BIRD Migration BIRD Migration A New Understanding JOHN H. RAPPOLE Johns Hopkins University Press Baltimore © 2022 Johns Hopkins University Press All rights reserved. Published 2022 Printed in the United States of America on acid-free paper 2 4 6 8 9 7 5 3 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218–4363 www.press.jhu.edu Library of Congress Cataloging-in-Publication Data Names: Rappole, John H., author. Title: Bird migration : a new understanding / John H. Rappole. Description: Baltimore : Johns Hopkins University Press, 2022. | Includes bibliographical references and index. Identifiers: LCCN 2021002624 | ISBN 9781421442389 (hardback) | ISBN 9781421442396 (ebook) Subjects: LCSH: Birds—Migration. Classification: LCC QL698.9 .R375 2022 | DDC 598.156/8—dc23 LC record available at https://lccn.loc.gov/2021002624 A catalog record for this book is available from the British Library. All illustrations not otherwise credited are the author’s. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at [email protected]. Among the most important discoveries for me con- cerning migratory birds is the central role played by the distaff members: not “subservient,” not “subordi- nate,” not “supporting,” but central. Male dominance, long considered the wellspring of avian interactions between the sexes, is found to be impotent, at least during the migrant breeding season. It is females who call the shots, with all due respect to Konrad Lorenz and many others. Accordingly, I dedicate this effort to the extraordinary women who have played central roles in my own life: my wife, Bonnie Carlson Rappole; my mother, Francesca Goodell Rappole; my daughter, Brigetta Rappole Stewart; and my sisters, Francesca Rappole Wellman Miller and Rosemary Rappole. This page intentionally left blank Contents Preface ix Acknowledgments xv CHAPTER 1 The Bird Migration Paradigm 1 CHAPTER 2 The Migrant Annual Cycle According to the Dispersal Theory 8 CHAPTER 3 Fall Migration 17 CHAPTER 4 Wintering Period 38 CHAPTER 5 Spring Migration 77 CHAPTER 6 Breeding Period 95 CHAPTER 7 Postbreeding Period 115 CHAPTER 8 Population Biology 127 CHAPTER 9 Origin and Evolution 162 CHAPTER 10 Biogeography 202 CHAPTER 11 Conservation 221 Coda 263 APPENDIX 1 Common and Scientific Names of Bird Species Mentioned in the Text 275 APPENDIX 2 A Critical Examination of the Assumptions in “Temperate Origins of Long-Distance Seasonal Migration in New World Songbirds” 279 APPENDIX 3 Notation Corrections for Alan Pine’s Multiple Carrying Capacity Equations from “Age-Structured Periodic Breeders” 295 Annotated Bibliography 297 Index 313 Preface Most people think they know a lot about bird migration. Almost everyone has seen many explanations for the phenomenon in a variety of media formats. However, these explanations are based on the premise that migration is all about weather (Figure P.1). It’s not. Certainly weather, es- pecially winds aloft, can be important in shaping some movements of some species some of the time, but it is not the ultimate cause. Migration is about competition and seasonal change in food resourc- es. It originates when young birds are forced to move away from the place where they were born in search of suitable habitat that is not already oc- cupied by competitors—habitat where they can find mates and raise more young than they would have been able to had they stayed at home. This movement, known as “dispersal,” is possible because of a superabundance of food. Without such a superabundance, the travelers would not get very far from home. Dispersal is the engine driving migration; seasonal change in food availability is the enabler. When food becomes superabundant— that is, available in greater supply than resident birds can consume due to seasonal climate change—the disperser can move and find new areas where it is possible to live and breed in the absence of, or at least in a re- duction of, competition. If the climate changes at some later date, causing a disappearance of the foods on which the disperser depends, then the dis- Preface ix