CANTHONINI (SCARABAEIDAE: SCARABAEINAE) OF GONDWANA: COMPARISON OF MALE GENITALIA, EVOLUTIONARY TRENDS AND PHYLOGENY BY CLAUDIA ALEJANDRA MEDINA URIBE SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE DOCTOR OF PHILOSOPHY (ENTOMOLOGY) DEPARTMENT OF ZOOLOGY AND ENTOMOLOGY FACULTY OF NATURAL AND AGRICULTURAL SCIENCES UNIVERSITY OF PRETORIA To my brother Jesús Andrés Medina Uribe (1978 – 2012) “Three things cannot be long hidden: the sun, the moon, and the truth” Buddha “Hay tres cosas que no pasan mucho tiempo ocultas: el sol, la luna y la verdad” Buda “The impassive skies were neutral, empty, still. Then something in the inscrutable darkness stirred; A nameless movement, an unthought Idea Insistent, dissatisfied, without an aim, something that wished but knew not how to be, teased the Inconscient to wake Ignorance. A throe that came and left a quivering trace, gave room for an old tired want unfilled, at peace in its subconscient moonless caveto raise its head and look for absent light, straining closed eyes of vanished memory, like one who searches for a bygone self and only meets the corpse of his desire” Savitri Canto 1 The Book of Beginning Sri Aurobindo “Los cielos impasibles eran neutros, vacíos, inmóviles entonces algo se movió en la tiniebla inescrutable; un movimiento sin nombre, una Idea impensada, insistente, insatisfecha, sin objeto, algo que sin saber como anhelaba ser, hostigo al Inconsciente para despertar la Ignorancia. Una angustia que vino y dejó una trémula huella dio lugar a que una vieja necesidad cansada e insatisfecha, en paz en su caverna subconsciente sin luna, alzara la cabeza y fuera en pos de una luz ausente, esforzando los ojos cerrados de una memoria desvanecida, como alguien que busca lo que fue en otro tiempo y solo encuentra el cadáver de su deseo” Savitri Canto 1 El libro de los comienzos Sri Aurobindo TABLE OF CONTENTS Table of Contents I Acknowledgements IV Agradecimientos VIII General Summary XII Resumen General XIV General Introduction (1–21) Systematics, Ecological, and Evolutionary Background 1 American Canthonines 5 African Canthonines 9 Madagascar Canthonines 11 Australian Canthonines 13 Oriental Canthonines 15 New Zealand Canthonines 16 The taxonomic status of the subfamily Scarabaeinae 17 Thesis Layout 20 References 21 Chapter I: Morphology and terminology of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) male genitalia (31–53) Abstract 32 Resumen 32 Introduction 32 Methods and materials 34 Results and discussion 35 Morphology of male genitalia in Scarabaeinae 35 Acknowledgements 51 References cited 51 I Chapter II: Systematics of the southern African genus Epirinus Reiche (Coleoptera: Scarabaeinae: Canthonini): descriptions of new species and phylogeny (54–70) Introduction 55 Material and Methods 55 Key to Species of Epirinus (modified from Scholtz and Howden 1987) 58 Epirinus pseudorugosos sp. n. 59 Epirinus hluhluwensis sp. n. 60 Epirinis ngomae sp. n. 61 Epirinus aquilus sp.n. 61 Epirinus sebastiani sp. n. 62 Epirinis minimus sp. n 63 Phylogenetic analysis of the genus Epirinus 64 List of characters and their states used in the cladistic analysis 65 Results of cladistic analysis 68 Acknowledgements 69 References 69 Chapter III: Evolutionary trends and phylogeny of the tribe Canthonini (71– 85) Abstract 71 Introduction 72 Historical aspects of the division and classification of the tribe Canthonini 72 Methods 74 Results and Discussion 75 Clade 1. Tribe Byrrhidiini new tribe 76 Diagnosis 76 Description 76 Type genus 77 Distribution 77 Remarks 77 II Clade 2. Circelliini new tribe 77 Diagnosis 78 Description 78 Type genus 79 Distribution 79 Remarks 79 Clade 3. Tribe Canthonini 79 Clade 4. Other tribes 80 Canthonini and Deltochilini: Are they different tribes? 81 Conclusions 84 General Conclusions 88 References 91 Appendices (96–126) Appendix 1. List of species examined 96 Appendix 2. List of morphological characters used for the phylogenetic analysis one. 105 Appendix 3. Matrix used for phylogenetic analysis one. 114 Appendix 4. List of morphological characters used for the phylogenetic analysis two. 119 Appendix 5. Matrix used for phylogenetic analysis two. 124 III ACKNOWLEDGMENTS Even though the Acknowledgments appear at the beginning of the text, they have to be written at the end, after the work is done. I am very happy finally to say “thank you” to so many people who have made possible the culmination of this work. I chose the long path to my Ph.D. My doctoral thesis went through many different stages over many years, and so many people encouraged, advised, and supported me physically and emotionally during this period of my life. The order in which I mention professors, colleagues, friends, family and institutions does not necessarily reflect the importance or quality of their help. The list is long, and I trust that I’ve left no one out. First, I want to generously thank my director, Prof. Clarke Scholtz, mainly for his patience with a somehow difficult student like I was. From the beginning he was enthusiastic about the project and his agile and effective diligence made possible and eased my arrival to African lands. I am grateful for his confidence in the project, his advice and his unconditional support. I also want to thank Prof. Chris Chimimba, Director of the Department of Zoology and Entomology of the University of Pretoria, for his crucial support in the completion of this thesis. I also received decisive support from Juan Carlos Bello, Coordinator of the Gestión de Información y Conocimiento Program of the Alexander von Humboldt Institute; his absolute support was important in the culmination of my thesis. Bruce Gill has been a very important influence on my evolving knowledge of dung beetles. When I finally came to realize the dimensions of my chosen thesis project, I understood why it was so difficult for me to convince him of my idea. He tried to persuade me to develop a more modest project, but Bruce! My stubbornness was stronger! Thanks for all your help! IV The time I spent at the University of Pretoria was stimulating and meaningful in my career as an entomologist, thanks in large part to the interaction with Vasily Grebennikov. From him, I learned enormously about the traditionally European style of pinning insects and storing specimens. He taught me the genitalia cleaning technique and helped me to improve my sketches. Vasily was a good partner and friend during my stay in Pretoria. My research greatly benefitted from his experience. Thank you, Vasia, for the “tea club”, the pizza and the chocolates at the end of the day! The interaction and friendship with Arturo González and Fredy Molano has been crucial to the development of my ideas about Canthonini tribe systematics. We shared many opinions and long working sessions. I am in debt to Arturo for his support during my leave of absence, and his valuable help in the revision and edition of the final draft of the thesis. The persistence of Fernando Vaz de Mello in the study of tropical dung beetles provided me with inspiration for the progress of the thesis. We discussed the evolution of the subfamily and shared the discoveries of male genitalia sclerites. I thank him for his answers to my many questions, specimen donation, as well as for critical literature. Ruth Müller, from the Transvaal Museum, supported my work in different ways; She made my stay at the collection easier and more pleasant and also facilitated the specimen loans. Ruth generously welcomed me to her home when I needed it; her hospitality was unlimited. I express my gratitude to Riaan Stals (National Collection, South Africa), for the loan of specimens. For the same reason I am also in debt to G. Montheith (Queensland Museum, Australia), Enrico Barbero (Torino University, Italy), Roig-Nuñet (IADIZA, Argentina), Francois Génier (Canadian Museum of Nature, Canada), Estella Monteresinos (Río Cuarto, Argentina), Patricia González (Republic University, Uruguay), and Eric Matthews (South Australian Museum, Australia). Several of them also donated specimens for genitalia dissections. V I am very grateful to Prof. Mario Zunino with whom I shared ideas about the evolution of beetle’s genitalia and for his help in reviewing the chapter on genitalia variation. I am also grateful to Keith Philips for his advice in cladistic analysis. Experiences exchanged with Shaun Forgie, Catherine Sole, Christian Deschodt, Adrian Davis and Ute Kryger were very rewarding for my study. I would also like to thank Ingrid Vis, the secretary of the Department of Zoology and Entomology for her logistic support. Also I am in debt to Carmen Jacobs and Cathy Barnard for all her help with the final process of the thesis. So many colleagues and friends helped me with this project in different ways. Thanks to: Alberto Ballerio, Xiomara Villalba, Johann Cárdenas, Diana Espitia, Astrid Pulido, Alejandro Lopera, Fernando Fernández, Mila Coca, and Rafael Miranda. John Miles, an unconditional friend, has contributed enormously to my academic formation with good books and discussions about systematics and evolution. I want to thank him for his constant interest in my academic projects and his valuable help in the editorial review of the final document. I am in debt to family -- my mother Rosa Matilde Uribe and my father Jairo Medina -- for their unconditional support. Their constant love and presence during my stay in South Africa stimulated me to persevere. For a long period of time during my doctorate studies, I was a nomad, with no permanent place to call home, so I lived with relatives and friends; I am almost certain that old thesis files still remain in their computers. Thanks to my brothers and sisters and their families: Felipe, Cecilia, Lila, Carolina, Andres, Jhon Jairo and Juan David. Thanks to my grandmother Rosita and my aunt Paty, for all their love and constant support. To Clara Uribe and her family for their hospitality in Popayán.