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we know of the way in which they perceive their environment and what determines how they
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respond to it at any given moment. It then focuses on why a particular behaviour takes the
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In offering straightforward explanations of some of the apparent curiosities which may have
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intrigued any casual observer of animals, Rory Putman draws on more than 20 years’ experience
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of teaching behaviour to undergraduate and adult education classes. He is Emeritus Professor of i
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Behavioural and Environmental Biology at the Manchester Metropolitan University and visiting
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Professor of Wildlife Welfare at the University of Utrecht. l B Rory Putman
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Understanding Animal Behaviour will be of special appeal to all h
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naturalists and everyone with an interest in animal behaviour.
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A highly readable and
accessible introduction to
current knowledge about the
way animals behave
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Comprehensive and authoritative P
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Beautifully illustrated throughout by
wildlife artist Catherine Putman
£18.99/$24.95
Whittles Publishing
Understanding Animal Behaviour
Published by
Whittles Publishing,
Dunbeath, Animal
Caithness KW6 6EG,
Scotland, UK
www.whittlespublishing.com
© 2018 Rory Putman
ISBN 978-184995-330-6
All rights reserved.
No part of this publication may be reproduced,
stored in a retrieval system, or transmitted,
in any form or by any means, electronic,
mechanical, recording or otherwise
without prior permission of the publishers.
Printed by
Understanding
Animal Behaviour
Rory Putman
Whittles Publishing
Contents
Preface ............................................................................................................ vii
Acknowledgements ....................................................................................... xi
1 Understanding animal behaviour ................................................................. 1
2 Behavioural reflexes ....................................................................................... 6
3 Linking responses and the co-ordination of more complex behaviours .. 14
4 Receiving stimuli and analysing the incoming information .................. 21
5 Motivation and its capacity to modify simple response patterns ........... 34
6 ‘Unpredictable’ behaviours: redirected and displacement activities ...... 44
7 Learning and its effects on behaviour ......................................................... 51
8 Behaviour genetics and the evolution of behaviour ................................ 57
9 Adaptiveness of behaviour: optimising the returns ................................. 73
10 Social organisation and social behaviour ................................................. 84
11 Reproductive behaviour and reproductive strategies ............................ 104
12 Mating behaviour and mate choice ......................................................... 112
13 Co-operative breeding and arguments about altruism ........................ 127
14 Territoriality and aggression ..................................................................... 136
15 How animals navigate ............................................................................... 148
16 ‘If we could talk to the animals…’ ............................................................ 155
Suggestions for further reading ............................................................... 173
Index of keywords ...................................................................................... 175
Index of species mentioned in the text .................................................... 177
Preface
It is a problem, it would seem, with any rapidly advancing science that successive professional
publications concern themselves increasingly with progressive refinement of the most
modern, or recent, of theories and somehow lose the context in which these are rooted.
Thus each generation of academic papers concerns itself only with the developments of
theory of the previous two or three years – and tends increasingly to debate their niceties
in isolation and out of the broader context of more general theory, which may remain
largely unaffected. However, each new publication becomes so concerned with dotting
‘i’s and crossing ‘t’s that, in paying attention to these minutiae, the main text is sometimes
forgotten or, at least, often erroneously, considered as general knowledge by those too close
to the finer detail.
As scientific publications become increasingly focused and erudite, more and more
involved in what might appear to be a semantic debate of the subtlest detail, so these
publications become inaccessible to the amateur or casual reader who merely seeks a
simple overview of more general theory. Even such general texts as may be available in
support of university courses often get bogged down in overmuch detail or complexity for
the non-specialist reader.
The greatest risk in attempting an overview for that non-specialist, however, and in
going back to basics, is that the text will be regarded as over-simplistic, naive or simply
out-dated, by those specialists who do then pick it up (or worse, review it!). For a number
of years I have been tempted to stick my neck out and compose such a book; for an equal
number of years I have shied away from the task, waiting for someone more qualified and
more articulate than myself to undertake the task. Yet maybe they, also, are too close to the
fine detail – or too nervous of the risk of critical review by professional colleagues!
What has finally prompted me to cast caution to the wind – and put onto paper this
basic introduction to animal behaviour for the interested non-specialist, was the recent
sad task of editing my late father’s papers. Amongst them was a series of philosophical
essays in which he had tried to make sense of some 80 years’ experience of life, as both
man and scientist, and offer to himself some conclusions about life, the universe and
everything. The essays were illuminating – but in many places sadly flawed by incomplete
knowledge or understanding. My father was a successful scientist (a nuclear physicist of
some reputation) and a lifelong naturalist – with a passionate interest in behaviour. Many
of the essays indeed dealt with an analysis of human behaviour patterns by comparison vii
with the behaviour of other animals, or at least interpreted within such a context; yet it
was in these essays, to my surprise, that the logic was often marred by misconception or,
at least, incomplete understanding of things which I myself would have assumed to be
common knowledge – especially to one such as my father who, albeit as an amateur, had
particular interest in behaviour and who read widely. If even his knowledge was so patchy
and incomplete, I had to argue, then there must indeed be many others who would be glad
of a simple book reviewing the very basics of our understanding of how and why animals
behave as they do.
The current book, then, is a development from a series of essays or lectures of my
own, tried and tested in numerous adult education courses given over the years to other
intelligent and interested non-specialists like my father. It is not an academic textbook –
indeed would be, and doubtless will be, scorned by many professionals as perhaps dated
and certainly as over-simplistic. It uses words and concepts which are intuitively easy
to grasp, even if they are in places oversimplified, in an attempt to offer to an amateur
audience an insight into the way animals behave, but couched within a framework and in
a conceptual language with which they can actually identify. I do not seek to patronise, but
to inform.
If, in my attempts to make it more accessible, I irritate professional behaviourists by
adopting devices or simplifications now frowned upon, or outmoded, I can only apologise
– but to an extent accept such criticism in advance, since this book is aimed not at them
but at that more general readership.
The book itself is presented in two main parts (although in practice these two sections
are effectively independent, and may indeed be read in either order). In the first I seek to
explore the ‘how’ of animal behaviour, taking a mechanistic approach to explore what we
know of the way in which animals perceive their environment and what determines how
they respond to it in the way that they do and what that response will be. In the latter part
of the book, the focus switches more to the ‘why’ of animal behaviour, asking not so much
how given responses are mediated through nerve and muscle systems, or how stimuli are
perceived in the brain, but rather what has moulded why a particular behaviour takes the
form that it does in its expression, what evolutionary forces have shaped – and continue
to shape – the detailed form of more complex behaviours. Why do animals forage in the
way they do? How may that foraging pattern be refined to optimal efficiency? Why do
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a (and why is there is such a bewildering variety of different ways of solving apparently the
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B same problem)? Why do some animals live as solitary individuals, while others live in
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di behaviours, but this seemed far too simple, and probably far too contentious, even if it
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viii years by those same ‘interested amateurs’.
But this volume is hopefully more than simply a non-technical book about animal
behaviour as such; or at least that is perhaps only part of the story. For the intention is
that it should also be highly visual – and copiously illustrated throughout, in support
and interpretation of the text. To an extent then, while the pictures are there to enhance
understanding of the written text, the text itself also takes on an additional role as a showcase
for some of the exquisite illustrations of my wife, wildlife artist Catherine Putman, who has
agreed to collaborate with me on the project. The book is intended, to a degree, as a joint
presentation of text and pictures, in celebration of the delights of a greater understanding of
animals and the way they behave.
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