The Rise of Birds This page intentionally left blank The Rise of Birds 225 Million Years of Evolution Second Edition Sankar Chatterjee Johns Hopkins University Press Baltimore © 2015 Johns Hopkins University Press Special discounts are available for bulk purchases of this book. For All rights reserved. Published 2015 more information, please contact Special Sales at 410- 516- 6936 or Printed in the United States of America on acid- free paper [email protected] .edu. 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press uses environmentally friendly Johns Hopkins University Press book materials, including recycled text paper that is composed 2715 North Charles Street of at least 30 percent post- consumer waste, whenever Baltimore, Maryland 21218- 4363 possible. www .press.jhu .edu Library of Congress Cataloging-in-Publication Data Chatterjee, Sankar. The rise of birds : 225 million years of evolution. — Second edition. pages cm Includes bibliographical references and index. ISBN 978-1-4214-1590-1 (hardcover : alk. paper) — ISBN 978-1-4214-1614-4 (electronic) — ISBN 1-4214-1590-9 (hardcover : alk. paper) — ISBN 1-4214-1614-X (electronic) 1. Birds—Evolution. 2. Birds, Fossil. 3. Birds—Flight. I. Title. QL677.3.C48 2015 598.13'8—dc23 2014017134 A catalog record for this book is available from the British Library. For Sibani, with my love This page intentionally left blank Contents Preface to the Second Edition ix Preface to the First Edition xiii 1 Mesozoic Pompeii 1 2 The Evolution of an Airframe 9 3 The Origin of Birds 19 4 Archaeopteryx: An Ancient Wing 57 5 Protoavis: A Precocious Triassic Bird 72 6 Basal Avialans: The Long- Tailed Birds 106 7 Pygostylia: The Short- Tailed Birds 113 8 Enantiornithes: Global Cretaceous Birds 123 9 Ornithuromorphs: The Prelude to Modern Birds 140 10 The End- Cretaceous Mass Extinction 158 11 The Avian Revolution Begins 175 12 The Origin of Flight 213 13 Eggs, Embryos, and Heterochrony 265 14 Feathers and Footprints 287 15 The Feeding Mechanism and Cranial Kinesis 304 16 Birds and Humans 333 Bibliography 343 Index 361 This page intentionally left blank Preface to the Second Edition Endowed with colorful plumages and beautiful songs, known groups of extant vertebrates, their origin, evolu- birds in flight symbolize spirits released from the bond- tion, and early adaptive radiation are poorly documented age of gravity. From the day that humans first looked up in the fossil record. Until recently, most knowledge of at the skies, birds have summoned a sense of wonder and the early history of birds and the evolution of their flight mystery, enchanting our earthbound ancestors with their was based on handful of diverse Mesozoic taxa widely freedom and song. They fly where they please and when separated in time and restricted to marine environments. they please. The power of flight has opened up to birds The rarity of bird fossils is generally attributed to the ex- a multilayered network of aerial highways and byways, treme fragility, lightness, pneumaticity, arboreal habits, enabling them to reach any place on our planet. They ex- and general smallness of their bones. However, there is a hibit a wide variety of sizes, shapes, colors, and habits. renaissance in the study of Mesozoic birds with the spec- Their awesome power of flight takes them to every con- tacular discoveries of amazing fossils from China during tinent, from pole to pole, and they fill every conceivable the past two decades. These new discoveries fill major niche. Birds have enriched our life with their free spirit, gaps in the evolutionary history of birds and provide beauty, and songs. They inform us about the rest of the critical insights into the origin of feathers, evolution of natural world, control insect population, disperse seeds, flight, and their phylogenetic relationships. and embody key values in human cultures. They have The Mesozoic has often been referred to as the “Dark played a key role in many important developments of Ages of Birds” because only a limited number of fossils evolutionary theories. Given their stark beauty, graceful such as Archaeopteryx from the Late Jurassic of Germany flight, exuberant activity, diurnal habits, dazzling feath- and Hesperornis and Ichthyornis from the Late Cretaceous ers, and melodious songs, it is hardly surprising that birds of Kansas filled the evolutionary gaps. Archaeopteryx was feature strongly in every culture in painting, poetry, and an icon in avian evolution, being regarded as the oldest music. Birds are the most successful terrestrial vertebrate, and most primitive known bird. In recent decades the abundant in both numbers of species and populations. narrative of bird evolution has broadened considerably About 300 billion birds, with 10,000 species, now inhabit with the discovery of more fossils from around the world. the Earth, as compared to 3,000 species of amphibians, Our story of bird evolution began 225 million years ago 6,000 species of reptiles, and perhaps 4,500 species of when Protoavis took to the air over tropical Texas forests mammals. at the very dawn of the age of dinosaurs and began to Birds are one of the oldest, most successful, and fasci- diverge in many forms and fashions. Soon after the dis- nating groups of flying vertebrates in our landscape. The covery of Protoavis, remarkable fossils of Cretaceous birds story of bird evolution as revealed by the study of fossils were unearthed in China, Spain, Mongolia, Australia, is important not only to paleontologists but also to those Madagascar, Argentina, and Antarctica. The present edi- engaged in other branches of science—geology, biology, tion represents a radical metamorphosis of the first edi- ecology, evolution, and natural history—as well as the tion of The Rise of Birds, published in 1997, amplified by the general public. Although living birds are one of the best- remarkable discovery of feathered dinosaurs and early