THE INTERNAI, MALE GENITALIA OF SELECTED GENERA OF MELANOPLINAE (ORTHOPTERA: ACRIDIDAE). by Spyros D. Skareas Depa rtment of Na tural Resource Sciences McGill University, Montreal June 1998 A thesis subrnitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements of the degree of Master of Science 0 Spyros D. Skartas 1998 National Library Bibliothèque nationale 1+1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Sewices senrices bibliographiques 395 Wellington Street 395. rue Wellington OmwaON KlA ON1 OnawaON K1AW Canada CaMda The author has granted a non- L'auteur a accordé une licence non exclusive licence ailowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seli reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfiche/film, de reproduction sur papier ou sur fonnat électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fkom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. ABSTRACT The internai male genitdia of selected genera of Melanoplinae (Onhoptera: Acndidae). M.Sc. Spyros D. Skareas Natural Resource Sciences The morphology of the intemal male genitaiia of selected genera of Melanoplinae (Onhoptera: Acrididae) was studied. A detailed cornparison of these structures was carried out, in an attempt to identiQ new characters that may be usehl in analyzing relationships between the members of the subfamily. Twenty two genera were examined, mostly of North American distribution, using one exernplar species for each of them. Standard dissection and drawing techniques were followed. The epiphallus and the apical parts of the aedeagus were found to be the most important taxonomie characters. The genera Apfenopedes and Buckeiîacris were highly divergent from the rest of the melanoplines, while the close relationship between Appalachia, Dendrotettix and Podisma was confirmed by genital characters. Internai male genitalic characters did not provide . significant support for many currendy recognized tribes and subtribes, suggesting that there is a great need for reclassification in the subfarnily. F~ÉSUME Les structures génitales mâles des genres sélectionnés de la sous-famille Mélanoplinae (Orthoptera: Acrididae) M.Sc. Spyros D.Skareas Sciences des Ressources Naturelles La morphologie des genitalia mle des genres sélectionnés de la sous-famille Mélanoplinae a été réalisée, afin de permettre d'identifier de nouveaux charactères qui pourraient être utiles dans l'analyse des liens entre les membres de cette sous-famille. Une espèce pour chacun des vingt-deux genres a été étudiée, majoritairement de provenance nord-américaine. Des techniques de dissection et de dessin coumantes ont été utilisées. II a été conclu que I'épiphaile et les parties apicales de l'édéage sont les charactères taxonomiques les plus importants. Les genres Aptenopedes et Brtckellacris étaient très divergents du reste des Mélanoplines. Le lien étroit entre Appalachia, Dendroteztix et Pudisma a été confirmé par l'examen des charactères génitaux. Les structures génitales mâles ne fournissent pas d'appui significatif pour plusieurs tribus et sous-tribus reconnues, indiquant qu'il est nécessaire de reclassifier la sous-famille. To Dr. Mazomenos & Dr Hsiung for introducing me to behavioral and systematic Entomology AKNOWLEDGEMENTS 1 would like to thank the Lyman Entomological Museum, for providing me the space and environment to work for my research as well as most of the material for my study. Moreover 1 would like to extend my gratitude to the Academy of Natural Sciences of Philadelphia and iis cüllection manager Donald Azurna, for the loans of specimens and the excellent cooperation we had these 2 years. 1 am furthemore grateful to the Onhopterists' Society, for providing me with hnds for my research. McGill University also provided a considerable amount of money through a bursary and several fee waivers; therefore 1 would like to express rny appreciation to the Department of Natural Resource Sciences. Overall, 1 would Iike to thank my research supervisor Dr. CC.Hsiung, to whom 1 will dways be gratefbl. Without his help and support this work would have been impossible to be completed. 1 want to thank him not only because of his scientific directions, and his cooperation as a supervisor, but also for his psychologicd support and advises in several personal problems I faced. Additionally, 1 would like to thank Dr. Vickery for his contribution to my work. No matter day or hour Dr. Vickery was always willing to answer my numerous questions, to help me out and to direct me the right way. His guidance duririg my studies was extremely valuable, and 1 feel fortunate to have been able to work with one of the world's leading authorities in Orthoptera. Moreover, 1 should also thank Dr. Wheeler, for his encouragement at my first conference presentation in Edmonton, his endless comrnents and corrections to rny written texts, as well as his personal cornments and suggestions about my work, which they al1 helped me become a better systematist. Apm for the professors directly associated with my work, 1 would first like to thank Prof. Lockwood of University of Wyoring, for his personal support from the first moment of our communication. ieff was the perfect colleague, who would dedicate his time answering my melanopline-related questions and encourage me to work harder. Thanks also to Prof. Nick Papadopoulos of Carleton University and his wife Irinela for providing their endless support and valuable assistance whenever 1 needed. In addition, 1 ought to thank Patrice Bouchard, Stephanie Boucher, Scott Brooks, Paul Cermak, TilIy Gaoh, Corey Keeler, Cyrena Riley, and Rachel Wallage for being both great colleagues and friends. Thanks aiso to Prosanta Chakrabarty for his continuous encouragement. Special acknowledgments also extend to Marie-France Julien for her her assistance in the French abstract, as well as for the psychological suppon during the last stressful period before my thesis submission. Finally, 1 would like to express my deepest gratitude to my fady in Greece, for supponing me both financially and psychologically during al1 this time 1 was away fiom home. TABLE OF CONTENTS Page * - .USTRACT II RESUME DEDICATION ACKNOWLEDGMENTS TABLE OF CONTENTS LIST OF FIGWS LIST OF TABLES INTRODUCTION 1 LITERATURE REVIEW 4 Systematics of the melanoplines 4 Phylogeny 5 The interna1 male genitalia as a systematic tooI in Acrididae and Melanoplinae MATERIALS AND METHODS Material examined Dissection and drawing Abbreviations and terminology. RESULTS Anatomy and fiinction of the copulatory structures Description of phailic structures for the genera studied DISCUSSION Range and distribution of phailic characters in the subfamily Melanoplinae Phylogenetic interpretations and suggestions for reclassification in the su b famil y Future research CONCLUSIONS REFERENCES ... Vlll LIST OF FIGURES Figure Page 1. Intemal male genitalia of Podisma pedesrris 67 2. Intemal male genitdia of A rgiacris rehni 69 3. Inted male genitalia of Asemopfus montanus 71 4 Intemal male genitaiia of Bradjnotes o. obesu 73 5. Internal male genitaiia of Buckeflacrisc. chifcofinae 75 6. Internal male genitaiia of Barytetfix h. hymphreysii 77 7. Internai male genitalin of Conalcaea miguelinata 79 8. Intemal male genitaiia of Dac~lotumbicolor picium 8 1 9. Intemal male genitalia of Hesperofettix virirüs praiensis 83 10. Internal male genitaiia of Dichropftis elongatus 85 11. Internai male genitalia of Aeoplides t. tumbulli 87 12. Int emal male genitalia of Aptenopedes sphenarioides 89 13. Internal male genitaiia of Hypochfora albû 9 1 14. Internai male genitaiia of Melanopius femzrrmbrz~m 93 15. Internal male genitaiia of Paroxya clavuliger 95 16. Internal male genitalia of Phoetaliotes nebrascer~sis 97 17. Internai male genitalia of Appafachia urcana 99 18. Internal male genitaiia of BohemanellaJ fi.isda 101 19. Internal male genitalia of Booneacris g. gfucialis 103 ix Figure Page 20 Intemal male genitalia of Dendroretrix queras 105 2 1 Internal male genitalia of Primnoa primnoa 1 O7 22 Internal male genitalia of Pmmnacris rainierensis 1O9 LIST OF TABLES Table Page 1. Classification of the subfamily Melanoplinae, as proposed by Rehn and Randel1 ( 1 963). I Il 2. List of genera and exemplar species studied in the present work. 113 3. Tenninoiogy used by different authors for the interna1 male genitalia of Melanoplinae. 115 4. Comparative table of the intemal male genitalia of the melanopline genera studied. Real proportions are rnaintained for convenience in cornparison. 1 18 Since the begiming of this century, male genitalia have been widely used as a useful tool in systematics in most insect orders (Eberhard 1985). However in grasshoppers, it was not until the middle 1930's that systematists included the copulatory structures in their studies. Especially after Dirsh (1956) who proposed a new higher classification in Acridoidea based on the phallic cornplex, more and more orthopterists adopted the inclusion of such structures in taxonomie papers, and today this is considered a standard procedure for