15 fish and aquatic resources series 15 fish cognition...‘’fish cognition and behavior is an...

27.5mm 176mm176mm

250mm

ISBN 978-1-4443-32216

9 781444 332216

CMYK

Fish and aquatic resources series 15

Fish cognition and Behavior

second editionedited by

culum Brown, Kevin Laland and Jens Krause

15

The study of animal cognition has until recently been largely confined to birds and mammals; a historical bias which has led to the belief that learning plays little or no part in the development of behavior in fishes and reptiles. Research in recent decades has begun to redress this misconception and it is now recognized that fishes exhibit a rich array of sophisticated behavior with impressive learning capabilities entirely comparable with those of mammals and other terrestrial animals.

Building on the very well-received First Edition, and covering all major areas of fish learning, all chapters in this brand new edition of Fish Cognition and Behavior have been revised and updated by an internationally recognized team of contributing authors, with the addition of three brand new chapters, covering personality traits and behavior, lateralization of cognitive functions in fish, and cognition and welfare.

This Second Edition of Fish Cognition and Behavior is essential reading for all fish biologists and ethologists, and contains much information of commercial importance for fisheries managers and aquaculture personnel. Libraries in all universities and research establishments where biological sciences, aquaculture and fisheries are studied and taught will find this book an extremely important addition to their shelves.

Reviews of the First Edition‘’Fish Cognition and Behavior is an essential read for anyone interested in fish welfare, psychology, sensory biology, neurobiology, conservation, and the application of pure research.’’

The Quarterly Review of Biology

Fish Cognition and Behavior … “will open the eyes of many readers to the importance of fish…in studying and understanding various aspects of animal cognition.”

Animal Behaviour

“Insightful and fascinating questions are presented together with useful suggestions for future research directions on fish cognition… sufficiently entertaining to be read cover to cover.”

Fish and Fisheries

“This is one of the best-edited and most valuable volumes I have encountered for those interested in the behavior of fish.”

Reviews in Fish Biology & Fisheries

About the EditorsCulum Brown is at the Department of Biological Sciences, Macquarie University, Sydney, Australia.

Kevin Laland is at the Centre for Social Learning and Cognitive Evolution, School of Biology, University of St Andrews, UK.

Jens Krause is at the Department of Biology and Ecology of Fishes, Leibniz-Institute of Freshwater Ecology and Inland Fisheries, and also at Humboldt University, both in Berlin, Germany.

Cover illustration by Robert Gibbings (1889-1958), St Brendan and the sea monsters. Medium: Woodcut. Date created: 1934. Collection: Christchurch Art Gallery Te Puna o Waiwhetu. Presented by Mrs Rosalie Archer, 1975. Copyright permission kindly granted by Reading University Library.

Cover design by Workhaus.

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BLBK374-FM BLBK374-Brown June 7, 2011 11:29 Copyeditor’s Name: Trim: 244mm X 172mm Char Count=


Fish Cognition and Behavior

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Fish and Aquatic Resources Series

Series Editor: Tony J. PitcherProfessor of Fisheries Policy & Ecosystem Restoration in Fisheries,Fisheries Centre, Aquatic Ecosystems Research Laboratory, University ofBritish Columbia, Canada

The Wiley-Blackwell Fish and Aquatic Resources Series is an initiative aimed at providing key booksin this fast-moving field, published to a high international standard.

The Series includes books that review major themes and issues in the science of fishes and theinterdisciplinary study of their exploitation in human fisheries. Volumes in the Series combine a broadgeographical scope with in-depth focus on concepts, research frontiers, and analytical frameworks.These books will be of interest to research workers in the biology, zoology, ichthyology, ecology, andphysiology of fish and the economics, anthropology, sociology, and all aspects of fisheries. They willalso appeal to non-specialists such as those with a commercial or industrial stake in fisheries.

It is the aim of the editorial team that books in the Wiley-Blackwell Fish and Aquatic ResourcesSeries should adhere to the highest academic standards through being fully peer reviewed and editedby specialists in the field. The Series books are produced by Wiley-Blackwell in a prestigious anddistinctive format. The Series Editor, Professor Tony J. Pitcher, is an experienced international author,and founding editor of the leading journal in the field, Fish and Fisheries.

The Series Editor, and Publisher at Wiley-Blackwell, Nigel Balmforth, will be pleased to discusssuggestions, advise on scope, and provide evaluations of proposals for books intended for the Series.Please see contact details listed below.

Titles currently included in the Series1. Effects of Fishing on Marine Ecosystems and Communities (S. Hall) 19992. Salmonid Fishes (Edited by Y. Altukhov et al.) 20003. Percid Fishes (J. Craig) 20004. Fisheries Oceanography (Edited by P. Harrison and T. Parsons) 20005. Sustainable Fishery Systems (A. Charles) 20006. Krill (Edited by I. Everson) 20007. Tropical Estuarine Fishes (S. Blaber) 20008. Recreational Fisheries (Edited by T. J. Pitcher & C. E. Hollingworth) 20029. Flatfishes (Edited by R. Gibson) 2005

10. Fisheries Acoustics (J. Simmonds & D. N. MacLennan) 200511. Fish Cognition and Behavior (Edited by C. Brown, K. Laland & J. Krause) 200612. Seamounts (Edited by T. J. Pitcher, T. Morato, P. J. B. Hart, M. R. Clark, N. Haggan &

R. S. Santos) 200713. Sharks of the Open Ocean (Edited by M. D. Camhi, E. K. Pikitch and E. A. Babcock) 200814. World Fisheries (Edited by R. E. Ommer, R. I. Perry, K. Cochrane & P Cury) 201115. Fish Cognition and Behavior, Second Edition (Edited by C. Brown, K. N. Laland & J. Krause)

2011

For further information concerning existing books in the series, please visit: www.wiley.com

To discuss an idea for a new book, please contact:Nigel Balmforth, Life Sciences, Wiley-Blackwell, 9600 Garsington Road, Oxford OX4 2DQ, UKTel: +44 (0) 1865 476501

Email: [email protected]

ii


Fish Cognition and Behavior

Edited by

Culum BrownDepartment of Biological Sciences, Macquarie University,

Sydney, Australia

Kevin LalandCentre for Social Learning and Cognitive Evolution, School of Biology,

University of St Andrews, UK

Jens KrauseDepartment of Biology and Ecology of Fishes, Leibniz-Institute of

Freshwater Ecology and Inland Fisheries, Berlin;

Humboldt University, Berlin

A John Wiley & Sons, Ltd., Publication

iii


First edition published 2006This edition first published 2011 C© 2011, 2006 by Blackwell Publishing Ltd.

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Designations used by companies to distinguish their products are often claimed as trademarks. Allbrand names and product names used in this book are trade names, service marks, trademarks orregistered trademarks of their respective owners. The publisher is not associated with any product orvendor mentioned in this book. This publication is designed to provide accurate and authoritativeinformation in regard to the subject matter covered. It is sold on the understanding that the publisheris not engaged in rendering professional services. If professional advice or other expert assistance isrequired, the services of a competent professional should be sought.

Library of Congress Cataloging-in-Publication Data

Fish cognition and behavior / edited by Culum Brown. – 2nd ed.p. cm. – (Fish and aquatic resources series)

Includes bibliographical references and index.ISBN 978-1-4443-3221-6 (hardcover : alk. paper) 1. Fishes–Behavior. 2. Fishes–Psychology.

3. Cognition in animals. I. Brown, Culum.QL639.3.F575 2011597–dc22

2011002188

A catalogue record for this book is available from the British Library.

This book is published in the following electronic formats: ePDF 9781444342505;Wiley Online Library 9781444342536; ePub 9781444342512; Mobi 9781444342529

Set in 10/13pt Times New Roman by Aptara R© Inc., New Delhi, India

1 2011

iv

http://www.wiley.com/wiley-blackwell


Contents

Preface and Acknowledgements xvSeries Foreword xviList of Contributors xix

1 Fish Cognition and Behaviour 1Brown, Laland and Krause

1.1 Introduction 11.2 Contents of this book 3

References 9

2 Learning of Foraging Skills by Fish 10Warburton and Hughes

2.1 Introduction 102.2 Some factors affecting the learning process 12

2.2.1 Reinforcement 122.2.2 Drive 122.2.3 Stimulus attractiveness 122.2.4 Exploration and sampling 142.2.5 Attention and simple association 142.2.6 Cognition 152.2.7 Memory systems and skill transfer 18

2.3 Patch use and probability matching 192.4 Performance 212.5 Tracking environmental variation 232.6 Competition 262.7 Learning and fish feeding: some applications 272.8 Conclusions 27

Acknowledgements 28References 29


vi Contents

3 Learned Defences and Counterdefences in Predator–Prey Interactions 36Kelley and Magurran

3.1 Introduction 363.2 The predator–prey sequence 38

3.2.1 Encounter 393.2.1.1 Avoiding dangerous habitats 393.2.1.2 Changing activity patterns 40

3.2.2 Detection 413.2.2.1 Crypsis 423.2.2.2 Sensory perception 42

3.2.3 Recognition 433.2.3.1 Associative learning 433.2.3.2 Learning specificity 443.2.3.3 Search images 453.2.3.4 Aposematism and mimicry 46

3.2.4 Approach 473.2.4.1 Pursuit deterrence 473.2.4.2 Gaining information about the predator 473.2.4.3 Social learning 473.2.4.4 Habituation 49

3.2.5 Evasion 493.2.5.1 Reactive distance and escape speed and trajectory 503.2.5.2 Survival benefits/capture success 50

3.3 Summary and discussion 51Acknowledgements 52References 53

4 Learning about Danger: Chemical Alarm Cues andThreat-Sensitive Assessment of Predation Risk by Fishes 59Brown, Ferrari and Chivers

4.1 Introduction 594.2 Chemosensory cues as sources of information 60

4.2.1 Learning, innate responses and neophobia 604.2.2 Learned predator recognition through conditioning

with alarm cues 624.3 Variable predation risk and flexible learning 62

4.3.1 Assessing risk in time 644.3.2 Sensory complementation and threat-sensitive learning 65

4.4 Generalisation of risk 664.4.1 Generalising of predator cues 664.4.2 Generalisation of non-predator cues 67

4.5 Predator recognition continuum hypothesis 684.5.1 Ecological selection for innate versus learned

recognition of predators 694.5.2 Ecological selection for generalised learning 69


Contents vii

4.6 Retention: the forgotten component of learning 704.7 Conservation, management and learning 72

4.7.1 Conditioning predator recognition skills 724.7.2 Anthropogenic constraints 734.7.3 Field-based studies 73

4.8 Conclusions 74Acknowledgements 74References 74

5 Learning and Mate Choice 81Witte and Nobel

5.1 Introduction 815.2 Sexual imprinting 82

5.2.1 Does sexual imprinting promote sympatricspeciation in fishes? 82

5.3 Learning after reaching maturity 835.4 Eavesdropping 84

5.4.1 Eavesdropping and mate choice 845.4.2 Benefits of eavesdropping 845.4.3 The audience effect 85

5.5 Mate-choice copying 875.5.1 Mate-choice copying – first experimental evidence

and consequence 885.5.2 Mate-choice copying – evidence from the wild 895.5.3 Mate-choice copying when living in sympatry

or allopatry 915.5.4 Mate-choice copying – the role of the early

environment 925.5.5 Quality of the model fish 93

5.6 Social mate preferences overriding genetic preferences 945.6.1 Indications from guppies 945.6.2 Indications from sailfin mollies 95

5.7 Cultural evolution through mate-choice copying 965.8 Does mate-choice copying support the evolution of a novel

male trait? 965.8.1 Theoretical approaches 975.8.2 Experimental approaches 98

5.9 Is mate-choice copying an adaptive mate-choicestrategy? 995.9.1 Benefits of mate-choice copying 995.9.2 Costs of mate-choice copying 100

5.10 Outlook 1015.11 Conclusions 102

References 102


viii Contents

6 Aggressive Behaviour in Fish: Integrating Information aboutContest Costs 108Hsu, Earley and Wolf

6.1 Introduction 1086.2 Information about resource value 1106.3 Information about contest costs 110

6.3.1 Assessing fighting ability 1116.3.2 Information from past contests 113

6.3.2.1 Winner and loser effects 1136.3.2.2 Individual recognition 1176.3.2.3 Social eavesdropping 117

6.3.3 Integrating different types of cost-relatedinformation 118

6.4 Physiological mechanisms 1196.5 Conclusions and future directions 126


7 Personality Traits and Behaviour 135Budaev and Brown

7.1 Introduction 1357.2 Observation and description of personality 137

7.2.1 Current terminology 1377.2.1.1 Shyness–boldness 1387.2.1.2 Coping styles 1407.2.1.3 Behavioural syndromes 140

7.2.2 Objectivity 1407.2.3 Labelling personality traits; construct validity 1427.2.4 Objective and subjective measurements

of personality 1427.2.5 Modern terminology and statistical

approaches 1457.3 Proximate causation 1467.4 Ontogeny and experience 1497.5 Is personality adaptive? 150

7.5.1 Frequency- and density-dependent selection 1507.5.2 State-dependent models 151

7.6 Evolution 1537.7 Wider implications 155

7.7.1 Fish production and reproduction 1557.7.2 Personality and population dynamics 155

7.8 Conclusions 156Acknowledgements 157References 157


Contents ix

8 The Role of Learning in Fish Orientation 166Odling-Smee, Simpson and Braithwaite

8.1 Introduction 1668.2 Why keep track of location? 1668.3 The use of learning and memory in orientation 1678.4 Learning about landmarks 1688.5 Compass orientation 1718.6 Water movements 1728.7 Inertial guidance and internal ‘clocks’ 1738.8 Social cues 1748.9 How flexible is orientation behaviour? 174

8.9.1 When to learn? 1748.9.2 What to learn? 1758.9.3 Spatial learning capacity 176

8.10 Salmon homing – a case study 1778.11 Conclusion 179


9 Social Recognition of Conspecifics 186Griffiths and Ward

9.1 Introduction 1869.2 Recognition of familiars 186

9.2.1 Laboratory studies of familiarity 1879.2.2 Mechanisms of familiarity recognition 1879.2.3 Functions of associating with familiar fish 1919.2.4 Familiarity in free-ranging fishes 1949.2.5 Determinants of familiarity 195

9.3 Familiarity or kin recognition? 1969.3.1 Kin recognition theory 1969.3.2 Evidence for kin recognition from laboratory studies 2009.3.3 Advantages of kin discrimination 2019.3.4 Kin association in the wild 2019.3.5 Explaining the discrepancies between laboratory

and field 2039.3.6 Kin avoidance 205

9.4 Conclusion 206References 207

10 Social Organisation and Information Transfer in Schooling Fish 217Ioannou, Couzin, James, Croft and Krause

10.1 Introduction 21710.2 Collective motion 21810.3 Emergent collective motion in the absence of external stimuli 219


x Contents

10.4 Response to internal state and external stimuli:Information processing within schools 22010.4.1 Collective response to predators 22010.4.2 Mechanisms and feedback in information transfer 22210.4.3 Information transfer during group foraging

and migration 22510.5 Informational status, leadership and collective

decision-making in fish schools 22510.6 The structure of fish schools and populations 22710.7 Social networks and individual identities 22910.8 Community structure in social networks 23210.9 Conclusions and future directions 233


11 Social Learning in Fishes 240Brown and Laland

11.1 Introduction 24011.2 Antipredator behaviour 24111.3 Migration and orientation 24411.4 Foraging 24711.5 Mate choice 24811.6 Aggression 24911.7 Trade-offs in reliance on social and asocial sources of information 25011.8 Concluding remarks 252


12 Cooperation and Cognition in Fishes 258Alfieri and Dugatkin

12.1 Introduction 25812.2 Why study cooperation in fishes? 25912.3 Cooperation and its categories 261

12.3.1 Category 1 – kin selection 26112.3.1.1 Cognition and kin selection 26112.3.1.2 Example of kin selected cooperation:

Cooperative breeding 26212.3.1.3 Example of kin selected cooperation:

Conditional territory defence 26212.3.2 Category 2 – reciprocity 263

12.3.2.1 Cognition and reciprocity 26412.3.2.2 Example of reciprocity: Egg trading 26512.3.2.3 Example of reciprocity: Predator inspection 26612.3.2.4 Example of reciprocity: Interspecific

cleaning behaviour 267


Contents xi

12.3.3 Category 3 – by-product mutualism 26812.3.3.1 Cognition and by-product mutualism 26812.3.3.2 Example of by-product mutualism:

Cooperative foraging 26912.3.4 Category 4 – trait group selection 270

12.3.4.1 Cognition and trait group selection 27012.3.4.2 Example of trait group selected

cooperation: Predator inspection 27012.4 Conclusion 271


13 Machiavellian Intelligence in Fishes 277Bshary

13.1 Introduction 27713.2 Evidence for functional aspects of Machiavellian intelligence 279

13.2.1 Information gathering about relationshipsbetween other group members 279

13.2.2 Predator inspection 28013.2.3 Group-living cichlids 28113.2.4 Machiavellian intelligence in cleaning mutualisms 283

13.2.4.1 Categorisation and individualrecognition of clients 283

13.2.4.2 Building up relationships betweencleaners and resident clients 284

13.2.4.3 Use of tactile stimulation by cleaners tomanipulate client decisions andreconcile after conflicts 284

13.2.4.4 Audience effects in response to imagescoring and tactical deception 285

13.2.4.5 Punishment by males during pairinspections 285

13.3 Evidence for cognitive mechanisms in fishes 28613.3.1 What cognitive abilities might cleaners need to

deal with their clients? 28613.3.2 Other cognitive mechanisms 287

13.4 Discussion 28813.4.1 Future avenues I: How Machiavellian is fish

behaviour? 28913.4.2 Future avenues II: Relating Machiavellian-type

behaviour to brain size evolution 29013.4.3 Extending the Machiavellian intelligence

hypothesis to general social intelligence 291Acknowledgements 291References 291


xii Contents

14 Lateralization of Cognitive Functions in Fish 298Bisazza and Brown

14.1 Introduction 29814.2 Lateralized functions in fish 300

14.2.1 Antipredator behavior 30014.2.1.1 Predator inspection 30114.2.1.2 Predator evasion 30214.2.1.3 Fast escape response 303

14.2.2 Mating behavior 30414.2.3 Aggression 30414.2.4 Shoaling and social recognition 30414.2.5 Foraging behavior 30614.2.6 Exploration and response to novelty 30614.2.7 Homing and spatial abilities 30714.2.8 Communication 307

14.3 Individual differences in lateralization 30814.3.1 Hereditary basis of lateralization 30814.3.2 Sex differences in lateralization 30914.3.3 Environmental factors influencing development

of lateralization 31014.3.4 Lateralization and personality 311

14.4 Ecological consequences of lateralization of cognitive functions 31214.4.1 Selective advantages of cerebral lateralization 31214.4.2 Costs of cerebral lateralization 31414.4.3 Maintenance of intraspecific variability in the

degree of lateralization 31614.4.4 Evolutionary significance of population biases

in laterality 31614.5 Summary and future research 317


15 Brain and Cognition in Teleost Fish 325Broglio, Gomez, Duran, Salas and Rodrıguez

15.1 Introduction 32515.2 Classical conditioning 327

15.2.1 Delay motor classical conditioning and teleostfish cerebellum 328

15.2.2 Role of the teleost cerebellum and telencephalicpallium in trace motor classical conditioning 330

15.3 Emotional learning 33115.3.1 Role of the medial pallium in avoidance

conditioning and taste aversion learning 33215.3.2 Teleost cerebellum and fear conditioning 334


Contents xiii

15.4 Spatial cognition 33615.4.1 Allocentric spatial memory representations in

teleost fishes 33715.4.2 Role of the teleost telencephalon in egocentric

and allocentric spatial navigation 34015.4.3 Map-like memories and hippocampal pallium in

teleost fishes 34515.4.4 Neural mechanisms for egocentric spatial

orientation 34715.5 Concluding remarks 349


16 Fish Behaviour, Learning, Aquaculture and Fisheries 359Ferno, Huse, Jakobsen, Kristiansen and Nilsson

16.1 Fish learning skills in the human world 35916.2 Fisheries 362

16.2.1 Spatial dynamics 36216.2.1.1 Learning skills and movement 36216.2.1.2 Social learning of migration pattern 36316.2.1.3 Implications of learning for

fisheries management 36616.2.2 Fish capture 367

16.2.2.1 Natural variations in spatial distributionand behaviour 369

16.2.2.2 Avoidance and attraction before fishing 36916.2.2.3 Before physical contact with the gear 36916.2.2.4 After physical contact with the gear 37116.2.2.5 Behaviour after escaping the gear and

long-term consequences 37216.2.3 Abundance estimation 374

16.3 Aquaculture 37516.3.1 Ontogeny 37516.3.2 Habituation, conditioning and anticipation 37616.3.3 Pavlovian learning – delay and trace conditioning 37816.3.4 Potential use of reward conditioning in aquaculture 37916.3.5 Operant learning 38216.3.6 Individual decisions and collective behaviour 383

16.4 Stock enhancement and sea-ranching 38416.5 Escapees from aquaculture 38816.6 Capture-based aquaculture 38916.7 Conclusions and perspectives 389



xiv Contents

17 Cognition and Welfare 405Sneddon

17.1 Introduction 40517.1.1 Fish welfare 40617.1.2 Preference and avoidance testing 40717.1.3 Behavioural flexibility and intraspecific variation 408

17.2 What is welfare? 40817.2.1 Sentience and consciousness 40917.2.2 Cognition and welfare 410

17.3 What fishes want 41017.3.1 Preference tests 411

17.3.1.1 Physical habitat 41117.3.1.2 Breeding 41317.3.1.3 Diet 41317.3.1.4 Social interactions 414

17.4 What fishes do not want 41617.5 Pain and fear in fish 41717.6 Personality in fish 42017.7 Wider implications for the use of fish 420

17.7.1 Aquaculture 42117.7.2 Fisheries 42517.7.3 Recreational fishing 42517.7.4 Research 42617.7.5 Companion fish 427

17.8 Conclusion 427Acknowledgements 429References 429

Species List 435Index 443


Preface and Acknowledgements

This is now the second edition of this book, which is a follow-up from our successful volumeof Fish and Fisheries dedicated to learning in fishes. All of the contributors to that volumeand our previous edition have updated their work in this second edition, and we have addedseveral more contributions covering a broad range of fish behaviour. It is encouraging to seea range of contributions from both established and emerging experts in fish behaviour. Theeditors would like to thank all of the contributors for their hard work and enthusiasm whilstproducing this volume. Such an undertaking would be far too big a task for one personalone, given the increasing volume of behavioural research conducted on fishes. There isalso a long list of reviewers whose comments have made valuable contributions to each ofthe chapters. We would like to thank Nigel Balmforth and his colleagues at Wiley-Blackwellfor their valuable support and Tony J. Pitcher for writing the Series Foreword.


Series Foreword

Many years ago, teaching a series of practical animal behaviour classes to undergraduates,I asked my students to try to train goldfish to feed on food pellets delivered from coloredtubes. It all worked well enough, the goldfish learned to feed from tubes painted with, toour eyes, subtly different colors, and after running the usual controls for light intensity,the students discovered that the fish had very effective color vision. After the classes, thegoldfish were returned to a stock aquarium and were left alone for a year, although someof them may have taken part in other experiments. The following year, it was evident rightat the start of the student practical that each goldfish remembered the exact color andlocation of its feeding tube from 1 year before, a remarkable cognitive feat from an animalthat is supposed to have only a 3-second memory, as satirized in the recent Pixar FindingNemo film.

Fishermen, anglers, and most of the general public encounter live fish only when theyare flopping helplessly, and apparently dumbly, on the boat deck or seashore. In suchcircumstances, it is hard to believe that fish are intelligent sentient beings: Even in theleast speciesist1 science fiction, in Douglas Adams’ (1979) otherwise splendid Hitchhiker’sGuide to the Galaxy for example, fish are merely food for whales (except for one smartautomaton, the universal translator known as Babelfish). On the other hand, watching fishhunting for food, engaging with mates, or raising young, aquarists and divers gain a verydifferent view of the behavioural complexities, elegant adaptations and cognitive abilitiesthat lie behind the actions of fish. Fish are endowed with a complex evolved neural andcognitive capacity that reflects the challenges faced by their ancestors, rather than anyphylogenetic proximity to humans. This is the scientific reality, and it is a subject of thisvolume in the Fish and Aquatic Resources series.

This is the second edition of the book Fish Cognition and Behavior, which grew out of a2003 special issue of the Wiley-Blackwell journal Fish and Fisheries (Brown et al. 2003),itself was built upon a pioneering review paper published in the early 1990s (Kieffer &Colgan 1992). In the second edition of this book, we have a set of 17 expanded and updatedchapters written by internationally renowned authors that review this important area. Thebook has been put together and edited by three of the world leaders in this field: CulumBrown from Macquarie University, Australia; Kevin Laland from St Andrews, Scotland;and Jens Krause from Humboldt University, Berlin.

1 “Speciesism” involves assigning different values or rights to beings on the basis of their species. The term wascoined by Richard D. Ryder in 1970 and is used to denote prejudice similar in kind to sexism and racism.


Series Foreword xvii

The editors point out that cognition includes perception, attention, memory formation,and executive functions related to information processing such as learning and problemsolving. Fish, it turns out, are not primitive in these respects. Bony fish have had, after all,over 60 million years for their genes to evolve the capacity to build and run fish brainsthat can deal flexibility with the diverse but volatile underwater environment—a time spanten times longer than our human line. Indeed, the editors of this book show that, far frombeing primitive automatons as had once been thought, fish “have evolved complex culturaltraditions, pursue Machiavellian strategies of manipulation, deception and reconciliation,can monitor the social prestige of others, and can cooperate during foraging, navigation,reproduction and predator avoidance.”

This volume presents fascinating, timely, and comprehensive “state of the art” reviewsof the cognitive abilities of fish, and readers will find the elements of a fresh synthesis inthis field. Therefore, it should find a home on the bookshelves and in the libraries of a broadset of practitioners and students concerned with fish evolution, behaviour, and ecology,including those, like myself, who might still wish to call themselves ichthyologists.

Professor Tony J. PitcherSeries Editor: Wiley-Blackwell Fish and Aquatic Resources Series

Fisheries Centre, University of British Columbia, Vancouver, Canada

References

Adams, D. (1979) The Hitchhiker’s Guide to the Galaxy. Heinemann, London, UK.Brown, C., Laland, K. & Krause, J. (2003) Special issue on learning in fishes: why they are smarter

than you think. Fish and Fisheries, 4(3), 197–288.Kieffer, J.D. & Colgan, P.W. (1992) The role of learning in fish behaviour. Reviews in Fish Biology

and Fisheries, 2, 125–143.


xviii


List of Contributors

Michael S. AlfieriBiology DepartmentViterbo University900 Viterbo DriveLa Crosse, WI 54601, USAEmail: [email protected]

Angelo BisazzaComparative Psychology Research GroupUniversity of PadovaPadova, ItalyEmail: [email protected]

Cristina BroglioLaboratorio de PsicobiologiaUniversidad de Sevillac/ Camilo Jose Cela s/n, 41018Sevilla, SpainEmail: [email protected]

Victoria A. BraithwaiteSchool of Forest Resources and

Department of BiologyPennsylvania State University

University ParkPA 16802, USAEmail: [email protected]

Culum BrownDepartment of Biological SciencesMacquarie UniversitySydney 2109, AustraliaEmail: [email protected]

Grant E. BrownDepartment of BiologyConcordia University7141 Sherbrooke St., W. MontrealQuebec, H4B 1R6, CanadaEmail: [email protected]

Redouan BsharyUniversite de Neuchatel,Rue Emile-Argand 11 CH-2007Neuchatel, SwitzerlandEmail: [email protected]

Sergey BudaevSevertsov Institute of Ecology and EvolutionRussian Academy of SciencesLeninsky prospect 33Moscow 119071, RussiaEmail: [email protected]

Douglas P. ChiversDepartment of BiologyUniversity of SaskatchewanSaskatoon, Saskatchewan

SK S7N 5E2, CanadaEmail: [email protected]

Iain D. CouzinDepartment of Ecology and EvolutionPrinceton UniversityPrinceton, NJ 08544-2016, USAEmail: [email protected]


xx List of Contributors

Darren P. CroftSchool of PsychologyExeter UniversityPerry RoadExeter, EX4 4QG, UKEmail: [email protected]

Lee A. DugatkinDepartment of BiologyUniversity of LouisvilleLouisville, KY 40208, USAEmail: [email protected]

Emilio DuranLaboratorio de PsicobiologiaUniversidad de Sevillac/ Camilo Jose Cela s/n, 41018Sevilla, SpainEmail: [email protected]

Ryan L. EarleyDepartment of Biological SciencesUniversity of AlabamaBox 870344Tuscaloosa, Alabama 35487, USAEmail: [email protected]

Anders FernoDepartment of BiologyUniversity of BergenPO Box 7800N-5020 Bergen, NorwayEmail: [email protected]

Maud C.O. FerrariDepartment of Biomedical SciencesWCVM, University of SaskatchewanSaskatoon, SK, CanadaEmail: [email protected]

Antonia GomezLaboratorio de PsicobiologiaUniversidad de Sevillac/ Camilo Jose Cela s/n, 41018Sevilla, SpainEmail: [email protected]

Sian W. GriffithsCardiff School of BiosciencesPO Box 915, CardiffWales, CF10 3TL, UKEmail: [email protected]

Yuying HsuDepartment of Life ScienceNational Taiwan Normal UniversityNo. 88, Section 4, Ting-Chou RoadTaipei 116, TaiwanEmail: [email protected]

Roger HughesSchool of Biological SciencesEnvironment CentreUniversity of Wales, BangorGwynedd, LL57 2UW, UKEmail: [email protected]

Geir HuseInstitute of Marine ResearchPO Box 1870-NordnesN-5817 Bergen, NorwayEmail: [email protected]

Christos C. IoannouDepartment of Ecology and EvolutionPrinceton UniversityPrinceton, NJ 08544-2016, USAEmail: [email protected]

Per Johan JakobsenDepartment of BiologyUniversity of BergenPO Box 7800N-5020 Bergen, NorwayEmail: [email protected]

Richard JamesDepartment of PhysicsUniversity of BathBath, BA2 7AY, UKEmail: [email protected]


List of Contributors xxi

Jens KrauseDepartment of Biology and Ecology

of FishesLeibniz-Institute of Freshwater Ecology

and Inland FisheriesBerlin, GermanyEmail: [email protected]

Jennifer L. KelleyCentre for Evolutionary BiologySchool of Animal BiologyThe University of Western AustraliaNedlands, WA 6009, AustraliaEmail: [email protected]

Tore S. KristiansenInstitute of Marine ResearchPO Box 1870-NordnesN-5817 Bergen, NorwayEmail: [email protected]

Kevin LalandCentre for Social Learning and

Cognitive EvolutionSchool of Biology

University of St AndrewsSt Andrews, Fife, KY16 9TS, ScotlandEmail: [email protected]

Anne E. MagurranGatty Marine LaboratoryUniversity of St AndrewsSt Andrews, Fife, KY16 8LB, ScotlandEmail: [email protected]

Jonatan NilssonInstitute of Marine ResearchPO Box 1870-NordnesN-5817 Bergen, NorwayEmail: [email protected]

Sabine NobelDepartment of BiologyUniversity of SiegenAdolf-Reichwein-Str. 2D-57068 Siegen, GermanyEmail: [email protected]

Lucy Odling-SmeeNature Publishing Group4 Crinan Street, LondonN1 9XW, UKEmail: [email protected]

Tony PitcherFisheries CentreThe University of British ColumbiaVancouver, BC, V6T 1Z4, CanadaEmail: [email protected]

Fernando RodrıguezLaboratorio de PsicobiologiaUniversidad de Sevillac/ Camilo Jose Cela s/n, 41018Sevilla, SpainEmail: [email protected]

Cosme SalasLaboratorio de PsicobiologiaUniversidad de Sevillac/ Camilo Jose Cela s/n, 41018Sevilla, SpainEmail: [email protected]

Lynne U. SneddonIntegrative BiologyUniversity of LiverpoolCrown StreetLiverpool, L69 7ZB, UKEmail: [email protected]

Kevin WarburtonSchool of Environmental SciencesFaculty of ScienceCharles Sturt UniversityThurgoona, New South Wales, AustraliaEmail: [email protected].

Ashley WardSchool of Biological SciencesUniversity of SydneyNew South Wales, AustraliaEmail: [email protected]


xxii List of Contributors

Klaudia WitteDepartment of BiologyUniversity of SiegenAdolf-Reichwein-Str. 2D-57068 Siegen, GermanyEmail: [email protected]

Larry L. WolfDepartment of BiologySyracuse University107 College Place, Life Sciences ComplexSyracuse, New York, 13244, USAEmail: [email protected]

BLBK374-01 BLBK374-Brown May 11, 2011 13:48 Copyeditor’s Name: Trim: 244mm X 172mm Char Count=

Chapter 1

Fish Cognition and Behaviour

Culum Brown, Kevin Laland and Jens Krause

1.1 Introduction

The field of animal cognition is the modern approach to understanding the mental capa-bilities of animals. The theories are largely an extension of early comparative psychologywith a strong influence of behavioural ecology and ethology. Cognition has been vari-ously defined in the literature. Some researchers confine cognition to higher order mentalfunctions including awareness, reasoning and consciousness. However, a more general def-inition of cognition also includes perception, attention, memory formation and executivefunctions related to information processing such as learning and problem solving. Thestudy of animal cognition has been largely confined to birds and mammals, particularlynon-human primates. This bias in the literature is in part due to the approach taken in the1950s when cognitive psychologists began to compare known human mental processes withother closely related species. This bias was reinforced by an underlying misconception thatlearning played little or no role in the development of behaviour in reptiles and fishes.

Throughout scientific history fishes have largely been viewed as automatons. Theirbehaviour was thought to be almost exclusively controlled by unlearned predispositions.Ethologists characterised their behaviour as a series of fixed action patterns released onexposure to appropriate environmental cues (sign stimuli). Whilst there is no doubt thatfishes are the most ancient form of vertebrates, they are only ‘primitive’ in the sense thatthey have been on earth for in excess of 500 million years and that all other vertebratesevolved from some common fish-like ancestor (around 360 million years ago). However, itis important to note that fishes have not been stuck in an evolutionary quagmire during thistime. Their form and function have not remained stagnant over the ages. On the contrary,within this time frame they have diversified immensely to the point where there are morespecies of fish than all other vertebrates combined (currently over 32,000 described species)occupying nearly every imaginable aquatic niche.

The erroneous view that both behavioural and neural sophistications are associated ina linear progression from fishes through reptiles and birds to mammals is largely due toa heady mix of outdated and unscientific thinking. Aristotle’s concept of Scala naturae

Fish Cognition and Behavior, Second Edition. Edited by Culum Brown, Kevin Laland and Jens Krause.C© 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.


2 Fish Cognition and Behavior

(the scale of nature) and a Christian fundamentalist view that man is the pinnacle of thenatural world have dominated conceptions of animal intelligence for millennia. However,Darwin’s theory of evolution is fundamentally inconsistent with a gradual progression ofbehavioural flexibility and cognitive complexity from ‘primitive’ to ‘advanced’ life forms,leading inevitably to humans at the peak (i.e. the wrong-headed notion of an evolutionaryladder). There is nothing progressive about Darwinian evolution, and any semblance ofprogression merely reflects our anthropocentric bias to track evolutionary lineages thatculminate in our species, and to evaluate other species by their similarity to ourselves. Thecognitive capabilities of a species will reflect the history of selection amongst its ancestors,rather than phylogenetic proximity to humanity.

Amongst the vertebrates, fishes have suffered the most from the common misconceptionof the evolutionary ladder. However, over the last few decades this fallacy has begun tobe redressed. Researchers now realised that, like the rest of the vertebrate kingdom, fishesexhibit a rich array of sophisticated behaviour and that learning plays a pivotal role in be-havioural development of fishes. Gone, or at least redundant, are the days where fishes werelooked down upon as pea-brained machines whose only behavioural flexibility was severelycurtailed by their infamous 3-second memory (a la Dory in Disney’s Finding Nemo). Asthis book will reveal, many fishes in fact have impressive long-term memories comparableto most other vertebrates (Brown 2001; Warburton 2003). Their neural architecture hasboth analogous and homologous components with mammals, and is capable of much thesame processing power (Broglio et al. 2003). Their cognitive capacity in many domainsis comparable with that of non-human primates (Bshary et al. 2002; Laland & Hopitt2003; Odling-Smee & Braithwaite 2003). Fishes have evolved complex cultural traditionsand pursue Machiavallian strategies of manipulation, deception and reconciliation (Bsharyet al. 2002; Brown & Laland 2003). They not only recognise one another, but can monitorthe social prestige of and dominance relations amongst others (McGregor 1993; Griffiths2003; Grosenick et al. 2007) and cooperate in a variety of ways during foraging, navigation,reproduction and predator avoidance (Huntingford et al. 1994; Johnstone & Bshary 2004;Fitzpatrick et al. 2006). It is clear that the recent developments in our understanding offish behaviour require a substantial reappraisal of their behavioural flexibility that warrantsfurther investigation.

Since the 1960s there has been a rapid increase in the number of papers published onlearning in fishes and those published since 1991 has risen dramatically (Fig 1.1). In theearly 1990s James Kieffer and Patrick Coglan published the first comprehensive reviewof the role of learning in the development of fish behaviour (Kieffer & Colgan 1992). Intheir review, they were able to draw on some 70 published papers on learning in fishes, avast improvement over previous works (Thorpe 1963; Gleitman & Rozin 1971). In 2003,we published a collection of reviews on the topic in a special issue of the journal Fishand Fisheries. The special issue contained eight reviews on various aspects of learning infishes referring to over 500 papers. In the first edition of this book, which contained 14chapters, many of these reviews were revised and extended. The second edition has beensignificantly expanded again, with revision of most chapters and the inclusion of three morechapters on laterality, personality and welfare consequences of cognition. This new editionnow examines the role of cognition in every major aspect of fish biology, from foragingand predator avoidance to fighting and social relationships.


Fish Cognition and Behaviour 3

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Fig. 1.1 The number of publications on fish learning and cognition since 1991 has increased substantially. Databased on key word search (fish, fishes, learn, learning and cognition) in Web of Science.

1.2 Contents of this book

Apart from this opening introduction, Chapter 2, by Kevin Warburton and Roger Hughes,investigates the role of learning in foraging behaviour, drawing on both psychological andbehavioural ecology literature. They suggest that learning and memory play significantroles in the foraging activities in fish and that memory, like many traits, seems to be highlyadapted to the specific requirements of each species. Interestingly, they suggest that insome circumstances forgetting might be just as important as remembering. The chapterhighlights that the similarities between vertebrate learning systems are far more strikingthan the differences and fishes rely on a wide array of learning mechanisms in their dailylives. The literature shows that learning is vital in many aspects of fish foraging behaviour,from the formation of foraging search images, to prey capture and handling. Warburtonand Hughes also outline various experiments that explore foraging theory and point outthat fishes are frequently ideal candidates for such research.

It is often assumed that anti-predator behaviour should have a significant unlearnedcomponent to it because fishes need to be able to escape predators from the moment theyhatch. The penalty for failure in this instance is death, so there is an expectation thatnatural selection will exert significant evolutionary pressure in this respect. Jennifer Kelleyand Anne Magurran point out in Chapter 3 that while this is the case to some degree,learning still plays a key role in the fine-tuning of predator recognition and responsesystems. In environments that are unpredictable from moment to moment and, perhapsmore importantly, from generation to generation, it is essential that prey species have somegeneral template for predator recognition, but that this template be flexible enough toenable fine-tuning to match the prevailing predatory threats. Kelly and Magurran discussthe various ways in which fishes learn about predators and the need for prey species to be



able to accurately assess potential risks and act accordingly. They cover the evolutionaryarms race between predators and prey highlighting the role learning plays in this race fromboth perspectives.

There are many ways in which prey can learn about predators without high-risk exposure,including the observation of conspecifics as they interact with, or detect, predators. Onesuch method is the reliance on predator odours and prey alarm cues that may be detectedfrom some distance and this is the focus of Chapter 4. Here, Grant Brown, Maud C.O. Ferrariand Douglas P. Chivers explore how fishes use chemical cues both to assess risk and tolearn about predators. There are obviously great fitness advantages to be had by the accurateassessment of risk, primarily because it frees the individual time budget from unnecessaryanti-predator behaviour (Lima & Dill 1990). Fishes not only learn from conspecifics butmay also respond to the alarm signals generated by heterospecifics that are part of the sameprey guild, thus enabling the recognition of predators and dangerous habitats alike. It isinteresting to note that fishes often undergo massive growth from larval to adult stages andin doing so pass through a series of predatory guilds each with its own specific threats. Inthis scenario, Brown, Ferrari and Chivers point out that learning may play a larger role inthe development of anti-predator behaviour than previously suspected.

In Chapter 5, Klaudia Witte and Sabine Nobel explore the role of learning in mate-choice decisions. In their review, Witte and Nobel examine the evidence for the influenceof imprinting during the critical period of early life-history stages on later mate-choicedecisions. They reveal that imprinting is most likely to occur in those species that showsome kind of extended parental care, such as the cichlids. However, it is also evident thatother social influences can also affect mate-choice decisions later in life. For example,naive male guppies can learn to discriminate between conspecifics and heterospecifics andalter their mating strategy to concentrate on courting conspecifics. Part of this alterationin behaviour may be mediated by their mating success and feedback from the femalesthey are attempting to court. Species recognition may be reinforced by learning in thoseareas where multiple closely related species coexist. Whilst mate choice often relies onsome predetermined innate recognition and preference system, Witte and Nobel reveal thatthese unlearned preferences can be overcome by learning and especially by copying themate-choice decisions of others. As discussed in many of the chapters, fishes are capable ofrelying on a mixture of eavesdropping and social information to help them make importantdecisions, and mate choice is no exception. Reliance on public information may enablefemales to gauge the quality and aggression levels of a potential mate without having tosuffer any negative consequences associated with the early stages of courtship.

Yuying Hsu, Ryan L. Earley and Larry L. Wolf examine the modulation of aggressionthrough prior experience in Chapter 6. Many factors combine to influence the outcomeof aggressive encounters, including size, motivation, prior residency and, as Hsu and hiscolleagues highlight, prior experience with fights can also play a large role. The outcome offights can have considerable consequences including access to food, mates or territories, soit is important to understand how experience can influence the outcomes of fights. Recentliterature suggests that fishes that have recently lost a fight are more likely to lose a secondencounter compared to winners, all else being equal. Therefore, an individual’s history mustbe considered when predicting the outcome of a fight at the present time. All of us know thatconfidence can influence our behaviour considerably and this is likely to be mediated both


Fish Cognition and Behaviour 5

through physiological as well as psychological mechanisms. Relying on both modellingand empirical data, Hsu et al. explore how previous experience combines or interacts toshape an individual’s present fighting capability.

Whilst Darwin and his immediate successors described animals as having personalities,this was characterised as anthropomorphic and fell out of favour, as a result of which,until fairly recently, discussions on animal personality have been something of a taboo.Perhaps there was a superficial acceptance that domestic animals such as dogs could havepersonality traits, but fishes? In Chapter 7, Sergey V. Budaev and Culum Brown explore therecent explosion in animal personality literature in which fishes have played a leading role.Owing in part to this fear of anthropomorphism, the literature relating to fish personalityhas been heavily fragmented with the adoption of alternative synonyms such as ‘copingstyle’, ‘behavioural syndrome’ and ‘boldness–shyness continuum’. This chapter representsthe first attempt to bring these streams of research together. The authors examine bothproximate and ultimate explanations of fish personality. Budaev and Brown conclude thatpersonality traits play a neglected role in evolution since individual variation is the breadand butter of natural selection. Personality traits not only are heritable but also have fitnessconsequences. The authors claim that examination of personality traits in fishes requiresa holistic view of behaviour in which multiple traits may be correlated with one anotheracross a range of contexts, and warn against too narrow a view that misses importantrelationships that constrain behavioural evolution.

In Chapter 8, Victoria Braithwaite and Lucy Odling-Smee explore the role of cognitionin spatial orientation, navigation and migration. The authors point out that, like mostanimals, the resources fish utilise are often widely separated in space. Many of thesebiologically important locations are relatively temporally and spatially stable and as suchcan be reliably found by learning and memory retrieval. As Warburton and Hughes pointedout in Chapter 2, here it is also the case that natural selection has favoured learningstrategies to closely match the needs of the species under consideration. Like in all animals,cue reliance is constrained by the species’ perception, and fishes display a huge array ofperceptual capabilities, many of which are only just beginning to be understood, such aselectroreception and UV vision. It is evident that fishes rely on a wide array of navigationcues and mechanisms, ranging from egocentric turns to the formation of cognitive maps, tomove accurately around their environments. Natural selection would favour the ability toselect the most efficient movement pathways possible so as to reduce any potential wasteof time and energy. Thus, accurate navigation is a key component to an individual’s fitnesslandscape. In the final part of their chapter, Braithwaite and Odling-Smee concentrate onlarge-scale migration in salmon as a case study, highlighting both the recall of long-termmemory and initial imprinting processes.

Sian Griffiths and Ashley Ward review the evidence for individual recognition inChapter 9. When closely examining social interactions, it is apparent that not all indi-viduals are treated equally by a given fish. For example, as discussed by Hsu et al. inChapter 9, closely related fishes often receive less aggression than non-relatives. Individualrecognition has several implications on multiple levels, including predicting species dis-persal patterns, which has conservation and fisheries management outcomes. But how dofishes recognise one another? Griffiths and Ward review the ever-increasing body of pub-lications that fishes not only recognise kin, but they can also distinguish between familiar



and unfamiliar individuals. This process seems to build up over 10–14 days although it mayvary from species to species. Being able to recognise, and preferentially associate with kinor familiar individuals, potentially has substantial direct and indirect fitness benefits. Forexample, there is evidence that shoals comprised of familiar individuals show more efficientschooling behaviour than those comprised of strangers. Such benefits may accrue due to anincrease in an individual fish’s ability to predict the response of familiar individuals acrossa variety of contexts. Individual recognition is germane to other aspects of fish behaviour,including cooperation (Chapter 12), exploitation of social cues and signals (MachiavellianIntelligence in Fishes; Chapter 13) and social learning (Chapter 11).

In Chapter 10, Christos Ioannou, Iain Couzin, Richard James, Darren Croft and JensKrause develop mathematical approaches and review current literature that links the be-haviour of individuals to the higher order properties at the group and population levels. It isevident that the behaviour of individuals within a social group is largely influenced by theirfellow group members. Through the rapid transfer of information between group members,shoals of fish often seem to behave as a single collective. However, a few individuals withina group can assert undue influence on the behaviour of the majority, particularly if these‘leaders’ are more motivated to perform some behaviour than the remainder of the shoal(i.e. they are more directed than the average). Such processes may have significant impacton the three-dimensional structure and movement of shoals. Moreover, because informa-tion is shared between group members, a shoal as a whole may be able to solve problemsmore efficiently than singletons (e.g. navigation), for example, by filtering environmentalnoise or collective detection and processing of external cues. In addition, examination ofassociation networks by Ioannou et al. can be utilised to predict the path through whichinformation is likely to be transferred within the group.

The transfer of information between individuals is reliant on social learning processes.Social learning refers to those situations where individuals acquire new information orbehaviour by observation of, or interaction with, others. Social learning can occur acrossa wide variety of contexts and appears to be a ubiquitous form of learning within fishes.Social learning often enables individuals to acquire information more rapidly and efficientlythan would be the case if they themselves had to explore their environment fully and learnvia trial and error. Traditionally, social learning was thought to be restricted to mammalsand birds, but in Chapter 11, Culum Brown and Kevin Laland explore the substantivebody of evidence showing the widespread existence of social learning in fishes. Sociallearning that occurs across generations (vertical or oblique transmission) can lead to theestablishment of localised, stable behavioural traditions that form the very roots of animalculture. Such cultural evolution can operate in tandem with biological evolution and theseprocesses interact in many interesting ways. Brown and Laland argue that social learning islikely to play a key role in the development of fish behaviour and point out that exploitationof such processes could be utilised in training regimes for fisheries and in conservationmanagement programmes such as restocking.

Cooperation between individuals has long been considered something of an enigmawithin evolutionary biology. If Darwinian fitness is all about out-competing others then onemight think all individuals ought to behave selfishly. This notion is central to many existingtheories such as the selfish herd hypothesis which is particularly pertinent to group-livinganimals such as fishes. However, it became clear that the evolution of cooperation could

15 fish and aquatic resources series 15 fish cognition...‘’fish cognition and behavior is an...

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