The Tinbergen Legacy. M.S. DAWKINS Research Lecturer Oxford University T.R. HALLIDAY R. DAWKINS. Edited by. Reader in Biology Open University.

Transcription

1 The Tinbergen Legacy

2 The Tinbergen Legacy Edited by M.S. DAWKINS Research Lecturer Oxford University T.R. HALLIDAY Reader in Biology Open University and R. DAWKINS Reader Oxford University CHAPMAN & HALL London New York Tokyo Melbourne Madras

3 Published by Chapman & Hall, 2-6 Boundary Row, London SE1 8HN Chapman & Hall, 2-6 Boundary Row, London SE1 8HN, UK Chapman & Hall, 29 West 35th Street, New York NY10001, USA Chapman & Hall Japan, Thomson Publishing Japan, Hirakawacho Nemoto Building, 7F, Hirakawa-cho, Chiyoda-ku, Tokyo 102, Japan Chapman & Hall Australia, Thomas Nelson Australia, 102 Dodds Street, South Melbourne, Victoria 3205, Australia Chapman & Hall India, R. Seshadri, 32 Second Main Road, CIT East, Madras , India First edition Chapman & Hall Typeset in 101/2 on 12pt Palatino by Mews Photosetting, Beckenham, Kent Printed in Great Britain by St Edmundsbury Press, Bury St Edmunds ISBN Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. 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. A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication data available Printed on permanent acid-free text paper, manufactured in accordance with the proposed ANSI/NISO Z X and ANSI Z

4 Contents Contributors Introduction Richard Dawkins 1. Early ethology: growing from Dutch roots Gerard P. Baerends 2. Study behavioural adaptations Nicholas B. Davies 3. From animals to humans Robert A. Hinde 4. War and peace revisited Felicity A. Huntingford 5. Animal communication: ideas derived from Tinbergen's activities John R. Krebs 6. The nature of culture Juan D. Delius 7. Niko Tinbergen, comparative studies and evolution Michael H. Robinson 8. The Tinbergen legacy in photography and film Lary Shaffer Afterword Aubrey Manning Index vii ix

5 Contributors Gerard P. Baerends Hoofdweg 265, 9765 CH Paterswolde, Netherlands Nicholas B. Davies Department of Zoology, University of Cambridge, UK Richard Dawkins Department of Zoology, University of Oxford, UK Marian S. Dawkins Department of Zoology, University of Oxford, UK Juan D. Delius Universitfit Konstanz, Konstanz, Germany Timothy R. Halliday The Open University, Milton Keynes, UK Robert A. Hinde Department of Zoology, University of Cambridge, UK Felicity A. Huntingford Department of Zoology, University of Glasgow, UK John R. Krebs Department of Zoology, University of Oxford, UK Aubrey Manning Department of Zoology, University of Edinburgh, UK Michael H. Robinson Smithsonian Institute, Washington DC, USA Lary Shaffer Department of Psychology, SUNY, Plattsburgh, USA

6 Introduction RICHARD DAWKINS A conference with the title 'The Tinbergen Legacy' was held in Oxford on 20th March, Over 120 of Niko Tinbergen's friends, family, colleagues, former students and people who had never met him in person converged at Oxford for what turned out to be a memorable day. To reflect the rather special atmosphere of the conference, we decided to begin this book with Richard Dawkins' opening remarks exactly as he gave them on that day. Welcome to Oxford. For many of you it is welcome back to Oxford. Perhaps even, for some of you, it would be nice to think that it might feel like welcome home to Oxford. And it is a great pleasure to welcome so many friends from the Netherlands. Last week, when everything had been settled except final, last minute arrangements, we heard that Lies Tinbergen had died. Obviously we would not have chosen such a time to have this meeting. I'm sure we'd all like to extend our deep sympathy to the family, many of whom, I'm happy to say, are at this meeting. We discussed what we should do and decided that, in the circumstances, there was nothing for it but to carry on. The members of the Tinbergen family that we were able to consult were fully in agreement. I think we all knew that Lies was an enormous support to Niko, but I think that very few of us really knew how much of a support she was to him, particularly during the dark times of depression. I should say something about this memorial conference and what led up to it. People have their own ways of grieving. Lies' way was to take literally Niko's characteristically modest instruction that he wanted no funeral or memorial rites of any kind. There were those

7 x Introduction of us who were fully sympathetic to the desire for no religious observance, but who nevertheless felt the need for some kind of rite of passage for a man whom we had loved and respected for so many years. We suggested various kinds of secular observance. For instance, the fact that there was such musical talent in the Tinbergen family led some of us to suggest a memorial chamber concert with readings or eulogies in the intervals. Lies made it very clear, however, that she wanted nothing of the kind and that Niko would have felt the same. So we did nothing for a while. Then, after some time had elapsed, we realised that a memorial conference would be sufficiently different from a funeral as not to count. Lies accepted this, and there came a time, during our planning of the conference, when she said that she hoped to attend the conference, although she later changed her mind about that, thinking, again with characteristic modesty and completely erroneously, that she would have been in the way. It is an enormous pleasure to welcome so many old friends. It is a tribute to Niko, and the affection that his old pupils felt for him, that so many of you are here today, converging on Oxford from, in some cases, very far away. The list of people coming is a galaxy of old friends, some of whom may not have set eyes on one another for 30 years. Just reading the guest list was a moving experience for me. We shall all of us have memories of Niko and of the group of his associates with whom we happen to be contemporary. My own begin when I was an undergraduate and he lectured to us, not at first on animal behaviour but on molluscs - for it was Alister Hardy's quaint idea that all the lecturers should participate in the 'Animal Kingdom' course which is one of the sacred cows of Oxford zoology. I didn't know, then, what a distinguished man Niko was. I think that if I had, I'd have been rather aghast at his being made to lecture on molluscs. It was bad enough that he gave up being a Professor in Leiden to become, by Oxford's snobbish custom, just plain 'Mr Tinbergen'. I don't remember much from those early mollusc lectures, but I do remember responding to his wonderful smile: friendly, kindly, avuncular as I thought then, although he must have been scarcely older than I am now. I think I must have been imprinted on Niko and his intellectual system then, for I asked my college tutor if I could have tutorials with Niko. I don't know how he managed to swing it, because I don't think Niko gave undergraduate tutorials as a rule. I suspect that I may have been the last undergraduate to have had tutorials with him. Those tutorials had an enormous influence on me. Niko's style as a tutor was unique. Instead of giving a reading list with some sort of comprehensive coverage of a topic, he would give a single, highly detailed

8 Introduction xi piece of work, such as a DPhil thesis. My first one, I remember, was a monograph by A.C. Perdeck, who I am happy to say is here today. I was asked simply to write an essay on anything that occurred to me as a result of reading the thesis or monograph. In a sense it was Niko's way of making the pupil feel like an equal - a colleague whose views on research were worth hearing, not just a student mugging up a topic. Nothing like this had ever happened to me before, and I revelled in it. I wrote huge essays that took so long to read out that, what with Niko's frequent interruptions, they were seldom finished by the end of the hour. He strode up and down the room while I read my essay, only occasionally coming to rest on whatever old packing case was serving him as a chair at the time, chain-rolling cigarettes and obviously giving me his whole attention in a way that, I'm sorry to say, I cannot claim to do for most of my pupils today. As a result of these marvellous tutorials, I decided that I very much wanted to do a DPhil with Niko. And so I joined the 'Maestro's Mob', and it was an experience never to be forgotten. I remember with particular affection the Friday evening seminars. Apart from Niko himself, the dominant figure at that time was Mike Cullen. Niko obstinately refused to let sloppy language pass, and proceedings could be stalled for an indefinite period if the speaker was not able to define his terms with sufficient rigour. These were arguments in which everybody became engaged, eager to make a contribution. If, as a result, a seminar wasn't finished at the end of the two hours, it simply resumed the following week, no matter what might have been previously planned. I suppose it may have been just the na'fvet6 of youth, but I used to look forward to those seminars with a sort of warm glow for the whole week. We felt ourselves members of a privileged 61ite, an Athens of ethology. Others, who belonged to different cohorts, different vintages, have talked in such similar terms that I believe that this feeling was a general aspect of what Niko did for his young associates. In a way, what Niko stood for on those Friday evenings was a kind of ultra-rigorous, logical commonsense. Put like that, it may not sound like much; it may seem even obvious. But I have since learned that rigorous commonsense is by no means obvious to much of the world. Indeed commonsense sometimes requires ceaseless vigilance in its defence. In the world of ethology at large, Niko stood for breadth of vision. He not only formulated the 'four questions' view of biology, he also assiduously championed anyone of the four that he felt was being neglected. Since he is now associated in peoples' minds with field studies of the functional significance of behaviour, it is worth

9 xii Introduction recalling how much of his career was given over to, for instance, the study of motivation. And, for what it is worth, my own dominant recollection of his undergraduate lectures on animal behaviour was of his ruthlessly mechanistic attitude to animal behaviour and the machinery that underlay it. I was particularly taken with two phrases of his - "behaviour machinery", and "equipment for survival". When I came to write my own first book, I combined them into the brief phrase "survival machine". In planning this conference, we obviously decided to concentrate on fields that Niko had been pre-eminent in, but we didn't want the talks to be only retrospective. Of course we wanted to spend some time looking back at Niko's achievements, but we also wanted people to pick up the torches that Niko had passed them, and run on with them towards the future. Torch-running behaviour, in new and exciting directions, bulks so large in the ethograms of Niko's students and associates that planning the programme was a major headache. "How on earth", we asked ourselves, "can we possibly leave out so-and-so? On the other hand, we have space for only six talks". We could have limited ourselves to Niko's own pupils - his scientific children, but this would have been to devalue his enormous influence via grandpupils and others. We could have concentrated on people and major areas not covered in the Festschrift volume edited by Gerard Baerends, Colin Beer and Aubrey Manning, but that too would have been a pity. In the end, it seemed almost not to matter which half a dozen of Niko's intellectual descendants stood up to represent the rest of us. And perhaps that is the true measure of his greatness.