MORPHOLOGICAL and ECOLOGICAL EVOLUTION in OLD and NEW WORLD FLYCATCHERS a Dissertation Presented to the Faculty of the College O
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MORPHOLOGICAL AND ECOLOGICAL EVOLUTION IN OLD AND NEW WORLD FLYCATCHERS A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Clay E. Corbin August 2002 This dissertation entitled MORPHOLOGICAL AND ECOLOGICAL EVOLUTION IN OLD AND NEW WORLD FLYCATCHERS BY CLAY E. CORBIN has been approved for the Department of Biological Sciences and the College of Arts and Sciences by Donald B. Miles Associate Professor, Department of Biological Sciences Leslie A. Flemming Dean, College of Arts and Sciences CORBIN, C. E. Ph.D. August 2002. Biological Sciences. Morphological and Ecological Evolution in Old and New World Flycatchers (215pp.) Director of Dissertation: Donald B. Miles In both the Old and New Worlds, independent clades of sit-and-wait insectivorous birds have evolved. These independent radiations provide an excellent opportunity to test for convergent relationships between morphology and ecology at different ecological and phylogenetic levels. First, I test whether there is a significant adaptive relationship between ecology and morphology in North American and Southern African flycatcher communities. Second, using morphological traits and observations on foraging behavior, I test whether ecomorphological relationships are dependent upon locality. Third, using multivariate discrimination and cluster analysis on a morphological data set of five flycatcher clades, I address whether there is broad scale ecomorphological convergence among flycatcher clades and if morphology predicts a course measure of habitat preference. Finally, I test whether there is a common morphological axis of diversification and whether relative age of origin corresponds to the morphological variation exhibited by elaenia and tody-tyrant lineages. The general results were this: 1) Morphology significantly predicted the foraging behavior in both NA and SA flycatchers. 2) North American and Southern African communities are concordant with respect to the ecomorphological relationships. 3) I found that there are lineage-specific positions in morphological space when examining synoptic morphological samples of Old and New World flycatcher clades. 4) There were fundamental differences in the orientations of morphological disparity of Old versus New world flycatchers. 5) The results of separate principal components and common principal component analyses reveal a larger morphological volume being occupied by older lineages, but 6) Macroevolutionary patterns exist within constituent clades that are inconsistent with a Brownian motion evolutionary hypothesis. Hence, the differences in morphological disparity observed in flycatchers are most likely due group-specific factors including genetic constraints, ecological limiting similarity and ecological opportunity. Phylogenetic constraints to morphological evolution are not universal throughout the history of flycatcher evolution. Furthermore, historical and/or stochastic factors were equal to ecological factors during flycatcher evolution. Approved: Donald B. Miles Associate Professor of Biological Sciences 5 ACKNOWLEDGEMENTS Foremost, I thank my wife Nancy. Her love, patience, and encouragement have been there from the start and continue to this day. My son Charlie (10 months old at the time of this writing) has added lots of encouragement for this project in a special way. Opie, as always, helped me keep things in perspective. These members of my family, above all, made my experiences at Ohio University great ones and I am dearly indebted to them. I want to thank my parents, Nancy’s parents and my grandmothers for lots of love, encouragement and financial support during this dissertation project. I could not have done it without them. Also, my siblings were there for me when I needed them and I appreciate their support. I especially want to thank Rocky for his time and encouragement on many occasions. His perspectives got me through a lot of tough times during research and writing. I want to thank my extended family in general for their support and high expectations. Don Miles gets a lot of credit for this project. Since my master’s thesis, he has influenced my worldview in positive ways and taught me to look at the ecology of organisms, especially birds, from an evolutionary perspective. John Scheibe also gets a lot of credit for pointing me in the right direction when it comes to comparative biology. These men taught me the importance legacy and chance current ecological communities. This project has been improved greatly by comments and suggestions from my proposal and dissertation committee members – Audrone Biknevicius, Mark Dybdahl, Jim Dyer, Molly Morris, Steve Reilly and Willem Roosenburg. I also want to thank the 6 Ohio University Evomorph group for helpful comments. Members from the laboratories of Audrone Biknevicius, Molly Morris, Willem Roosenburg, Larry Witmer and J. Van Remsen Jr. have also lent support and helpful comments and suggestions. Thanks to T. Smith and two anonymous reviewers of the “elatody” manuscript (Chapter 4) for helpful comments and suggestions. Also, I would like to thank the natural history museums and staff of the Transvaal Museum (T. Cassidy), Louisiana State University (J. V. Remsen Jr.), University of Kansas (M. Robbins), Field Museum (D. Willard), and National Museum of Natural History (J. Dean) for museum loans and access to their collections. Thanks to Kevin de Queiroz, Mark Lensmeyer and Shaun Pharr for putting me up (putting up with me) and feeding me while collecting museum data. For help while in the field, I thank Stacey McKinnen, Sara Kozup, Dave Rushworth, Alan Kemp, Robert Ricklefs and the Transvaal Museum staff. I thank Laurie Dries, Scott Moody, Jim Robbins, John Scheibe, Jayc Sedlmayr and Martin Wikelski for helpful discussions about the research. Pete Larson and Chris Klingenberg helped considerably with common principal components and bootstrapping programs. Finally, thanks to my office mates: Rick Essner, Lance McBrayer for helpful comments on my research and appropriate ribbing. 7 TABLE OF CONTENTS Abstract................................................................................................................................3 Acknowledgements..............................................................................................................5 Table of Contents.................................................................................................................7 List of Tables .......................................................................................................................9 List of Figures....................................................................................................................12 List of Appendices .............................................................................................................15 Chapter 1. Introduction to the Dissertation........................................................................16 Background and Interests ......................................................................................17 Chapter Summaries ...............................................................................................20 Chapter 2. Concordance among North American and Southern African Flycatcher Communities......................................................................................................................24 Introduction............................................................................................................25 Methods..................................................................................................................28 Results....................................................................................................................38 Discussion..............................................................................................................43 Chapter 3. Convergence in the Adaptive Radiations of Old and New World Flycatchers ........................................................................................................................51 Introduction............................................................................................................52 8 Methods..................................................................................................................55 Results....................................................................................................................61 Discussion..............................................................................................................70 Chapter 4. Comparative Morphological Disparity of Elaenia and Tody-tyrant flycatcher Clades ................................................................................................................................77 Introduction............................................................................................................78 Methods..................................................................................................................80 Results....................................................................................................................87 Discussion..............................................................................................................92 Chapter 5. General Conclusions and Future Directions....................................................98 Literature Cited ................................................................................................................101