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Hhihhwhihiiwiiimmll 3 % Llhilihhhwhhh HHIHHWHIHIIWIIIMmll llHILIHHHWHHH 3% THESiS l\\\\\\\\\ Mll\\\\\\\\\\\\\\\\\\ \\l \\\\\\\\\\1\\\2\\\\\\\\l 3 1293 This is to certify that the dissertation entitled PHYLOGENY AND EVOLUTION OF FORAGING SPECIALIZATION IN THE TYRANT FLYCATCHERS presented by Jeffrey S. Birdsley has been accepted towards fulfillment of the requirements for Ph. D . degree in Zoology 953227 ' Major professor W Dang/Q06 [798/ MS U is an Affirmative Action/Equal Opportunity Institution 0-12771 LIBRARY Michigan State University PLACE IN RETURN Box to remove this checkout from your record. To AVOID FINE return on or before date due. MAY BE RECALLED with earlier due date if requested. DATE DUE DATE DUE DATE DUE 1/” Wu PHYLOGENY AND EVOLUTION OF FORAGING SPECIALIZATION IN THE TYRANT FLYCATCHERS By Jeffrey S. Birdsley A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY W. K. Kellogg Biological Station and Department of Zoology 1998 ABSTRACT PHYLOGEN Y AND EVOLUTION OF FORAGING SPECIALIZATION IN THE TYRANT FLYCATCHERS By Jeffrey S. Birdsley Little is known about the evolutionary implications of ecological specialization. Specialized species have long been considered to be evolutionary "dead-ends", having lower potential to evolve into different fonns than do their more generalized relatives. I tested this idea by performing a phylogenetic reconstruction of the evolution of specialization and generalization in foraging tactics in the tyrant flycatchers (Aves: Tyrannidae). I first performed a phylogenetic analysis of nearly all of the approximately 100 tyrannid genera using all available morphological and behavioral data in the literature. I then mapped foraging behavior onto the resulting most-parsimonious trees. In the flycatchers, specialized lineages have not been dead-ends but have given rise to different forms as often as have generalized lineages. A possible explanation is that, although specialized foraging tactics may be accompanied by specialization of some morphological structures which are inefficient for many other tactics, some of the specialist’s remaining morphology may not be so specialized. These structures may often allow expansion of the specialist’s foraging repertoire, particularly to new tactics which don’t require use of the specialized structures. Results of the phylogenetic analysis support three of the five previously proposed tyrannid assemblages, as well as several lower level relationships. No higher level relationships are well supported but there is some support for a basal placement of the flatbills and tody-tyrants. Nasal septum characters that past workers have considered conservative and phylogenetically informative are supported as important synapomorphies of the kingbird assemblage and a restricted Empidonax assemblage. A nasal capsule character provides equivocal support for a Myiarchus assemblage without Attila. Several foraging tactics are supported as homologous within large groups of taxa, demonstrating a strong historical component to foraging ecology. ACKNOWLEDGEMENTS I wish to thank the members of my guidance committee Torn Getty, Don Beaver, Fred Dyer, Don Straney, and Scott Winterstein. I thank my advisor Tom Getty for pushing me to find the important questions to ask. The quality of this dissertation was greatly improved as a result. I thank Tom and the rest of the faculty at Kellogg Biological Station for providing a truly incredible amount of financial and equipment support. This work would not have been possible without years of unrestricted access to KBS’s excellent computer facilities. Thanks to John Gorentz for years of help and some last-minute assistance to improve the quality of my presentation. Carolyn Hammarskjold did much invaluable library legwork for me, especially during the long months I spent writing this dissertation. You can stop worrying Carolyn, I finally took that vacation. I wish to thank a host of KBS and MSU colleagues and friends for inspiration, ideas, support and companionship. In particular, Andy Turner, Chris Rogers, Casey Huckins, Jeff White, Jeff Dudycha, and Rich Leschen were always there to talk and were very helpful in the development of this research and of my approach to biology. Rob Olendorf, Jill Fisher, Sandy Halstead, Kellie Ellis, Doug Jakubiak, Jackie Smith, Elizabeth Smiley, Beth Capaldi, John Wallace, Steve Fradkin, Mike Rondinelli, Lisa Horth, Puja Batra, Emily Lyons, and Steven Mom'ssey all helped make this journey possible and enjoyable. Finally and most importantly, I must thank Becky Fuller for the encouragement and ‘push’ I needed to see this through. You’re awesome Becky, keep it up! TABLE OF CONTENTS LIST OF TABLES .................................................................................. v LIST OF FIGURES ................................................................................ vi CHAPTER 1 INTRODUCTION .................................................................................. l CHAPTER2 PHYLOGENY OF THE TYRANT FLYCATCHERS BASED ON MORPHOLOGY AND BEHAVIOR .......................................................................................... 3 INTRODUCTION .................................................................................. 3 METHODS ........................................................................................... 5 RESULTS AND DISCUSSION .................................................................. 8 Flatbill and tody-tyrant assemblage ..................................................... 15 Kingbird assemblage ..................................................................... 16 Empidonax assemblage .................................................................. l7 Myiarchus assemblage ................................................................... l9 Elaenia assemblage ....................................................................... l9 Problematic genera ........................................................................ 20 Homology of nasal septum characters .................................................. 22 Summary ................................................................................... 24 CHAPTER3 THE EVOLUTIONARY IMPLICATIONS OF ECOLOGICAL SPECIALIZATION 26 INTRODUCTION ................................................................................. 26 METHODS ......................................................................................... 27 Reconstructing the evolution of foraging behavior ................................... 27 Hypothesis testing ........................................................................ 29 RESULTS .......................................................................................... 29 DISCUSSION ..................................................................................... 33 Specialization is not a cul-de-sac ........................................................ 33 Evolution of tyrannid foraging behaviors .............................................. 38 Conclusion ................................................................................. 39 APPENDIX ........................................................................................ 42 LIST OF REFERENCES ......................................................................... 61 iv LIST OF TABLES Table l - The average number of unambiguous changes from each foraging behavior state into any of the others as reconstructed on trees estimated with foraging behavior included. Note that cells on the diagonal from upper left to lower right contain the frequency of stasis for each state ..................................................... 34 Table 2 - Relative rates of stasis and change for each foraging behavior state ............. 35 Table 3 - Relative rates of stasis and change for pooled generalist states and for pooled specialist states ......................................................................... 36 Table 4 - Relative rates of stasis and change for pooled generalist states and for pooled specialist states as reconstructed on trees estimated with foraging behavior excluded ................................................................................. 36 LIST OF FIGURES Figure 1 - Strict consensus tree derived from all most-parsimonious trees. Synapomorphies providing unambiguous support in all most-parsimonious trees are numbered. Numbers in circles indicate support for each node in the form of a decay index giving the number of extra steps required to show a node as unsupported. Higher index values indicate more robust clades. Descriptions of characters in Appendix ............................................................... 10 Figure 2 - Adams consensus tree derived from all most-parsimonious trees. Asterisks denote clades which are not present in any of the most—parsimonious trees 13 Figure 3 - Evolution of tyrannid foraging behavior as reconstructed on one of the most parsimonious trees chosen randomly from the set of trees estimated with foraging behavior included. In the legend, generalist foraging behavior states are listed in lower-case letters, specialist states in all capitals ................... 31 vi Chapter 1 INTRODUCTION Little is known about the evolutionary implications of ecological specialization. Specialized species have long been considered to be evolutionary "dead—ends", having lower potential to evolve into different forms than do their more generalized relatives. Simpson (1953) and Rensch (1959) promoted the ideas that evolution
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