BEHA VIOR OF MARINE ANIMALS. VOLUME 5 Current Perspectives in Research

SHOREBIRDS Breeding Behavior and Populations

BEHA VI OR OF MARINE ANIMALS Current Perspectives in Research

Series Editors: Howard E. Winn and Bori L. OHa

Volume 1 Invertebrates Edited by Howard E. Winn and Bori L. Olla

Volume 2 Vertebrates Edited by Howard E. Winn and Bori L. Olla

Volume 3 Cetaceans Edited by Howard E. Winn and Bori L. Olla

Volume 4 Marine Birds Edited by Joanna Burger, Bori L. Olla, and Howard E. Winn

Volume 5 Shorebirds: Breeding Behavior and Populations Edited by Joanna Burger and Bori L. Olla

Volume 6 Shorebirds: Migration and Foraging Behavior Edited by Joanna Burger and Bori L. Olla

BEHAVIOR OF MARINE ANIMALS • VOLUME 5 Current Perspectives in Research

SHOREBIRDS Breeding Behavior and Populations

Edited by

Joanna Burger Rutgers University Piscataway, New Jersey

and

Bori L. Olla Oregon State University Newport, Oregon and Northwest and Alaska Fisheries Center Seattle, Washington

PLENUM PRESS • NEW YORK AND LONDON

Library of Congress Cataloging in Publication Data

Winn, Howard Elliott, 1926-Behavior of marine animals; current perspectives in research.

Vol. 4 edited by J. Burger, B. L. ~Ua, and H. E. Winn; v. 5-OUa.

Includes bibliographical references and indexes.

by J. Burger and B. L.

CONTENTS: v. 1. Invertebrates.-v. 2. Vertebrates.-(etc.}-v. 5. Shorebirds, breeding behavior and populations.

1. Marine fauna-Behavior-CoUected works. I. ~Ua, Bori L., joint author. II. Burger, Joanna, joint author. III. Title. QLl21.W5 591.5'2636 79-167675 ISBN 978-1-4684-4693-7 ISBN 978-1-4684-4691-3 (eBook) DOl 10.1007/978-1-4684-4691-3

©1984 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1984

A Division of Plenum Publishing Corporation 233 Spring Street, New York, N. Y. 10013

All rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

CONTRIBUTORS

Joanna Burger Department of Biology and Bureau of Biological Research, Rutgers University, Piscataway, New Jersey 08854

P. R. Evans Department of Zoology, University of Durham, Durham DHI 3LE, England

Michael Gochfeld Department of Environmental and Community Medicine, UMDNJ-Rutgers Medical School, Piscataway, New Jersey 08854

Marshall A. Howe U . S. Fish and Wildlife Service, Patuxent Wildlife Research Center, Laurel, Maryland 20708

Joseph R. Jehl, Jr. Hubbs-Sea World Research Institute, Mission Bay, San Diego, California 92109

David B. Lank Department of Biology, University of North Dakota, Grand Forks, North Dakota 58202

Sarah Lenington Institute of Animal Behavior, Rutgers University, Newark, New Jersey 07102

Edward H. Miller Vertebrate Zoology Division, British Columbia Provincial Museum, and Biology Department, University of Victoria, Victoria, British Columbia V8V lX4, Canada

Lewis W. Oring Department of Biology, University of North Dakota, Grand Forks, North Dakota 58202

M. W. Pienkowski Department of Zoology, University of Durham, Durham DHI 3LE, England

Stanley E. Senner Hawk Mountain Sanctuary Association, Kempton, Pennsylvania 19529

v

vi Contributors

Jeffrey R. Walters Allee Laboratory of Animal Behavior, The University of Chicago, Chicago, Illinois 60637. Present address: Zoology Department, North Carolina State University, Raleigh, North Carolina 27695

INTRODUCTION

Among birds, shorebirds provide some of the more unique opportunities to examine basic problems in behavior, ecology, and evolution. This is in large measure due to the diversity, both behaviorally and ecologically, of a group closely related taxonomically and distributed throughout the world. The overall aim of these two volumes is to provide a representative selection of current research being conducted on shorebird behavior and ecology.

Traditionally, marine birds have included those species that breed in large colonies on offshore islands along coasts (see Volume 4 of this series). Although shorebirds have generally not been considered within this group, the fact that almost 40% of the species breed along coasts and more than 60% often or always spend the nonbreeding season in coastal habitats more than justifies their inclusion as marine birds (at least those species that totally or partially depend upon the marine environment). Their inclusion markedly increases species diversity in marine birds since shorebirds add about 217 species to the 280 that are traditionally thought of as marine.

Over the last twenty years there has been a veritable explosion in studies on shorebirds. Prior to this, there were few studies of breeding behavior and only cursory examinations of their migratory patterns. The increased interest in shorebirds was not the least caused by the increased awareness of the diversity and plasticity of their social systems. More pragmatically, the increased accessibility to researchers of Arctic breed­ing grounds as well as increase in the availability of fixed winged planes and helicopters for censusing populations and monitoring movements dur­ing the nonbreeding season also did much to foster the research effort.

Advances in shorebirds research have been of vital interest to those studying other animal groups, especially in the area offoraging behavior, with many of the paradigms and theories developed for optimal foraging theory being readily applicable to, and testable in shorebird species. Un­like the traditional marine species, their prey (primarily invertebrates) can be readily sampled both qualitatively and quantitatively. Further, since they forage primarily in estuaries it is possible to follow individuals

vii

viii Introduction

in the nonbreeding season. The variability imposed by tidal regimes only make the system more valuable for field testing foraging theory.

Future studies on shorebirds will continue to center on examining breeding behavior and ecology, migration, and foraging behavior. How­ever, more emphasis will be placed on examining the entire life cycle of particular species, comparing the success of populations that differ in breeding sites, breeding and nonbreeding habitats, and migration routes. Finally, the mechanisms that control behavioral and ecological plasticity in shorebirds should provide some of the more interesting avenues for future research.

J. Burger B. Olla

PREFACE

Most chapters in these two volumes on the behavior of shorebirds include a balance between literature review, original data and synthesis. The re­search approaches of the authors range from in-depth study of a single species to a comparative one involving groups of species that are tax­onomically or ecologically related.

The subject has been divided into two volumes, one dealing with breeding behavior and populations, and the second with migration and foraging behavior. The first two chapters of Volume 5 serve as an intro­duction to the two volumes and address the issues of shorebird classifi­cation and whether shorebirds are marine animals. Most of the remaining chapters in Volume 5 concern aspects of breeding behavior including breeding site fidelity, polyandrous mating systems, communication, pa­rental behavior, and antipredator behavior. Two other aspects discussed in Volume 5 are population dynamics and conservation of shorebirds.

Volume 6 covers behavior aspects of shorebirds during the non­breeding season and includes abiotic factors affecting migration, migra­tory behavior in the Western Pale arctic and New World, foraging and activity patterns, food supplies, and spacing patterns of nonbreeding mi­grants. Taken together these two volumes provide paradigms for the fur­ther study of organisms that move freely between marine and terrestrial ecosystems, as well as summarizing current research with a particular group of birds. Some other important areas of shorebirds behavior (gen­eral mating systems, sexual dimorphism) have not been included because other reviews of these topics exist.

In recent times man's encroachment has been particularly strong on the beaches and shorelines of marine habitats used extensively by shore­birds. A volume elucidating aspects of their breeding, migration, and win­tering behavior and ecology is particularly useful to managers and con­servationists involved with coastal planning and protection of marine av­ifaunas. Their accessibility for study in breeding and nonbreeding habitats permits the identification of those factors which playa role in controlling popUlation levels, a prerequisite for the application of rational manage­ment plans. For ornithologists, ecologists and behaviorists the two vol-

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x Preface

urnes provide a review of the concepts concerning shorebird behavior and the elucidation of the basic mechanisms affecting behavior.

We are especially grateful to Michael Gochfeld, Betty Green, Jill Grover, Brook Lauro, Cindy Paszkowski and Gary Braun for their help in indexing, and to the authors of Chapter 1 in Volume 5 for providing a shorebird classification that could be used as a basis for the other chapters in the two volumes.

J. Burger B.Olla

CONTENTS

Chapter 1 The Classification of the Shorebirds of the World

I. II.

Chapter 2

Michael Gochfeld, Joanna Burger, and Joseph R. Jehl, Jr.

Introduction ............................................... . List of the Taxa of Shorebirds ........................... . References

1 3

14

Shorebirds as Marine Animals

Joanna Burger

I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 II. Marine Avifaunas .......................................... 23

III. Shorebirds in Marine Ecosystems ........................ 26 IV. Habitat Selection in Shorebirds .......................... 29

A. Taxonomic Summary ................................ 29 B. Comparisons and Discussion........................ 71 References ................................................. 72

Chapter 3 Population Dynamics of Shorebirds

P. R. Evans and M. W. Pienkowski

I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 II. Recruitment and Mortality on the Breeding Grounds ... 85

A. Population Density ................................... 86 B. Nesting Attempts..................................... 87 C. Clutch Size ........................................... 88 D. Hatching Success .................................... 89 E. Fledging Success ..................................... 90

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xii Contents

F. Adult Mortality during the Breeding Season ....... 92 G. Y ear-to-Year Variation in Breeding Output ........ 92

III. Adult Survival during the Nonbreeding Season ......... 93 A. Mortality during Migration .......................... 94 B. Mortality during Molt ................................ 95 C. Mortality in the Nonbreeding Areas................ 96

IV. Survival of Immature Birds and Age of First Breeding 105 A. Survival of Young Birds after Fledging ............ 105 B. Age of First Breeding ................................ 109

V. Are Shorebird Populations Regulated? ................... 110 A. Reproductive Output ................................. 111 B. Mortality in the Nonbreeding Areas................ 114 C. Epilogue .............................................. 117 References ................................................. 118

Chapter 4 Breeding Area Fidelity, Natal Philopatry, and the Social Systems of Sandpipers

Lewis W. Oring and David B. Lank

I. Introduction ............................................... . A. Why Return? ........................................ . B. Why Disperse? ...................................... .

II. Breeding Area Fidelity ................................... . A. Species Differences ................................. . B. Sex Bias ............................................. . C. Effects of Age and Breeding Success .............. .

III. Natal Philopatry .......................................... . A. Species Differences ................................. . B. Sex Bias ............................................. .

IV. Late-Season Movements of Spotted Sandpipers ....... . V. Conclusion ................................................ .

References

Chapter 5 The Evolution of Polyandry in Shorebirds

Sarah Lenington

125 126 127 127 127 130 135 138 139 140 142 143 145

I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 II. Forms of Polyandry in Shorebirds ....................... 150

III. Theories for the Evolution of Polyandry................. 151 A. Simple Predation Model ............................. 152

Contents

IV.

Chapter 6

B. Replacement Clutch Model ......................... . C. Energetic Model ..................................... . Conclusion ................................................ . References

xiii

154 157 164 164

Communication in Breeding Shorebirds

Edward H. Miller

I. Introduction... . ............................................ 169 II. Communication in Breeding Shorebirds ................. 172

A. External Characteristics and Appearance .......... 172 B. Studies on Display Organization .................... 180 C. Acoustic Communication............................ 189 D. Ecological and Social Effects ....................... 205 E. Evolutionary and Phylogenetic Considerations .... 220

III. Concluding Remarks ...................................... 231 References ................................................. 232

Chapter 7 The Evolution of Parental Behavior and Clutch Size in Shorebirds

Jeffrey R. Walters

I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 II. Parental Behavior.......................................... 244

A. Studies of Lapwings ................................. 244 B. Parental Behaviors of Other Shorebirds............ 261 C. Discussion............................................ 266

III. Clutch Size ................................................. 267 A. Species That Feed Young ........................... 268 B. Species That Do Not Feed young.................. 270 C. Discussion. . .. . .. . . .. . . ... . .... . .. . . . . . .. . . . .. . . . . . . . . 279 Appendix: Ethogram ...................................... 281 References ................................................. 281

Chapter 8 Antipredator Behavior: Aggressive and Distraction Displays of Shorebirds

Michael Gochfeld

I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 A. Categories and Nomenclature of Nest Protection

Behavior ............................................ 291

xiv Contents

B. Response to an Approaching Predator ........... 291 C. Avian Distraction Behavior ....................... 292

II. The Context in Which Nest Protection Behavior Occurs .................................... ............. ... 295 A. Temporal and Spatial Factors and Habitat ....... 298 B. Appropriate Stimuli ................................ 299

III. Descriptions of Nest Protection Behavior and Displays .................................................. 299 A. Nest-Departure Behavior and Response

Distance ............................................ 300 B. Distraction Displays and Related Behaviors ..... 303 C. Aggressive Antipredator Behavior ................ 321 D. Response, Approach, Display, and Luring

Distances ........................................... 326 E. Motor Patterns or Acts............................ 328 F. Stereotypy.......................................... 330 G. Reentrapment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331 H. Transitions among Distraction Display

Components ........................................ 331 IV. Factors Influencing Nest Protection Behavior ........ 333

A. Characteristics of Individual ...................... 334 B. Characteristics of Intruder ........................ 344 C. Characteristics of the Context .................... 347 D. Observer Variability ............................... 347 E. Discussion of Variation............................ 348

V. Social Aspects of Nest Protection Behavior........... 348 A. Mobbing Behavior: Social Aspects ............... 349 B. Distraction Behavior: Social Aspects ............ 351 C. Discussion of Social Aspects ..... . . . . . . . . . . . . . . . . 351

VI. Efficacy and Costs of Various Types of Antipredator Behavior .................................................. 351 A. Nest-Departure Behavior: Efficacy............... 352 B. Aggressive Behavior: Efficacy .................... 352 C. Distraction Behavior: Efficacy .................... 352 D. Costs of Antipredator Behavior........ ... ........ 353

VII. Origins of Predator-Induced Aggression and Distraction Displays ...................................... 354 A. Decision Making ................................... 357 B. Parental Investment ............................... 358 C. The Role of Fear ................................... 358 D. Consciousness...................................... 359 E. Conflict Hypotheses ............................... 359 F. Conclusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362

Contents xv

VIII. Perspectives for Future Research ...................... 363 A. The Displaying Bird ............................... 364 B. The Nature and Response of the Intruder ....... 366 C. Nature of the Context ............................. 367 D. V ocal Components ................................. 368 E. Methodologic Considerations ..................... 368 F. The Future ......................................... 369 References ............................................... 369

Chapter 9 Conservation of Nearctic Shorebirds

Stanley E. Senner and Marshall A. Howe

1. Introduction.. . . .. . . . . . .. . . .. . .. . . .. . . . . . .. . .. . . ... . . .. . . . 379 II. Sensitive Periods in the Annual Cycle ................. 380

A. Breeding Season ................................... 380 B. Migration Period ................................... 382 C. Winter Season ...................................... 388

III. Threats to Shorebird Populations. .. . . . . . .. . . .. . . . .. . . .. 390 A. Hunting............................................. 391 B. Disturbance......................................... 392 C. Environmental Contaminants ..................... 394 D. Habitat Loss and Degradation .................... 396

IV. Management of Shorebird Populations ................. 400 A. Legal-Political Tools ............................... 400 B. Habitat Manipulation .............................. 405 C. Species Reintroduction ............................ 407 D. Population Estimation and Monitoring ........... 407

V. Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409 A. Critical Area Concept ............................. 409 B. Conclusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 414 References ............................................... 415

Species Index ............................................. 423

Subject Index ............................................. 433