Parental Care in Mammals PDF
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Parental Care in Mammals Parental Care in Mammals Edited by David J. Gubernick and Peter H. Klopfer Duke UniversitY Durham, North Carolina PLENUM PRESS • NEW YORK AND LONDON Library of Congress Cataloging in Publication Data Main entry under title: Parental care in mammals. Includes index. 1. Mammals-Behavior. 2. Parental behavior in animals. I. Gubernick, David J. II. Klopfer, Peter H. [DNLM: 1. Behavior, Animal. 2. Maternal behavior. 3. Paternal behavior. QL763 R228) QL739.3.P37 599.05'6 80-36692 ISBN-13: 978-1-4613-3152-0 e-ISBN-13: 978-1-4613-3150-6 001: 10.1007/978-1-4613-3150-6 © 1981 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1981 A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N. Y. 10011 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 Marc Bekoff, Department of Environmental, Population, and Organismic Biol ogy, Behavioral Biology Group, University of Colorado, Boulder, Colorado 80309 Bennett G. Galef, Jr., Department of Psychology, McMaster University, Ham ilton, Ontario, Canada L8S 4K1 DavidJ. Gubernick, Department of Zoology, Duke University, Durham, North Carolina 27706. Present address: Department of Psychology, Indiana Uni versity, Bloomington, Indiana 47405 Lawrence V. Harper, Department of Applied Behavioral Sciences, University of California, Davis, Davis, California 95616 Myron A. Hofer, Albert Einstein College of Medicine at Montefiore Hospital, Bronx, New York 10467 Devra G. Kleiman, Department of Zoological Research, National Zoological Park, Smithsonian Institution, Washington, D.C. 20008 Peter H. Klopfer, Zoology Department, Duke University, Durham, North Car olina 27706 James R. Malcolm, Department of Zoological Research, National Zoological Park, Smithsonian Institution, Washington, D.C. 20008 James J. McKenna, Department of Anthropology and Sociology, Pomona Col lege, Claremont, California 91711 Jay S. Rosenblatt, Institute of Animal Behavior, Rutgers University, Newark, New Jersey 07102 Leonard A. Rosenblum, Primate Behavior Laboratory, Department of Psychia try, Downstate Medical Center, Brooklyn, New York 11203 Harold I. Siegel, Institute of Animal Behavior, Rutgers University, Newark, New Jersey 07102 Bruce B. Svare, Department of Psychology, State University of New York at Albany, Albany, New York 12222 Karyl B. Swartz, Department of Psychological Sciences, Purdue University, West Lafayette, Indiana 47907 v Foreword The editors of this volume have honored me by their invitation to write its Fore word, an invitation extended because of my editing a book on the maternal behav ior of mammals in 1963. Much as I would like to think that I had opened a new area of study-and so played some part in the appearance of this fine new collec tion of chapters-the facts are quite otherwise. That in 1963 I could assemble the efforts of many distinguished investigators shows that the topic had already engaged their attention, and had for some years past. But even then, the topic had origins extending much farther into the past, to mention only Wiesner and Sheard's book Maternal Behavior in the Rat of 1933. Nevertheless, in 1963 it seemed to me that the study of maternal care in mammals had lagged behind the study of other kinds of social behavior. The present volume does much to establish parental care of the young as a topic central to an understanding of the relation between ontogeny and phylogeny, to the development of the young, to the social organization of the species, and to its preservation. It may now be seen not only as interesting but as a most signifi cant pattern of behavior among mammals. This volume clearly shows that in the intervening years the topic has indeed been a lively one, and the progress made must be characterized as truly remark able. Some of the contributions to knowledge were foreshadowed in the earlier volume, such as the process of weaning, evidence of maternal aggression, the part the young play in obtaining the care they require, the role of fathers and unrelated members of the group in ministering to the needs of the young, and the role of parental care in the social organization of the group. But what a wealth of new information has been amassed in these few years when each of these topics is made the primary focus of study! Several of the contributions wrestle with the concept of attachment, a term used only by Harlow in the 1963 volume but which since then has received much attention. Now, however, several contributors to the current volume are question ing its usefulness as an explanatory concept. Proximity of the young to its care takers provides much more than protection from predators; it provides food, warmth, and so on, but more important, stimulation, opportunities to learn, and vii viii FOREWORD models for becoming members of the social group. The disrupting effects of sep aration of the young from mother and other members of the group are also shown to vary with many other factors. Under the hands of these skilled investigators, the seemingly magical property of the social bond is yielding to sharply analytic attempts at explanation. The advances in neurology, biochemistry, and endocrinology from 1963 on have also provided new techniques for the study of processes. Further, the ultra sonic sounds of rodent pups represent a new finding, as does knowledge about pheromones and their effects on both rodent pups and their mothers. Last but by no means least, a new vocabulary of evolutionary concepts now distinguishes the present collection of chapters. That is not to say that the contributors of the 1963 volume were unaware of the evolution of parental care and its adaptiveness, but they did not have at hand the explicit organizing concepts introduced in the inter vening years. I refer to such concepts as parental investment, inclusive fitness, and kin selection, as well as cost-benefit analysis, the proximate and ultimate functions of behavior, and optimal strategies. The chapters of this book, then, provide not only new information on an important class of behaviors, but also a fresh and stimulating framework in which they can be examined. Rich indeed are the delineations of questions still to be answered. The chapters, furthermore, provide models not only for the investiga tion of parental care in other mammals (dare I now include the human mam mal ?), but also for the investigation of other related areas of social behavior. I am confident that all readers will profit as have I, for in these chapters they will find challenges to inform and sustain their own efforts for years to come. Harriet L. Rheingold University of North Carolina at Chapel Hill Preface Since the publication in 1963 of Harriet L. Rheingold's classic volume Maternal Behavior in Mammals, the study of parental care has witnessed remarkable prog ress in theoretical orientation, conceptual advances, and an empirical data base. Past research had focused predominantly on the immediate causation of maternal behavior and the development of maternal-filial relationships. While these topics continue to stimulate fruitful investigations, a new generation of com plimentary research questions has emerged. First, we have witnessed a new awareness of the infant as an active, potent force in the shaping, maintenance, and expression of parental behavior. Gone are the simpler notions that mothers auton omously deliver care to their passive, ineffectual infants. In fact, we now recognize that the allocation of parental care is not restricted to the mother alone and that other individuals can also exert a profound influence on the developing infant. These newly recognized sources include male parental care, allomothering, sibling interactions, and the social context of parental care. A significant factor in the emergence of these "new" questions and issues has been the application of principles and concepts of evolutionary biology toward understanding the nature of parental care and the offspring's relationships with other individuals. Indeed, the profound influence of the evolutionary perspective can be seen in the original contributions presented in this volume. In the first chapter, Klopfer discusses why parental care evolved and what factors might influence the evolution of various parental care strategies. He also raises the question of whether, and in what sense, patterns of parental care are even adaptive (a question also raised by McKenna, Chapter to, in his consider ation of allomothering). Chapters 2-7 are primarily concerned with the mother-infant relationship, while Chapters 8-11 deal with the broader social network of the infant. The hor monal and nonhormonal factors involved in the onset and maintenance of mater nal behavior are examined in Chapter 2 by Rosenblatt and Siegel. They discuss and review an extensive literature about these functions and present a theoretical framework for regarding the principal phases in the regulation of maternal behavior. Next, Hofer (Chapter 3) examines the contributions parents make to the ix x PREFACE development of their offspring. He provides compelling evidence that the sensory stimulation provided by the mother regulates the physiology and behavior of the developing infant. Although parents obviously contribute to the development of their young, the singular view of the infant as a passive recipient of parental care is no longer acceptable. As an active participant in parent-infant relationships, the young can affect the caregiver's behavior. Harper (Chapter 4) draws together a vast and diverse literature of such offspring effects and discusses these within an evolution ary, ecological, and developmental framework. One dramatic example of such offspring effects is that the suckling stimula tion of the young can regulate the mother's aggression toward conspecifics. Despite the fact that defense of the young is a common and widespread feature of parental care, it has received little systematic study. The psychobiology of maternal aggres sion in mammals is an emerging field which is discussed and reviewed by Svare (Chapter 5). The process of weaning has also received comparatively little analysis or the oretical treatment despite its importance in the transition of an infant from a dependent to an independent existence. Galef (Chapter 6) presents a provocative analogy of the infant as a parasite of its mother and discusses weaning within this context. The specificity of parent-infant relationships (i.e., attachment) is discussed next by Gubernick in Chapter 7. He addresses the questions of "What is attach ment?" and "Why form attachments?" The presence of parent and infant attach ment in only some mammals raises the question of what conditions favor attachment, and these conditions are also examined. In Chapter 8, Bekoff considers alloparenting among siblings and discusses the conditions favoring sibling interactions and sibling recognition. In many mam malian groups, males provide some form of care for the young, yet such male parental investment has not been well studied. Kleiman and Malcolm (Chapter 9) discuss the evolution of male parental care, the various types of such male parental behavior, and its influence on social organization. They also present a summary of those mammals displaying male parental investment. Like male parental behavior, allomothering is also widespread among mam mals. McKenna (Chapter 10) discusses the evolution and presumed functions of allomothering and uses the common Indian langur monkey as a case study. He also raises the issue that allomaternal behavior, as well as parental behavior in general, is not necessarily adaptive, and in some cases allomothering may actually be harmful to the infant. In the final chapter, Swartz and Rosenblum discuss the social context of parental behavior with special emphasis on infant socialization in primates. They examine the influences various social agents, such as those included in other chap ters, may have on the social development of the infant. PREFACE xi We would like to express our appreciation to all the authors for their effort and contributions and to the publishers for their encouragement and support. We hope that this volume will stimulate further research and the evolution of new questions. J. David Gubernick and Peter H. Klopfer Durham
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