Unicentre CH-1015 Lausanne http://serval.unil.ch Year : 2012 PERSONALITY AND MELANIN-BASED COLOURATION TRAITS IN BARN OWLS, TAWNY OWLS AND KESTRELS VAN DEN BRINK Valentijn VAN DEN BRINK Valentijn, 2012, PERSONALITY AND MELANIN-BASED COLOURATION TRAITS IN BARN OWLS, TAWNY OWLS AND KESTRELS Originally published at : Thesis, University of Lausanne Posted at the University of Lausanne Open Archive. http://serval.unil.ch Droits d’auteur L'Université de Lausanne attire expressément l'attention des utilisateurs sur le fait que tous les documents publiés dans l'Archive SERVAL sont protégés par le droit d'auteur, conformément à la loi fédérale sur le droit d'auteur et les droits voisins (LDA). A ce titre, il est indispensable d'obtenir le consentement préalable de l'auteur et/ou de l’éditeur avant toute utilisation d'une oeuvre ou d'une partie d'une oeuvre ne relevant pas d'une utilisation à des fins personnelles au sens de la LDA (art. 19, al. 1 lettre a). 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Departement d’Ecologie et d’Evolution PERSONALITY AND MELANIN-BASED COLOURATION TRAITS IN BARN OWLS, TAWNY OWLS AND KESTRELS Thèse de doctorat ès sciences de la vie (PhD) présentée à la Faculté de biologie et de médecine de l’Université de Lausanne par Valentijn VAN DEN BRINK Master en Biologie de l’Université de Groningen, Pays-Bas Jury Prof. Alexandre Reymond, Président Prof. Alexandre Roulin, Directeur de thèse Prof. Niels J. Dingemanse, expert Prof. Julien Gasparini, expert Lausanne 2012 Departement d’Ecologie et d’Evolution PERSONALITY AND MELANIN-BASED COLOURATION TRAITS IN BARN OWLS, TAWNY OWLS AND KESTRELS Thèse de doctorat ès sciences de la vie (PhD) présentée à la Faculté de biologie et de médecine de l’Université de Lausanne par Valentijn VAN DEN BRINK Master en Biologie de l’Université de Groningen, Pays-Bas Jury Prof. Alexandre Reymond, Président Prof. Alexandre Roulin, Directeur de thèse Prof. Niels J. Dingemanse, expert Prof. Julien Gasparini, expert Lausanne 2012 Voor Anneke Contents Contents .................................................................................................................................................. 4 Résumé .................................................................................................................................................. 11 Summary ............................................................................................................................................... 13 Acknowledgements ............................................................................................................................... 15 Introduction ........................................................................................................................................... 17 The development of polymorphism .................................................................................................. 17 Maintenance of polymorphism ......................................................................................................... 17 Adaptive functions of polymorphism ................................................................................................ 18 Colour polymorphism ........................................................................................................................ 19 Sources for colouration ..................................................................................................................... 20 Carotenoids ................................................................................................................................... 20 Melanin .......................................................................................................................................... 21 Polymorphism in melanin-based colouration ................................................................................... 22 Pleiotropy in the melanocortin system ............................................................................................. 22 Box 1. Structure, mechanisms, and functions of the melanocortin system ......................................... 24 Structure and mechanism of pigmentation ...................................................................................... 24 MC1R colour production ................................................................................................................... 24 Other functions ................................................................................................................................. 25 Individual behavioural traits or personalities.................................................................................... 23 Box 2 Animal personality ....................................................................................................................... 27 Evolution of animal personality research .......................................................................................... 27 Concepts, definitions and terminology ............................................................................................. 28 Behavioural “axes” ............................................................................................................................ 28 Underlying mechanisms .................................................................................................................... 29 Future challenges .............................................................................................................................. 29 Need for investigating underlying mechanisms ................................................................................ 30 Melanic colouration in relation to individual behaviour ................................................................... 30 Box 3 –Study species ............................................................................................................................. 31 1. The barn owl (Tyto alba) ............................................................................................................... 31 Appearance and colouration ......................................................................................................... 31 Population and range .................................................................................................................... 31 4 Feeding habits ............................................................................................................................... 32 Reproduction ................................................................................................................................. 32 Life span ......................................................................................................................................... 32 Study population ........................................................................................................................... 33 2. The Eurasian kestrel (Falco tinnunculus) ....................................................................................... 33 Appearance and Colouration ........................................................................................................ 33 Population and range .................................................................................................................... 34 Feeding habits ............................................................................................................................... 34 Reproduction ................................................................................................................................. 34 Life span ......................................................................................................................................... 35 Study population ........................................................................................................................... 35 3. The tawny owl (Strix aluco) ........................................................................................................... 35 Appearance and colouration ......................................................................................................... 35 Population and range .................................................................................................................... 36 Feeding habits ............................................................................................................................... 36 Reproduction ................................................................................................................................. 36 Life span ......................................................................................................................................... 37 Study population ........................................................................................................................... 37 Predictions about relationship between colouration and behaviour ............................................... 39 Thesis outline .................................................................................................................................... 40 1 Melanic color-dependent antipredator behavior strategies in barn owl nestlings ...................... 43 Abstract ............................................................................................................................................. 44 Introduction ....................................................................................................................................... 45 Methods ............................................................................................................................................ 48 Study species ................................................................................................................................. 48 General methods ........................................................................................................................... 49 Hissing behavior ............................................................................................................................ 49 Tonic immobility ............................................................................................................................ 51 Breathing rate ................................................................................................................................ 52 Docility ........................................................................................................................................... 52 Covariation of the measured behavioral traits ............................................................................. 53 Statistical methods ........................................................................................................................ 53 Results ............................................................................................................................................... 54 5 Hissing behavior ............................................................................................................................ 54 Tonic immobility ............................................................................................................................ 55 Breathing rate ................................................................................................................................ 57 Docility ........................................................................................................................................... 57 Covariation between behavioral antipredator traits .................................................................... 58 Discussion .......................................................................................................................................... 58 Boldness-shyness ........................................................................................................................... 59 Consistency of behavior ................................................................................................................ 60 Melanin-based coloration and stress ............................................................................................ 61 Selection pressure on stress response .......................................................................................... 62 Acknowledgements ........................................................................................................................... 62 2. Melanin-based colouration in juvenile kestrels (Falco tinnunculus) covaries with anti-predatory personality traits ................................................................................................................................... 63 Abstract ............................................................................................................................................. 64 Introduction ....................................................................................................................................... 65 Methods ............................................................................................................................................ 67 Study species and study area ........................................................................................................ 67 Body measurements and assessment of plumage traits .............................................................. 68 Determination of melanin content from feathers ........................................................................ 69 Tonic Immobility Test .................................................................................................................... 70 Breathing rate ................................................................................................................................ 71 Docility ........................................................................................................................................... 71 Stick grabbing ................................................................................................................................ 72 Lifting by claws .............................................................................................................................. 72 Statistical procedures .................................................................................................................... 72 Results ............................................................................................................................................... 75 Melanin content ............................................................................................................................ 75 Main experiment in 2009 .............................................................................................................. 75 Minor experiment in 2010............................................................................................................. 77 Discussion .......................................................................................................................................... 77 Behaviour and feather melanin content ....................................................................................... 78 Behavioural syndromes ................................................................................................................. 78 Differences between results obtained in 2009 and 2010 ............................................................. 79 Similarity and differences between kestrels and other species .................................................... 80 6 Acknowledgements ........................................................................................................................... 81 3. Melanin-based colouration predicts natal dispersal in the barn owl Tyto alba ........................... 83 Abstract ............................................................................................................................................. 84 Introduction ....................................................................................................................................... 85 Methods ............................................................................................................................................ 87 Study species and study area ........................................................................................................ 87 Cross-fostering .............................................................................................................................. 88 Assessment of colouration ............................................................................................................ 89 Individual traits .............................................................................................................................. 89 Measuring dispersal distance ........................................................................................................ 90 Breeder density and dispersal distances ....................................................................................... 91 Natal dispersal of parents and their offspring and individual dispersal tendency ........................ 91 Habitat characteristics ................................................................................................................... 92 Statistical methods ........................................................................................................................ 92 Results ............................................................................................................................................... 95 Natal dispersal ............................................................................................................................... 95 Comparison of natal dispersal distances of parents and offspring ............................................... 96 Breeding dispersal ......................................................................................................................... 97 Within-individual repeatability in dispersal behaviour ................................................................. 99 Dispersal distance and reproductive success ................................................................................ 99 Discussion .......................................................................................................................................... 99 Potential mechanisms explaining colour-specific dispersal ........................................................ 101 Acknowledgements ......................................................................................................................... 103 4. Pheomelanin-based colouration signals nest defence behaviour in tawny owls (Strix aluco) ... 105 Abstract ........................................................................................................................................... 106 Introduction ..................................................................................................................................... 107 Methods .......................................................................................................................................... 110 Study species and site ................................................................................................................. 110 Colour measurements ................................................................................................................. 111 Brood size manipulation .............................................................................................................. 111 Long-term analyses of female nest-defence during daylight hours ............................................ 112 Nest predation ............................................................................................................................. 112 Playback experiment during daylight hours in 2010 ................................................................... 113 7