SYSTEMATICS OF NEW WORLD CUCKOOS (AVES, CUCULIDAE) AND THE EVOLUTION OF BROOD PARASITISM Janice Maryan Hughes A thesis submitted in confomiity with the requirements for the degree of Doctor of Philosophy Graduate De partment of Zoology , University of Toronto O Copyright by Janice Maryan Hughes 1997 National Library Biblioth6que nationale I*l of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Weilingtm Street 395. nie Wellingtm Ottawa ON K1A ON4 CMawaON K1AON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantid extracts fkom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Canada -- -- ABSTRA CT Systernatics of New World Cuckoos (Aves, Cuculidae) and the Evolution of Brood Parasitism Degree in Doctor of Philosophy, 1997 Janice Maryan Hughes Department of Zoology University of Toronto Monophy ly of the cuckoos (Cuculidae) was demonstrated through the description of 14 osteological synapomorphies. Only three apparent synapomorphies united the otherwise anatomically divergent turacos (Musophagidae), Hoatzin (Opisrhocomus hoazin), and Cuculidae. Therefore, I recomrnended their placement in separate, but adjacent orders. Cuckoo phylogeny reconstructed using 135 osteological characters differed markedly from currently accepted classifications. Al1 brood parasitic cuckoos, including New World obligate parasites Tapera and Dromococcyx and facultative parasites Coccyzus, formed a clade. This implies that the ancestral Coccyirts was an obligate parasite, and is consistent with many be havioral adaptations to parasitism exhibited by this genus. Tribe Saurothenni (Phaenicophaeinae) was erected to house al1 nonparasitic, arboreal. solitady breeding New World cuckoos. The Old World genus Carpococcyx was removed from the New World Neomorphinae. Three subfamilies, comprised of nonparasitic. terrestrial cuckoos of Old World and New World distribution, occupied basal positions on the phylogeny. A new classification of the Cuculidae was proposed. I demonstrated that two species of facultative brood parasites, the Black-billed (Coccyziis erythoprhalmus) and Yellow-billed (C.americanus) cuckoos, produce eggs that fully or nearly match the eggs of over 70% of their reported host species, a proportion significantly greater than if hosts were being selected at random from a potential host pool. This supports an hypothesis of egg mimicry, and impiies an historically intense relationship between parasite and host. Factors responsible for the loss of obligate parasitism in this genus may also have contributed to the general paucity of obligate parasitism in New World cuckoos. Cornpetitive exclusion or resistance to invasion by parasitic cowbirds (Molotlirusspp.) should be considered. The phylogeny of the genus Coccyzrts was reconstmcted using skeletal characters, cytochrome b gene sequences, and extemal morphology. The optimal topology suppons a Neotropical origin for the genus. Several vicariant and colonization events were responsible for the current distribution of South Amencan endemic species. The Mangrove Cuckoo (C.minor) had two major northward routes out of South Amerïca. Yellow-billed and Black-bilied cuckoos are highly divergent, and each represents a separate invasion of North America from South America. .. Ill ACKNO WLEDGEMENTS First, I would like to thank rny academic advisor, Jon Barlow, for sharing his vast knowledge of the avian world, his sense of humor, and love of music. Most of all, 1 appreciate his fortitude and good advice. Yes, Jon, I was listening. 1 also thank the members of my advisory cornmittee: Bob Murphy, for those long talks on our two shared interests of phylogenetics and photography, and Roger Hanse11 for consistently presenting a direction to my work that had not yet occurred to me. 1 thank Allan Baker and Bob Murphy for providing laboratory facilities, and Oliver Haddrath, Mark Peck, Dilara Ally, and Dawn Marshall for technical assistance. 1 am particularly grateful to Oskana Borowik for her expertise with the DNA sequences. I thank D. J. Agro, L. R. Bevier, and 1. I. Lovette (Academy of Natural Sciences, Philadelphia), G. F. Bmowclough, C. Blake, and P. Sweet (American Museum of Natunl History. New York), R. M. Zink and J. T. KIicka (Bell Museum of Natural History, St. Paul), B. C. Livezey and R. Panza (Carnegie Museum of Naturai History, Pittsburgh), D. E. Willard and J. M. Bates (Field Museum of Natural History, Chicago), T. Webber and D. W. Steadman (Flonda Museum of Natural History, Gainesville), J. V. Rernsen, D. L. Ditunan, and F. H. Sheidon (Louisiana State University Museum of Zoology, Baton Rouge), R. C. Banks and R. Browning (United States National Museum of Natural History, Washington, D.C.). C. Cicero and N. K. Johnson (University of California Museum of Vertebrate Zoology, Berkeley), and R. B. Payne. L. L. Payne, and J. Hinshaw (University of Michigan Museum of Zoology, Ann Arbor) for their invaluable assistance in providing the many specimens required for this study. I thank Jim Dick, Oliver Haddrath, Brad Millen, Glenn Murphy, and Mark Peck for iheir assistance in the Royal Ontario Museum collections. In addition, 1 am grateful to the many people in the Department of Zoology who have helped to straighten my road: Sheila Freeman. Dr. Harold Harvey, Sonia McKenUe, Mary Delac, Liz Tudor-Mulroney, Dr. Dan Brooks, and Dr. Jim Rising. This work was supported by grants from the Naturai Sciences and Engineering Research Council to me and Jon C. Barlow (grant A-3472). Portions of chapter 2 of this dissertation have been published previousiy, and copyright release for reproduction herein has been obtained from the Canadian Journal of Zoology. J. C. Barlow, M. G. Brooker, R. 1. C. Hansell, L. F. Kiff, D. R. Kozlovic. S. M. Lanyon, D. A. McLennan, M. T. Murphy, R. W. Murphy, and K. C. Parkes provided helpful comments on some aspects of this dissertation. Field work in Texas would not have been possible without the kindness and hospitality of the Texas Parks and Wildlife Department, in particular Jack "Looks like it's gonna rain" Kilpatric, Lonesome Dave Dvorak, Bob West. Robert Culpepper, Rick MacIntyre, and EM House. My sincere thanks to my brother, Mark Hughes, for providing the illustrations accompanying this manuscript and, of course, to my daughter, Eliana, for always giving me something else to think about. Finally, 1 am most deeply indebted to my husband, Ron. for his encouragement, patience, and support through Iong periods of neglect, for listening to unceasing 'cuckoo-talk' to the exclusion of al1 other conversation. and for starting the whole thing many years ago with Lefty the Canary. 1 dedicate this manuscript to the memory of rny father, Arthur Edward Hughes. How grateful 1 am that his bedtime stories retold the taies of Charles Darwin. Richard Leakey, and Tuzo Wilson, and not Peter Rabbit. TABLE OF CONTENTS .. ABSTRACï .......................................................... 11 ACKNOWLEDGEMENTS ................................................ iv DEDICATION......................................................... vi LISTOFTABLES...................................................... xi LISTOFFIGURES..................................................... xii LIST OF APPENDICES .................................................. xiv PREFACETO THE DISSERTATION........................................ xv G ENERAL INTRODUCTION .............................................. 1 CHAPTER1: Monophyly and phylogeny of the cuckoos (Cuculidae) based on osteological characters .............................................. Introduction ................................................... Monophyly ............................................. Phy logeny and classification ................................ Materials and methods ........................................... Monophyly .............................................. Phylogeny ............................................... Taxa and specimens ................................. Outgroup selection.................................. Character analysis .................................. Tree denvation and analysis .......................... Results ....................................................... Monophyly .............................................. Phylogeny ............................................... vii Cuckoos .......................................... Outgroups ........................................ Character consistency ............................... Discussion .................................................... Monophyly .............................................. Hoatzin ........................................... Phylogeny ............................................... Single evolution of brood parasitism ................... Inclusion of Cocc_vtus in the Cuculinae