SSStttooonnnyyy BBBrrrooooookkk UUUnnniiivvveeerrrsssiiitttyyy The official electronic file of this thesis or dissertation is maintained by the University Libraries on behalf of The Graduate School at Stony Brook University. ©©© AAAllllll RRRiiiggghhhtttsss RRReeessseeerrrvvveeeddd bbbyyy AAAuuuttthhhooorrr... The origins of diversity in frog communities: phylogeny, morphology, performance, and dispersal A Dissertation Presented by Daniel Steven Moen to The Graduate School in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Ecology and Evolution Stony Brook University August 2012 Stony Brook University The Graduate School Daniel Steven Moen We, the dissertation committee for the above candidate for the Doctor of Philosophy degree, hereby recommend acceptance of this dissertation. John J. Wiens – Dissertation Advisor Associate Professor, Ecology and Evolution Douglas J. Futuyma – Chairperson of Defense Distinguished Professor, Ecology and Evolution Stephan B. Munch – Ecology & Evolution Graduate Program Faculty Adjunct Associate Professor, Marine Sciences Research Center Duncan J. Irschick – Outside Committee Member Professor, Biology Department University of Massachusetts at Amherst This dissertation is accepted by the Graduate School Charles Taber Interim Dean of the Graduate School ii Abstract of the Dissertation The origins of diversity in frog communities: phylogeny, morphology, performance, and dispersal by Daniel Steven Moen Doctor of Philosophy in Ecology and Evolution Stony Brook University 2012 In this dissertation, I combine phylogenetics, comparative methods, and studies of morphology and ecological performance to understand the evolutionary and biogeographical factors that lead to the community structure we see today in frogs. In Chapter 1, I first summarize the conceptual background of the entire dissertation. In Chapter 2, I address the historical processes influencing body-size evolution in treefrogs by studying body-size diversification within Caribbean treefrogs (Hylidae: Osteopilus ). In this chapter I combine analyses of resource use, community assembly, phylogenetics, and rates of body-size evolution within Osteopilus to examine support for the influence of past competition on body-size evolution within the genus. In Chapter 3, I develop an approach to quantify the relative importance of in situ evolution (ISE) within a region and ecologically conservative dispersal (ECD) from outside that region to better understand the evolutionary and biogeographical processes that influence community iii structure in Middle American hylid treefrogs. I also test whether colonization of the region (Middle America) is related to climatic similarity of invaded areas to ancestral areas of colonizers, and whether temporal staggering of colonization is related to subsequent evolution within the region. Last, I determine whether species that are ecologically similar can co-occur in communities and whether ecological differences are necessary for a species or lineage to colonize the region. Finally, in Chapter 4 I examine the evolution of microhabitat use, morphology, and performance in three assemblages of frogs to ask whether these processes (ISE and ECD) are important on a worldwide scale across a large clade. I examine the consequence for morphological and performance evolution of both cross-continental, ecologically conservative dispersal of lineages, as well as microhabitat diversification within a single clade in a single geographic location. Overall, these studies suggest that the ecological, morphological, and performance diversity we see in a given location is a mixture of both ISE and ECD, even at the global scale. iv To my family – you may not always understand why I do what I do, but you have invested an awful lot in my happiness and your immense love has been clear at every step of the way. It means the world to me. v Table of Contents Title page i Signature page ii Abstract iii Dedication v List of tables viii List of figures ix Acknowledgements x Statements of permission for copyrighted work xiii Chapter 1: Introduction 1 Chapter 2: Phylogenetic evidence for Competitively-driven divergence: Body-size evolution in Caribbean treefrogs (Hylidae: Osteopilus ) 9 Introduction 10 Materials and methods 17 Results 32 Discussion 38 Acknowledgements 51 Literature cited 52 Tables 69 Figures 73 Chapter 3: Community assembly through evolutionary diversification and dispersal in Middle American treefrogs 77 vi Introduction 78 Materials and methods 82 Results 102 Discussion 108 Acknowledgements 115 Literature cited 115 Tables 123 Figures 128 Chapter 4: Convergence and conservatism in the evolution of ecology, morphology, and performance across continents in frogs 135 Introduction 136 Materials and methods 140 Results 174 Discussion 181 Acknowledgements 188 Literature cited 189 Tables 205 Figures 213 Bibliography 219 Appendix 1: Specimen numbers from Chapter 2 256 Appendix 2: Chapter 2 supplementary methods and data 257 Appendix 3: Chapter 3 supplementary methods and data 309 Appendix 4: Chapter 4 supplementary data 357 vii List of Tables Table 2.1 69 Table 2.2 71 Table 2.3 72 Table 3.1 123 Table 3.2 124 Table 3.3 126 Table 3.4 127 Table 4.1 205 Table 4.2 207 Table 4.3 208 Table 4.4 209 Table 4.5 210 Table 4.6 211 Table 4.7 212 Table A2.1 284 Table A2.2 295 Table A2.3 296 Table A2.4 298 Table A2.5 303 Table A2.6 305 Table A3.1 320 Table A3.2 331 Table A3.3 341 Table A3.4 348 Table A3.5 349 Table A3.6 350 Table A4.1 357 Table A4.2 361 viii List of Figures Figure 2.1 73 Figure 2.2 74 Figure 2.3 75 Figure 2.4 76 Figure 3.1 128 Figure 3.2 130 Figure 3.3 131 Figure 3.4 134 Figure 4.1 213 Figure 4.2 214 Figure 4.3 215 Figure 4.4 216 Figure 4.5 217 Figure 4.6 218 Figure A2.1 307 Figure A2.2 308 Figure A3.1 352 Figure A3.2 353 Figure A3.3 354 Figure A3.4 355 Figure A3.5 356 ix Acknowledgments This work represents an effort that was supported by many family, friends, and colleagues throughout its entirety. I cannot thank everyone, so to those I do not explicitly address here, I thank you and ask for your understanding. I first thank my advisor, John Wiens, for showing me by example how to ask interesting questions, address them creatively in my research, and effectively communicate my message by writing clearly. His incredible ability to motivate and bring out the best in his students are what brought me to Stony Brook and what helped me finish. I can only hope that other students can see and appreciate what John has to offer his students, which is to say everything that he has. Thank you, John. To my committee: Douglas Futuyma, Stephan Munch, and Duncan Irschick. Doug encouraged me to be scholar, to always think about all possible angles of a research question, whether I would explicitly address such aspects in a given project or not. Steve gave me the confidence to develop my mathematical and statistical skills, and he believed in my efforts when I did not myself. Duncan helped bring me from being someone who knew nothing about performance to someone who at least knows a bit, assisting in designing my studies, determining which data to collect and how, and helping to interpret my quite complex results. I thank Craig Stockwell, who introduced me to basic research while I was an undergraduate at North Dakota State University, and Jim Grier for molding me into a graduate student long before I started graduate school. Jim Grier, Whit Gibbons, Bob Reed, and Chris Winne showed me that being a herpetologist really was as awesome as it seemed. I thank Martha Nolan, Iris Roth, Lee Stanley, Donna DiGiovanni, Fumio Aoki, and Irene for help in solving various logistical and academic problems during my time as a graduate student. I have had many labmates help with labwork, research design, and manuscript review: Windsor Aguirre, Saad Arif, Jeni Dixon, Paola Espinosa-Hidalgo, Caitlin Fisher-Reid, Hua Xia, Carl Hutter, Ken Kozak, Kate Kuczynski, Shea Lambert, Jure Skejic, Sarah Smith, Patrick Stephens, and Carolina Ulloa-Garzón. Outside of my laboratory, I have many to thank for giving suggestions, helping with data collection, and loaning equipment: Dean Adams, Greg Brown, Elisa Cabrera-Guzmán, Michael Collyer, Rebecca Grella, Joe Lachance, Roy McDiarmid, Dianna Padilla, Juancho Parra, Jim Rohlf, Simón Velasquez-Silva, Jorge Velasquez-Tibatá, and George Zug. I would like to express my sincere gratitude to those individuals and institutions that hosted the international fieldwork that the latter part of this dissertation involved, kindly lending much time and expertise in the field. In particular I thank Che Jing, Zhang Ya-ping, and Rao Ding-qi (Kunming Institute of Zoology) for Chinese fieldwork. In Colombia I was greatly helped by Andrew Crawford (Universidad de Los Andes); Eliana Jiménez (Universidad Nacional de Colombia Sede Amazonia); and Rodolfo Mesa, Adriana Aguilar, and Goran Mihajlovic (Reserva Tanimboca). Australian x fieldwork was generously assisted by Matt Greenlees, Iris Bleach, Greg Brown, Michelle Franklin, and Rick Shine (University of Sydney). I owe special thanks to Matt Greenlees for coordinating permits, ethics approval, and fieldwork in Australia, and in particular for traveling to Fogg Dam for 10 days to help initiate work at that location, making my time there the smoothest and most productive of all three fieldwork locations. I was able to conduct much of this work due to the generosity of many folks in the museum community, who allowed access to specimens, granted me loans, or donated