Systematics of the Genus Ptychadena Boulenger
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University of Texas at El Paso DigitalCommons@UTEP Open Access Theses & Dissertations 2010-01-01 Systematics of the genus Ptychadena Boulenger, 1917 (Anura: Ptychadenidae) from Democratic Republic of the Congo Katrina Marie Weber University of Texas at El Paso, [email protected] Follow this and additional works at: https://digitalcommons.utep.edu/open_etd Part of the Biodiversity Commons, Biology Commons, Evolution Commons, and the Genetics Commons Recommended Citation Weber, Katrina Marie, "Systematics of the genus Ptychadena Boulenger, 1917 (Anura: Ptychadenidae) from Democratic Republic of the Congo" (2010). Open Access Theses & Dissertations. 2612. https://digitalcommons.utep.edu/open_etd/2612 This is brought to you for free and open access by DigitalCommons@UTEP. It has been accepted for inclusion in Open Access Theses & Dissertations by an authorized administrator of DigitalCommons@UTEP. For more information, please contact [email protected]. SYSTEMATICS OF THE GENUS PTYCHADENA BOULENGER, 1917 (ANURA: PTYCHADENIDAE) FROM DEMOCRATIC REPUBLIC OF THE CONGO KATRINA M. WEBER Department of Biological Sciences APPROVED: Eli Greenbaum, Ph.D., Chair Max Shpak, Ph.D. Jasper Konter, Ph.D. Patricia D. Witherspoon, Ph.D. Dean of the Graduate School Copyright © by Katrina M. Weber 2010 Dedication This thesis is dedicated to my mother and father, my continual support system, who showed me how to learn for the sake of learning. I have become the person I am today because of you. Also to Shawn T. Dash, I may not have always been appreciative of your assistance but this never would have gotten done without your help. SYSTEMATICS OF THE GENUS PTYCHADENA BOULENGER, 1917 (ANURA: PTYCHADENIDAE) FROM DEMOCRATIC REPUBLIC OF THE CONGO by KATRINA M. WEBER, B.Sc. THESIS Presented to the Faculty of the Graduate School of The University of Texas at El Paso in Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE Department of Biological Sciences THE UNIVERSITY OF TEXAS AT EL PASO December 2010 Acknowledgements I would like to start by acknowledging my thesis committee, without which this thesis would never have been finished: Dr. Eli Greenbaum, who accepted me as a graduate student and introduced me to a new side of biology, Dr. Max Shpak for helping me with statistical analyses and providing new ideas, and Dr. Jasper Konter, who filled in the many gaps in my geological knowledge. I would also like to thank all the other faculty members that have helped me along the way: Dr. Elizabeth Walsh, who has been a caring mentor since my time as an undergraduate and is always willing to provide help and advice and Dr. Vanessa Lougheed, who patiently explained statistics when I knew next to nothing. Omar Hernandez, Dr. Carolina Lema, and Ana Betancourt in the UTEP DNA Analysis Core Facility (NIH grant #5G12RR008124) completed hundreds of DNA sequencing requests for me. Also, the UTEP Graduate School gave me two Graduate Student Research Grants, which paid for all the supplies needed for DNA sequencing. Field work conducted by Dr. Eli Greenbaum for this study was supported by National Geographic Research and Exploration Grant #8556–08. Garin Cael provided invaluable research assistance at the Royal Museum for Central Africa in Tervuren, Belgium. Cesar Villanueva taught me all the lab procedures I know and was always the one to troubleshoot lab problems. The other members of the Greenbaum lab: Chris Anderson and Federico Valdez provided support and advice throughout my project. To my friends and fellow graduate students: Jennifer Martinez, Kevin Floyd, and Shawn Dash, thank you for endless lunch discussions, project advice, and general encouragement. v Abstract This thesis increases the scope of previous phylogenetic analyses that revealed high levels of genetic differentiation within the anuran genus Ptychadena. Herein, I increase sampling of Central African populations of Ptychadena by over six times to examine their relationships to other African populations and to search for cryptic species. This study represents the most comprehensive molecular phylogeny of Ptychadena to date. A total of three mitochondrial (12S, 16S, and cyt b) and two nuclear (RAG1 and rhodopsin) genes were sequenced for 67 specimens of Ptychadena. Maximum parsimony, maximum likelihood, and Bayesian analyses were conducted. These analyses revealed a great deal of genetic diversity in the genus Ptychadena, suggesting that many species have been separated into distinct genetic lineages that will be recognizable as distinct species as some point. There are five genetically distinct lineages of P. mascareniensis sensu lato, three of which occur in the DRC. Multivariate analyses of morphometric and meristic data did not reveal any distinct morphological differences between any of the clades except P. mascareniensis sensu stricto in Madagascar. Based on habitat structuring, two of these distinct lineages, a savannah-dwelling lineage and a forest-dwelling lineage, should be recognized as distinct species under the names P. nilotica and P. marchei, respectively. The formation of the Albertine Rift combined with subsequent climatic shifts that increased habitat heterogeneity likely led to the establishment of these savannah and forest forms of P. mascareniensis. vi Table of Contents Acknowledgements.............................................................................................. v Abstract .............................................................................................................. vi Table of Contents............................................................................................... vii List of Tables ...................................................................................................... ix List of Figures ..................................................................................................... x Chapter 1: General Introduction ........................................................................... 1 1.1 Amphibian Declines .............................................................................. 1 1.2 Introduction to Democratic Republic of the Congo ................................ 4 1.3 Historical Geology and Climate of Africa .............................................. 5 1.4 Introduction to Ptychadena.................................................................... 7 1.5 Research Objectives ............................................................................ 14 Chapter 2: Materials and Methods...................................................................... 15 2.1 Specimen Acquisition.......................................................................... 15 2.2 Molecular analysis .............................................................................. 15 2.3 Species Concepts ................................................................................. 20 2.4 Morphological Analysis ...................................................................... 20 Chapter 3: Results of Molecular Study ............................................................... 22 3.1 Phylogenetic Analyses ........................................................................ 22 3.2 Combined Dataset ............................................................................... 25 3.3 Genetic diversity in Ptychadena .......................................................... 29 3.4 Genetic diversity in Ptychadena mascareniensis.................................. 30 3.5 Haplotype Networks ............................................................................ 34 Chapter 4: Results of Morphological Analyses ................................................... 40 4.1 Diagnostic characters in Ptychadena mascareniensis ........................... 40 4.2 Multivariate Statistical Analyses ......................................................... 46 Chapter 5: General Discussion and Conclusions ................................................ 57 5.1 Genetic diversity in Ptychadena .......................................................... 57 5.2 Genetic diversity in Ptychadena mascareniensis.................................. 58 5.3 Taxonomic status of Ptychadena mascareniensis ................................ 60 vii 5.4 Biogeography ...................................................................................... 66 5.5 Conservation Implications ................................................................... 72 5.6 Conclusions and future directions ........................................................ 72 Literature Cited .................................................................................................. 74 Appendices ........................................................................................................ 93 Appendix A: Locality Information for All Specimens ............................... 93 Appendix B: Morphometric and Meristic Characters Used for Morphological Analyses ........................................................................................... 99 Appendix C: Complete Measurement Data for All Ptychadena Specimens ...................................................................................... 102 Curriculum Vita ............................................................................................... 118 viii List of Tables Table 1.4.1: Summary of the supraspecific taxonomic history of the family Ptychadenidae.. ................11 Table 2.2.1: Primers used for sequencing mitochondrial and nuclear genes ..........................................16