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The Elucidation of the &Ina M D a Complex THEELUCIDATION OF THE &INA MDACOMPLEX Raoul HarIey Bain A thesis submitted in confonnity with the requirements for the degree of Master of Science Graduate Department of Zoology University of Toronto O Copyright by Raoul Harley Bain 1998 National Library Bibliothèque nationale 191 ofCanada du Canada Acquisitions and Acquisitions et Bibliographic Services services bibliographiques 395 Wellington Street 395, rue Wellington Ottawa ON KIA ON4 Ottawa ON K1A ON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/fïlm, de reproduction sur papier ou sur format électronique. The author retauis ownersship 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. The Elucidation of the Rma livida Complex Raoul Harley Bain, Master of Science Degree, 1998 Department of Zoology, University of Toronto ABSTRACT 1 investigated the suspected polytypic nature of the green cascade frog, Rana livida, from southeast Asia with a multi-step approach. My initial anatomical study of specimens from Vietnam resulted in the recognition of three distinct species and five morphological groups. I * tested the strength of these groupings with morphometrics and cellular DNA content analysis. Congruence among the results of morphology, morphometrics, and cellular DNA content, led me to suspect the presence of further cryptic species. 1 tested this hypothesis with allozyme electrophoresis, resolving 15 presumptive loci. Fixed allelic differences suggested the presence of four additional species. They also suggested that one of the new species may be two distinct species, but the evidence collected here is equivocal. 1 elucidated seven species of the Rana livida complex. 1believe that much more cryptic diversity of this complex and other cascade ranids await discovery. This highlights the need for rigorous taxonomic and phylogenetic analyses of dl cascade ranids. ACKNOWLEDGMENTS 1 would like to first thank my supervisor Bob Murphy. As rnost know, one is not part of the Murphy lab, it is part of you. 1thank him for al1 the mernorable experiences and lessons taught (some knowingly, most not). I especially thank Bob for ROMMY. Though, admittedly hesitant at first, it became more and more apparent to me, as we sweated in the Pianetariurn, that it would be worth every drop (Oh!). I would also like to thank rny cornmittee, Bob, Doug Currie, Hans Sues, and Dan Brooks. The cornments that Ham and Dan lent to earlier drafts helped tremendously. There are many whose technical help 1 gratefully acknowledge. For discussions of things morphometric, I would like to thank Pedro Peres-Neto, Paul Galpern, and Debra Yunnan. For teaching me allozyme electrophoresis, I thank Ross MacCulIoch, Darlene Upton, Jinzhong Fu, and Bob. 1thank Ross for performing the DNA Content analyses. I thank Amy Lathrop for teaching me how to x-ray. Valuable assistance was also given in the fonn of translations. For this I thank Chun-mei Huang, Jinzhong Fu, Wu Xiao-Chun, Johann Lindell, Nasreen Rahman, and Amy's dictionaries. 1 thank Tanya Trepanier for allowing me to use her laser pnnter. 1 also thank Bob and Amy for allowing me to use their photographs in this thesis and in talks I have given. 1 would also like to thank Linda Ford of the Amencan Museum of Natural History for her assistance. 1would like to thank some colleagues in Vietnam. First, 1 thank Prof. Dr. Cao van Sung, Director of the Instiîute of Ecology and Biological Resources (IEBR), Hanoi for dl of his efforts. For invaluable field assistance, 1 would Mce to thank Dang Tuan Dat, vice director of the Institute of Epidemiology, Ban Ma Thuot, Pham Duc Tien (IEBR),and, of course Ms. Ho Thu Cuc also of the IEBR. I also thank Nikolai Orlov, Brad Hubley dong with Amy, Bob, Doug, and students of the BI0 308 field course who helped collect specimens. 1 wish to pointedly ignore acknowledging mefloquine. I am lucky to have been part of the ROM. There are many who make this a great place and 1 am grateful to have met thern. For their humour and wisdom 1 would like to thank Doug Cume, Jon Barlow, Chris Darling, Burton Lim, Josh Feltharn, Alison Stuart, Andy Bennett, John Swann, Marty Rouse, Judy Edwards-Davies, Antonia Guidotti, Colette Baril, and Kevin Doyle. In a past life at the Museum, I was involved with Palaeobiology. For tremendous experiences and special times with that department, 1 would tmly like to thank, Tan Nicklin, Ted Ecclestone, Tim Fedak, Ryosuke Motani, Kevin Seymour, Catherine Skrabec, Hans Sues, MicheIe Bobyn, and Ian Momson. I would also like to make special mention to Chris McGowan for his support, his boyish awe, and devilish humour. 1 would like to acknowledge my friends, though I will surely Ieave some out. From the Department of Zoology 1 thank Amaya Ortigosa, Paul GaIpern, Ian Dworkin, Pedro Peres-Neto, Nasreen Rahman, Fernando Portella de Luna Marques, Kristy Chna, and Sarah Kalhok. Liz Tudor-Mulroney was exceptionally helpful in the grad office. A special thank- you to my lab mates, past and present: Darlene Upton, Jinzhong Fu, and Amy Lathrop for great times in Our various office-cubicIes around the ROM. Heartfelt thanks to Diana Dobson, Ted Ecclestone, Tim Fedak, Kate Holmes, Blair Klayman, Mike Leibovici, Amy Meckler, Jory Nash, Andrew Pifko, Max Westhead, Deana Vardy, and Ilana Zylbennan for their support and encouragement (late night or otherwise). 1 thank my loving family. My sister Hilary and rny brothers Cale and Zachary aiways help keep things in perspective and ensure that part of me stays twelve years old. My parents, though not dways understanding why ("You're going where? To do what?), have aIways supported me in the end ("lt would give us so much pleasure if you would let us buy you a pair of hiking boots"). 1 am very thanldul. Of course, 1know that none of this would be possible without my grandparents. Final thanks to Sarah Kalhok for al1 of her love and help. She has been my biggest booster, my master typesetter, and my best critic the whole way through. This project was supported in part by two University of Toronto Special Open Masters Fellowships, and two Department of Zoology Pure and Applied Sciences Grants. Field and lab work was supported by an NSERC (A3 148) and ROM Foundation Grants to Robert W. Murphy as well as an American Museum of Natural History Collection Study Grant to myself. This thesis was catered by Blaze Catering. Now to the very heart of wonder. Because species diversity was created prior to humanity, and because we evolved within it, we have never fathomed its limits. As a consequence, the living world is the naturd domain of the most restless and paradoxicd part of the human spirit. Our sense of wonder grows exponentially .. E.O.Wilson vii TABLE OF CONTENTS THE ELUCIDATION OF THE RANA LlVIDA CQMPLEX ........................................ i .. ABSTRACT...... ..... ... .. ..... .................... .. ... .. ... ............ .. ..... .. ... ..... .*. .. .....a. ... .. .m... .. m.. ....... .. lI .. ACKNOWLEDGMENTS .. * .. .. .. ........ TABLE OF CONTENTS. .... ..... ............................... ............... ....................... ............ viii .. LIST OF TABLES ........... .... ... ........ .... .. ...... .. ... ...... .. ... .. ...... .. ... ..... ... ... .. ...... ...................... XII LIST OF FIGURES .. .. .. ... .. .. .. .. .. .. .. .. .. .. .. .. .. xiv LIST OF APPENDICES.... .. .. .. .. .. .. -. .. xv CHAPTER 1: RANA LIVTDA AND PTS IMPORTANCE TO BIODIVERSITY: ......... ..., .......... 1 BACKGROUND........................... .. .......................................................................... ....... ........ 1 RANA LMDA: CONFUSIONABOUT SPECIES IDENTiTY ..................... .................... .............. 4 Rana livida............................................ ....... .................. ,... ............ ,......................... ,.,.............. 4 Rana grarninea... .. ... , . .. .. .. 6 Rana 2epof.ipes............................... ... ....................................................... ........ , ....... 6 Rana sinica ...,, ........................................................... .......................................................... 7 RANA LIVIDA: GENERICAND SUBGENERIC CONFUSION ...................................................8 Runa. .. ., .. , , . .. .. ., .. .. .. .. .. .. .. .. .. .. .. .. .9 Hylarana .. , . .. .. .. .. .. .. , . .. .. , . .. ., . .. .. .. .. .9 Odorrana .......................................................... , ..................................................................... 11 Eburana .......................,......... , .....,...... .......... ......... ............................................................... 12 RANALMDA: A SPECIES cOMPLEX? ............................... .................. ......... .... ................ .... 13 RANA LNIDA: A SPECIES COMPLEX? .....................,......*........................................... ........... 13 THISSTUDY
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