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CURRENT ORNITHOLOGY VOLUME 16 Current Ornithology Editorial Board C. Davison Ankney, University of Western Ontario, London, Ontario; Canada Gregory F. Ball, Johns Hopkins University, Baltimore, Maryland Steven R. Beissinger, University of California, Berkeley, California Cynthia Carey, University of Colorado, Boulder, Colorado Robert C. Fleischer, National Zoological Park, Washington, D.C. Ellen D. Ketterson, Indiana University, Bloomington, Indiana Patricia Monaghan, Glasgow University, Glasgow, Scotland Theunis Piersma, Netherlands Institute for Sea Research {NIOZj, Texel and University of Groningen, The Netherlands Irene M. Pepperberg, Massachusetts Institute of Technology, Cambridge, Massachusetts Christopher M. Perrins, University of Oxford, Oxford, England Stephen L. Rothstein, University of California, Santa Barbara, California Ken Yasukawa, Beloit College, Beloit, Wisconsin 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 16 Edited by VAL NOLAN JR. Indiana University Bloomington, Indiana and CHARLES F. THOMPSON Illinois State University Normal, Illinois SPRINGER SCIENCE+BUSINESS MEDIA, LLC The Library of Congress cataloged the first volume of this title as foUows: 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. 1. Johnston, Richard F. QL671.C87 598' .05-dc19 84-640616 (8509) AACR 2 MARC-S ISBN 978-1-4613-5443-7 ISBN 978-1-4615-1211-0 (eBook) DOI 10.1007/978-1-4615-1211-0 ©2001 Springer Science+Business Media New York Originally published by Kluwer Academic/Plenum Publishers, New York in 2001 Softcover reprint of the hardcover 1s t edition 2001 10 9 8 7 6 5 4 3 2 1 A C.I.P. record for this book is available from the Library of Congress AU 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 wriUen permission from the Publisher CONTRIBUTORS CHARLES R. BROWN, Department of Biological Sciences, University of Tulsa, Tulsa, Oklahoma 74104 MARY BOMBERGER BROWN, Department of Biological Sciences, University of Tulsa, Tulsa, Oklahoma 74104 AMBER E. BUDDEN, School of Biological Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, United Kingdom PETER O. DUNN, Department of Biological Sciences, University of Wisconsin, Milwaukee, Milwaukee, Wisconsin 53201 RUSSELL GREENBERG, Smithsonian Migratory Bird Center, Washing ton, D.C. 20008 JUHA MERILA, Department of Population Biology, Evolutionary Biol ogy Center, Uppsala University, SE-752 36 Uppsala, Sweden CLAUDIA METTKE-HOFMANN, Forschungsstelle fUr Ornithologie der Max-Planck-Gesellschaft, D-82346 Erling/A ndechs, Germany BEN C. SHELDON, Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, Oxford OX1 3PS, United Kingdom B. IRENE TIELEMAN, Zoological Laboratory, University of Groningen, 9750 AA Haren, The Netherlands v vi CONTRIBUTORS FRAN<;ms vEZINA, Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia V5A 1S6, Canada LINDA A. WHITTINGHAM, Department of Biological Sciences, Univer sity of Wisconsin, Milwaukee, Milwaukee, Wisconsin 53201 JOSEPH B. WILLIAMS, Department of Evolution, Ecology and Organ ismal Biology, Ohio State University, Columbus, Ohio 43210 TONY D. WILLIAMS, Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia V5A 1S6, Canada JONATHAN WRIGHT, School of Biological Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, United Kingdom PREFACE The objective of this volume is to publish comprehensive, authoritative, and timely reviews of topics of interest to the general ornithological community as well as to students of other vertebrates. We believe that the seven chapters of this volume meet that objective admirably. They testify to the continuing need for a series that publishes papers dealing with any area of active research on birds, unconstrained by restrictions on depth and breadth of coverage and therefore on length. The first chapter in this volume focuses on the least understood of avian social systems, coloniality. In Chapter 1, Charles R. Brown and Mary Bomberger Brown bring their extensive experience with colonial species to bear on what they regard as the central difficulty facing students of this second most common form of avian social organization. This is the inability to find generalizations that satisfactorily explain either its origin or maintenance. Following a review of earlier attempts to develop a general explanation, the authors describe the value of phylogenetic analyses that are designed to provide information on how often coloniality has evolved. The authors point out that repeated evolu tion of coloniality among different lineages suggests that it is a response to particular ecological conditions. They cast a skeptical eye on the widely held belief that coloniality is a result of limitation of nest-sites and suggest a number of approaches to test this hypothesis. In their discussion of the various costs and benefits of coloniality that have been identified, the authors distinguish between causative factors that first led to the evolution of coloniality and maintenance factors that have contributed to its continuance. Their consideration of costs and benefits leads to the conclusion that many of the conventional views surround ing the origin and maintenance are based on much less evidence than is generally appreciated. This paucity of evidence and the results of vii viii PREFACE phylogenetic analyses lead them to warn that a general explanation for the origin of coloniality is unlikely to emerge. To identify factors con tributing both to origin and maintenance, they urge future studies to include (1) measurement of reproductive success in colonies of different sizes, which combines the effects of both costs and benefits, (2) investi gation of causes of variation in colony size, and (3) documentation of phenotypic and genotypic differences among individuals in colonies of different sizes. Amber E. Budden and Jonathan Wright's review (Chapter 2) of beg ging by nestling birds is an important analysis of the many theoretical and empirical studies of conflicts between parent and young, parent and parent, and offspring and offspring (intra-brood and across-broods). These interactions usually occur at a fixed location, the nest, and they offer an unusually straightforward opportunity to observe, manipulate, and quantify critical activities at definable and more or less standard ized stages of life. For these reasons, possibly, begging has interested a variety ofresearchers: modelers and game theoreticians, theoretical and empirical students of signaling and communication, population and developmental biologists, and behavioral ecologists interested in such problems as mating and social systems. Focus on signaling has led to an emphasis on trade-offs between costs (expenditure of energy, attraction of predators) and benefits, with ramifications into honesty of signaling, brood size and volume of sound, extent and effects of within-brood relatedness (including signaling by brood parasites), and many other subjects. Somewhat overlapping the approach that stresses signaling as critical to the evolution of begging is the view that begging originated as a mechanism for prevailing in sibling competition, with parents simply taking advantage of information provided by the competition and using it on occasion as the basis for discriminating among the competitors. This complex and multi-faceted literature has been badly in need of analysis, organization, and the imposition of structure; and authors Budden and Wright have filled this need. They relate the purely theoretical literature, with its simplifying assumptions, to the empirical data and their complexities. They conclude their review with a discus sion of the probability that nestlings learn to adjust their begging efforts to the expected payoff, i.e., with facultative behavior that presumably is related to resource availability and associated factors. They suggest that in this dynamic respect, begging may be analogous to foraging and that understanding begging may be advanced by bringing to bear insights derived from optimal foraging studies. Intraspecific and interspecific variations in responses of animals to stimuli never previously encountered have interested comparative psy- PREFACE ix chologists and ethologists for decades. In their chapter on neophobia and neophilia in birds (Chapter 3), Russell Greenberg and Claudia Mettke-Hofmann analyze the extensive literature, both avian and non avian, but do so from a neglected perspective, i.e., the ecological conse quences of such responses. The authors argue that neophobia (aversion to the novel) and neophilia (the tendency to approach and explore it) are probably important driving forces in avian evolution: behavioral shifts arising out of new environmental experiences precede selection for adaptations that enhance fitness resulting from the shifts. A theme of this chapter is the theoretical importance of distinguishing between two competing hypotheses: the argument that neophobia and neophilia are simply opposite extremes of a one-axis continuum and the alternative hypothesis, espoused by the authors, that neophobia and neophilia are the products of motivationally independent systems, i.e., that both mo tivational states can exist simultaneously and interact in the same indi vidual. The authors describe and analyze methods for determining the relative importance of the two states and the complications introduced by such factors as hunger, details ofthe environment, and innate biases. A section on adult birds deals with familiarization, social facilitation (with attention to the possible role of dominance), and genetic and neurobiological bases of interspecific variation; it also reviews studies of the importance of sex, individual differences in temperament, and seasonal change. A section on juveniles describes the typical ontogeny of exploratory tendencies and behavior: highest toward the end of fledg ling life, followed by canalization of behavior and a shift to neophobia. Many problems are most easily and fruitfully studied in naive individ uals or those whose experience has been controlled, and research that follows the individual throughout life can explore the effects on adults oftheir early behavior and experience. Other topics covered are compar ative (taxonomic, island vs. mainland species) and Greenberg's ecology based Neophobia Threshold Hypothesis, which aims to synthesize many studies by relating environmental background to behavioral char acteristics. A valuable summary emphasizes gaps in our understanding and, by asking twelve questions, points out future research possibilities. The chapter will stand as a valuable synthesis of a subject with far reaching ramifications. The investigator's ability to identify and associate individual par ents and offspring makes birds excellent subjects for the study of quan titative genetics of natural populations, the topic of Chapter 4 by Juha Merila and Ben C. Sheldon. During the almost fifteen years since pub lication of the last review of field studies of avian quantitative genetics, the number of published works has soared, and this led the authors to x PREFACE set three goals for themselves. The first was to give an overview of the field, both in methodology and taxonomic coverage. The second was to provide a critical analysis of the strengths and weaknesses of different approaches that are used to study natural populations. In describing the methods available to estimate heritability and genetic correlation, Me rila and Sheldon emphasize each method's assumptions and the conse quences of measurement error. The third goal was to illustrate how avian systems could be used to study genetic variation within and among populations, thereby contributing to our general understanding of evolution and adaptation. A review of 167 studies, in which heri tabilities or genetic correlations were measured in a range of traits , leads to the conclusion that avian research has contributed to conceptual advances that apply to a wide range of organisms. Despite these ad vances in understanding, however, we still know little about the quan titative genetics of sexually selected and life history traits. The impor tance of considering confounding environmental and maternal effects on estimates of quantitative genetic parameters also receives attention, as do the consequences of extra-pair paternity. The authors conclude by pointing out that the goal of quantitative genetic studies is a better understanding of evolutionary trajectories of populations, and they de scribe two approaches they regard as particularly promising for future research: modeling individual animals to make use simultaneously of all information in a pedigree, and integration of quantitative and mo lecular genetic information. This chapter will be an important contribu tion to quantitative geneticists, whatever their level of experience and degree of familiarity with the subject. Since the discoveries that extra-pair fertilizations are widespread in birds and that the level of paternal effort varies considerably both across species and among conspecific individuals, the relationship between paternity and male care of offspring has attracted a great deal of atten tion. The problem, treated here by Linda A. Whittingham and Peter O. Dunn in Chapter 5, is especially interesting because expenditure of time and energy on paternal care (e.g., incubation, feeding young) is gener ally accepted as being incompatible with devoting time and energy to gaining fertilizations (mating effort). Each type of effort is expected to enhance male fitness, so understanding how and why males of different species, or different males of the same species (within or across popula tions), or the same individual at different nesting attempts vary in their allocation of parental effort engages students with many interests, e.g., mating and social systems. The allocation problem cries out for model ing, and many models have been produced. Similarly, because the pre dictions of the models are in many respects amenable to empirical
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