The Causes and Consequences of Fission-Fusion Dynamics in Female Northern Long-Eared Bats (Myotis Septentrionalis)
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THE CAUSES AND CONSEQUENCES OF FISSION-FUSION DYNAMICS IN FEMALE NORTHERN LONG-EARED BATS (MYOTIS SEPTENTRIONALIS) by Krista J Patriquin Submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy at Dalhousie University Halifax, Nova Scotia May 2012 © Copyright by Krista J Patriquin, 2012 DALHOUSIE UNIVERSITY DEPARTMENT OF BIOLOGY The undersigned hereby certify that they have read and recommend to the Faculty of Graduate Studies for acceptance a thesis entitled “THE CAUSES AND CONSEQUENCES OF FISSION-FUSION DYNAMICS IN FEMALE NORTHERN LONG-EARED BATS (MYOTIS SEPTENTRIONALIS)” by Krista J Patriquin in partial fulfilment of the requirements for the degree of Doctor of Philosophy. Dated: May 23,2012 External Examiner: _________________________________ Research Supervisor: _________________________________ Examining Committee: _________________________________ _________________________________ Departmental Representative: _________________________________ ii DALHOUSIE UNIVERSITY DATE: May 23, 2012 AUTHOR: Krista J Patriquin TITLE: THE CAUSES AND CONSEQUENCES OF FISSION-FUSION DYNAMICS IN FEMALE NORTHERN LONG-EARED BATS (MYOTIS SEPTENTRIONALIS) DEPARTMENT OR SCHOOL: Department of Biology DEGREE: PhD CONVOCATION: October YEAR: 2012 Permission is herewith granted to Dalhousie University to circulate and to have copied for non-commercial purposes, at its discretion, the above title upon the request of individuals or institutions. I understand that my thesis will be electronically available to the public. The author reserves other publication rights, and neither the thesis nor extensive extracts from it may be printed or otherwise reproduced without the author’s written permission. The author attests that permission has been obtained for the use of any copyrighted material appearing in the thesis (other than the brief excerpts requiring only proper acknowledgement in scholarly writing), and that all such use is clearly acknowledged. _______________________________ Signature of Author iii TABLE OF CONTENTS LIST OF TABLES .......................................................................................................... vii LIST OF FIGURES ..........................................................................................................ix ABSTRACT.......................................................................................................................xi LIST OF ABBREVIATIONS USED............................................................................. xii ACKNOWLEDGEMENTS .......................................................................................... xiii CHAPTER 1 INTRODUCTION……………………………………………………….1 1.1 Rationale....................................................................................................... 1 1.2 Thesis Objectives ......................................................................................... 4 1.3 References..................................................................................................... 5 CHAPTER 2 ROOST-SWITCHING BY FEMALE NORTHERN LONG-EARED BATS IS EXPLAINED BY CHANGES IN AMBIENT CONDITIONS……………...8 2.1 Introduction ................................................................................................. 8 2.2 Methods ...................................................................................................... 11 2.2.1 General Methods........................................................................................... 11 2.2.2 Variation in Roost-use with Year, Reproductive Period, and Ambient Conditions.................................................................................................... 16 2.2.3 Roost fidelity and switching ......................................................................... 19 2.3 Results......................................................................................................... 21 2.3.1 General description of roost characteristics and ambient conditions............ 21 2.3.2 Inter-annual roost-use, roost fidelity and ambient conditions ...................... 22 2.3.3 Intra-annual roost-use, roost-switching and ambient conditions .................. 27 2.3.3.1Reproductive period................................................................................... 27 2.3.3.2 Daily roost-switching................................................................................ 32 2.4 Discussion ................................................................................................... 33 2.5 References................................................................................................... 39 iv CHAPTER 3 DO SOCIAL NETWORKS OF FEMALE NORTHERN LONG- EARED BATS VARY WITH REPRODUCTIVE PERIOD AND AGE?..................48 3.1 Introduction ............................................................................................... 48 3.2 Methods ...................................................................................................... 51 3.2.1 Capture and Marking .................................................................................... 51 3.2.2 Analyses........................................................................................................ 53 3.2.2.1 Associations ............................................................................................. 53 3.2.2.2 Temporal patterns of associations............................................................. 55 3.2.2.3 Network analyses...................................................................................... 56 3.2.2.4 Nature of associations: Effects of reproductive period and age ............... 57 3.3 Results......................................................................................................... 59 3.3.1 Associations ..................................................................................................... 60 3.3.1.1 Temporal pattern....................................................................................... 61 3.3.2 Reproductive Period......................................................................................... 63 3.3.3 Age................................................................................................................... 63 3.4 Discussion ................................................................................................... 69 3.5 References................................................................................................... 74 CHAPTER 4 FEMALE NORTHERN LONG-EARED (MYOTIS SEPTENTRIONALIS) THAT ROOST TOGETHER ARE RELATED……………88 4.1 Introduction ............................................................................................... 88 4.2 Methods ...................................................................................................... 90 4.2.1 Capture and marking..................................................................................... 90 4.2.2 DNA Extraction and genotyping .................................................................. 91 4.2.3 Statistical analyses ........................................................................................ 91 4.3 Results......................................................................................................... 93 4.4 Discussion ................................................................................................. 100 4.5 References................................................................................................. 102 CHAPTER 5 THE POTENTIAL FOR CULTURE IN BATS…………………….109 5.1 Defining Culture ...................................................................................... 109 v 5.1.1 Groups Differ in Socially-Learned Behaviours .......................................... 110 5.1.2 Behaviours are Social ................................................................................. 112 5.1.3 Behaviours are Learned Specifically through Teaching and Imitation ...... 114 5.1.4 Summary..................................................................................................... 115 5.2 Is There Evidence for Culture in Bats?................................................. 115 5.2.1 Groups differ in socially-learned vocalizations that may be normative and symbolic .............................................................................................. 117 5.2.2 Groups differ in socially-learned foraging behaviours that may be normative and symbolic.............................................................................. 120 5.2.3 Groups differ in socially-learned roosting behaviours that may be normative and symbolic.............................................................................. 122 5.2.4 Summary..................................................................................................... 125 5.3 Possible Implications of Bat Culture ..................................................... 126 5.4 Conclusions............................................................................................... 129 5.5 References................................................................................................. 129 CHAPTER 6 CONCLUSIONS……………………………………………………...161 6.1 Causes and Consequences of Fission-Fusion Dynamics....................... 161 LIST OF REFERENCES..............................................................................................164 APPENDIX A PUBLICATIONS..................................................................................207