CURRENT ORNITHOLOGY VOLUME 10

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1 CURRENT ORNITHOLOGY VOLUME 10

2 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.

3 CURRENT ORNITHOLOGY VOLUME 10 Edited by DENNIS M. POWER Santa Barbara Museum of Natural History Santa Barbara, California PLENUM PRESS NEW YORK AND LONDON

4 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 X - Current ornithology. 1. Ornithology-Periodicals. I. Johnston, Richard F. QL671.C87 598'.05-dc19 [8509] AACR 2 MARC-S Suggested citation: Current Ornithology, Vol. 10 (D. M. Power, ed.). Plenum Press. New York. ISBN-13: e-isbn-13: : 10_1007/ 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 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

5 CONTRIBUTORS HAL CASWELL, Biology Department, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 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 Turku, Finland J. DAVID LIGON, Department of Biology, The University of New Mexico, Albuquerque, New Mexico CARL D. MARTI, Department of Zoology, Weber State University, Ogden, Utah DAVID B. McDONALD, Archbold Biological Station, Lake Placid, Florida 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 v

6 vi CONTRIBUTORS KERRY N. RABENOLD, Department of Biological Sciences, Purdue University, West Lafayette, Indiana J. M. STARCK, Zoologisches Institut, Eberhard Karls-Universitat Tiibingen, D-7400 TUbingen 1, Germany

7 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

8 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 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. Santa Barbara, California Dennis M. Power

9 CONTENTS CHAPTER 1 THE RoLE OF PHYLOGENETIC HISTORY IN THE EVOLUTION OF CONTEMPORARY AVIAN MATING AND PARENTAL CARE SYSTEMS J. DAVID LIGON 1. Introduction The Phylogenetic Approach The Interplay Between Adaptation and Constraint Legacies of Evolutionary History Paternal Care in Ratites and Tinamous Parental Care Systems and Habitats Occupied Phylogeny and Taxonomy Foraging Ecology Mating System Theory Evolutionary Precursors of Parental Care The Case for Phylogenetic Inertia Conclusions Clutch Size in Shorebirds Egg-Formation Ability Incubation Ability Predation on Nests Phylogenetic Inertia Parental Behavior Classical Polyandry in Shorebirds Kinds of Polyandry The Distribution of Classical Polyandry The Diversity of Mating Systems in the Shorebirds 17 ix

10 x CONTENTS 4.4. Phylogenetic History as One Critical Factor Promoting Polyandry Clutch Size Constancy and Mating System Diversity The Value of Early Broods The Relationship Between Double Clutching and Classical Polyandry An Assessment of Ecological Hypotheses for the Evolution of Classical Polyandry in Shorebirds Classical Polyandry in a Cuckoo Classical Polyandry: General Conclusions Cavity Nesting and Roosting by Coraciiform, Piciform, and Trogoniform Birds Energetic Characteristics Slow Development Rates and Nest Predation The Significance of Fecal Sacs Cavity Nesting and Roosting: Conclusions Nocturnal Incubation by Males in Cuckoos and Woodpeckers Cuckoos Woodpeckers Nocturnal Incubation by Male Cuckoos and Woodpeckers: Conclusions Phylogenetic Components of Cooperative Breeding The Stimulus-Response Nature of Provisioning Behavior by Helpers Phylogenetic History and the Taxonomic Distribution of Cooperative Breeding Australian Fairy-Wrens Convergence of Cooperative Breeding Systems Cooperative Breeding: Conclusions General Conclusions References CHAPTER 2 TROPHIC STRUCTURE OF RAPTOR COMMUNITIES: A THREE-CONTINENT COMPARISON AND SYNTHESIS CARL D. MARTI, ERKKI KORPIMAKI, AND FABIAN M. ]AKSIC 1. Introduction Materials and Methods Geographic Coverage

11 CONTENTS xi 2.2. Sources of Data Analytical Techniques Results Assemblage-Level Patterns Guild-Level Patterns Species-Level Patterns Conclusions Assemblage-Level Patterns Guild-Level Patterns Species-Level Patterns Relationships of Prey Distribution to Raptor Trophic Characteristics Need for Future Research Appendices References CHAPTER 3 MATRIX METHODS FOR AVIAN DEMOGRAPHY DAVID B. McDONALD AND HAL CASWELL 1. Introduction Setting Up Matrix-Based Models Population Projection Projection Interval and Timing of Census Assumptions and a Note on the Meanings of "Adult" Deriving the Coefficients in the Graph or Matrix Calculating Demographic Parameters The Stable Stage Distribution Reproductive Value >t, the Population Growth Rate Calculating the Eigenvectors and Eigenvalues The Characteristic Equation Generation Times Fitness, Sensitivities, and Elasticities Sensitivity Analysis-Absolute Changes Elasticity Analysis-Proportional Changes Variances and Confidence Limits An Approximation to the Variance Applications to Field Studies and Existing Models Conceptual Links to Some Previous Models

12 xii CONTENTS 6.2. Graphs for Incomplete Data and Other Uses of the z-1fansform Difference Equations and the Life Cycle Graph Nonbreeder Loops for Albatrosses and Condors 'Ifuncated Life Cycle Analysis Estimating Parameters by Assuming Zero Growth A Case Study-Florida Scrub Jays Constructing a Stage-Classified Life Cycle Graph Results of the Analysis Effect of 'Ifuncating the Life Span Summary and Synthesis The Uses of Matrix Population Models Suggestions for Field Workers Synthetic and Comparative Analyses Application to Conservation Problems Appendix: Glossary of Terms References CHAPTER 4 NOCTURNAUTY IN COLONIAL WATERBIRDS: OCCURRENCE, SPECIAL ADAPTATIONS, AND SUSPEcrED BENEFITS RAYMOND McNEIL, PIERRE DRAPEAU, AND RAYMOND PmROTII 1. Introduction Nocturnal Habits Spheniscifo~es.. : Procellariiformes Pelecaniformes Alcidae Rynchopidae and Laridae Marsh Birds Special Adaptations Procellariiformes and Sphenisciformes Pelencaniformes and Charadriiformes Marsh Birds Suspected Benefits... " Procellariiformes, Sphenisciformes, Pelecaniformes, and Alcidae Rynchopidae and Laridae Marsh Birds

13 CONTENTS xiii 5. Discussion and Conclusions 222 References CHAPTER 5 LATITUDINAL GRADIENTS IN AVIAN SPECIES DIVERSITY AND THE RoLE OF LONG-DISTANCE MIGRATION KERRY N. RABENOLD 1. Introduction Methods Spruce-Fir and Deciduous-Forest Censuses Species Lists from Venezuela and Virginia Results Spruce-Fir Communities Temperate Deciduous Forests Long-Term Censuses Temperate/Tropical Comparison Discussion Diversity Gradients within North America Comparisons of Temperate and Tropical Avifaunas Disentangling Ecological and Historical Explanations Summary References CHAPTER 6 EVOLUTION OF AVIAN ONTOGENIES J. M. STARCK 1. Introduction Classification of Avian Ontogenies Previous Research in Precocial and Altricial Birds Phylogenetic Analysis of the Precocial-Altricial Spectrum Ontogeny and Time Embryonic Period Postnatal Period Adult Period Conclusions on Ontogeny and Time

14 xiv CONTENTS 3. Eggs, Clutch Size, and Incubation Postnatal Growth and Ontogeny Mathematical Description of Growth Curves Adaptation of Growth Rates Ecological Constraints on Developmental Mode The Embryonic Development of Integument The Feathers of Precocial and Altricial Hatchlings Previous Work on Feather Development External Development of Feathers Histology of Feather Development Conclusions on Feather Development The Development of the Skeleton Previous Work on the Development of Skeleton Development of the Vertebral Column, Ribs, and Sternum Development of Wing and Pectoral Girdle Development of Leg and Pelvic Girdle Development of Neural and Visceral Cranium Conclusions on the Development of the Skeleton The Development of the Brain Previous Work on the Development of the Brain Brain Development in Precocial Species-Barred Buttonquail Brain Development in Precocial Species-Muscovy Duck Brain Development in Altricial Species-Rock Dove Brain Development in Altricial Species-Budgerigar Brain Development in Altricial Species-Java Sparrow Conclusions on Brain Development in Precocial and Altricial Birds The Development of the Intestine Previous Studies on the Development of the Intestine Preliminary Growth Data on the Alimentary Tract Conclusion~ and Summary References INDEX

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