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United States National Museum Bulletin 265

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United States National Museum Bulletin 265 ,^««.«i«,,a SMITHSONIAN INSTITUTION MUSEUM O F NATURAL HISTORY UNITED STATES NATIONAL MUSEUM BULLETIN 265 The Evolutionary History of the Avian Genus Chrysococcyx HERBERT FRIEDMANN Director^ Los Angeles County Museum of Natural History- Research Associate, Smithsonian Institution SMITHSONIAN INSTITUTION PRESS WASHINGTON, D.C. 1968 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the different subjects. The Proceedings, begun in 1878, are intended for the publication, in separate forin, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, sepa- rate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been pubUshed in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 265 of the Bulletin series. Frank A. Taylor Director, United States National Museum Contents Acknowledgments vii Introduction 1 Phylogenetic relationships 3 Migratory behavior 25 Courtship behavior 34 Features of brood parasitism in Chrysococcyx 36 Host selection and its evolution 36 Host specificity 39 Egg morphism 66 Mode of egg-laying by adult cuckoos 81 Mode of deposition 81 Interval between eggs 85 Number of eggs 87 Removal of host eggs by adult cuckoos 87 Host-parasite nestling relations 91 Eviction of nest-mates by nestling cuckoos 96 Fledgling feeding by adult cuckoos 98 Summary and conclusions 103 Appendix 108 The existing species and their distribution 108 The plumages of Chrysococcyx flavigularis 112 References 113 Index 129 V Acknowledgments The present study is the second in a series which, when completed, hopefully may provide the background for a better understanding of the phenomenon of parasitic breeding in the Cuculidae. The first paper in this series [Friedmann, 1964] dealt with the crested cuckoos, forming the genus Clamator. The genus covered in the present paper contains three times as many species as Clamator and occupies a much wider portion ot the land areas of the Old World. It is the first attempt to study all the glossy cuckoos as a group. In the past the African species have been dealt with by specialists on African birds, the Australian and Malaysian ones separately by other regional students, and the Asiatic ones by still other writers. Until a compre- hensive and comparative study of all of the glossy cuckoos was made, it was impossible to compare meaningfully the known facts about the life histories of these birds. As in the earlier study of Clamator, the attempted appraisal of the phylogenetic relationships of the 12 species of Chrysococcyx demanded careful examination of large amounts of preserved material of each species. A research grant from the Frank M. Chapman Me- morial Fund of the American Museum of Natural History enabled me to spend 19 very productive da5'-s at the British Museum (Natural History) in London and also to examine the extensive material in the American Museum of Natural History. These two collections comprise by far the bulk of the pertinent material. The specimens in the United States National Museum in Washington were also studied. To the custodians of these bird collections I express my thanks for their cooperation in making these materials available to me. Museum study specimens were examined to ascertain the nature, the frequency, and the distribution within and between the dozen members of the genus of the types of plumage patterns, plumage phases, and kinds and degrees of variation within these patterns and phases. This survey was made with an eye to any phylogenetic suggestions that might be adduced from the many hundreds of specimens studied. By correspondence with the unfailingly cooperative curators of their respective collections and, in a few instances, by loans of selected, critical specimens, I have been able to make use of the data on per- tinent material in the museums of Bulawayo, Durban, Nairobi, and Vm U.S. NATIONAL MUSEUM BULLETIN 2 65 Pretoria, in Africa; of Bombay, Singapore, and Manila, in Asia; of Sydney and Adelaide, in Australia; of Berlin, Copenhagen, Genoa, Leiden, Milan, Paris, Stockholm, Tmin, and Vienna, in Em-ope; and of Ann Arbor, Cambridge, Chicago, and Los Angeles, in the United States. To the officials of these museums who have helped me with data or specimens in the past 25 years I offer my thanks. The following individuals also have assisted with contributed field observations, with egg-specimen data, and with other relevant information: S. A. Ali, L. Auber, R. K. Brooke, A. C. Cameron, A. H. Chisholm, G. Duve, T. Farkas, N. Favaloro, N. C. Fearnley, M. T. Goddard, F. Haverschmidt, Iv. A. Hindwood, A, Hoogerwerf, L. Hyem, M. P. S. Irwin, C. H. Jerome, F. Johnston, R. Kreuger, R. Liversidge, C. J. Marinkelle, A. R. McGUl, A. R. McEvey, W. Meise, H. M. Miles, P. Millstein, B. V. Neuby-Varty, J. Ottow, J. Paludan, E. Pike, C. R. S. Pitman, O. P. M. Prozesky, R. A. Reed, E. H. Sedgwick, D. L. Serventy, C. J. Skead, R. H. N. Smithers, G. Symons, R. E. Symons, V. G. L. van Someren, C. J. Vernon, J. G. Williams, and J, M. Winterbottom. Mr. K. A. Hindwood kindly helped me with the nomenclature of Australian birds and also put me in touch with a number of the Australian contributors, whose names are listed above. Others who assisted me in various ways in the past but who will not see this publication are the late E. Ashby, E. C. S. Baker, J. P. Chapin, and A. Roberts. My personal field experience with Chrysococcyx, although limited to three of the four species that occur in Africa, formed my original basic interest in this group and played a contributing role in the present study. This field experience was made possible by grants from the National Research Council, the American Philosophical Society, the John Simon Guggenheim Memorial Foundation, and the Smith- sonian Institution. Permission to reproduce a map (fig. 4) from Fundamentals oj Ornithology by van Tyne and Berger was granted by the publishers, John Wiley & Sons. The colored plates of the plumages of the glossy cuckoos were pre- pared for this publication by Evelina R. Templeton, scientific illus- trator of the Los Angeles County Museum of Natural History. For her patience with endless changes in the manuscript and for her careful typing of the final copy, my thanks are due my secretary, Myrna L. Patrick. The Evolutionary History of the Avian Genus Chrysococcyx Introduction The genus Chrysococcyx is one of a number of genera of cuckoos that are brood parasites in their reproductive habits. Their phylog- eny, differentiation, dispersal, and ethology— all of which are re- viewed in a comparative, yet detailed, way for the first time in this paper— are of interest in themselves not only as a special "case history" in the development of parasitism, but also for the light they throw on the broad, general problems of evolution of parasitic reproduction in the cuckoos as a family and in birds in general. As I pointed out in an earlier paper on Clamator (1964) the various genera of parasitic cuckoos reveal divergent paths of development in their mode and degree of speciahzation as parasites. Chrysococcyx, for example, agrees with Cuculus in the evicting habit in the early nestUng stage, a phenomenon not present in Clamator; it agrees with both Cuculus and Clamator in the habit of host-egg removal by the adult cuckoos from parasitized nests. Among its dozen included species Chrysococcyx presents a broad range of host-parasite situa- tions, from those found in species like malayanus and osculans with only a very small number of regular hosts to those shown by others like lucidus (plagosus), basalis, Idaas, and caprius with many dozens of host species apiece. Chrysococcyx agrees more (but not rigidly) with Cuculus than with Clamator in the intensity of parasitism on its usual hosts, as measured by the incidence of multiple to single eggs in individual nests of the latter. Among the most interesting and intriguing biological problems exhibited by the glossy cuckoos are: 1. The atavistic retention of courtship feeding by the adult male with the correlated fledgiinghke behavior of the female; 2. the still more atavistic (but not infrequent) fledgling, and even nestling, feeding by adults of these parasitic breeders; 3. the tendency, def- initely begun but far from perfected, toward primary differentiation in host selection by sympatric species of the genus; 4. the achieve- 1 2 U.S. NATIONAL MUSEUM BULLETIN 2 65 ment of various degrees of similarity to host eggs by the eggs of different species of glossy cuckoos, and the lack of continuity of pattern found in the development of this favorable adaptation in the phylogeny of the genus Chrysococcyx; 5. the fact, related to the preceding, that various glossy cuckoos show different degrees of egg morphism or the lack of it (one phenotypic egg) ; 6. the enormous diversity of migratory behavior in different segments of the genus and even in regional, geographic portions of the total population of some of the species; 7.
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