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Systematics and Cophylogenetics of Toucans and Their Associated Louisiana State University LSU Digital Commons LSU Doctoral Dissertations Graduate School 2003 Systematics and cophylogenetics of toucans and their associated chewing lice Jason David Weckstein Louisiana State University and Agricultural and Mechanical College, [email protected] Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_dissertations Recommended Citation Weckstein, Jason David, "Systematics and cophylogenetics of toucans and their associated chewing lice" (2003). LSU Doctoral Dissertations. 1376. https://digitalcommons.lsu.edu/gradschool_dissertations/1376 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Doctoral Dissertations by an authorized graduate school editor of LSU Digital Commons. For more information, please [email protected]. SYSTEMATICS AND COPHYLOGENETICS OF TOUCANS AND THEIR ASSOCIATED CHEWING LICE A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Biological Sciences by Jason D. Weckstein B.S., University of Michigan, 1993 M.S., University of Minnesota, 1997 December 2003 ACKNOWLEDGEMENTS This dissertation benefited from the help of many people and institutions. I thank my advisor Fred Sheldon, for all of his help and support during my time at LSU. His door was always open for assistance and advice. Also, his comments on many grant proposals and the chapters of this dissertation were invaluable. My graduate committee, Van Remsen, Mark Hafner, Mike Hellberg, and James Oard were also always available for advice and discussions and made helpful comments on the dissertation. I was very fortunate to be surrounded by a wonderful group of fellow graduate students during my time at LSU. In no particular order they include: Alex Aleixo, Dan Lane, Ben Marks, Rob Moyle, Chris Witt, Kazuya Naoki, Mario Cohn-Haft, Mark McRae, Dan Christian, Rob Faucett, Alison Styring, Jessica Light, David Reed, and Thomas Valqui. They helped me to develop as a scientist by reading my manuscripts and grant proposals, by discussing project ideas, listening to practice talks, and giving lab advice. They were also great friends who I could always count on to encourage me to take a break for a movie, dinner, or a fun day of fishing or birding. Also, I could not have asked for better expedition companions than Alex Aleixo, Ben Marks, and Rob Moyle. Kevin Johnson was always a supportive friend who contributed significantly to this project through a number of lab visits, many long phone calls discussing louse biology, and by reading and commenting on grant proposals and parts of this dissertation. He, Dale Clayton, and R. J. Adams taught me a great deal about louse biology and laboratory methods. I would also like to thank Kevin and Dale for inviting me to take part in the Untangling Coevolutionary History symposium at the 2002 joint meetings of the Society for the study of Evolution and Society of Systematic Biologists from which Chapter 4 ii was developed. Chapter 4 also benefited significantly from reviews by Vince Smith and Adrian Paterson. Several folks were especially helpful in the lab and in preparing for expeditions. Tyler Ortego was an excellent lab assistant who helped to produce a significant amount of the data presented here. None of the DNA sequence data would exist without Nannette Crochet’s hard work loading and running samples on the automated DNA sequencer. Steve Cardiff and Donna Dittmann not only taught me to skin birds the “LSU way,” but they also helped out in the frenzy of organizing collecting expeditions. Many people helped with the loan and collection of tissues and louse specimens. Donna Dittmann always helped to expedite my in-house tissue loan requests. David Oren, Museu Paraense Emilio Goeldi, Mark Robbins, University of Kansas Museum of Natural History, and Leo Joseph, Academy of Natural Sciences generously loaned tissues to me for this project. In addition, many generous bird collectors, including Alex Aleixo, Dan Christian, Dale Clayton, Rob Faucett, Dan Lane, Ben Marks, Rob Moyle, Kazuya Naoki, John O’Neill, Thomas Valqui and Chris Witt, collected lice for this project in localities I was unable to visit on my own. The louse chapters also benefited from expeditions to South Africa and Ghana where Tamar Cassidy, Transvaal Museum, Blackie Swart, Mpumulanga Parks Board, Brian Colahan, Free State Environmental Affairs and Tourism, John Mason and Patrick Adjewodah, Nature Conservation Research Centre, Mike Adu-Nsiah, James Braimah, James Oppong, and Joseph Amponsah, Ghana Wildlife Division provided much needed assistance with collecting permits, export permits, and in-country support. iii Laboratory and field work for this dissertation were supported by NSF Doctoral Dissertation Improvement Grant DEB-0104919, the Frank M. Chapman Memorial Fund of the American Museum of Natural History, Sigma Xi, the American Orithologists’ Union, the T. Vinton Holmes Endowment, LSU Museum of Natural Science, LSUMNS Graduate Student Bird-a-thon, LSU Graduate School, and the LSU Department of Biological Sciences. This dissertation would not have been possible without the love, encouragement, and support of my parents and family. I want to especially thank Stacey Hoffer Weckstein for being such a supportive loving friend for so many years. She moved all the way to Louisiana for us and endured many lonely nights while I typed away in the office or while I was on an expedition. Her love, encouragement, support, and adventurous companionship made all of this hard work much easier. iv TABLE OF CONTENTS ACKNOWLEDGEMENTS………………………………………………………ii ABSTRACT……………………………………………………………………...vi CHAPTER 1 INTRODUCTION…………………………………………………....1 2 MOLECULAR PHYLOGENETICS AND CONVERGENCE IN THE COLORATION OF RAMPHASTOS TOUCANS…………..13 3 MOLECULAR PHYLOGENETICS OF THE DEGEERIELLA (INSECTA: PHTHIRAPTERA) COMPLEX: IS THE GENUS AUSTROPHILOPTERUS MONOPHYLETIC?...................................49 4 BIOGEOGRAPHY EXPLAINS COPHYLOGENETIC PATTERNS IN TOUCAN CHEWING LICE………………………..76 5 CONCLUSIONS……………………………………………………...99 REFERENCES…………………………………………………………………....107 APPENDIX I TABLE OF UNCORRECTED PROPORTIONAL DISTANCES AMONG PAIRS OF TOUCAN SPECIES ANALYSED FOR CHAPTER 2…………………………………………………………..119 II TABLE OF UNCORRECTED PROPORTIONAL DISTANCES AMONG PAIRS OF LOUSE SPECIES ANALYSED FOR CHAPTER 3…………………………………………………………..127 VITA……………………………………………………………………………....132 v ABSTRACT Historically, comparisons of host and parasite phylogenies have concentrated on cospeciation. However, many of these comparisons have demonstrated that the phylogenies of hosts and parasites are seldom completely congruent, suggesting that phenomena other than cospeciation play an important role in the evolution of host- parasite assemblages. Other coevolutionary phenomena, such as host switching, parasite duplication (speciation on the host), sorting (extinction), and failure to speciate can also influence host-parasite assemblages. In this dissertation I explore several aspects of the evolutionary history of Ramphastos toucans and their ectoparasitic chewing lice using molecular phylogenetic and cophylogenetic reconstructions. First, using mitochondrial DNA sequences, I reconstructed the phylogeny of the Ramphastos toucans. I used this phylogeny to assess whether the striking similarity in plumage and bare-part coloration of sympatric Ramphastos is due to convergence or shared ancestry. Ancestral character state reconstructions indicate that that at least half of the instances of similarity in plumage and bare-part coloration between sympatric Ramphastos are due to homoplasy. Second, using mitochondrial and nuclear protein-coding DNA sequences, I reconstructed the phylogeny of ectoparasitic toucan chewing lice in the Austrophilopterus cancellosus subspecies complex, and compared this phylogeny to the phylogeny of the hosts to reconstruct the history of coevolutionary events in this host-parasite assemblage. Three salient findings emerged. (1) reconstructions of host and louse phylogenies indicate that they do not branch in parallel and that their cophylogenetic history shows little or no significant cospeciation. (2) members of monophyletic Austrophilopterus toucan louse lineages are not necessarily restricted to monophyletic host lineages. Often, closely vi related lice are found on more distantly related, but sympatric, toucan hosts. (3) the geographic distribution of the hosts apparently plays a role in the speciation of these lice. These results suggest that for some louse lineages, biogeography may be more important than host associations in structuring louse populations and species. This is particularly true in cases where host life history (e.g. hole-nesting) or parasite life history (e.g. phoresis) might promote frequent host switching events between syntopic host species. These findings highlight the importance of integrating biogeographic information into cophylogenetic studies. vii CHAPTER 1: INTRODUCTION No region of the planet harbors more biological diversity than the Neotropics (Stotz et al., 1996). Although many biologists have studied the phylogenetics, biogeography, and population genetics of lowland Neotropical vertebrates (e.g. Hackett, 1996; Mariaux and Braun, 1996; Silva and Patton, 1998; Bates et al., 1999), few have tried to understand these same evolutionary patterns
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