DRIVING COGNITION AN INVESTIGATION OF THE FACTORS IN DARWIN'S FINCHES. Dissertation. Irmgard Teschke

Transcription

1 AN INVESTIGATION OF THE FACTORS DRIVING COGNITION IN DARWIN'S FINCHES Dissertation zur Erlangung des akademischen Grades des Doktors der Naturwissenschaften (Dr. rer. nat.) an der Universitat Konstanz Mathematisch-Naturwissenschaftliche Sektion Fachbereich Biologie vorgelegt von Irmgard Teschke Tag der miindlichen Priifung: 5. Juli, Referent: Dr. Christoph Kleineidam 2. Referent: Prof. Dr. Martin Wikelski 3. Referent: Prof. Dr. Thomas Bugnyar

2 page TABLE OF CONTENTS LIST OF FIGURES AND TABLES page ix GENERAL INTRODUCTION page J CHAPTER 1 Sometimes Tool-use is Not the Key: No Evidence for Cognitive Adaptive Specializations in Tool-using Woodpecker Finches page 17 CHAPTER 2 Physical Cognition and Tool-use: Performance of Darwin's Finches in the Two-Trap Tube Task page 35 CHAPTER 3 CHAPTER 4 Investigating the relationship between extractive foraging and learning abilities in two species of Darwin's Finches page 49 The Tale of the finch: adaptive radiation and behavioural flexibility page 73 GENERAL DISCUSSION page 91 SUMMARY page 99 ZUSAMMENFASSUNG page 101 ACKNOWLEDGEMENTS page 103 REFERENCES page 105 APPENDICES: APPENDIX 1 History of experimental subjects page 115 APPENDIX 2 Details of methodology and statistical procedures of Chapter 1 page 117 APPENDIX 3 Data and detailed statistical results of Chapter APPENDIX 4 Testing conditions, experiment order and time elapsed between Experiments 1-2 of Chapter 2 page 128 APPENDIX 5 Counterbalanced design of the LNR and P reversal conditions of Chapter 3 page 131

3 viii The factors driving cognition in Darwin's finches APPENDIX 6 Supplementary movies of subjects performing in Experiments 2-3 of Chapter 1 see enclosed CD-rom LIST OF PUBLICATIONS page 132 AUTHOR'S CONTRIBUTIONS/ ABGRENZUNG DER EIGENLEISTUNG page 133 ADDRESSES OF CO-AUTHORS page 134 ERKLARUNG DER SELBSTANDIGEN ANFERTIGUNG page 135 A CD-ROM containing supplementary Movies (Appendix 6) is enclosed along sis. with the the Please note that Chapter 4 has already been published. Therefore, in my thesis this chapter is generally cross-referenced as "Tebbich et al. 2010" whereas all other chapters reference by chapter number. For example, to reference my published chapter, I would write "see Tebbich et al. 2010" whereas for Chapter 1 (unpublished), I would write "see Chapter 1". Chapter 4 is included in its published form with some minor formatting modifications so that it fits the style of this thesis. Furthermore, Figures 1.1. and 1.2 were originally published in Tebbich et al. 2010, but to avoid repetition and to keep the flow of the thesis these figures appear in Chapter 1 and are not included again in Chapter 4.

4 FIGURES AND TABLES GENERAL INTRODUCTION unnumbered Figure Woodpecker finch using a tool to probe into a substrate page 12 CHAPTER 1 Figure 1.1 The seesaw apparatus page22 Figure 1.2 The cane task apparatus.page 23 Figure 1.3 The box opening task apparatus.page 24 Figure 1.4 Logistic regression models of success probability across trial number for two-choice learning tasks (reversal learning, seesaw and cane tasks) -page 28 Figure 1.5 Cane task transfer performance page 30 Table 1.1 Test statistics comparing perseverance by group phases of the seesaw and cane tasks and both phases reversal task for the initial of the page29 Table 1.2 Descriptive statistics quantifying perseverance by group for the initial phases of the seesaw and cane tasks and both phases of the reversal task page 30 CHAPTER 2 Figure 2.1 Schematic illustrations of two-trap tube apparatus -page 41 Figure 2.2 Figure 2.3 Performance of all birds in the initial phase of the modified two-trap tube with pre-inserted stick page 43 Performance of all birds in the initial phase of the modified two-trap tube without pre-inserted stick page 46 CHAPTER 3 Figure 3.1 Schematic illustration of the perseverance (P) and learned non-reward (LNR) reversal concepts -page 67 Figure 3.2 Objects used in measuring novelty reactions -page 68 Figure 3.3 Between-species comparison of LNR and P reversal learning performance -page 62 Figure 3.4 Within-species comparison learning performance of LNR and P reversal.page63

5 X The factors driving cognition in Darwin's finches Figure 3.5 Figure 3.6 in the first and second Comparison of learning performance acquisition learning phase experienced by species -page 64 Comparison of neophobia to a control feeding latency for woodpecker finches and small tree finches PaSe 65 Figure 3.7 Consistency ofneophobicresponses -Pa8e 66 Table 3.1 Table 3.2 Spearman's correlations between (a) neophobia learning and (b) neophilia and reversal learning and reversal in the three reversal conditions page 67 Relationship between the number of positive reinforcements of each of the three reversal in the acquisition phase conditions and neophobia page 68 CHAPTER 4 Table 4.1 Comparison of reversal performance between pigeons, three corvid species and three species of Darwin's finches page 100 APPENDICES Table A1 Subject's experiment participation and history -page 115 Table A3.1 Raw data used in the assessment of the effect of apparatus malfunction occurrences on success in the seesaw task for each bird page 123 Table A3.2 Generalized linear mixed model (binomial error) showing the effect of group, trial number and condition on success probability for the Seesaw and cane tasks Page 124 Table A3.3 Generalized linear mixed model (binomial error) results showing the effect of group and trial number on success probability for both phases ofthereversaltask.page 124 Table A3.4 Post hoc tests for cane, seesaw and reversal task models -page 125 Table A3.5 Summary statistics of point of steepest slope by group for the cane and seesaw tasks and both phases ofthe reversal task -page 125 Table A3.6 Summary statistics of percent errors by group for the cane and seesaw tasks andboth phases of the reversal task page 125 Table A3.7 Statistical results of the comparison of point of steepest slope for the cane and seesaw task and both phases of the reversal task by group page 126 Table A3.8 Statistical results of the comparison of percent errors for the cane and seesaw task and both phases ofthe reversal task by group page 126 Table A4 Details of testing conditions, experiment order and time elapsed between Experiments 1-2 (Chapter 2).page 128

6 Figures and tables xi Table A5 Counterbalanced design of P and LNR conditions (Chapter 3) showing the rewarded (+) and non-rewarded (-) colour stimuli for each acquisition and reversal phase page 131