Handbook of Avian Hybrids of the World EUGENE M. McCARTHY OXFORD UNIVERSITY PRESS Handbook of Avian Hybrids of the World This page intentionally left blank Handbook of Avian Hybrids of the World EUGENE M. MC CARTHY 3 2006 3 Oxford University Press, Inc., publishes works that further Oxford University’s objective of excellence in research, scholarship, and education. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugual Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Copyright © 2006 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data McCarthy, Eugene M. Handbook of avian hybrids of the world/Eugene M. McCarthy. p. cm. ISBN-13 978-0-19-518323-8 ISBN 0-19-518323-1 1. Birds—Hybridization. 2. Birds—Hybridization—Bibliography. I. Title. QL696.5.M33 2005 598′.01′2—dc22 2005010653 987654321 Printed in the United States of America on acid-free paper For Rebecca, Clara, and Margaret This page intentionally left blank For he who is acquainted with the paths of nature, will more readily observe her deviations; and vice versa, he who has learnt her deviations, will be able more accurately to describe her paths. —Francis Bacon, Novum Organum (1620) This page intentionally left blank Preface his book is intended to provide basic information about each of the thousands Tof types of reported avian crosses, to provide access to documenting literature, and to familiarize readers with the nature of avian hybridization. I have long been interested in hybridization, especially its role in evolutionary processes. I began this book because I wanted to make a thorough survey of hybridization within the con- text of a well-studied class of organisms. I thought birds would be ideal. There was plenty of data available, much as yet ungathered. Available surveys were far out of date. Moreover, more information on geographic distribution is available for birds than for almost any other major group of organisms. Detailed information on spa- tial distribution greatly facilitates the identification of hybrid zones. My general approach to the subject has been exploratory. I therefore often include not only prima facie evidence, but also facts merely suggesting the occurrence of hybridiza- tion. My intention in writing this book was to learn more about the subject myself, and to help and encourage others to study it, too. The construction of a reference on avian hybridization is an enterprise fraught with hazard. There are many pitfalls related to spelling, nomenclature, and classifi- cation, as well as an ever-present risk of factual error, often exacerbated by con- flicting reports. For each cross, various details must be accurately determined, to the extent that information is available: Are the hybrids fertile? Are they viable? What do they look like? Do the birds participating in the cross come into natural contact? If so, where? Are there additional reports of the cross that have been omit- ted? While I have done my utmost to eliminate mistakes, any book dealing with a subject of such complexity is liable to error. In addition, the literature on avian hybridization is so vast that I am sure that I have inadvertently omitted some reported crosses. The bibliography lists more than 5,000 references, but some for- eign language publications and articles in out-of-the-way breeding journals have no doubt escaped my attention. Nevertheless, I believe this book does constitute a step forward in the understanding of avian hybridization. This page intentionally left blank Acknowledgments arious portions and drafts of the manuscript were circulated for comments and VI am grateful here to my fellow University of Georgia scientists Paul Mack and J. P. Schmidt. I also appreciate the helpful suggestions of two anonymous review- ers, and the validation reviewer, W. S. Moore. The reference staff at the University of Georgia Science Library, Lucy Rowland, Diana Hartle, and Monica Pereira in par- ticular, have provided unstinting and invaluable assistance, as has our Computer Services Specialist, Dave Brown, here in the Genetics Department. I am indebted to many people for their generosity in answering correspondence and supplying information and articles relevant to this work, namely: Bill Aragon, Curator of Birds, Albuquerque Biological Park; Jonathan Barnaby, Department for Environment, Food and Rural Affairs, UK; Mark Bremer, National Aviary; Michael Brooks, SAFRING Avian Demography Unit; Nick Fox, International Wildlife Consultants Ltd.; Anthony Ganesh, Singapore Zoological Gardens; Jim Granlund, Michigan Audubon Society; Holly Green, Registrar, Indianapolis Zoo; Kai Grosch, Max-Planck-Forschungsstelle für Ornithologie; David Hancock, Hancock Wildlife Research Center; Johan Ingels; Laszlo Janossy, Curator, Budapest Zoo; James Kamstra; Daisy Ling, Dep. Exec. Director, Jurong Bird Park, Singapore; Tim Melling, British Ornithological Union; Jemima Parry-Jones, National Birds of Prey Centre, UK; Lucia Liu Severinghaus, Academica Sinica, Taipei; Daniel Shearing, Editor, Foreign Bird League 1991–2003; Michael Sorenson, Boston University; Per Gustav Thingstad, Museum of Natural History and Archaeology, Trondheim; William Todd, Assistant Curator of Birds, Houston Zoo; Francis Toldi; Ülo Väli, Estonian Agricultural University; Renate van den Elzen, Alexander Koenig Research Institute and Museum of Zoology; Vince Walkosak, Arizona-Sonora Desert Museum; Carolyn Walsh, Memorial University of Newfoundland. My good friend Stuart Katz has read numerous draft manuscripts and provided me with many keen insights and valuable suggestions. Finally I thank my wife, Rebecca, who throughout has provided me with her advice, criticism, encour- agement, and patience, as well as editorial assistance. This page intentionally left blank Contents Introduction Rollers 117 Courol 117 Using this Book 3 Motmots 117 Identifying Hybrids 14 Todies 118 Hybrid Zones 18 Kingfishers 118 Hybrid Populations and Taxa 27 Kookaburras 119 Causes of Hybridization 31 Bee-eaters 119 Underreporting 33 Mousebirds 120 Future Research 37 Cuckoos and Coucals 120 Parrots and Their Allies 121 Cross Accounts Cockatoos 141 Ostriches, Rheas, and Kiwis 39 Swifts 143 Tinamous 39 Crested-Swifts 143 Brush-turkeys and Scrubfowls 39 Hummingbirds 144 Upland Game Birds 41 Turacos 155 New World Quails 65 Barn Owls 155 Screamers 67 Typical Owls 156 Swans, Geese, and Ducks 67 Nightjars, Frogmouths, Honeyguides 102 and Nighthawks 158 Woodpeckers 102 Pigeons and Doves 158 Barbets and Tinkerbirds 110 Bustards 165 Toucans and Aracaris 112 Cranes 165 Jacamars 114 Rails 167 Puffbirds 114 Sandgrouses 168 Hornbills 114 Plovers, Sandpipers, and Their Allies 169 Hoopoes 115 Jacanas 171 Woodhoopoes 115 Oystercatchers 171 Scimitar-bills 116 Stilts and Avocets 172 Trogons 116 Pratincoles 172 xiv Contents Jaegers and Skuas 173 Cuckoo-shrikes 233 Gulls, Terns, and Noddies 173 Fantails and Drongos 233 Alcids 179 Ioras 234 Eagles, Hawks, and Old World Bushshrikes and Helmetshrikes 234 Vultures 180 Waxwings 235 Falcons 184 Thrushes 235 Grebes 187 Old World Flycatchers 241 Boobies and Gannets 188 Starlings and Mynas 243 Cormorants 188 Mimids 245 Herons and Egrets 189 Nuthatches 246 Flamingos 192 Wrens, Tree-Creepers, Ibises and Spoonbills 192 and Gnatcatchers 247 Pelicans 193 Tits, Chickadees, and Their Allies 248 New World Vultures 194 Swallows 253 Storks 194 Regulids 255 Penguins 194 Bulbuls and Greenbuls 255 Loons 195 Cisticolids 258 Shearwaters, Petrels, and Prions 196 White-eyes 260 Albatrosses 198 Old World Warblers 261 Pittas 199 Larks 265 Asities 199 Sunbirds and Their Allies 266 Tyrant Flycatchers 199 Old World Sparrows 268 Manakins 204 Pipits and Wagtails 270 Ovenbirds and Their Allies 205 Weavers and Their Allies 272 Australo-Papuan Treecreepers 210 Estrildine Finches 275 Lyrebirds 210 Indigobirds and Whydahs 294 Bowerbirds 211 Fringillid Finches 295 Fairywrens 211 Hawaiian Finches 307 Honeyeaters 212 Wood Warblers 307 Pardalotids 214 Tanagers, Cardinals, and Australo-Papuan Robins 215 Their Allies 316 Australo-Papuan Babblers 216 New World Blackbirds 333 Shrikes 216 Vireos and Greenlets 219 Appendices Quail-thrushes 220 1 Canary Crosses 339 Sittellas 220 2 Dubious Reports 345 Whistlers and Shrike-thrushes 221 3 Internet Citations 349 Jays and Magpie-Jays 221 Crows, Ravens, Magpies, Works Cited 355 and Nutcrackers 224 Birds of Paradise 228 Artamids 232 Index of English and Scientific Figbirds and Orioles 233 Names 515 Handbook of Avian Hybrids of the World This page intentionally left blank Introduction andbook of Avian Hybrids of the World attempts to list all avian crosses reported Hin the scientific literature and/or on the Internet (the vast majority of the documentation is of the former type). Quite a few personal communications are also included. It is the broadest survey of its kind to date, listing not only crosses occur- ring under natural conditions,