinae and Chironominae combined form the sister group of the proposed new subfamily Prodiamesinae, and that these three subfamilies combined are a monophyletic group. The anagenesis and kladogenesis of the treated genera suggest that Zalutschia Lip. forms the apomorphic sister group of Trissocladius Kief f., that Oliveria gen.n. forms the sister group of Hydrobaenus Fries comb.n., and the four genera combined are possibly a monophyletic group. Baeoctenus gen.n. and Freemaniella gen.n. are closely related to the above genera. Sixteen new species and two new subspecies are described: Baeoctenus bicolor, Hydrobaenus laticaudus, Hydrobaenus scapulapilosus, Hydrobaenus virgo, Hydrobaenus hudsoni, Hydrobaenus martini, Hydrobaenus pilipodex, Hydrobaenus spinnatis, Hydrobaenus calvescens, Hydrobaenus tumidisty lus, Hydrobaenus conformis Zabradorensis, Zalutschia furcarea, Zalutschia vockerothi, Zalutschia Zingulata, Zalutschia lingulata pauca, Zalutschia pusa, Zatutschia trigonacies and Bryophaenocladius impectinus. In addition, 25 species have been redescribed. Both descriptions and redescriptions include male, female, pupa, and larva whenever possible. Six new synonyms are given: Hydrobaenus pilipes (Mall.) (syn. Trissocladius grandis Kieff.), Hydrobaenus johannseni (Subl.) (syn. Trissocladius domus (Subl.) and Trissocladius hamiltoni Saeth.), Paratrisso- cladius excerptus (Walk.) comb.n. (syn. Paratrissocladius fluviatilis (Goetgh.), Bryophaenocladius Thien. (syn. Clinocladius Subl. and Cantomyia Rob.). Most redescribed species are in new generic combinations. Résumé SPETHER, O. A. 1976. Revision of Hydrobaenus, Trissocladius, Zalutschia, Paratrissocladius, and some related genera (Diptera: Chironomiade). Bull. Fish. Res. Board Can. 195: 287 p. L'auteur présente une révision taxonomique des genres Hydrobaenus Fries, Trisso- cladius Kieff., Zalutschia Lipina, Paratrissocladius Zavl.el et de quelques-uns de leurs 2 plus proches parents, avec commentaires écologiques et mentions de leur répartition. Il discute brièvement de la méthodologie et de la morphologie. Il examine les relations phylétiques de quelques sous-familles de Chironomidae, relations qui semblent indiquer que les Orthocladiinae et les Chironominae se combinent pour former le groupe soeur de la nouvelle sous-famille qu'il propose, celle des Prodiamesinae, et que ces trois sous-familles combinées sont un groupe monophylétique. L'anagénèse et la kladogénèse des genres traités donnent à croire que Zalutschia Lip. forme le groupe soeur apomorphe de Trissocladius Kieff., qu'Oliveria gen.n. forme le groupe soeur d'Hydrobaenus Fries comb.n. et que ces quatre genres sont possiblement un groupe monophylétique. Baeoctenus gen.n. et Freemaniella gen.n. sont étroitement apparentés aux genres ci-haut mentionnés. On décrit seize espèces et deux sous-espèces nouvelles: Baeoctenus bicolor, Hydrobaenus laticaudus, Hydrobaenus scapulapilosus, Hydrobaenus virgo, Hydrobaenus hudsoni, Hydrobaenus martini, Hydrobaenus pilipodex, Hydrobaenus spinnatis, Hydro- baenus calvescens, Hydrobaenus tumidistylus, Hydrobaenus conformis labradorensis, Zalutschia furcarca, Zalutschia vockerothi, Zalutschia lingu/ata, Zalutschia Ungutata pauca, Zalutschia pusa, Zalutschia trigonacies et Bryophaenocladius impectinus. De plus, on redécrit 25 espèces. Les descriptions et les redescriptions incluent toutes deux le mâle, la femelle, la pupe et la larve là où la chose est possible. L'auteur donne six synonymes nouveaux: Hydrobaenus pilipes (Mall.) (syn. Trissocladius grandis Kief f.), Hydrobaenus johannseni (Subl.) (syn. TrissocZadius domus (Subl.) et Trisso- eladius hamiltoni Saeth.), Paratrissocladius exeerptus (Walk.) comb.n. (syn. Para- trissocladius fluviatilis (Goetgh.), Bryophaenocladius Thien. (syn. Clinocladius Subl. et Cantomyia Rob.). La plupart des espèces red&crites sont placées dans de nouvelles combinaisons génériques. 3 Introduction Inadequate knowledge of the specific identities of chironomids taken in bottom samples has hitherto prevented the development of lists of indicator communities for Nearctic lakes, with the exception of the ultraoligotrophic communities described by Oliver (1963, 1964). The finding and description of all stages of several important members of oligotrophic and mesotrophic communities (Saather 1973b, 1975a, b, c) has made it possible to develop preliminary lists of indicator communities for North America (Saether 1975e). One genus containing species that are members of indicator communities in Europe is Trissocladius Kieff. sensu Brundin (Brundin 1949, 1956a, b). This genus, however, appeared to be heterogeneous, with Nearctic species as dissimilar as T. obsepta (Webb) and T. pilipes (Mall.). Futhermore, as Mozley (1970: 446) showed, the larva and pupa of Hydrobaenus Zugubris Fries were not generically distinguishable from some species of Trissocladius Kieff. sensu Brundin. The statement by Brundin (1956a: 19), originating from Goetghebuer (1914: 20), about the presence of "more or less" striated ventromental plates in Hydrobaenus Zugubris was shown to be incorrect. This made it necessary to examine the relationship of Hydrobaenus sensu Brundin to TrissocZadius sensu Brundin. Other genera and species possibly related to Trisso- cladius sensu Brundin have also been examined. The opinion that there are probably no true Holarctic species except for arctic and subarctic chironomids (expressed for instance by Fittkau 1961: 960) has been invalidated (Winker et al. 1968). The proportion of species which are Holarctic, however, seems to vary considerably from one genus to another. In Heterotrissocladius there apparently is only one true Holarctic species, HeterotrissocZadius marcidus