Paleogene Fossil Birds Gerald Mayr Paleogene Fossil Birds ISBN 978-3-540-89627-2 e-ISBN 978-3-540-89628-9 DOI 10.1007/978-3-540-89628-9 Library of Congress Control Number: 2008940962 © 2009 Springer-Verlag Berlin Heidelberg This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: WMXDesign GmbH, Heidelberg, Germany Printed on acid-free paper springer.com Gerald Mayr Forschungsinstitut Senckenberg Sektion Ornithologie Senckenberganlage 25 D-60325 Frankfurt am Main Germany e-mail: Gerald.Mayr@senckenberg.de Preface Since birds are predominantly diurnal and often quite vociferous animals, their behavior and ecological requirements are probably better studied than those of any other vertebrate group. Detailed knowledge of their evolutionary history is, how- ever, still limited to a small circle of specialists, and there is a widespread belief that the avian fossil record is poor. This is certainly true if the abundance of bird bones is compared with that of mammalian teeth, which are robust enough to survive even rough depositional environments and collection techniques. In many fossil locali- ties complete skeletons and postcranial elements of birds are, however, not much rarer than those of other small land vertebrates. Numerous avian fossils in collec- tions worldwide have remained further unstudied for decades, so the significant underrepresentation of birds in vertebrate paleontology seems to be due to a low number of specialists rather than a low number of fossils. Concerning certain geological periods and geographic areas, our knowledge of the early evolutionary history of birds is anything but poor. In fact, so many new fossils were described during the past two decades that it becomes increasingly difficult for a single person to cover the whole field of paleornithology. This book gives an account of the evolution of modern birds in the first half of the Cenozoic, aiming not only at specialists in the field of paleornithology, but also at ornithologists and paleontologists in need of detailed information, either for the calibration of molecular data or to set Paleogene faunas into a full context. Given the current pace of new discoveries, I am not cherishing the illusion that this survey will remain up to date for a long time. I do hope, however, that the overall frame- work outlined for the early diversity and evolution of modern birds will form a stable basis for future studies, and that the readers will find the book a useful source for their own research. Frankfurt am Main Gerald Mayr October 2008 v Acknowledgements I am indebted to Sven Tränkner for taking the photographs, Cécile Mourer- Chauviré for discussions on the fossil birds from the Quercy fissure fillings, and Albrecht Manegold for insightful comments on parts of the manuscript. For provid- ing photographs of fossil specimens, I thank Herculano Alvarenga, Julia Clarke, Ewan Fordyce, James Goedert, Peter Houde, Dan Ksepka, Bent Lindow, Cécile Mourer-Chauviré, Norbert Micklich, and Ilka Weidig. Access to fossil specimens was kindly provided by Walter Boles, Elvira Brahm, Sandra Chapman, Michael Daniels, Dino Frey, Norbert Hauschke, Meinolf Hellmund, Peter Houde, Norbert Micklich, Cécile Mourer-Chauviré, Wolfgang Munk, Burkhard Pohl, Stephan Schaal, Thierry Smith, and Basil Thüring. I am further obliged to Dieter Czeschlik for enabling this book project, and Anette Lindqvist and Thavamani Saravanan for their efforts in the production of this book. Above all, however, I thank my wife, Eun-Joo, for her patience and moral support during the preparatory stage of this work. vii Contents 1 Introduction 1 2 Stratigraphy and Major Fossil Localities 5 2.1 Europe 5 2.2 Asia 7 2.3 North America 8 2.4 Central and South America 9 2.5 Africa 10 2.6 Australia, New Zealand, and Antarctica 10 3 Higher-Level Phylogeny of Extant Birds 13 4 Mesozoic Neornithes 19 5 Palaeognathous Birds 25 5.1 †Lithornithidae 26 5.2 †Palaeotididae, †Remiornithidae, and †Eleutherornithidae 28 5.2.1 †Palaeotididae 28 5.2.2 †Remiornithidae 29 5.2.3 †Eleutherornithidae 30 5.3 †Eremopezidae 31 5.4 Rheidae (Rheas) 32 5.5 Casuariidae (Emus and Cassowaries) 33 5.6 Putative Ratite from the Eocene of Antarctica 33 6 Galloanseres 35 6.1 Galliformes (Landfowl) 35 6.1.1 †Gallinuloididae 36 6.1.2 †Paraortygidae 40 6.1.3 †Procrax, †Archaealectrornis, and †Palaeonossax 41 6.1.4 †Quercymegapodiidae 41 ix x Contents 6.1.5 Megapodiidae (Megapodes) 42 6.1.6 Phasianidae (Grouse, Quails, Pheasants, and Allies) 42 6.2 †Gastornithidae 44 6.3 †Dromornithidae 47 6.4 Anseriformes (Waterfowl) 48 6.4.1 Anhimidae (Screamers) 48 6.4.2 Anseranatidae (Magpie Geese) 49 6.4.3 †Presbyornithidae 51 6.4.4 Anatidae (Ducks, Geese, and Swans) 53 6.5 †Pelagornithidae (Bony-Toothed Birds) 55 7 Aquatic and Semiaquatic Taxa 61 7.1 Fregatidae (Frigatebirds) and Suloidea (Gannets, Boobies, Cormorants, and Anhingas) 61 7.1.1 †Protoplotidae 62 7.1.2 Fregatidae (Frigatebirds) 63 7.1.3 Sulidae (Gannets and Boobies) 64 7.1.4 Phalacrocoracidae (Cormorants) and Anhingidae (Anhingas) 65 7.2 †Plotopteridae 67 7.3 Sphenisciformes (Penguins) 70 7.4 Gaviiformes (Loons) 75 7.5 Procellariiformes (Tubenoses) 76 7.6 Scopidae (Hamerkop), Balaenicipitidae (Shoebill), and Pelecanidae (Pelicans) 80 7.7 Ardeidae (Herons) 80 7.8 †Xenerodiopidae 81 7.9 Threskiornithidae (Ibises) 81 7.10 Ciconiidae (Storks) 84 7.11 †Prophaethontidae and Phaethontidae (Tropicbirds) 84 8 Charadriiformes (Shorebirds and Allies) 87 8.1 Lari (Gulls, Auks, and Allies) 88 8.2 Charadrii (Plovers and Allies) 89 8.3 Scolopaci (Sandpipers and Allies) 90 9 “Core-Gruiformes” (Rails, Cranes, and Allies) 93 9.1 †Messelornithidae and †Walbeckornis 93 9.2 Ralloidea (Finfoots and Rails) 96 9.3 Gruoidea (Trumpeters, Limpkins, and Cranes) 99 9.3.1 †Parvigruidae 100 9.3.2 †Geranoididae 101 9.3.3 †Eogruidae 102 9.3.4 Aramidae (Limpkins) and Gruidae (Cranes) 103 Contents xi 10 Phoenicopteriformes (Flamingos) and Podicipediformes (Grebes) 105 11 Columbiformes (Doves and Sandgrouse), Cuculiformes (Cuckoos), and Other Neoavian Taxa of Uncertain Affi nities 111 11.1 Columbiformes (Doves and Sandgrouse) 111 11.2 Opisthocomiformes (Hoatzin) 112 11.3 †Foratidae 112 11.4 Musophagiformes (Turacos) 113 11.5 Cuculiformes (Cuckoos) 113 11.6 †Pumiliornis and †Morsoravis 114 11.7 †Parvicuculidae 115 11.8 Otididae (Bustards), Eurypygidae (Sunbittern), Rhynochetidae (Kagu), and Mesitornithidae (Mesites) 116 12 “Caprimulgiformes” and Apodiformes (Nightjars and Allies, Swifts, and Hummingbirds) 119 12.1 †Fluvioviridavidae 119 12.2 Steatornithidae (Oilbirds) 124 12.3 Podargidae (Frogmouths) 124 12.4 †Protocypselomorphus 125 12.5 †Archaeotrogonidae 126 12.6 Nyctibiidae (Potoos) and Caprimulgidae (Nightjars) 128 12.6.1 Nyctibiidae 128 12.6.2 Caprimulgidae 128 12.7 Aegothelidae (Owlet-Nightjars) and Apodiformes (Swifts and Hummingbirds) 129 12.7.1 †Eocypselidae 130 12.7.2 †Aegialornithidae 132 12.7.3 Hemiprocnidae (Tree Swifts) and Apodidae (True Swifts) 133 12.7.4 Trochilidae (Hummingbirds) 135 13 Cariamae (Seriemas and Allies) 139 13.1 †Phorusrhacidae 139 13.2 †Idiornithidae and †Elaphrocnemus 142 13.3 †Bathornithidae 146 13.4 †Ameghinornithidae 148 13.5 †Salmilidae 150 13.6 †Gradiornis 152 xii Contents 14 “Falconiformes” (Diurnal Birds of Prey) 153 14.1 Falconidae (Falcons) 153 14.2 †Masillaraptor 155 14.3 †Teratornithidae and Cathartidae (New Word Vultures) 156 14.4 †Horusornithidae 158 14.5 Sagittariidae (Secretary Birds), Pandionidae (Ospreys), and Accipitridae (Hawks and Allies) 158 14.5.1 Sagittariidae 158 14.5.2 Accipitridae and Pandionidae 159 15 Strigiformes (Owls) 163 15.1 †Berruornis and †Sophiornithidae 163 15.2 †Protostrigidae 164 15.3 †Ogygoptyngidae 165 15.4 †Necrobyinae, †Palaeoglaucidae, and †Selenornithinae 166 16 Arboreal Birds 169 16.1 Leptosomidae (Courols) 169 16.2 Coliiformes (Mousebirds) 171 16.2.1 †Sandcoleidae 172 16.2.2 Coliidae 172 16.3 Psittaciformes (Parrots) 177 16.3.1 †Psittacopes and Allies 177 16.3.2 †Quercypsittidae 179 16.3.3 †Halcyornithidae (“Pseudasturidae”) 180 16.3.4 †Messelasturidae 183 16.4 †Zygodactylidae and Passeriformes (Passerines) 184 16.4.1 †Zygodactylidae 185 16.4.2 Passeriformes 189 16.5 Trogoniformes (Trogons) 191 16.6 Bucerotes (Hornbills, Hoopoes, and Woodhoopoes) 191 16.7 Coraciidae/Brachypteraciidae (Rollers and Ground Rollers) 194 16.7.1 †Primobucconidae 195 16.7.2 †Eocoraciidae and †Geranopteridae 195 16.8 Alcediniformes (Bee-Eaters, Kingfi shers, Todies, and Motmots) 197 16.9 Piciformes (Jacamars, Puffbirds, Woodpeckers, and Allies) 199 16.9.1 †Sylphornithidae 200 16.9.2 Pici 201 16.10 †Gracilitarsidae 201 16.11 †Cladornithidae 203 Contents xiii 17 Paleogene Avifaunas: Synopsis of General Aspects 205 17.1 Continental Avifaunas of the Northern Hemisphere 205 17.1.1 Biogeography 205 17.1.2 Climatic Cooling and Avifaunal Turnovers 208 17.2 Continental Avifaunas of the Southern Hemisphere 209 17.2.1 Biogeography 209 17.2.2 Extant Southern Hemisphere “Endemics” in the Paleogene of the Northern Hemisphere 213 17.3 Ecological Interactions 214 17.3.1 Mammalian Evolution and Terrestrial Avifaunas 214 17.3.2 The Impact of Passerines on the Diversity of Paleogene Avian Insectivores 216 17.3.3 Marine Avifaunas 217 Appendix 221 References 227 Index 255 [...]... distribution found in Paleogene fossil birds The book aims at bringing some of this information together, and many of the following data are based on first-hand examination of fossil specimens In the first chapter, the most important fossil localities for Paleogene birds are introduced I then outline current hypotheses on the higher-level phylogeny of birds and summarize the Mesozoic fossil record of Neornithes... Chapter 2 Stratigraphy and Major Fossil Localities In this chapter some of the major sites which yielded Paleogene fossil birds are briefly introduced to avoid redundancies in the taxonomic sections Dyke et al (2007, p 341) stated that “aquatic environments of preservation dominate the early Paleogene avian fossil record, because these were the habitats in which more modern birds lived at the time of the... least in the Northern Hemisphere, the Paleogene fossil record of birds is no longer much short of the mammalian one concerning the number of well-represented higher-level taxa In some renowned fossil localities, such as the London Clay in England and the Green River Formation in North America, remains of birds are even much more abundant than mammalian remains Numerous Paleogene avian taxa were described... without modern counterparts, and allow intriguing insights into the past diversity of long-vanished avifaunas G Mayr, Paleogene Fossil Birds, DOI: 10.1007/978-3-540-89628-9_1, © Springer-Verlag Berlin Heidelberg 2009 1 2 1 Introduction In the present book the Paleogene fossil record of birds is detailed for the first time on a worldwide scale I have developed the idea for such a project for several years,... et al 2002), there are no published bird fossils from the Paleogene of Central America The Cenozoic avian fossil record of South America was summarized by Tonni (1980), but many Paleogene fossils were described after this study The earliest specimens stem from late Paleocene (Itaboraian) marl fillings of the Bacia Calcária of Itaboraí in Brazil The described fossils from this site belong to a palaeognathous... Namibia According to Mourer-Chauviré et al (1996), G Mayr, Paleogene Fossil Birds, DOI: 10.1007/978-3-540-89628-9_5, © Springer-Verlag Berlin Heidelberg 2009 25 26 5 Palaeognathous Birds the fossils of this species suggest that didactyl ostriches originated in Africa before the Miocene, from where they dispersed into Eurasia in the Neogene The earliest fossil Tinamidae are from the Miocene of Argentina (Bertelli... often fragmentary bones 2.6 Australia, New Zealand, and Antarctica Likewise, very little is known about the Paleogene avifaunas of Australia, and most fossil birds from this period were described after the reviews by Vickers-Rich (1991) and Boles (1991) The continent has no Paleocene fossil record of birds, and the only Eocene avian specimens stem from the deposits of the early Eocene Tingamarra Local Fauna... Quaternary fossil record and except for penguins, Paleogene birds are also rare in New Zealand (Worthy and Holdaway 2002) The Eocene and Oligocene localities where penguins have been found were summarized by Simpson (1971) More recently, well-preserved remains of fossil stem group representatives of the Sphenisciformes were found in the Paleocene (ca 58–61 Ma) Waipara Greensand (Slack et al 2006) The Paleogene. .. Procellariiformes Earlier authors assumed that various extant birds whose hindlimbs lack the ambiens muscle are closely related, and a group including these birds was informally termed “higher land birds by Olson (1985) In its usual composition, this 18 3 Higher-Level Phylogeny of Extant Birds assemblage includes the Strigiformes (owls), Coliiformes (mousebirds), Trogoniformes (trogons), Piciformes, Passeriformes... all nonneornithine birds became extinct at the Cretaceous Paleogene boundary (Feduccia 1995, 2003; van Tuinen et al 2006), the factual evidence for this hypothesis is rather weak, given the poor early Paleocene fossil record of birds At least Qinornis paleocenica Xue, 1995 from the early Paleocene of China may represent a nonneornithine avian lineage, which survived the Cretaceous Paleogene extinction . Paleogene Fossil Birds Gerald Mayr Paleogene Fossil Birds ISBN 978-3-540-89627-2 e-ISBN 978-3-540-89628-9 DOI 10.1007/978-3-540-89628-9 Library. G. Mayr, Paleogene Fossil Birds, DOI: 10.1007/978-3-540-89628-9_1, 1 © Springer-Verlag Berlin Heidelberg 2009 2 1 Introduction In the present book the Paleogene fossil record of birds is detailed. the most important fossil localities for Paleogene birds are introduced. I then outline current hypotheses on the higher-level phylogeny of birds and summarize the Mesozoic fossil record of Neornithes.