Table Of ContentParental 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
