Table Of ContentThe Behaviour
of Pinnipeds
The Behaviour
of Pinnipeds
Edited by
Deane Renouf
Professor, Department of Psychology and
Ocean Sciences Center,
Memorial University of Newfoundland, Canada
S
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
First edition 1991
© 1991 Springer Science+Business Media Dordrecht
Originally published by Chapman and Hall Ltd in 1991
Softcover reprint ofthe hardcover lst edition 1991
Typeset in 10/12pt Trump Mediaeval by
Mayhew Typesetting, Bristol
ISBN 978-94-0lO-5373-0
AlI rights reserved. No part of this publication may be reproduced
or transmitted, in any form or by any means, electronic,
mechanical, photocopying, recording or otherwise, or stored in any
retrieval system of any nature, without the written permission of
the copyright holder and the publisher, application for which shall
be made to the publisher.
The publisher makes no representation, express or implied, with
regard to the accuracy of the information contained in this book
and cannot accept any legal responsibility or liability for any errors
or omissions that may be made.
British Library Cataloguing in Publication Data
The behaviour of pinnipeds.
1. Mamrnals: Seals
1. Renouf, Deane
599.745
ISBN 978-94-010-5373-0 ISBN 978-94-011-3100-1 (eBook)
DOI 10.1007/978-94-011-3100-1
Library of Congress Cataloging-in-Publication Data
The Behaviour of pinnipeds I edited by Deane Renouf. - 1st ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-94-0lO-5373-0
1. Pinnipedia - Behavior. 1. Renouf, Deane, 1950-
QL737.P6B44 1991
599. 74'5-dc20 90-48768
CIP
To Bora, Tustin, Luke, Colin and Laura who encourage
the intrusion of projects like this into the otherwise
smooth running of a family.
To Meem, facilitator and peace-maker.
Contents
Contributors xi
Preface xiii
Acknowledgements xv
Foreword xvii
1 Determinants of mating systems in the Otariidae
(Pinnipedia) 1
1.1 Introduction 1
1.2 Overview of current theory 2
1.3 Types of mating system among Otariids 5
1.4 Variation in the form of Otariid mating systems 21
1.5 Conclusions 31
Acknowledgements 34
References 35
2 Pinniped mating system on land, ice and in the water
Emphasis on the Phocidae 45
2.1 What do we expect in pinnipeds and other
mammals? 47
2.2 What do we find in· mammals? 48
2.3 What determines the distribution of females and
the degree of polygyny? 51
2.4 Do the unique features of pinnipeds give an
advantage to one sex and affect the form that
mating systems take? 54
2.5 What are the consequences of mating on land
verses in the water? 57
2.6 Conclusions 58
References 59
viii Contents
3 Behavioural ecology of pinniped neonates 66
3.1 Introduction 66
3.2 Life history characteristics 67
3.3 The lactation period 89
3.4 Weaning 107
3.5 Behaviour and diet of weaned pups 110
3.6 Discussion 114
Acknowledgements 117
References 117
4 Communication in pinnipeds, with special reference to
non-acoustic signalling 128
4.1 Introduction 128
4.2 Signal structure and variation 130
4.3 Some concepts in communication 159
4.4 Combat behaviour and the communication of
threat 185
4.5 Ecological variation and time/activity budgets 196
4.6 Evolutionary patterns 208
4.7 Concluding comments 213
Acknowledgements 215
References 215
5 Physiology of behaviour in pinnipeds 236
5.1 Introduction 236
5.2 The physiology of reproductive behaviour 236
5.3 Maternal behaviour 249
5.4 Motor development and play 262
5.5 Immune system 266
5.6 Diving 267
5.7 Metabolism 270
5.8 Activity patterns 272
5.9 Thermoregulation 274
5.10 Moult 281
5.11 Orientation 282
5.12 Conclusion 285
Acknowledgements 286
References 286
6 Reproductive and foraging energetics of pinnipeds:
Implictions for life history patterns 300
6.1 Introduction 300
6.2 Reproductive biology 301
6.3 Maternal investment 306
6.4 Foraging behaviour 327
Contents ix
6.5 Summary and conclusions 337
Acknowledgements 338
References 338
7 Sensory reception and processing in: Phocidae and
Otariidae 345
7.1 Introduction 345
7.2 Methodological caveats in psychophysical threshold
measurement 347
7.3 Vision 348
7.4 Hearing 358
7.5 Tactile sensation: Vibrissae 373
7.6 Chemoreception 379
7.7 Magnetic reception 379
7.8 Discussion 381
7.9 Umwelten: Beyond the receptor 385
References 387
Index 396
Contributors
Burney J. Le Boeuf
Department of Biology and Institute of Marine Sciences, University
of California, Santa Cruz, California, USA 95064
Daryl J. Boness
Department of Zoological Research, National Zoological Park,
Smithsonian Institution, Washington, DC, USA 20008
W.D. Bowen
Bedford Institute of Oceanography, Department of Fisheries &
Oceans, Dartmouth, Nova Scotia, Canada B2Y 4A2
Daniel P. Costa
Joseph M. Long Marine Laboratory, Institute of Marine Sciences,
University of California, Santa Cruz, California, USA 95060
Edward H. Miller
Biology Department, University of Victoria, P.O. Box 1700,
Victoria, British Columbia, Canada V8W 2Y2
and
Royal British Columbia Museum, Victoria, British Columbia,
Canada V8V lX4
Deane Renouf
Ocean Sciences Centre, Memorial University of Newfoundland, St.
John's, Newfoundland, Canada AlB 3X9
Douglas Wartzok
Department of Biological Sciences, Indiana University-Purdue
University, Fort Wayne, IN, USA 46805
Preface
We teach our students of behavioural science that one first defines a
research problem, and then the most appropriate animal is selected
to investigate hypotheses. The reverse order of events is improper: a
particular class of animals should not be studied for its own sake. In
the case of the Pinnipeds (seals, sea lions, fur seals and walruses) the
organism and the problem are essentially the same. The research
questions presented in this volume in one way or another relate to
survival in two worlds, the ocean for foraging, and the terrain at its
edge or frozen above it for breeding.
The evolution of Pinniped behaviour and the mechanisms which
underlie it are a consequence of having to cope with two seemingly
incompatible sets of environmental constraints. The physiological
adaptations for concomitant functioning in two media with very
different physical characteristics have produced correlated behav
ioural modifications. The energetic demands of reproduction and
foraging are idiosyncratic because each activity occurs on opposite
sides of the air/water interface. As a result, the mating system must
reconcile aquatic design for such functions as locomotion and
thermoregulation, with the terrestrial requirements for successful
pupping. Similarly, the ecology of this dual habitat prescribes the
rules governing the behaviour of the neonate and its interactions
with its mother. Sensation, perception and cognition * and conse
quently the vehicle for communication must operate properly in air
and underwater either by using different mechanisms in each
medium, or by modifying systems which are designed for use in one
medium to operate effectively in the other.
Throughout the chapters which follow, whenever possible the
• This volume was to have a chapter entitled 'Learning and cognition in pinnipeds: a
quarter century of research' by Ronald J. Schusterman, but at the eleventh hour he
was unfortunately unable to provide it.
xiv Preface
lines of distinction between Otariidae (sea lions and fur seals) and
Phocidae (true seals) are clearly drawn. Too often, either
inadvertently or as a result of a paucity of information, the
behavioural literature has treated the two families collectively as if
they were a single group. In fact the differences between phocids and
otariids reflect variation in the extent of each family's re-adaptation
to the marine habitat. Therefore, contrasts between the two groups
illuminate aspects of adaptation. Behavioural investigations of the
third pinniped family, the Odobenidae (walrus) are unfortunately few,
but they are described when available.
The intention of this collection is to examine the behaviour of
Pinnipeds from the point of view of evolution and mechanism. The
history of these two schools of thought is riddled with controversy,
and in the 1960s could have been described as the American versus
the European positions in what was a rather lively warfare. (Consider
the battles in the scientific journals between the late Daniel Lehrman
and Konrad Lorenz and their respective proponents.) Fortunately
rapprochement has occurred since then, though occasionally one
hears behavioural ecologists and physiologists/psychologists arguing
from the floor of conferences, when in fact they are agreeing. This
volume is a demonstration of this rapprochement, wherein the
present and historical functions as well as the operating machinery
of Pinniped behaviour are presented as mutually important
components of its explanation.
Unless stated otherwise, the nomenclature used in this book is
that of Harrison and Ridgeways' 1981 Handbook of Marine
Mammals, volumes 1 and 2. Where possible, all units have been
changed to conform with the SI metric system.
