Tinbergian Practice, themes and variations: the field and laboratory methods and practice of the Animal Behaviour Research Group under Nikolaas Tinbergen at Oxford University. Graeme Robert Beale Ph.D. Thesis The University of Edinburgh, 2008 I confirm that the content of this thesis is entirely my own work, and that all sources, quotations and pictures have been acknowledged and referenced. (Graeme Beale) 26th April 2009. Graeme Beale, 05/04/09 Table of Contents Acknowledgements.......................................................................................................2 Abstract.........................................................................................................................4 Introduction to the thesis...............................................................................................5 Section 1: Tinbergen, man, and animal observer..................................................... 7 Section 1.1 Tinbergen's pre–Oxford Studies....................................................... 8 Section 2: Notes on reading Tinbergen’s fieldnotes...............................................14 Section 3: Selected literature review......................................................................16 Section 3.1 Burkhardt and Ethology from Evolution to Ecologies...................17 Section 3.2 Laboratory and Field...................................................................... 19 Section 3.3 Research Schools and Ethology..................................................... 21 Section 3.4 Ethology and Anthropomorphism ................................................25 Section 4: Outline of this Thesis............................................................................ 26 Chapter One: Tinbergen and the background to his Ethology....................................29 Introduction............................................................................................................ 29 Section 1: Tinbergen's Route to Ethology..............................................................35 Section 2: Dissecting the field, Tinbergen's ideas of field methods.......................39 Section 2.1 Tinbergen's Writing on Method...................................................... 40 Section 2.2 The Importance of the Comparative Method and the Comparative Morphology heritage......................................................................................... 45 Section 2.3 The development of ethograms as fundamental objects of comparative study..............................................................................................50 Section 3: “As the Gulls might say”...................................................................... 53 Section 4: The Tale of Casanova............................................................................63 Section 4.1 Social Drama.................................................................................. 66 Section 4.2 Ascribing Character to birds ..........................................................68 Conclusion: Tinbergen's own Ethology................................................................. 73 Chapter Two: Tinbergian early years: the field...........................................................75 Introduction............................................................................................................ 75 Section 1: “Ethologists are Scientists who love the animals they use in their research”.................................................................................................................76 Section 2: Two of Tinbergen's earliest students, Hinde and Moynihan ................84 Section 2.1 Hinde's way out of the woods.........................................................85 Section 2.2 Moynihan's path through the dunes................................................90 Conclusion: From the field to the Ethogram........................................................101 Chapter Three: The Early Tinbergen Laboratory......................................................105 Introduction.......................................................................................................... 105 Section 1: Queer Fish, Vibrating Fruit flies, and Bored Pilchards.......................106 Section 1.1 Desmond Morris, Sticklebacks and Ethograms.......................107 Section 1.2 Margaret Bastock's Drosophila linking behaviour and genetics .....................................................................................................................117 Section 1.3 Mike Cullen bringing pilchards to the laboratory....................126 Section 2: Exchange between laboratory and field.............................................. 132 Section 2.1 Naturalising the Laboratory in practice and observation style 132 Section 2.2 Images and Ideas moving between laboratory and field..........137 Conclusion: Watching Nature in the Laboratory..................................................146 Chapter Four: The later Tinbergians in laboratory and field....................................149 Introduction: In search of the adaptationists........................................................ 149 Section 1: Watching nature in the later field school ............................................150 Section 1.1 Esther Cullen and adaptation to the cliffs.....................................150 Section 1.2 Colin Beer's egg work ................................................................. 158 Section 1.3 Kruuk and Tinbergen's egg–shell work........................................ 162 Section 1.4 Blurton–Jones returns to old territory...........................................167 Section 1.5 Heather McLannahan and the return to chick studies..................171 Section 2: Watching Nature in the later laboratory ............................................. 175 Section 2.1 Morris, laboratory studies of bird behaviour, and serendipity,.....176 Section 2.2 Dawkins and the domestic chick.................................................. 185 Section 2.3 Delius and the Vivisectionist turn.................................................190 Conclusion ...........................................................................................................197 Chapter Five: The Tinbergians and the Public..........................................................201 Introduction.......................................................................................................... 201 Section 1: Tinbergian Public Communication of their Animal Behaviour Research ..............................................................................................................................203 Section 1.1 Field description as it appeared in the academic literature...........203 Section 1.2 Contrasting the published academic accounts with the fieldnotes ......................................................................................................................... 206 Section 1.3 Contrasting the popular science with the academic output..........211 Section 1.4 Contrasting the field and the popular science accounts................213 Section 1.5 Field life described in fieldnotes and popular science..................216 Section 2: Tinbergen's motivations to write for the public...................................219 Section 2.1 Teaching the ornithological world to observe as he did ..............222 Section 2.2 Justifying ethology....................................................................... 225 Section 3: Science of the People, for the People..................................................228 Section 3.1 Humans as a part of the animal world.......................................... 229 Section 3.2 Tinbergen's writing on homo sapiens at the species level............234 Section 3.3 Tinbergen on the individual human predicament.........................241 Conclusions.......................................................................................................... 248 Conclusions...............................................................................................................251 The argument chapter by chapter......................................................................... 252 Closing argument................................................................................................. 257 Bibliography............................................................................................................. 259 Unpublished, Archival and Interview Material....................................................259 Bibliography of Published sources.......................................................................260 Appendix A: Tinbergen's Students studied in the thesis........................................... 279 Chapter Two.................................................................................................... 279 Chapter Three.................................................................................................. 280 Chapter Four....................................................................................................280 Appendix B: Tinbergen's Students a (nearly) Complete List ...................................283 Tinbergen's Dutch Students..................................................................................283 Tinbergen's Oxford Students................................................................................283 Acknowledgements This thesis was made possible by an ESRC 1+3 grant, which covered both