Learn and Let Learn: Comparative Cognition and Social Learning in Bats Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften vorgelegt von Theresa Maria Anna Clarin an der Mathematisch-Naturwissenschaftliche Sektion Fachbereich Biologie Tag der mündlichen Prüfung: 16.07.2014 1. Referent: Prof. Dr. Martin Wikelski 2. Referent: Assistant Prof. Dr. John Ratcliffe for PD Dr. Björn M. Siemers Die geistigen Fähigkeiten der Flatterthiere sind keineswegs so gering, als man gern annehmen möchte, und strafen den auf ziemliche Geistesarmuth hindeutenden Gesichtsausdruck Lügen. [...] Alle Flatterthiere zeichnen sich durch einen ziemlich hohen Grad von Gedächtnis und einige sogar durch verständige Ueberlegung aus. (Brehm , A.E. 1864 . Illustrirtes Thierleben: eine allgemeine Kunde des Thierreichs ). The intellectual abilities of the bats are by no means as low as one would think and give the lie to the expression of their faces, which is indicative of a certain poverty of mind. […] All bats stand out by a rather high degree of memory and some even by the presence of reasoning powers. TABLE OF CONTENTS SUMMARY ...................................................................................................................................................... 1 ZUSAMMENFASSUNG (GERMAN SUMMARY) ................................................................................................. 3 GENERAL INTRODUCTION ............................................................................................................................... 5 ABOUT THE STUDY OF COGNITION IN ANIMALS .............................................................................................................. 6 COMPARATIVE RESEARCH ON COGNITION ..................................................................................................................... 6 THE EVOLUTION OF COGNITION : THEORIES ................................................................................................................... 7 BATS AS STUDY SPECIES ............................................................................................................................................ 9 BULGARIA AS A STUDY SITE ...................................................................................................................................... 14 THESIS OUTLOOK ................................................................................................................................................... 16 CHAPTER 1 (FORAGING ECOLOGY PREDICTS LEARNING PERFORMANCE IN INSECTIVOROUS BATS) ............... 17 MATERIALS AND METHODS ..................................................................................................................................... 21 RESULTS .............................................................................................................................................................. 25 DISCUSSION .......................................................................................................................................................... 28 SUPPORTING INFORMATION ..................................................................................................................................... 32 ACKNOWLEDGEMENTS ............................................................................................................................................ 32 CHAPTER 2 (SOCIAL LEARNING WITHIN AND ACROSS SPECIES: INFORMATION TRANSFER IN MOURSE-EARED BATS) ........................................................................................................................................................... 35 MATERIALS AND METHODS ..................................................................................................................................... 37 RESULTS .............................................................................................................................................................. 44 DISCUSSION .......................................................................................................................................................... 48 ACKNOWLEDGEMENTS ............................................................................................................................................ 52 CHAPTER 3 (DO BATS EXPERIENCE A TRADE-OFF BETWEEN ENERGY CONSERVATION AND LEARNING?) ....... 53 MATERIALS AND METHODS ..................................................................................................................................... 55 RESULTS .............................................................................................................................................................. 59 DISCUSSION .......................................................................................................................................................... 61 ACKNOWLEDGEMENTS ............................................................................................................................................ 63 CHAPTER 4 (EFFECT OF DIFFERENCT LIGHT CONDITION ON MYOTIS MYOTIS FORAGING BEHAVIOR) ............ 65 MATERIALS AND METHODS ..................................................................................................................................... 67 RESULTS .............................................................................................................................................................. 73 DISCUSSION .......................................................................................................................................................... 78 GENERAL DISCUSSION .................................................................................................................................. 81 MAIN FINDINGS ..................................................................................................................................................... 82 IMPLICATIONS ....................................................................................................................................................... 84 FUTURE DIRECTIONS .............................................................................................................................................. 86 ACKNOWLEDGEMENTS ................................................................................................................................. 89 REFERENCES ................................................................................................................................................. 91 LIST OF FIGURES ......................................................................................................................................... 109 LIST OF TABLES ........................................................................................................................................... 111 AUTHOR CONTRIBUTIONS .......................................................................................................................... 113 CURRICULUM VITAE .................................................................................................................................... 115 LIST OF PUBLICATIONS ................................................................................................................................ 117 PART OF THIS THESIS ............................................................................................................................................. 117 NOT PART OF THIS THESIS ...................................................................................................................................... 117 LIST OF CONFERENCE CONTRIBUTIONS ....................................................................................................... 119 ORAL PRESENTATIONS .......................................................................................................................................... 119 POSTER .............................................................................................................................................................. 119 CONFERENCE PARTICIPATION WITHOUT CONTRIBUTION .............................................................................................. 119 PARTICIPATION IN THE IMPRS-OB CURRICULUM ......................................................................................... 121 COURSES ............................................................................................................................................................ 121 OTHER IMPRS-OB EVENTS ................................................................................................................................... 121 EXTRA-CURRICULAR OUTREACH AND OTHER ACTIVITIES ............................................................................. 123 A CD containing the supplementary movie to Chapter 1 (S1) is enclosed along with the thesis. The movie is also available at: http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0064823#s5 SUMMARY In this thesis, I present work on cognitive aspects of bat behavior. In my first study (Chapter 1: “Foraging ecology predicts learning performance in insectivorous bats”), I investigated the relationship between foraging ecology and cognitive performance in insectivorous European bats. I found that learning performance and behavioral flexibility in a foraging task can be predicted by