Acoustic Variability and Its Biological Significance in Nocturnal Lemurs
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Acoustic variability and its biological significance in nocturnal lemurs Von der Naturwissenschaftlichen Fakultät der Gottfried Wilhelm Leibniz Universität Hannover zur Erlangung des Grades Doktorin der Naturwissenschaften Dr. rer. nat. genehmigte Dissertation von Dipl.-Biol. Pia Braune geboren am 06.08.1973 in Peine 2007 Referentin: Prof. Dr. Elke Zimmermann Korreferent: Prof. Dr. Uwe Jürgens Tag der Promotion: 12.09.2007 Contents 1 Contents 1 Abstract............................................................................................................................ 3 2 Zusammenfassung .......................................................................................................... 4 3 General introduction ...................................................................................................... 5 3.1 Animal communication........................................................................................... 5 3.2 Acoustic variability on the inter- and intra-species level........................................ 6 3.3 Malagasy lemurs...................................................................................................... 8 3.4 Model species of nocturnal lemurs.......................................................................... 9 3.5 Intra-specific variation in acoustic communication of two species of nocturnal lemurs .................................................................................................................... 11 3.6 Inter-specific variation and species-specificity in acoustic communication of mouse lemurs......................................................................................................... 12 3.7 Aim of the study.................................................................................................... 14 4 Study 1 - Spacing and group coordination in a nocturnal primate, the golden brown mouse lemur ( Microcebus ravelobensis ): the role of olfactory and acoustic signals............................................................................................................................. 16 4.1 Introduction ........................................................................................................... 17 4.2 Methods................................................................................................................. 18 4.2.1 Study site and data sampling................................................................... 18 4.2.2 Marking behaviour.................................................................................. 21 4.2.3 Sound recording and analysis.................................................................. 21 4.3 Results ................................................................................................................... 24 4.3.1 Spacing.................................................................................................... 24 4.3.2 Behaviour during dispersal and reunion ................................................. 25 4.3.3 Marking behaviour.................................................................................. 25 4.3.4 Vocal behaviour ...................................................................................... 26 4.3.5 Context of acoustic signals...................................................................... 26 4.3.6 Trill structure........................................................................................... 27 4.3.7 Group specific signatures of trills ........................................................... 28 4.4 Discussion ............................................................................................................. 29 4.4.1 Spacing.................................................................................................... 29 4.4.2 Marking behaviour.................................................................................. 29 4.4.3 Vocal behaviour ...................................................................................... 30 4.4.4 Group-specific acoustic signatures ......................................................... 31 4.5 Conclusion............................................................................................................. 32 5 Study 2 - Loud calling, spacing, and cohesiveness in a nocturnal primate, the Milne Edwards' sportive lemur ( Lepilemur edwardsi )............................................... 33 5.1 Introduction ........................................................................................................... 34 5.2 Methods................................................................................................................. 35 5.2.1 Study site and animals............................................................................. 35 5.2.2 Data collection and processing................................................................ 36 5.2.3 Statistical analysis................................................................................... 39 2 Contents 5.3 Results ................................................................................................................... 40 5.3.1 Sleeping associations .............................................................................. 40 5.3.2 Home ranges and overlap........................................................................ 40 5.3.3 Loud calling behaviour ........................................................................... 41 5.3.4 Sex and individual identity in loud calls................................................. 42 5.4 Discussion ............................................................................................................. 47 5.4.1 Spacing and cohesion within and between male-female pairs................ 47 5.4.2 Functions of loud calling......................................................................... 48 5.4.3 Loud calling between pair partners, and its evolution among primates.. 49 5.5 Conclusions ........................................................................................................... 50 6 Study 3 - Specific acoustic divergence in communication of cryptic species of nocturnal mammals: social calls and their perception in Malagasy primates (Microcebus ssp.) ........................................................................................................... 51 6.1 Introduction ........................................................................................................... 52 6.2 Methods................................................................................................................. 53 6.2.1 Recording and analysis of advertisement calls ....................................... 53 6.2.2 Playback experiments.............................................................................. 54 6.3 Results ................................................................................................................... 57 6.3.1 Interspecific comparison of advertisement calls ..................................... 57 6.3.2 Behavioural responses to advertisement and short whistle stimuli......... 59 6.4 Discussion ............................................................................................................. 61 6.4.1 Species-specific structure in advertisement calls.................................... 62 6.4.2 Species-specific call recognition............................................................. 62 7 General discussion ........................................................................................................ 64 7.1 Factors influencing the acoustic variability on the inter-and intra-species level .. 64 7.1.1 Inter-specific acoustic variability............................................................ 65 7.1.2 Inter- and intra-specific acoustic variability............................................ 66 7.1.3 Intra-specific acoustic variability............................................................ 68 7.2 Intra-specific acoustic variability in mouse and sportive lemurs.......................... 70 7.3 Inter-specific acoustic variability in cryptic mouse lemurs and its biological significance............................................................................................................ 72 7.3.1 The effect of sympatry and allopatry ...................................................... 74 7.3.2 Evolutionary aspects of inter-species call variability in mouse lemurs .. 75 7.4 Inter-specific comparison of the acoustic variability between mouse and sportive lemurs ...................................................................................................... 76 7.4.1 Biological function of loud calls – the effect of social organisation....... 78 7.5 Evolution of long distance calls in primates ......................................................... 80 7.6 Concluding remarks .............................................................................................. 82 8 References...................................................................................................................... 83 9 Acknowledgements ....................................................................................................... 98 10 Professional development........................................................................................... 100 Abstract 3 1 Abstract The aim of this thesis was to examine the acoustic variability and its biological significance in nocturnal lemurs from an evolutionary