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Long Calls Mediate Male-Male Competition in Bornean Orangutans

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Long Calls Mediate Male-Male Competition in Bornean Orangutans Long Calls Mediate Male-Male Competition in Bornean Orangutans: an Approach Using Automated Acoustic Localization Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr. sc. nat.) vorgelegt der Mathematisch-naturwissenschaftlichen Fakultät der Universität Zürich von Brigitte Spillmann von Villnachern und Zofingen AG Promotionskommission Prof. Dr. Carel van Schaik (Leitung der Dissertation) Prof. Dr. Marta Manser Dr. Erik P. Willems Dr. Maria A. van Noordwjik Zürich, 2017 To my parents Table of Content THESIS SUMMARY……………………………………………………………………………………………………...7 ZUSAMMENFASSUNG……………………………………………………………………………………………….11 ACKNOWLEDGEMENTS……………………………………………………………………………………………15 GENERAL INTRODUCTION Aims of the thesis...............................................................................................................................18 Communication…………………………………………………………………………………...18 Intra-sexual selection…………………………………………………………………………..20 Orangutans………………………………………………………………………………………….22 Contents of the thesis……………………………………………………………………………………..23 Chapter 1: Validation of an acoustic localization system (ALS)………………24 Chapter 2: Confrontational assessment as an intra-sexual competition mechanism in Bornean orangutans……………………..………………………………..25 Chapter 3: Automated caller recognition procedure……………………………...26 Chapter 4: Long calls as indicators of contest motivation..……………………..27 References……………………………………………………………………………………………………..29 CHAPTER 1: VALIDATION OF AN ACOUSTIC LOCATION SYSTEM TO MONITOR BORNEAN ORANGUTAN (Pongo pygmaeus wurmbii) LONG CALLS Abstract…………………………………………………………………………………………………………36 Introduction…………………………………………………………………………………………………..37 Methods………………………………………………………………………………………………………...40 Study area and recording……………………………………………………………………...40 Long call extraction……………………………………………………………………………...41 Validation of extraction performance……………………………………………………42 Area covered by microphone array……………………………………………………….42 Time of arrival differences……………………………………………………………………43 Localization of long call source……………………………………………………………...44 Validation of acoustic location………………………………………………………………45 Results…………………………………………………………………………………………………………..46 1 Success of recognition model………………………………………………………………..46 Coverage area of ALS……………………………………………………………………………46 Triangulation: Localization of long calls with Sound Finder…………………….47 Discussion……………………………………………………………………………………………………..49 Factors that account for errors in the acoustic location results………………..50 Is the acoustic location system accurate enough? ..................................................51 Acknowledgements………………………………………………………………………………………..52 References……………………………………………………………………………………………………..53 CHAPTER 2: CONFRONTATIONAL ASSESSMENT IN THE ROVING MALE PROMISCUITY MATING SYSTEM OF THE BORNEAN ORANGUTAN Abstract…………………………………………………………………………………………………………58 Introduction…………………………………………………………………………………………………..59 I: Number of males………………………………………………………………………………62 II: Call rate…………………………………………………………………………………………..62 III: Long call response tendency and the proportion of responding males………………………………………………………………………………...63 Methods………………………………………………………………………………………………………...64 Study area…………………………………………………………………………………………...64 Male roving behavior…………………………………………………………………………...65 Data from the acoustic localization system (ALS)………………………………….65 Ecological factors………………………………………………………………………………...65 Number of males in the study area……………………………………………………….66 Average long call rate per male…………………………………………………………….66 Proportion of long call responses and of responders……………………………..66 Number of sexually attractive females………………………………………………….67 Statistical analysis……………………………………………………………………………….67 Results…………………………………………………………………………………………………………..68 Male roving behavior…………………………………………………………………………...68 I: Social and ecological correlates of male presence……………………………...68 II: Average long call rate per male………………………………………………………...69 III: Long call interactions………………………………………………………………….….70 a: Proportion of long call responses………………………………………..….70 2 b: Proportion of responding males……………………………………………..71 Discussion…………………………………………………………………………………………………..…72 Acknowledgements…………………………………………………………………………………..……76 References………………………………………………………………………………………………..…...78 CHAPTER 3: WHO SHALL I SAY IS CALLING? VALIDATION OF A CALLER RECOGNITION PROCEDURE IN BORNEAN FLANGED MALE ORANGUTANS (Pongo pygmaeus wurmbii) LONG CALLS Abstract…………………………………………………………………………………………………………86 Introduction…………………………………………………………………………………………………..87 Methods……………………………………………………………………………………………………...…91 Study area………………………………………………………………………………………...…91 Caller recognition……………………………………………………………………………..…91 High quality recordings……………………………………………………………..91 Recordings from the ALS…………………………………………………………...92 Results………………………………………………………………………………………………………......94 High quality recordings……………………………………………………………..94 Recordings from the ALS…………………………………………………………...95 Discussion……………………………………………………………………………………………………..97 Acknowledgements………………………………………………………………………………………..99 References………………………………………………………………………………………………..….100 CHAPTER 4: LONG CALLS AS INDICATORS OF CONTEST MOTIVATION Abstract……………………………………………………………………………………………………….106 Introduction……………………………………………………………………………………………...…107 Testing confrontational assessment on an individual level with Indices of long call activity…………………………………………………………………110 Is long call production energetically costly?.........................................................110 Impact of poor social knowledge………………………………………………………..110 Does long call activity indicate contest motivation………………………………111 Is fluctuating fighting motivation reflected in long call activity…………….111 3 Methods………………………………………………………………………………………………………112 Study area…………………………………………………………………………………………112 Long call data from focal follows………………………………………………………...112 Long call data from the acoustic localization system (ALS)…………………..112 Identification of ALS callers…………………………………………………………..……113 Resident – Visitor distinction…………………………………………………………..…114 Fruit availability……………………………………………………………………………..…115 Sexually attractive females…………………………………………………………………115 Non-responder versus responder………………………………………………….……115 Elo rating………………………………………………………………………………………..…115 Statistical analysis……………………………………………………………………………..116 Results Individual-level confirmation of confrontational assessment using long call activity…………………………………………………………...……………117 Is long call production energetically costly? ..........................................................118 Impact of poor social knowledge…………………………………………………………119 Does long calling activity indicate fighting motivation? ...................................120 Fluctuation of individuals’ long call activity…………………………………………121 Discussion……………………………………………………………………………………………………123 The importance of using an ALS ………………………………………………………127 Acknowledgements …......…………………………………………………………………………...…129 References…………………………………………………………………………………………………...130 CONCLUSION AND PERSPECTIVES……………………………………………………………………….…137 References…………………………………………………………………………………………………...144 SUPPLEMENTARY MATERIAL Supplementary Material Chapter 1 I Recorders and power supply………………………………………………………...….149 II Study area covered with recorder grid (ALS)………...…………………………151 Supplementary Material Chapter 2 I Empirical validation of the use of ALS as a proxy for male presence..…153 II Latency of long call responses……………………………...…………………….……154 4 Supplementary Material Chapter 4 I Focal follow data……………………………………………………….…………………..…155 II Identification of long calling males from the ALS………………...……………157 III Resident – Visitor distinction…………………..………………………………..……158 CURRICULUM VITAE………………………………………………………………………………………………159 CO-AUTHORED PUBLICATION ABSTRACTS……………………………………………………………..163 5 6 Thesis summary Sexual selection theory explains the occurrence of traits that evolved to function in relation to mating behavior. These traits are divided into intra-sexual selection for traits that are advantageous in competition among same-sex individuals or inter-sexual selection for traits that make an individual more attractive to mating partners. Theory therefore predicts that the distribution of females in space - which serves to maximize access to resources and safety - is crucial for males’ social strategies to get access to fertile mates. The extent to which a male can monopolize mating access to females strongly affects his competitive strategy, along the continuum of scramble and contest competition, with some interesting variations. In this thesis I concentrate on the role of a sexually selected long-distance vocalization, the long call of Bornean flanged male orangutans (Pongo pygmaeus wurmbii) in intra-sexual competition. Orangutans are exceptional among non-human primates because of their semi-solitary social organization. All other primates are gregarious and live in mixed-sex groups, whereas the only other solitary species are nocturnal. Orangutans show a roving-male promiscuous mating system: females live alone in relatively small home ranges compared to males and (flanged) males roam and converge at, and compete for matings with, receptive females. This social system is hard to observe for humans, which has made it hard to develop and test hypotheses about the nature of their intra-sexual competition mechanisms and the way individuals assess their relative fighting ability in respect to other rivals in an area. For this reason, I developed a passive acoustic localization system (ALS) that reveals male presence through their call activity in space
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