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Phylogenetic Relationships Within the Batagur Complex (Testudines: Emydidae: Batagurinae) Jean M

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Phylogenetic Relationships Within the Batagur Complex (Testudines: Emydidae: Batagurinae) Jean M Eastern Illinois University The Keep Masters Theses Student Theses & Publications 1993 Phylogenetic Relationships Within the Batagur Complex (Testudines: Emydidae: Batagurinae) Jean M. Capler This research is a product of the graduate program in Zoology at Eastern Illinois University. Find out more about the program. Recommended Citation Capler, Jean M., "Phylogenetic Relationships Within the Batagur Complex (Testudines: Emydidae: Batagurinae)" (1993). Masters Theses. 2114. https://thekeep.eiu.edu/theses/2114 This is brought to you for free and open access by the Student Theses & Publications at The Keep. It has been accepted for inclusion in Masters Theses by an authorized administrator of The Keep. For more information, please contact [email protected]. THESIS REPRODUCTION CERTIFICATE TO: Graduate Degree Candidat.es who have written formal theses. SUBJECT: Permi~sion to reproduce theses. The University Library is r~c;:eiving a number of requests from other institutions asklng permission to reproduce dissertations for inclusion in thelr library holdings. Although no copyr~ght laws are involved, we feel that professional courtesy demands that permission be obtained from the author before we allow theses to be copied. Please sign one of the following statements: Booth Library of Eastern Illinois University has my permission to lend my thesis to a reputable college or university for the purpose of copying it for inclusion in that institution's library or research holdings. Date I respectfully request Booth Library of Easter,n Illinois University not ~llow my thesis be reproduced because ---~~~~--~~~~~---........ Date Author m L Phylogenetic Relationships Within The Batagur Complex (Testudines: Emydidae: Batagurinae) (TITLE) BY Jean M. Capler THESIS SUBMITIED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master Of Science IN THE GRADUATE SCHOOL, EASTERN ILLINOIS UNIVERSITY CHARLESTON, ILLINOIS 1993 YEAR I HEREBY RECOMMEND THIS THESIS BE ACCEPTED AS FULFILLING 17 ~ \'\93 DA . ,. " . ~· PHYLOGENETIC RELATIONSHIPS WITHIN THE BATAGUR COMPLEX {TESTUDINES: EMYDIDAE: BATAGURINAE) ABSTRACT Relationships between 10 species of the batagurine genera Batagur, Callaqur, Kachuga, Hardella, and Morenia are discussed based on a cladistic analysis of 35 morphological characters. Ocadia sinensis (Emydidae: Batagurinae) was used as the outgroup species. Four cladograms were produced with a length of 59.0 steps and a 0.75 consistency index. In contrast with previous studies, Hardella and Morenia do not appear as a monophyletic clade. Instead, Hardella is included with the remaining ingroup taxa (exclusive of Morenia) based on five synapomorphies. The present genus Kachuga was determined to be paraphyletic, having excluded the present genera Batagur and Callagur. The pangshura subgroup of the genus (consisting of the smaller members of the genus -- K· smithi, K· tecta, and K· tentoria) is distinguished by six synapomorphies and appears to be a sister group of the clade formed by an unresolved polychotomy including Bataqur, Callagur, and the kachuga subgroup (consisting of the larger members of the genus --K. dhongoka, K· kachuga, and K· trivittata). Batagur and Callagur are included with the kachuga subgroup based on two synapomorphies. A possible taxonomic revision suggested is to elevate the pangshura group to generic rank and include Batagur, Callagur, and the remaining Kachugas as a separate genus Batagur. Dedicated to the memory of my mom, who taught me to love the natural world around me, and to my dad, who taught me to believe in myself. I love you both. ACKNOWLEDGMENTS There are numerous persons and institutions which deserve many thanks for assistance in the completion of this project. Financial support for this project was provided by the Graduate School of Eastern Illinois University through a directed research assistantship. I greatly appreciate the generous loan of study specimens from Dr. Peter c. H. Pritchard, The American Museum of Natural History, The British Museum of Natural History, and the University of Florida. I must thank Dr. John L. Carr for his invaluable insight and suggestions, and for introducing me to PAUP. Without his help, especially at the beginning of this project, I might never have gotten past the point of staring helplessly at the PAUP logo on the computer screen. Many thanks go to my committee members: Dr. Charles Costa for many hours spent drawing cladograms and making slides (I knew there was a reason I had a physiologist on my committee); Dr. Kipp Kruse for use of his computer in the initial stages of the project; and Dr. Michael Goodrich for helping me find my way through a maze of taxonomic rules and regulations. I thank all three for critically reading this dissertation and for their general support, encouragement, and helpful suggestions. Much friendship and stress-relief was provided by my friends and fellow graduate students during the completion of this project, especially Sean Jenkins, Sally Erwin, Ken Stetina, Pat Sullivan, Margaret Gaseor, Charity and Dave Coates, Darryl and Tina Coates, Kaye Surratt, Jeanette Brown, Scott and Marcy Kight, and Vicki Sherman. A special thanks goes to Lori Pierce for the loyal friendship and saintly patience she displayed by listening to endless hours of my ramblings about cladistics, dead turtles, and stubborn computers. I must thank my family: my parents, .Frank and Dorothy Capler, my sister, Michelle, and my brother, Tom for their years of encouragement through the research and writing of this dissertation. Their love and support has been a constant source of strength for me. I offer my deepest thanks to Linda Radtke, a very special woman whom I met in the last year of this project but without whom I might not have completed it. Her steadfast faith in me helped me to persevere through the setbacks and finally get this thing written. In this past year, she has managed to successfully walk that fine line between encouragement and nagging. Thank you, Linda. Finally, I owe an immeasurable debt of gratitude to my advisor and mentor, Dr. Edward o. Moll, for his excellent and patient guidence throughout the completion of this project. I am enormously impressed by his expertise in his field and inspired by his dedication to his work. He is the person who taught me what research is all about and enlightened me regarding the fascinating and noble nature of his beloved turtles. Many thanks. TABLE OF CONTENTS Introduction ..•.•......•. p. 1 Historical Overview. p. 1 Species Accounts. p. 3 Materials and Methods. p. 9 Results .••• p. 14 Discussion. p. 18 The Hardella-Morenia Question .. p. 20 The kachuga-pangshura Question. p. 21 The Batagur-Callagur-Kachuga Question. p. 22 Taxonomic Revision. p. 24 Literature Cited. p. 32 List of Tables. p. 37 Appendices ..... p. 46 List of Figures. p. 68 INTRODUCTION Historical Overview Relationships of the subfamily Batagurinae {Emydidae: Testudines) have proven problematic due to the paucity of material available for study and the apparent high degree of homoplasy in the group. In his classic paper on the taxonomy of emydid turtles published in 1964, McDowell partitioned the family Emydidae into two subfamilies, the predominately New-World Emydinae and the predominately Old-World Batagurinae. However, later workers {Hirayama, 1984; Gaffney and Meylan, 1988) have suggested that McDowell's Batagurinae may be polyphyletic. Hirayama {1984) divided the subfamily into a primary palate group and a secondary palate group, the latter forming a clade with the Testudinidae. The secondary palate group corresponds to McDowell's "broad triturating surface" group, including his Hardella, Batagur, and Orlitia {sic) Complexes. Based on Hirayama's work, Gaffney and Meylan {1988) proposed that McDowell's Emydinae, Batagurinae, and Testudininae should be elevated to family rank, with the newly formed "Bataguridae" comprising two subfamilies {Batagurinae and Geoemydinae). McDowell {1964) introduced the designation Batagur complex to include species of the genera Batagur, Callagur, Kachuga, Ocadia, Hieremys, Malayemys, and Chinemys, all members of Hirayama's secondary palate group and Gaffney and Meylan's Batagurinae. Loveridge and Williams {1957) previously had suggested Morenia was a close relative as 1 (1984), Moll (1986), and Gaffney and Meylan (1988) suggests that only Batagur, Callagur, and Kachuga, along with Hardella and Morenia represent a monophyletic lineage. Relationships within this complex group, especially within the Kachuga, require resolution. Over the taxonomic history, the Kachuqa have been considered as: 1.) a monophyletic genus comprised of two subgenera, Kachuga and Pangshura (Gray, 1855; Moll, 1986); 2.) a monophyletic genus comprised of two distinct species groupings -- one comprised of larger riverine species E· trivittata, E· kachuga, and E· dhongoka and a second which included the smaller, lotic-adapted K· smithi, K· tecta, K· tentoria, and K· sylhetensis (McDowell, 1964); 3.) a monophyletic genus without species groupings (Boulenger, 1889, 1890); and 4.) two separate genera Pangshura and Kachuga (Gunther, 1864; Gray, 1869). The situation is further complicated when the positions of Callagur and Batagur are also considered. McDowell noted the close relationship between K· kachuga, E· trivittata, and Callagur, suggesting that they might be considered a single superspecies. He suggested no affinity between Batagur and the rest of this complex. However, Hirayama (1984) hypothesized and Gaffney and Meylan (1988) accepted
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