Helpers At Birds' Nests : A Worldwide Survey of title: Cooperative Breeding and Related Behavior author: Skutch, Alexander Frank. publisher: University of Iowa Press isbn10 | asin: 0877456747 print isbn13: 9780877456742 ebook isbn13: 9781587292255 language: English subject Birds--Behavior, Cooperative breeding in animals. publication date: 1999 lcc: QL698.3.S55 1999eb ddc: 598.1563 subject: Birds--Behavior, Cooperative breeding in animals. Page iii Helpers at Birds' Nests A Worldwide Survey of Cooperative Breeding and Related Behavior An Expanded Edition Alexander F. Skutch Illustrated by Dana Gardner Foreword by Stephen T. Emlen Page iv University of Iowa Press, Iowa City 52242 Copyright © 1999 by the University of Iowa Press All rights reserved Printed in the United States of America http://www.uiowa.edu/~uipress No part of this book may be reproduced or used in any form or by any means, electronic or mechanical, including photocopying and recording, without permission in writing from the publisher. All reasonable steps have been taken to contact copyright holders of material used in this book. The publisher would be pleased to make suitable arrangements with any whom it has not been possible to reach. Printed on acid-free paper Library of Congress Cataloging-in-Publication Data Skutch, Alexander Frank, 1904- Helpers at birds' nests: a worldwide survey of cooperative breeding and related behavior/by Alexander F. Skutch; illustrated by Dana Gardner; foreword by Stephen T. Emlen.An expanded ed. p. cm. Includes bibliographical references and index. ISBN 0-87745-674-7 (pbk.) 1. BirdsBehavior. 2. Cooperative breeding in animals. I. Title. QL698.3.S55 1999 598.1563dc21 98-51782 99 00 01 02 03 P 5 4 3 2 1 Page v To Ian Rowley, pioneer in the quantitative study of cooperative breeding Page vii Contents Foreword to the 1999 Edition by Stephen T. Emlen ix Preface to the 1999 Edition xiii 1. Introduction 1 2. Oceanic Birds 7 3. Precocial Birds 12 4. Diurnal Raptors 17 5. Rails, Coots, and Gallinules 21 6. Pigeons and Doves 26 7. Parrots 28 8. Cuckoos and Anis 31 9. Hoatzins 39 10. Swifts 43 11. Wood-hoopoes 46 12. Todies 51 13. Kingfishers 54 14. Bee-eaters 63 15. Hornbills 68 16. Puffbirds 73 17. Barbets 76 18. Toucans 79 19. Woodpeckers 83 20. Ovenbirds 92 Page viii 21. American Flycatchers 96 22. Cotingas 100 23. New Zealand Wrens 103 24. Swallows 105 25. Crows and Jays 109 26. Mudnest Builders 130 27. Australian Butcher-birds and Bell-magpies 136 28. Titmice 138 29. Long-tailed Tits and Bushtits 142 30. Nuthatches 148 31. Treecreepers 151 32. Wrens 154 33. Gnatcatchers and Old World Warblers 160 34. Wren-warblers 163 35. Old World Flycatchers 170 36. Thrushes 173 37. Babblers and Allies 178 38. Mockingbirds, Thrashers, and Catbirds 186 39. Accentors 189 40. Wood-swallows 191 41. Shrikes 195 42. Helmet-Shrikes 199 43. Starlings 201 44. Honeyeaters 206 45. White-eyes 211 46. Wood-warblers 213 47. Tanagers and Honeycreepers 216 48. New World Blackbirds and Orioles 222 49. Grosbeaks, Finches, and Sparrows 227 50. Weavers and Sparrows 234 51. The Significance of Interspecific Helping 245 52. Characteristics of Cooperative Breeders 252 53. Benefits and Evolution of Cooperative Breeding 260 Bibliography 271 Index 287 Page ix Foreword to the 1999 Edition STEPHEN T. EMLEN In 1930, a young Haverford College undergraduate found himself in Honduras as a member of a small bird-collecting expedition from the Philadelphia Academy of Natural Sciences. One day, while collecting his quota of bird specimens, he spotted a strange, tentlike structure adjacent to a bulky bird nest in the forest. He correctly deduced that it was a bird blind. In time, a second young man appeared and introduced himself as Alexander Skutch. Skutch was working at the time as a plant pathologist for the United Fruit Company. Having completed his routine chores for the day, he was now going to spend the rest of the afternoon observing the behavior of Rufous-breasted Spinetails at their nest. When the two men discovered their mutual enthusiasm for birds and bird-watching, Skutch invited the visitor into his blind. The visitor was John T. Emlen, Jr., my father, who would later become a pioneering ornithologist and animal behaviorist at the University of Wisconsin. John Emlen was utterly fascinated by what he saw that day. He would later write in his diary: "Skutch had inserted a small paintbrush, dipped in yellow paint, into the roof of the tunnel-like entrance to the bird's nest. As we watched, the female came, incubated the Page x two eggs, and left with a neat, yellow stripe down her back, a readily distinguishable marker for future individual recognition." This story illustrates, in one small way, the collective debt that many of us owe Alexander Skutch. By his example, he showed that studies of avian behavior could be a legitimate aspect of ornithology and that by focusing on birds as individuals, one could discover many previously unknown aspects of their social behavior. Alexander Skutch was a pioneer, not only by studying bird behavior but also by bringing the uniqueness of tropical birds to the attention of the world. In 1935, long before the study of bird behavior was fashionable among North American ornithologists, Skutch published a paper in the Auk, the scientific journal of the American Ornithologists' Union, appropriately titled "Helpers at the Nest." In that paper, he described his observations at the nests of three tropical species: Brown Jays, Banded- backed Wrens, and Bushtits. He reported seeing individuals, in addition to the mated pair, regularly bringing food to the nestlings or feeding the breeding female as she incubated the eggs. Skutch was years ahead of his time with these observations. It would be a quarter of a century before others began reporting similar observations and thirty-five years before the study of helping at the nest (now more commonly called cooperative breeding) would become a major focus of evolutionary studies of social behavior. At the time of the publication of this first helping paper, Skutch had already been living in the tropics for seven years. In addition to his work for the United Fruit Company, he had traveled through Panama, Honduras, Guatemala, and Costa Rica studying botany and collecting plant specimens for herbariums and museums to the north. But his real interest was birds, and he observed their behaviors with a patience, a passion, and a dedication for detail unparalleled at the time. From his earliest travels to the present, a period spanning some seventy years, Skutch has always been, first and foremost, an excellent observer of natural history. Despite his fascinating early observations, interest in the topic of helping behavior among birds lay dormant until the mid 1960s. Then, two major advances in our conceptual understanding of social behavior occurred that ushered the question of helping onto center stage in the theater of avian research. One advance was ecological; the other was genetic. On the ecological side, ornithologists such as David Lack, Gordon Orians, Jerram Brown, and John H. Crook argued convincingly that ecological factors play a major role in shaping the forms of animal societies. For the first time, it became possible to forecast (in advance of conducting a study) whether a particular species would be solitary or gregarious, whether it would pair monogamously or polygamously, and whether it would form year-round social groups. The study of bird behavior was on its way to becoming a predictive science. At about the same time, on the genetic side, evolutionary biologist William D. Hamilton was coming to terms with the paradox presented by helping behavior: namely, how could such seemingly altruistic behaviors evolve if those individuals exhibiting the unselfish behaviors did not themselves breed? On the surface, helping behavior seemed to defy the tenets of natural selection. Hamilton elegantly demonstrated in his theory of kin selection that behaviors such as helping are indeed in a helper's selfish interest, as long as the helper is a close genetic relative of the individual(s) it helps. The reasoning is simple: a close ge-