CURRENT ORNITHOLOGY VOLUME 10

Current Ornithology

Editorial Board

George F. Barrowclough, American Museum of Natural History.

New York. New York

Enrique H. Bucher, Centro de Zoologia Aplicada.

Cordoba. Argentina

Ellen D. Ketterson, Indiana University. Bloomington. Indiana

Robert E. Ricklefs, University of Pennsylvania.

Philadelphia. Pennsylvania

Stephen I. Rothstein, University of California.

Santa Barbara. California

John A. Wiens, Colorado State University. Fort Collins. Colorado

A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher.

CURRENT ORNITHOLOGY VOLUME 10

Edited by DENNIS M. POWER Santa Barbara Museum of Natural History Santa Barbara, California

PLENUM PRESS • NEW YORK AND LONDON

The Library of Congress cataloged the first volume of this title as follows:

Current ornithology.-Vol. 1-New York: Plenum Press, c1983-

v.: ill.; 24 cm. Annual. Editor: Richard F. Johnston. ISSN 0742-390X - Current ornithology. 1. Ornithology-Periodicals. I. Johnston, Richard F.

QL671.C87 598'.05-dc19 [8509]

Suggested citation: Current Ornithology, Vol. 10 (D. M. Power, ed.). Plenum Press. New York.

84-640616 AACR 2 MARC-S

ISBN-13: 978-1-4615-9584-7 e-ISBN-13: 978-1-4615-9582-3 001: 10_1007/978-1-4615-9582-3

© 1993 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1993 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

HAL CASWELL, Biology Department, Woods Hole Oceanographic Institu­tion, Woods Hole, Massachusetts 02543

PIERRE DRAPEAU, Departement de Sciences Biologiques, Universite de Montreal, Montreal, Quebec, Canada H3C 3J7

FABIAN M. JAKSI(~, Department of Ecology, Catholic University, Santiago, Chile

ERKKI KORPIMAKI, Laboratory of Ecological Zoology, Department of Biology, University of Thrku, SF-20500 Turku, Finland

J. DAVID LIGON, Department of Biology, The University of New Mexico, Albuquerque, New Mexico 87131

CARL D. MARTI, Department of Zoology, Weber State University, Ogden, Utah 84408

DAVID B. McDONALD, Archbold Biological Station, Lake Placid, Florida 33852-2057.

RAYMOND McNEIL, Departement de Sciences Biologiques, Universite de Montreal, Montreal, Quebec, Canada H3C 3J7

RAYMOND PIEROTTI, Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701

v

vi CONTRIBUTORS

KERRY N. RABENOLD, Department of Biological Sciences, Purdue Uni­versity, West Lafayette, Indiana 47907

J. M. STARCK, Zoologisches Institut, Eberhard Karls-Universitat Tiibingen, D-7400 TUbingen 1, Germany

PREFACE

Ornithology continues to make significant contributions to the advance­ment of many areas of science. This volume of Current Ornithology demon­strates that fact very clearly. It includes contributions that address phy­logeny, mating, parental care, trophic structure, demography, behavioral ecology, species diversity, and ontogeny. This volume also fosters an international exchange of ideas. We have chapters by authors working in the United States, Canada, Chile, Finland, and Germany.

J. David Ligon, University of New Mexico, opens the volume by reviewing the influence of phylogenetic history on mating and parental care. He provides a number of examples supporting the thesis that certain reproductive traits have become fixed in different lineages, which the purely ecological or adaptationist approach to reproductive systems may ignore. Male incubation, clutch size, polyandry, cavity nesting, and cooper­ative breeding are among the topics addressed in this thought-provoking paper.

Carl D. Marti, Erkki Korpimaki, and Fabian M. Jaksic have collaborated by comparing the trophic structure of raptor communities over three continents. Marti works out of Weber State University in Utah, Korpimaki is at the University of Turku in Finland, and Jaksic is based at Catholic University in Santiago, Chile. They compare a variety of food and niche metrics among and within continents, as well as at the assemblage, guild, and species levels. Their monumental bibliography on raptor diet and feeding behavior is international in scope and probably the largest of its kind published anywhere.

The collaboration by David B. McDonald, University of Florida, and Hal Caswell, Woods Hole Oceanographic Institution, tills new ground for this series. It is a "methods" paper, advocating the use of matrix algebraic procedures in the study of avian population statistics. The authors analyze data on Florida Scrub Jays to provide an example of how demographic

vii

viii PREFACE

analyses are carried out. We hope their chapter will stimulate the applica­tion of matrix methods. Reviewers, we would like your comments on whether papers on techniques should be a regular part of this series.

We generally think of birds as diurnal. That thinking is changed by Raymond McNeil and Pierre Drapeau, University of Montreal, and Ray­mond Pierotti, University of Arkansas, who address nocturnality in colo­nial waterbirds. These authors address why certain species visit the nest at night, why some waterbirds forage extensively at night, and what the agents of natural selection favoring nocturnality might be. A massive list of references is assembled and summarized in helpful tables, which should be valuable sources of information for anyone working .on nocturnality or colonial waterbirds.

Kerry N. Rabenold, Purdue University, revisits the interesting problem that in eastern North America, avian species diversity is the reverse of the general trend of increasing diversity that we see from temperate to tropical zones. His findings point out that regional ecological characteristics, such as periods of high productivity, are important in northern forests, while evolutionary history plays a role in the tropics.

J. M. Starck, Eberhard Karls-UniversWit in Thbingen, Germany, has produced a very fine and comprehensive analysis of the evolution of avian ontogenies. This is a major integration of physiological, ecological, and morphological data. The author provides ample illustrations and identifies a number of major trends in the evolution of avian ontogenies.

We continue to solicit reviews, syntheses, and position papers ad­dressing current topics and active areas of research in avian biology. Interested authors may contact any member of the editorial board or send a letter and prospectus to me at the Santa Barbara Museum of Natural History, Santa Barbara, California 93105.

I am grateful to the editorial board for suggesting potential contribu­tions and for their advice on the suitability of submitted manuscripts. Special thanks and appreciation are due the authors, who worked dili­gently on their chapters.

Dennis M. Power Santa Barbara, California

CONTENTS

CHAPTER 1

THE RoLE OF PHYLOGENETIC HISTORY IN THE EVOLUTION OF

CONTEMPORARY AVIAN MATING AND PARENTAL CARE SYSTEMS

J. DAVID LIGON

1. Introduction ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.1. The Phylogenetic Approach ........................... 3 1.2. The Interplay Between Adaptation and Constraint. . . . . . . 4 1.3. Legacies of Evolutionary History. . . . . . . . . . . . . . . . . . . . . . . 6

2. Paternal Care in Ratites and Tinamous ...................... 7 2.1. Parental Care Systems and Habitats Occupied. . . . . . . . . . . 7 2.2. Phylogeny and Taxonomy.... .. ................ ....... 8 2.3. Foraging Ecology ..................................... 8 2.4. Mating System Theory... .... ...... ... .... . .... ....... 8 2.5. Evolutionary Precursors of Parental Care. . . . . . . . . . . .. . . . 9 2.6. The Case for Phylogenetic Inertia ...................... 10 2.7. Conclusions.......................................... 11

3. Clutch Size in Shorebirds ................................. 11 3.1. Egg-Formation Ability. . . .. . . .. . . . . . . .. . . .. . . . . . . . . . . . . 13 3.2. Incubation Ability .................................... 13 3.3. Predation on Nests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 3.4. Phylogenetic Inertia .................................. 13 3.5. Parental Behavior. . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . .. . . . 14

4. Classical Polyandry in Shorebirds .......................... 14 4.1. Kinds of Polyandry ................................... 15 4.2. The Distribution of Classical Polyandry. . . . . . . . . .. . . . .. . 16 4.3. The Diversity of Mating Systems in the Shorebirds 17

ix

x CONTENTS

4.4. Phylogenetic History as One Critical Factor Promoting Polyandry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

4.5. Clutch Size Constancy and Mating System Diversity. . . . . 18 4.6. The Value of Early Broods ............................. 20 4.7. The Relationship Between Double Clutching and Classical

Polyandry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 4.8. An Assessment of Ecological Hypotheses for the

Evolution of Classical Polyandry in Shorebirds. . . . . . . . . . 21 5. Classical Polyandry in a Cuckoo ........................... 22 6. Classical Polyandry: General Conclusions ................... 24 7. Cavity Nesting and Roosting by Coraciiform, Piciform, and

Trogoniform Birds ........................................ 25 7.1. Energetic Characteristics .............................. 27 7.2. Slow Development Rates and Nest Predation..... . ...... 28 7.3. The Significance of Fecal Sacs ......................... 29 7.4. Cavity Nesting and Roosting: Conclusions .............. 30

8. Nocturnal Incubation by Males in Cuckoos and Woodpeckers 30 8.1. Cuckoos ............................................. 31 8.2. Woodpeckers .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 8.3. Nocturnal Incubation by Male Cuckoos and Woodpeckers:

Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 9. Phylogenetic Components of Cooperative Breeding .......... 34

9.1. The Stimulus-Response Nature of Provisioning Behavior by Helpers ........................................... 34

9.2. Phylogenetic History and the Taxonomic Distribution of Cooperative Breeding ................................. 36

9.3. Australian Fairy-Wrens ................................ 37 9.4. Convergence of Cooperative Breeding Systems .......... 37 9.5. Cooperative Breeding: Conclusions. . . . . . . . . . . . . . . . .. . . . 39

10. General Conclusions ...................................... 39 References ............................................... 40

CHAPTER 2

TROPHIC STRUCTURE OF RAPTOR COMMUNITIES: A THREE-CONTINENT

COMPARISON AND SYNTHESIS

CARL D. MARTI, ERKKI KORPIMAKI, AND FABIAN M. ]AKSIC

1. Introduction ............................................... 47 2. Materials and Methods ..................................... 49

2.1. Geographic Coverage. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

CONTENTS xi

2.2. Sources of Data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 2.3. Analytical Techniques ................................. 49

3. Results .................................................... 51 3.1. Assemblage-Level Patterns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 3.2. Guild-Level Patterns ................................... 72 3.3. Species-Level Patterns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

4. Conclusions ............................................... 97 4.1. Assemblage-Level Patterns ....... . . . . . . . . . . . . . . . . . . . . . . . 97 4.2. Guild-Level Patterns ................................... 98 4.3. Species-Level Patterns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 4.4. Relationships of Prey Distribution to Raptor Trophic

Characteristics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 100 4.5. Need for Future Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 101 Appendices ............................................... 102 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 115

CHAPTER 3

MATRIX METHODS FOR AVIAN DEMOGRAPHY

DAVID B. McDONALD AND HAL CASWELL

1. Introduction ............................................... 139 2. Setting Up Matrix-Based Models ............................ 142

2.1. Population Projection .................................. 142 2.2. Projection Interval and Timing of Census ................ 144 2.3. Assumptions and a Note on the Meanings of "Adult" ..... 144 2.4. Deriving the Coefficients in the Graph or Matrix ......... 145

3. Calculating Demographic Parameters ........................ 149 3.1. The Stable Stage Distribution. . . . . . . . . . . . . . . . . . . . . . . . . .. 149 3.2. Reproductive Value .................................... 149 3.3. >t, the Population Growth Rate .......................... 150 3.4. Calculating the Eigenvectors and Eigenvalues ............ 150 3.5. The Characteristic Equation ............................ 151 3.6. Generation Times. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 152

4. Fitness, Sensitivities, and Elasticities ........................ 153 4.1. Sensitivity Analysis-Absolute Changes. . . . . . . . . . . . . . . .. 154 4.2. Elasticity Analysis-Proportional Changes. . . . . . .. . . . . . .. 154

5. Variances and Confidence Limits. . . . . . . . . . . . . . . . . . . . . . . . . . .. 155 5.1. An Approximation to the Variance ...................... 155

6. Applications to Field Studies and Existing Models ........... 156 6.1. Conceptual Links to Some Previous Models. . ... . ........ 157

xii CONTENTS

6.2. Graphs for Incomplete Data and Other Uses of the z-1fansform ........................................... 159

6.3. Difference Equations and the Life Cycle Graph........... 163 6.4. Nonbreeder Loops for Albatrosses and Condors .......... 164 6.5. 'Ifuncated Life Cycle Analysis .......................... 166 6.6. Estimating Parameters by Assuming Zero Growth......... 168

7. A Case Study-Florida Scrub Jays........................... 168 7.1. Constructing a Stage-Classified Life Cycle Graph ......... 169 7.2. Results of the Analysis ................................. 173 7.3. Effect of 'Ifuncating the Life Span...... .... . . ... ..... . .. 175

8. Summary and Synthesis .................................... 177 8.1. The Uses of Matrix Population Models .................. 177 8.2. Suggestions for Field Workers .......................... 178 8.3. Synthetic and Comparative Analyses.................... 178 8.4. Application to Conservation Problems ................... 179 Appendix: Glossary of Terms ............................... 180 References. . . .. ... . .. ..... ..... . ... . . ...... . ... .... ..... . .. 183

CHAPTER 4

NOCTURNAUTY IN COLONIAL WATERBIRDS: OCCURRENCE, SPECIAL ADAPTATIONS, AND SUSPEcrED BENEFITS

RAYMOND McNEIL, PIERRE DRAPEAU, AND RAYMOND PmROTII

1. Introduction............................................... 187 2. Nocturnal Habits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 191

2.1. Spheniscifo~es .. :.................................... 191 2.2. Procellariiformes ...................................... 192 2.3. Pelecaniformes ........................................ 200 2.4. Alcidae ............................................... 201 2.5. Rynchopidae and Laridae .............................. 204 2.6. Marsh Birds ........................................... 207

3. Special Adaptations ........................................ 212 3.1. Procellariiformes and Sphenisciformes .................. 212 3.2. Pelencaniformes and Charadriiformes ................... 214 3.3. Marsh Birds ........................................... 215

4. Suspected Benefits .................................... " . .. 216 4.1. Procellariiformes, Sphenisciformes, Pelecaniformes, and

Alcidae ............................................... 216 4.2. Rynchopidae and Laridae .............................. 220 4.3. Marsh Birds ........................................... ·221

CONTENTS xiii

5. Discussion and Conclusions 222 References. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. 232

CHAPTER 5

LATITUDINAL GRADIENTS IN AVIAN SPECIES DIVERSITY AND THE RoLE

OF LONG-DISTANCE MIGRATION

KERRY N. RABENOLD

1. Introduction ............................................... 247 2. Methods .................................................. 250

2.1. Spruce-Fir and Deciduous-Forest Censuses .............. 250 2.2. Species Lists from Venezuela and Virginia. . . . . . . . . . . . . .. 252

3. Results .................................................... 254 3.1. Spruce-Fir Communities ............................... 254 3.2. Temperate Deciduous Forests ........................... 256 3.3. Long-Term Censuses ................................... 260 3.4. Temperate/Tropical Comparison. . . . . . . . . . . . . . . . . . . . . . . .. 260

4. Discussion ................................................ 265 4.1. Diversity Gradients within North America ............... 265 4.2. Comparisons of Temperate and Tropical Avifaunas ....... 267 4.3. Disentangling Ecological and Historical Explanations. . . .. 268

5. Summary.................................................. 271 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 272

CHAPTER 6

EVOLUTION OF AVIAN ONTOGENIES

J. M. STARCK

1. Introduction .............................................. 275 1.1. Classification of Avian Ontogenies ..................... 276 1.2. Previous Research in Precocial and Altricial Birds. . . . . .. 280 1.3. Phylogenetic Analysis of the Precocial-Altricial Spectrum 282

2. Ontogeny and Time. ... .... . . ... ..... ... . .... ...... . . ..... 286 2.1. Embryonic Period .................................... 286 2.2. Postnatal Period ...................................... 290 2.3. Adult Period ......................................... 291 2.4. Conclusions on Ontogeny and Time. ...... .... ......... 292

xiv CONTENTS

3. Eggs, Clutch Size, and Incubation .......................... 293 4. Postnatal Growth and Ontogeny..... . . ... .................. 294

4.1. Mathematical Description of Growth Curves ............ 295 4.2. Adaptation of Growth Rates ........................... 296

5. Ecological Constraints on Developmental Mode ............. 302 6. The Embryonic Development of Integument ................. 304

6.1. The Feathers of Precocial and Altricial Hatchlings ....... 304 6.2. Previous Work on Feather Development. . . . . . . . . . . . . . . .. 305 6.3. External Development of Feathers. . . . . . . . . . . . . .. . . . . . .. 306 6.4. Histology of Feather Development . . . . . . . . . . . . . . . . . . . . .. 307 6.5. Conclusions on Feather Development .................. 313

7. The Development of the Skeleton .......................... 316 7.1. Previous Work on the Development of Skeleton ......... 316 7.2. Development of the Vertebral Column, Ribs, and Sternum 317 7.3. Development of Wing and Pectoral Girdle .............. 320 7.4. Development of Leg and Pelvic Girdle .................. 324 7.5. Development of Neural and Visceral Cranium. . . . . . . . . .. 329 7.6. Conclusions on the Development of the Skeleton. . . . . . .. 331

8. The Development of the Brain ............................. 333 8.1. Previous Work on the Development of the Brain . . . . . . . .. 334 8.2. Brain Development in Precocial Species-Barred

Buttonquail .......................................... 337 8.3. Brain Development in Precocial Species-Muscovy

Duck ................................................ 339 8.4. Brain Development in Altricial Species-Rock Dove . . . .. 342 8.5. Brain Development in Altricial Species-Budgerigar .... 344 8.6. Brain Development in Altricial Species-Java Sparrow. .. 346 8.7. Conclusions on Brain Development in Precocial and

Altricial Birds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 351 9. The Development of the Intestine .......................... 354

9.1. Previous Studies on the Development of the Intestine . . .. 354 9.2. Preliminary Growth Data on the Alimentary Tract....... 355

10. Conclusion~ and Summary ................................ 356 References ............................................... 359

INDEX....................... .............. ...... ..... ........ 367