National Library Bibliothèque nationale .+. of Canada du Canada Acquisitions and Direction des acquisitions el Bibliographie Services Branch des services bibliographiques 395 Wellingloo Street 395. rue Wellington Ottawa. Ontario Ottawa (Onlano) K1AON4 K1AON4 NOTICE AVIS The quality of this microform is La qualité de cette microforme heavily dependent upon the dépend grandement de la qualité quality of the original thesis de la thèse soumise au submitted for microfilming. microfilmage. Nous avons tout Every effort has been made to fait pour assurer une qualité ensure the highest quality of supérieure de reproduction. reproduction possible. If pages are missing, contact the S'il manque des pages, veuillez university which granted the communiquer avec l'université degree. qui a conféré le grade. Some pages may have indistinct La qualité d'impression de print especially if the original certaines pages peut laisser à pages were typed with a poor désirer, surtout si les pages typewriter ribbon or if the originales ont été university sent us an inferior dactylographiées à l'aide d'un photocopy. ruban usé ou si l'université nous a fait parvenir une photocopie de qualité inférieure. Reproduction in full or in part of La reproduction, même partielle, this microform is governed by de cette microforme est soumise the Canadian Copyright Act, à la Loi canadienne sur le droit R.S.C. 1970, c. C·30, and d'auteur, SRC 1970, c. C-30, et subsequent amendments. ses amendements subséquents. Canada PHYLOGENETIC STUDY OF APOCRITA • (HYMENOPTERA) WITH EMPHASIS ON WING VENATION by Alain Roy Department ofNatural Resource Sciences McGill University, Montreal • May 1995 A thesis submitted to the Faculty ofGraduate Studies and Research in partial fulfilment ofthe degree requirements ofthe degree of Master ofScience. © Alain Roy, 1995 • National Library Bibliothèque nationale .+. of Canada du Canada Acquisitions and Direction des acquisitions et Bibliographie Services Branch des services bibliographiques 395 Wellinglon Street 395, rue Wellington Ottawa. Ontario Ottawa (Ontano) K1AON4 K1AON4 rour frlo VOtre rofdrence Our '110 No/rlJ r~'drenco The author has granted an L'auteur a accordé une licence irrevocable non-exclusive licence irrévocable et non exclusive allowing the National Library of permettant à la Bibliothèque Canada .to reproduce, loan, nationale du Canada de distribute or sell copies of reproduire, prêter, distribuer ou hisjher thesis by any means and vendre des copies de sa thèse in any form or format, making de quelque manière et sous this thesis available to interested quelque forme que ce soit pour persons. mettre des exemplaires de cette thèse à la disposition des personnes intéressées. The author retains ownership of' L'auteur conserve la propriété du the copyright in hisjher thesis.' droit d'auteur qui protège sa Neither the thesis nor substantial thèse. Ni la thèse ni des extraits extracts from it may be printed or substantiels de celle·ci ne otherwise reproduced without doivent être imprimés ou hisjher permission. autrement reproduits sans son autorisation. ISBN 0-612-12264-6 Canada • Short title: Phylogeny of Apocrita (Hymenoptera) with emphasis on wing venation • • • ABSTRACT: A phylogenetic character analysis ofthe wing vcnation within Apocrila (Hymenoptera) was conducted in order to examine thc phylogenctie signil1cance ofwing characters. The venational data were then combincd with publishcd studies ofother character suites to produce a hypothesis of phylogcnetic relationships among apocritan superfamilies and fal11ilies. Whereas thc c1adograms obtained from the collection ofpapers on character suites provide strong support for the sister group relationship between Orussidae and Apocrila, the ones derived from wing venation did not establish such a relationship, although they provided more support for relationships within Apocrita. A ground plan is proposed for the wing venation ofeach taxon exal11incd; for Apocrita, it is close to the groundplan established for Aculeata. The variability ofwing venation is higher between and within apocritan taxa than in SYl11phyta. This research confirms that groupings such as the Symphyta, Siricoidea, Evanioidea • and possibly the Proctotrupoidea are artificial. • 1 RÉSUMÉ: • Cette étude compare la nervation alaire des taxons d'Hyménoptères apocrites ainsi que des Symphytes afin d'en établir l'importance phylogènétique. Ces données sur la nervation alaire permirent d'établir différents arbres phylogénétiques qui furent comparés à ceux provenant d'autres groupes de caractères. Ces données, une fois combinées, ont permis d'établir une hypothèse globale basée sur des groupes de caractères. Bien que ces derniers produisent des résultats permettant de supporter la relation groupe soeur entre Orussidae et Apocrita, les cladogrames dérivant des matrices de données sur la nervation alaire possèdent une meilleure résolution parmi les familles d'apocrites. Les caractéristiques de base des ailes des différents groupes sont décrites et celles des Apocrites demeurent très proches de celles des Aculéates. La nervation alaire varie plus chez les Apocrites que chez les Symphytes. Les résultats de cette recherche 60nfirment que les regroupements comme les Symphytes, Siricoidea, • les Evanioidea et possiblementies Proctotrupoidea sont artificiels. • II ACKNOWLEOGMENTS: • 1 wish to recognize Dr. D.J. Lewis (Departmcnt ofNatural Resourcc Sciences) and especially Dr. M.J. Sharkey for tlleir supervision during the projee!. 1 wish to thank Dr. C.C. Hsiung for making the Lyman Museum Collection available for study and Drs. J.T. Huber, G.A.P. Gibson, H. Goulet and L. Masner ofthe Center for Land and Biological Resourees Researeh for providing me with specimens from the Canadian National Colleetion and for the useful information they provided on different hymenopteran taxa. 1 aiso wish to thank Drs. D. Camlean (University ofCalifornia at Davis), J.M. Carpenter (Ameriean Museum ofNatural I-listory), Ml'. M. Huben, L.S. Kimsey (University ofCalifomia at Davis), F. Ronquist (Uppsala University) and S.R. Shaw (university ofWyoming) for the information they provided on their • taxa ofspecialization. 1 am indebted to Dr. P.M. Sanborne who originally accepted me for graduate studies and supported the research through a researeh tùnd from McGili University. Dr. T. Wheeler was very helpful in making numerous corrections and suggestions during the writing ofthis thesis. Valuable suggestions were also given by Ml'. A. Aranguren and Ms. A. Audet on various parts ofthe tex!. • III TABLE OF CONTENTS: • Page ABSTRACT · . .. .. .. l RÉSUMÉ ·. .. .. ..-. II ACKNOWLEDGMENTS .............. .. III LIST OF TABLES ... ... ... .. .. VI LIST OF FIGURES ................. VII INTRODUCTION . ..... .. 1 LITERATURE REVIEW ........ 7 • Analysis of character suites 7 Willg v.enation . 12 Relationships of apocritan taxa 13 MATERIALS AND MET~ODS o' 17 RESULTS AND DISCUSSION 23 Character analysis, fore wing 23 Character analysis, hind wing 33 Cladistic analysis 39 CONCLUSION · . .... .. .. .. 53 • REFERENCES 56 • APPENDIX l Wing drawings 65 APPENDIX 2 Analysis of non-wing characters 85 APPENDIX 3 Analysis of the combined data 113 • • y LIST OF TABLES: • Page Table 1. Taxa recognized by Huber (1993) 2 Table 2. Wing characters of Apocrita (with state of the veins indicated) •••••••• 42 Table 3. wing characters of Apocrita (with presence or absence of the veins only) •••• •• 45 Table 4. Data matrix of non-wing character suites of Apocrita ••• . .. ... .. 108 Table 5. Data matr.ix of combined data on character suites .. .. ... 114 • LIST QF FIGURES: • Page Fig.l Consensus cladogram of unweighted characters from data matrix 1 43 Fig.2 Consensus cladogram of weighted characters from data matrix 1 44 Fig.3 Consensus cladogram of unweighted characters from data matrix 2 ..... .... 46 Fig.4 Consensus cladogram of weighted characters from data matrix 2 47 Fig.5 Ground plan of the wing venation of Xyelidae • 66 Fig.6 Ground plan of the wing venation of Megalodontoidea 67 Fig.7 Ground plan of the wing venation of Tenthredinoidea .... 67 Fig.8. Ground plan of the wing venation of Siricidae 68 Fig.9 Ground plan of the wing venation of Xiphydriidae ..... ..,.. 68 • Fig.10 Ground plan of the wing venation of • Cephidae 69 Fig.ll Ground plan of the wing venation of Orussidae 69 Fig.12 Ground plan of the wing venation of Aculeata 70 Fig.13 Ground plan of the wing venation of Braconidae 71 Fig.14 Ground plan of the wing venation of Ichneumonidae 71 Fig.15 Ground plan of the wing venation of • Trigonalidae 72 Fig.l6 Ground plan of the wing venation of Stephanidae 72 Fig.l7 Ground plan of the wing venation of Evaniidae 73 Fig.l8 Ground plan of the wing venation of Megalyridae 73 Fig.l9 Ground plan of the wing venation of • Aulacidae 74 Fig.20 Ground plan of the wing venation of • Gasteruptiidae 74 Fig.21 Ground plan of the wing venation of Cynipidae 75 Fig.22 Ground plan of the wing venation of Ibaliidae 75 Fig.23 Ground plan of the wing venation of Figitidae 76 Fig.24 Ground plan of the wing venation of Liopteridae 76 Fig.25 Ground plan of the wing venation of • Charipidae 77 Fig.26 Ground plan of the wing venation of Eucoilidae 77 Fig.27 Ground plan of the wing venation of Ceraphronidae 78 Fig.28 Ground plan of the wing venation of Megaspilidae 78 Fig.29 Ground plan of the wing venation of • Chalcidoidea 79 Fig.30 Ground plan of the wing venation of • Diapriidae 79 Fig.31 Ground plan of the wing venation of Vanhorniidae 80 Fig.32 Ground plan of the wing venation of