THEIR FOSSIL T EW RELATIVES & D A FN EVOLUTIONARY O G R HISTORY D& “Sets a new standard. . . . With Antón’s imagery, Wang and Tedford’s Dogs is nothing TEXT BY XIAOMING WANG & RICHARD H. TEDFORD ILLUSTRATIONS BY MAURICIO ANTÓN short of wonderful.” AMERICAN PALEONTOLOGIST Xiaoming Wang and Richard H. Tedford combine their research with Mauricio Antón’s impeccable reconstructions to present a remarkable portrait of canids over D the past 40 million years. Wang and Tedford cull their history from the most recent scientific research conducted on the vast collections of the American Museum of O Natural History and other leading institutions. With their rich fossil record, diverse G adaptations to various environments, and different predatory specializations, canids are an ideal model organism for the mapping of predator behavior and morpho- S logical specializations. They also offer an excellent contrast to felids, which remain entrenched in extreme predatory specializations. The innovative illustrated approach of this book transforms the science of paleontology into a thrilling visual experience, and it forms the perfect accompaniment to an extremely important branch of animal and fossil study. COLUMBIA ET UNIVERSITY VH “An easy-to-read text, accompanied by Antón’s marvelous illustrations.” CHOICE OE PRESS LI UR NEW YORK TF “A definitive, readable treatment of the evolution of the canine clan.” FOSSIL NEWS IO www.cup.columbia.edu O S N XIAOMING WANG is a curator of vertebrate paleontology at the Natural History Mu- PRINTED IN THE U.S.A. ASI RL COVER DESIGN: MARTIN HINZE seum of Los Angeles County and has been studying the evolutionary history of the Y R COVER IMAGE: MAURICIO ANTÓN family Canidae for the past twenty years. In collaboration with Richard H. Tedford, HE IL he has published three volumes on the fossil canids of North America. SA TT OI RV RICHARD H. TEDFORD is curator emeritus in the Department of Vertebrate Paleon- E Y S tology at the American Museum of Natural History. & MAURICIO ANTÓN is a paleontological artist based at the National Museum of Natural Sciences, Madrid. Among his books are, with Jordi Agustí, Mammoths, Sabertooths, and Hominids: 65 Million Years of Mammalian Evolution in Europe and, with Alan Turner, The Big Cats and Their Fossil Relatives: An Illustrated Guide to Their Evolution and Natural History and Evolving Eden: An Illustrated Guide to the Evolution of the African Large Mammal Fauna. COLUMBIA DOGS DOGS THEIR FOSSIL RELATIVES AND EVOLUTIONARY HISTORY ILLUSTRATIONS BY XIAOMING WANG + RICHARD H. TEDFORD  |  MAURICIO ANTO´N COLUMBIA UNIVERSITY PRESS NEW YORK columbia university press Publishers Since 1893 New York Chichester, West Sussex Copyright © 2008 Columbia University Press Illustrations copyright © 2008 Mauricio Antón Paperback edition, 2010 All rights reserved Library of Congress Cataloging-in-Publication Data Wang, Xiaoming, 1957– Dogs : their fossil relatives and evolutionary history / Xiaoming Wang and Richard H. Tedford ; illustrations by Mauricio Antón. p. cm. Includes bibliographical references and index. isbn 978-0-231-13528-3 (cloth : alk. paper) — isbn 978-0-231-13529-0 (pbk. : alk. paper) — isbn 978-0-231-50943-5 (e-book) 1. Dogs. 2. Canis, Fossil. 3. Dogs—Evolution. I. Tedford, Richard H. II. Antón, Mauricio. III. Title. ql737.c22w36 2008 599.77'2—dc22 2007052328 Columbia University Press books are printed on permanent and durable acid-free paper. Printed in the United States of America c 10 9 8 7 6 5 4 3 2 1 p 10 9 8 7 6 5 4 3 2 1 book design by vin dang + jacket design by martin hinze contents preface vii acknowledgments ix 1 Methods of Study and the Place of Dogs in Nature 1 2 The Origin of Canids and Other Doglike Carnivorous Mammals 7 3 Diversity: Who Is Who in the Dog Family 23 4 Anatomy and Function: How the Parts Work 69 5 Hunting and Social Activity 103 6 Changing Environments and Canid Evolution 117 7 Going Places: Braving New Worlds 139 8 Domestic Dogs 153 appendix 1: Canid Species and Classification 169 appendix 2: Phylogenetic Tree of the Family Canidae 177 glossary 181 further reading 187 index 209 preface the long association of humans and the domestic dog has attracted considerable interest in the biology and natural history of the Canidae (dog fam- ily), among both specialists and the general public. Several recent events have con- verged to inspire further interest in canids. New discoveries of early domestic dogs living in Israel and Germany more than 14,000 years ago push back the fossil record and provide tantalizing evidence of initial interactions between early humans and their first domestic animal. Meanwhile, molecular studies indicate that China, not Europe or the Middle East, may have been the center of first domestication. More- over, humans’ fascination with large predators such as wolves and hunting dogs has continued to draw public attention to a variety of issues ranging from predatory behavior to ecology and conservation. The conservation and reintroduction of gray wolves in Europe and North America and of red wolves in the southeastern United States have promoted public debate regarding the need to protect large predators in order to achieve a balanced ecological community. Against this background of sustained public interest in everything about dogs, the unparalleled Frick Collection (American Museum of Natural History) of North American Cenozoic canids has been the subject of three monographs by Xiaoming Wang (curator of Vertebrate Paleontology), Richard Tedford (curator emeritus of Vertebrate Paleontology), and Beryl Taylor (curator of the Frick Collection) (Wang 1994; Wang, Tedford, and Taylor 1999; Tedford, Wang, and Taylor in press). The wealth of information provided in these monographs reveals a vast array of extinct canids in North America, the continent of their origin, and their subsequent expan- sion to the Old World about 5 to 7 million years ago (Ma) and to South America around 3 Ma. These immigration events ultimately helped to make canids one of the most widely spread living carnivores in the world and the dominant predators on some continents. This book attempts to capture the excitement of recent advances in our understanding of the natural history of this family of carnivorans. The book is viii • preface richly illustrated, both to provide an unprecedented visual reference for the special- ists and to make it accessible and interesting to a wide nontechnical audience. The family Canidae was the earliest group that still has extant members to arise within the order Carnivora. During their history of more than 40 million years, canids have also achieved a longevity and diversity unrivaled by any other group of carnivores. Their success is also exemplified by a worldwide distribution and achievement of top predator status in modern North and South America, Australia, and northern Eurasia—only in Africa are they less dominant among large predators. It is argued that the propensity for canids to form large, social hunting groups and the associated development of their brain were crucially important in the domestica- tion process—humans learned to form a mutually beneficial relationship with such an intelligent carnivoran that was preadapted to life among a different species. The importance of the domestication of various animals in early human societies cannot be overestimated, and it may have been first inspired by the domestication of dogs. The evolutionary history of canids is a history of successive radiations (rapid ex- pansions of diversity within a group of organisms, often in response to environmen- tal changes or new resources) repeatedly occupying a broad spectrum of niches, ranging from large, pursuit predators to small omnivores and possibly even to her- bivores. Three such radiations were first recognized by Richard Tedford (1978), each represented by a distinct subfamily. Two archaic subfamilies, Hesperocyoninae and Borophaginae, thrived in the middle to late Cenozoic from about 40 to 2 Ma. All extant canids belong to the final radiation, the subfamily Caninae, which achieved its present diversity only in the past few million years. Previous diversities of canids were intimately related to environmental changes during the past 40 million years. Large, wolflike pursuit predators were in particular the direct consequence of the opening up of the landscape. A worldwide change from a warm and humid climate to an increasingly colder and drier climate resulted in large-scale transformation of forested environments to open grasslands world- wide. These changes were reflected in the terrestrial vertebrate community by an increasing cursoriality (the ability to run fast and for a long time) among predators and prey. This greater ability to run is clearly reflected in the fossil history of canids, which shows increasingly erect standing postures, progressively lengthened limb bones, and restricted movements of the joints. Humans’ fascination with canids is derived from two main sources: our love affair with our dogs and our reverence of their top predator status in much of the world. Such a high interest in a group of mammals is generally rare and offers a great op- portunity to engage the public in paleontology, functional morphology, evolution, and behavioral ecology. acknowledgments we thank our publishers at Columbia University Press, Patrick Fitzger- ald and Robin Smith, and editor Irene Pavitt for their encouragement and helpful guidance. We are indebted to Lars Werdelin and Blaire Van Valkenburgh for their critical reviews, which brought to our attention records that we missed and other factual errors. A thorough editing by copyeditor Annie Barva has greatly improved the language and expression of this book. Alejandra Lora of the American Museum of Natural History typed part of the manuscript for publication and helped in nu- merous other ways to facilitate communications. A substantial part of the mono- graphic treatments of the three subfamilies of canids were funded by two grants from the National Science Foundation (DEB-9420004 and DEB-9707555) and a Frick Postdoctoral Fellowship from the American Museum of Natural History. Xiaoming Wang thanks his wife, Yanping, and his son, Alex, for their support and understanding throughout the preparation of this book. Richard Tedford is grateful for the patience of Elizabeth and Vivien during the 30 years this study took to com- plete. Mauricio Antón thanks his wife, Puri, and his son, Miguel, for their patience during this long process.