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Song Dissertation SYSTEMATICS OF CYRTACANTHACRIDINAE (ORTHOPTERA: ACRIDIDAE) WITH A FOCUS ON THE GENUS SCHISTOCERCA STÅL 1873: EVOLUTION OF LOCUST PHASE POLYPHENISM AND STUDY OF INSECT GENITALIA DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Hojun Song, M.S. ***** The Ohio State University 2006 Dissertation Committee: Approved by Dr. John W. Wenzel, Advisor Dr. Norman F. Johnson ______________________________ Dr. Johannes S. H. Klompen Advisor Graduate Program in Entomology Copyright by Hojun Song 2006 ABSTRACT The systematics of Cyrtacanthacridinae (Orthoptera: Acrididae) is investigated to study the evolution of locust phase polyphenism, biogeography, and the evolution of male genitalia. In Chapter Two, I present a comprehensive taxonomic synopsis of the genus Schistocerca Stål. I review the taxonomic history, include an identification key to species, revise the species concepts of six species and describe a new species. In Chapter Three, I present a morphological phylogeny of Schistocerca, focusing on the biogeography. The phylogeny places the desert locust S. gregaria deep within the New World clade, suggesting that the desert locust originated from the New World. In Chapter Four, I review the systematics of Cyrtacanthacridinae and present a phylogeny based on morphology. Evolution of taxonomically important characters is investigated using a character optimization analysis. The biogeography of the subfamily is also addressed. In Chapter Five, I present a comprehensive review the recent advances in the study of locust phase polyphenism from various disciplines. The review reveals that locust phase polyphenism is a complex phenomenon consisting of numerous density-dependent phenotypically plastic traits. The evolution of locust phase polyphenism is investigated from a phylogenetic perspective. I show that different components of locust phase polyphenism evolved independently and often phylogenetically conserved. In Chapter ii Six, I address some philosophical issues in the study of genital evolution. I show that the fundamental assumptions in the field of genital evolution are often violated and need to be tested more rigorously. iii To Vitaly Mikhailovich Dirsh (1904-1982), An acridologist and a member of the phallic cult What the chewing locust left, the swarming locust has eaten; What the swarming locust left, the crawling locust has eaten; And what the crawling locust left, the consuming locust has eaten (Joel 1:4, NKJV) iv ACKNOWLEDGMENTS My childhood dream was to be an entomologist. This dream has now become a reality and I owe it all to my mentors, friends, colleagues, and family. First and foremost, I would like to sincerely thank my advisor, Dr. John W. Wenzel, for helping me to become a scientist. He is the best mentor I have ever met, always willing to spare his time and always looking out for students’ best interest. After six years under his guidance, I sound a lot like him and I am proud of it! I would also like to thank the members of my committee, Dr. Norman F. Johnson and Dr. Hans Klompen for providing guidance and support throughout my graduate career. In particular, I am grateful to Dr. Johnson for allowing me to use the collection and for discussions about taxonomy in general. My understanding of grasshoppers and locusts improved greatly thanks to Dr. Greg Sword. He welcomed me into his arms (in Sidney, Montana) when I was a budding graduate student and he has continued to encourage me throughout my career. Dr. Theodore J. Cohn is one of the most generous and kind orthopterists I have ever known. Whenever I needed help in pursuing my study, he was willing to help. I would like to express my gratitude for the late Dr. Reginald F. Chapman. He was the one who was v truly interested in the evolution of Schistocerca and helped me shaping my ideas in many ways. He will be immensely missed. I am grateful to my lab mates during my stay, Marc Branham, Sibyl Bucheli, Ryan Caesar, Eric Dotseth, Todd Gilligan, Chi-Feng Lee, Kurt Pickett, and Joe Raczkowski, and for putting up with my insanity. I am especially thankful to my office mate Sibyl for taking care of me like a brother. You made my stay ever more enjoyable. I also like to thank the members of the Museum of Biological Diversity for stimulating discussion and friendship. My study would not have been complete without help of the curators the museums from where I borrowed specimens. Especially, I thank Dr. Dan Otte and Dr. Jason Weintraub (ANSP) and Dr. Lacey Knowles and Mark O’Brien (UMMZ), Judith Marshall (BMNH) for allowing me to work at their collections. Dr. John Capinera provided invaluable information on the ecology of grasshoppers during the fieldwork in Florida. Department of the Army kindly allowed me to conduct a field study in Oklahoma Fort Sill. Also, I would like to thank Dr. Zenón Cano-Santana (UNAM), CONABIO, Secretaría de Marina, Armada de México for making a trip to Socorro Island possible. José Luis Castillo, Marcos Flores, Ivan Hernandez, and Enrique Arias provided invaluable assistance in the field. Dr. David B. Weissman and Dr. Ludividina Barrientos- Lozano provided help in shaping the Socorro research into a manuscript. My graduate study was supported by the National Science Foundation Graduate Research Fellowship, OSU Alumni Grants for Graduate Research and Scholarship, OSU Herta Camerer Gross Fellowship, and Tinker Field Research Grant. vi On a personal note, I want to thank my kids at KUMC for helping me realize the meaning of life. I am grateful to my parents, Young-Gil Song and Jee-Yeon Koo, for giving me a chance and supporting me throughout my life. I want to shout out to my little brother Wonjun for encouraging me. My degree is really a fruit of my family’s prayers. I am indebted to my soul mate and best friend, Haeran Park, for believing in me. After six years of studying grasshoppers and locusts, I feel like I only scratched the surface. It has been a humbling experience, but I remain hopeful because there are still many wonderful things out there for me to discover. vii VITA March 31, 1977 …………………………… Born – Seoul, Korea 2000 ………………………………………. B.S. Entomology, Cornell University 2002 ………………………………………. M.S. Entomology, The Ohio State University 2002-2005 ………………………………… N.S.F. Graduate Research Fellow, The Ohio State University PUBLICATIONS Research Publication 1. Song, H., D. B. Weissman, L. Barrientos-Lozano, and Z. Cano-Santana. In Press. Locust Island. American Entomologist. 2. Song, H. In Press. Phylogenetic perspectives on the evolution of locust phase polyphenism. Journal of Orthoptera Research. 3. Song, H. 2006. Description of Schistocerca cohni n. sp. and redescription of S. socorro (Dirsh) (Orthoptera: Acrididae: Cyrtacanthacridinae) from Mexico. Zootaxa 1150: 43-52. 4. Weissman, D. B., H. Song, and L. Barrientos-Lozano. 2004. Locust Outbreak on Socorro Island, Islas Revillagigedo, México. TecnoINTELECTO 1: 119-121. 5. Song, H. 2004. On the origin of the desert locust Schistocerca gregaria (Forskål) (Orthoptera: Acrididae: Cyrtacanthacridinae). Proceedings of the Royal Society of London B. 271: 1641-1648. 6. Song, H. 2004. Revision of the Alutacea Group of genus Schistocerca (Orthoptera: Acrididae: Cyrtacanthacridinae). Annals of the Entomological Society of America 97: 420-436. viii 7. Song, H. 2004. Post-adult emergence development of genitalic structures in Schistocerca Stål and Locusta L. (Orthoptera: Acrididae). Proceedings of the Entomological Society of Washington 106: 181-191. 8. Liebherr, J. K., and H. Song. 2002. Distinct ground beetle (Coleoptera: Carabidae) Assemblages within a New York State wetland complex. Journal of the New York Entomological Society 110: 127-141. FIELDS OF STUDY Major Field: Entomology Specialization: Systematics of Acrididae, especially Cyrtacanthacridinae. ix TABLE OF CONTENTS Abstract ………….………………………………………………………………………. ii Dedication ………….…………………………………………………………………… iv Acknowledgments ………….………………………...…………………………………. v Vita ………….…………………..…………………………………………………….. viii List of Tables …………………..……………………………………..…………..……. xv List of Figures ………………..……………………………………..…………...……. xvii Chapters: 1. Introduction ……………………………………………………………………… 1 2. Systematics of the locust genus Schistocerca Stål 1873 (Orthoptera: Acrididae: Cyrtacanthacridinae) …………………………………………………………….. 6 2.1. Introduction …………………………………………………………………. 6 2.2. Concept of the genus ……………………………………………………...… 7 2.3. Taxonomic history of the genus …………………………………………… 10 2.4. Biology and ecology of Schistocerca ……………………………………... 16 2.5. Variation of color ………………………………………………………….. 18 2.6. Identification key to Schistocerca species ………………………………… 21 2.7. A partial revision of the genus Schistocerca ………………………………. 37 2.7.1. Materials and methods ………………………………..………... 38 2.7.2. Taxonomic treatments ………………………………………..… 39 2.8. Taxon description ………………………………………………………….. 81 x 2.9. Conclusion ……………………………………………………………….. 147 3. Phylogeny, biogeography, and evolution of the locust genus Schistocerca Stål 1873 (Orthoptera: Acrididae: Cyrtacanthacridinae) ………………………….. 149 3.1. Introduction ………………………………………………………………. 149 3.2. Biogeography of Schistocerca and controversies ………………………... 150 3.3. Materials and methods ………………………………………………..….. 155 3.3.1. Taxon sampling ……………………………………………….. 155 3.3.2. Character sampling …………………………………………… 159 3.3.3. Phylogenetic analysis …………………………………………. 169 3.4. Results ……………………………………………………………………. 172 3.5. Discussion ……………………………………………………………...…
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