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Relationships and Evolution of Flamingos (Aves: Phoenicopteridae)

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Relationships and Evolution of Flamingos (Aves: Phoenicopteridae) STORRS L. OLSON and ALAN FEDUCCIA SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 316 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropo/ogy Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world cf science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 316 Relationships and Evolution of Flamingos (Aves: Phoenicopteridae) Storrs L. Olson and Alan Feduccia SMITHSONIAN INSTITUTION PRESS City of Washington 1980 ABSTRACT Olson, Storrs L., and Alan Feduccia. Relationships and Evolution of Fla- mingos (Aves: Phoenicopteridae). Smithsonian Contributions to Zoology, number 316, 73 pages, 40 figures, 2 tables, 1980.—Previous evidence supposedly showing a relationship between flamingos and either storks (Ciconiiformes) or ducks (Anseriformes) is re-examined in light of the recent hypothesis deriving flamingos from shorebirds of the order Charadriiformes. Anatomical charac- ters used to indicate relationship between flamingos and storks are shown to consist entirely of primitive "non-anseriform" traits found in several other orders, including Charadriiformes. Most of the presumed anseriform charac- ters of flamingos also occur in the Charadriiformes, as do all of the characters of flamingos that do not occur in either Ciconiiformes or Anseriformes. The distinctive life history and behavior of flamingos is demonstrated as being very similar to that of the Recurvirostridae (Charadriiformes), particularly the Australian Banded Stilt {Cladorhynchus leucocephalus), but is unlike that of storks or ducks. The appendicular myology of Cladorhynchus is described and is found to be quite similar to that of flamingos, whereas neither is close to storks. The thigh muscle M. iliotibialis medialis, heretofore considered unique to flamin- gos, was discovered in Cladorhynchus but not in other Recurvirostridae. Evi- dence from osteology, natal down, oology, and internal parasites strongly supports a charadriiform derivation of flamingos; pterylosis does not contra- dict such a relationship; and knowledge of the early evolution of flamingos and Anseriformes offers a logical explanation for their sharing similar mallo- phagan parasites. The earliest certain flamingo, from the early Middle Eocene of Wyoming, is described herein as a new monotypic genus and species that was intermediate in size and morphology between the Recurvirostridae and modern flamingos. Other aspects of paleontology of flamingos are discussed. Evolutionary steps in the development of filter feeding in birds are outlined. The structure of the feeding apparatus of flamingos is shown to be entirely different from that of the Anseriformes, but is strikingly convergent towards that of baleen whales. Morphological and behavioral precursors for filter feeding are shown to occur in the Charadriiformes but not in the Ciconi- iformes. Flamingos (Phoenicopteridae) clearly belong in the order Charadri- iformes, suborder Charadrii, immediately following the Recurvirostridae. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavemosa (Linnaeus). Library of Congress Cataloging in Publication Data Olson, Storrs L Relationships and evolution of flamingos (Aves, Phoenicopteridae) (Smithsonian contributions to zoology ; no. 316) Bibliography: p. 1. Flamingos—Evolution. 2. Flamingos—Classification. 3. Birds, Fossil. 4. Birds—Evolu- tion. 5. Birds—Classification. I. Feduccia, J. Alan, joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no 316 QL1.S54 no. 316 [QL696.C56] 591s [598'.34] 80-607058 Contents Page Introduction 1 Acknowledgments 3 Review of Previous Anatomical Information 3 Life History and Behavior 9 Myology 15 Appendicular Myology of Cladorhynchus 17 Pterylosis 31 Natal Down 34 Oology 34 Parasitology 36 Biochemistry 39 Osteology 39 Skull 39 Postcranial Skeleton 40 Paleontology 43 A New Stilt-like Flamingo from the Middle Eocene of Wyoming 47 Order CHARADRIIFORMES Huxley, 1867 48 Suborder CHARADRII Huxley, 1867 48 Family PHOENICOPTERIDAE Bonaparte, 1831 48 Juncitarsus, new genus 48 Juncitarsus gracillimus, new species 48 Aspects of Evolution of Filter Feeding 58 Conclusions 66 Literature Cited 69 111 Relationships and Evolution of Flamingos (Aves: Phoenicopteridae) Storrs L. Olson and Alan Feduccia Introduction made no innovations in generic usage in the present paper. The living flamingos (Phoenicopteridae) con- The proper phylogenetic position of the Phoen- stitute a small and easily recognized group of four icopteridae has long been a perplexing problem to six species of large, brightly colored waterbirds in avian systematics. Because of the conflicting with extremely long legs and neck and a uniquely nature of the taxonomic evidence presented so shaped bill adapted for filter-feeding. They are far, the relationships of these birds have never colonial and characteristically inhabit highly sa- been satisfactorily resolved. Flamingos are most line bodies of shallow water. The existing species often placed with storks, herons, and ibises in the are placed in three genera that are separable into order Ciconiiformes. This assemblage consists of two groups on bill morphology: (1) Phoenicopterus, large, long-legged waterbirds having a long neck, which has a more primitive filtering apparatus, "desmognathous" palate, and usually altricial and (2) Phoenicoparrus and Phoeniconaias, which are young. Olson (1979) has recently suggested that more specialized (Jenkin, 1957). The last two the order may be an entirely artificial collection genera are separated from each other only by the of unrelated families. The association of flamin- presence or absence of the hind toe, which is very gos with storks and their supposed allies was reduced in flamingos in any case, so that contin- probably made originally solely on the basis of ued recognition of the genus Phoeniconaias is prob- their large size and general proportions. This ably unwarrented. We have not detected char- allocation was later "supported" by a body of acters of generic value in the postcranial skeleton evidence that we shall show to be altogether that will separate the two groups differing in fallacious. feeding adaptations. For purposes of extrafamilial Other workers noted that the lamellate bill, comparisons, modern flamingos can be regarded webbed feet, mallophagan parasites, and some as essentially monogeneric, although we have general aspects of the behavior of flamingos seemed to point toward a relationship with ducks Storrs L. Olson, Department of Vertebrate Zoology, National Museum and geese (Anseriformes). Controversy has sub- of Natural History, Smithsonian Institution, Washington, sequently centered on whether flamingos are DC. 20560. Alan Feduccia, Department of Zoology, University of North Carolina, Chapel Hill, North Carolina 27514. more closely related to the storks or to the ducks. 1 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Continual equivocation has resulted in some au- may be attributable to both having been derived thors elevating the flamingos to an order of their from the Charadriiformes, rather than indicating own (Phoenicopteriformes), an action that merely that they are closely related to each other. Nev- avoids the question of relationships. ertheless, Presbyornis coincidentally pointed the All comparisons of flamingos have traditionally way towards the proper association of flamingos been made with Ciconiiformes on one hand and with the Charadriiformes. Anseriformes on the other, but as we shall see, In the course of our investigations we found most of the characters cited as showing relation- that all the evidence relating to the systematics of ship also occur outside those two groups. Many flamingos
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