Antler size provides an honest signal of male phenotypic quality in roe deer.

Abstract

Identifying factors shaping secondary sexual traits is essential in understanding how their variation may influence male fitness. Little information is available on the allocation of resources to antler growth in territorial ungulates with low sexual size dimorphism. We investigated phenotypic and environmental factors affecting both absolute and relative antler size of male roe deer in three contrasting populations in France and Sweden. In the three populations, we found marked age-specific variation in antler size, with an increase in both absolute and relative antler size between yearling and prime-age stages, followed by a decrease (senescence) for males older than 7 years. Antler size increased allometrically with body mass. This increase was particularly strong for senescent males, suggesting the evolution of two reproductive tactics: heavy old males invested particularly heavily in antler growth (potentially remaining competitive for territories), whereas light old males grew small antlers (potentially abandoning territory defense). Finally, environmental conditions had little effect on antler size: only population density negatively affected absolute antler size in one of the three populations. Antler size may therefore provide an honest signal of male phenotypic quality in roe deer. We discuss the implications of these results in terms of territory tenure and mating competition.

Full text

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UNIVERSITE PAUL SABATIER - TOULOUSE III
SWEDISH UNIVERSITY OF AGRICULTURAL SCIENCES
THESIS
to obtain the degree of
Docteur de l’Université Paul Sabatier - Toulouse III
Doctor of the Swedish University of Agricultural Sciences
Discipline: Behaviour, Ecology and Evolution
Presented and defended in public by:
Cécile VANPE
On the 21th of June 2007
MATING SYSTEMS AND SEXUAL SELECTION IN UNGULATES
New insights from a territorial species with low sexual size dimorphism:
the European roe deer (Capreolus capreolus)
Ph.D. supervisors:
Mark Hewison (CEFS – INRA Toulouse, France)
Henrik Andrén (Grimsö Wildlife Research Station - SLU, Sweden)
JURY
Stéphane Aulagnier
Marco Festa-Bianchet
Anna Qvarnström
Dominique Allainé
Etienne Danchin
Mark Hewison
Henrik Andrén
Petter Kjellander
Jean-Michel Gaillard
Université Paul Sabatier - Toulouse III
Université de Sherbrooke
Uppsala University
Université Claude Bernard – Lyon I
CNRS Toulouse
INRA Toulouse
Swedish University of Agricultural Sciences
Swedish University of Agricultural Sciences
CNRS Lyon
Chairman
Opponent/Referee
Examiner/Referee
Examiner
Examiner
Co-director
Co-director
Invited
Invited
Laboratoire Comportement et Ecologie de la Faune Sauvage – INRA, Castanet-Tolosan, France
Grimsö Wildlife Research Station, Swedish University of Agricultural Sciences, Riddarhyttan, Sweden

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"Of the branches of biological science to which Charles
Darwin's life-work has given us the key, few, if any,
are as attractive as the subject of sexual selection"
(Fisher 1915)
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ACKNOWLEDGEMENTS
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true-false “French” supervisor. Merci pour ta confiance, ton soutien et tes encouragements
quotidiens, tes précieux conseils, ta disponibilité sans limites et ta patience (toc toc, c’est
encore moi !!), ton enthousiasme et ta bonne humeur à toutes épreuves. Merci aussi de
m’avoir permis d’améliorer mon anglais (je pense que tes efforts ont porté leurs fruits !) et de
t’être soucié de mon avenir professionnel. Merci enfin d’avoir accepté d’être mon chauffeur
de nombreuses fois entre chez Jean-Michel et le CEFS (Mea culpa, je n’ai jamais demandé
mon autorisation de conduire les véhicules de service !). Bien au-delà de mon sujet de thèse,
j’ai appris beaucoup à tes côtés (notamment durant les longues heures de voiture en tête à tête,
les pauses café ou les soirées arrosées) sur les ongulés, l’écologie comportementale et la
biologie des populations en général, mais aussi sur les rouages complexes du monde de la
recherche, ta vision du métier de chercheur, le rock anglais, le sport, et bien d’autres choses…
I could not wish for a better supervisor !
First and foremost, I would like to express all my gratitude towards Mark Hewison, my
•
supervisor. Tack so mycket ! Without you, my field work at Bogesund would be impossible !
Thank you for your warm welcome in Sweden, your precious help for all aspects of my life in
Sweden (from field-work at Bogesund to holidays in Lapland, from Swedish food to rented
car, from administrative documents to credit card and mobile phone), for your great ideas for
naturalist trips (grouse, crane, bear, wolf)... Despite the distance between us and your busy
life (between field-work, research, children, pets, moose hunting, ...), you managed to be a
good supervisor ! I have excellent memories of my stay in Sweden thanks to you ! The wolf
howling during one of my last days in Sweden has remained engraved in my memory...
Second, I would like to warmly thank Petter Kjellander, my Swedish true-false co-
•
supervisor, to have been the precious link with the Swedish University of Agricultural
Sciences, and to help me in administrative problems of my joint-agreement.
Henrik Andrén should be thanked to have accepted to be my official Swedish co-
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comments, corrections and advice on my work, for his help in statistics, for his support and
his encouragements. Merci d’avoir pris le temps de m’expliquer les rouages de la recherche,
de m’avoir également fait partagé ta vision du métier de chercheur (une vision un peu
différente de celle de Mark ?) et de t’être soucié de mon avenir professionnel. Plus qu’un
collaborateur de la thèse, tu as joué largement le rôle d’un co-directeur.
A special thanks to Jean-Michel Gaillard for his extensive and helpful suggestions,
•
R programmation, his support and his advice. Merci pour ta disponibilité sans limites, ton
aide précieuse et ton soutien !
I am also particularly grateful to Nicolas Morellet for his extensive help in statistics and
•
helpful suggestions on my thesis project: Mark Hewison, Henrik Andrén, Petter Kjellander,
that I have already thanked, but also Jacob Höglund, Jean-Michel Gaillard, Jean-François
Cosson, and Stéphane Aulagnier. Fate decreed that we could not get all together !
Thank you to all the members of my Ph.D. committee for their precious advices and
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Qvarnström, Etienne Danchin, Dominique Allainé, Jean-Michel Gaillard, Mark Hewison,
I would like to thank all the members of my Ph.D jury Marco Festa-Bianchet, Anna
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Henrik Andrén, Petter Kjellander, Jacob Höglund, Stéphane Aulagnier. Thank you to Marco
Festa-Bianchet and Anna Qvarnström to have accepted to review my thesis as “referees”.
•
assistants who have helped me to collect the field data at Bogesund during the spring-
summer 2004 and 2005: Pauline Delos, Markus Olsson, Anna Sandhammar, Maria
Tybergsson, Marci Johnson, Kathryn Schwadere, Mathieu Sarrasa, Jonas Bergson, Anders
Janermo. A special thanks to Pauline and Mathieu to have put up with me during 5 months,
to have been willing to get up very early at dawn and to go to bed very late after dusk to
radio-track roe deer, to have shared my life on a day-to-day basis, my stress, my joice and my
sadness, my meals, my holidays in Lapland, Vesterälen and Norway (it was quite a saga !),
my hiking on the tracks of Bogesund, my drive to the library of Waxhölm (the precious
umbilical cord with the boyfriend, the family, the lab, and Mark) or to the supermarket...
Without you, my stay in Sweden would have been really lonely. I be deeply indebted to you !
I am also grateful to Markus and Jonas (and their numerous dogs) for their great and hard
work on the field, their friendship, their support and endless good mood. You have been great
guides to discover Swedish culture. Best wishes guys !!!! Jonas, I didn’t forget the rotten fish
experience ! You are welcome to France to have a test of snakes and frogs !!! Thank you
Anders for your precious help and advice to catch fawns. Thank you Maria for your great
work on vegetation sampling and Per Grängstedt for vegetation weighting. Thank you Anna
and the US team Katy and Marci (Do not forget my French lessons) for your help with
radio-tracking. I would like finally to thank Mark Hewison, Nicolas Morellet, Bruno
Cargnelutti, Petter Kjellander and Fredrik Knape to have done fawn captures in 2006.
Mark, Nico and Bruno, I was happy that you discover my field-work study site and “my” roe
deer.
Another very important factor contributing to my work has been the numerous field
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samples in thousands !). Merci de m’avoir expliquer et montrer patiemment comment réaliser
les extractions d’ADN, les PCR, les génotypages, les relectures. Merci pour ton accueil
pendant mes séjours au CBGP. Thanks a lot to Jean-François Cosson for his advice on
molecular analyses and welcome at the CBGP. The following people also contributed to lab
work: Pia and Petter Kjellander for extractions and genotypages, Bruno Lourtet and Jean-
Marc Angibault for the preparation of DNA samples.
Maxime Galan deserve special thanks for his great work on molecular analyses (DNA
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advice on niche and K-select analyses.
I am grateful to Michel Goulard for his helps in statistics and Clément Calenge for his
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Fontaines, and especially to Guy Van Laere, Daniel Delorme and François Klein.
Thank you to the ONCFS to have allowed me to analyse data from Chizé and Trois
I am also grateful to a large number of people for help of different sorts and their support:
•
Stéphane Aulagnier (the director of the lab) pour son accueil au laboratoire et pour m’avoir
permis de m’initier à l’enseignement.
Aurélie Coulon, Marion Delon, Nina Giotto, Cathy Boleat, Emmanuelle Richard and
Sarah Benhaiem who have shared my office and my days of work. Merci Aurélie pour ton
soutien, ton aide, tes conseils, tes encouragements, ton accueil chez toi quand j’étais SDF, nos
People of CEFS – INRA Toulouse:
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discussions, et surtout ton amitié. Ma thèse n’aurait pas été si agréable sans ta joyeuse
présence à mes côtés ! Bon vent à toi pour la suite !
Pascale Dufour and Lucette Gabet (the secretaries) to have patiently solved my
administrative problems. Merci Pascale pour ton accueil au labo, l’animation du labo, les
sorties culturelles, les discussions sur tout et rien aux pauses café et au restau de l’INRA, ton
soutien quotidien.
People of the team “Ecologie des Populations”, pour tous les bons moments partagés au
labo, en salle café, durant les captures à Fabas, lors de colloques ou de fêtes: Mark Hewison
(thanks again !), Nicolas Morellet (merci encore pour ton soutien et ton aide), Hélène
Verheyden (merci pour les sorties au ski, tes conseils sur les analyses d’habitat et tes
encouragements), Tanguy Daufresne (merci pour nos sorties naturalistes, nos échanges sur les
voyages, ton soutien), Emmanuel Serrano (merci pour nos échanges sur la sélection de
modèle, le flamenco, les bouquetins ibériques, tes encouragements et tes conseils pour les
postdocs), Jean Joachim (merci de m’avoir dépannée de nombreuses fois en informatique, les
« échanges ornithologiques » et pour la capture d’oiseaux), Bruno Cargnelutti (merci pour ta
bonne humeur communicative et de m’avoir initiée au radiotracking), Jean-Marc Angibault
(merci pour tes conseils sur le terrain), Dominique Pépin (bonne retraite et bon vent à toi!),
Bruno Lourtet, Joël Merlet.
"Students and co": Thomas Marguet (merci pour les nombreuses initiatives de sorties à
Toulouse), Mélanie Nemoz (« bat woman », merci pour les sorties en ski de rando et nos
échanges sur les voyages, la montagne et bien d’autres choses…), Nora Khames (merci pour
ton amitié, nos échanges sur la Kabylie et le Liban, de m’avoir initiée aux danses, à la cuisine,
à la musique et autres coutûmes kabyles), Nicolas Kidjo (merci d’avoir partagé avec moi les
durs et intenses moments de la fin de thèse), Carine Pastorelli et Nina Giotto (merci pour
votre joie de vivre communicative et votre chaleureuse présence à mes côtés), Nicolas Villeret
(merci pour tes conseils et ton soutien), and cie.
Others: Georges Gonzalez (merci pour nos échanges sur les ongulés de l’Himalaya et d’Asie
Central, et sur la conservation de la faune en général), Nicolas Cebe (merci pour les bons
plans naturalistes), Denis Picot (merci pour ta joyeuse présence au restau de l’INRA, aux
pauses café et durant nos sorties restau à Toulouse), Jean-François Gérard (merci pour nos
discussions sur la faune et le reste), Cathy Cibien et Marie-Pierre Winterton (merci pour votre
soutien et de m’avoir fait partagé votre vision sur les aspects plus appliqués de la gestion de la
faune et des milieux), Eric Bideau, Jean-Luc Rhames, and cie.
•
Henrik Andrén, Anders Janermo, Jean-Michel Roberge, Jonas and Johan, the secretary pour
leur accueil au labo.
People of the Grimsö Wildlife Research Station: Petter & Pia Kjellander, Olof Liberg,
•
secourue lors de mon accident de travail.
People of CBGP – INRA Montpellier pour leur accueil au labo et pour m’avoir
•
incroyable.
Dominique Galy (secrétaire SEVAB) pour son chaleureux soutien et sa disponibilité
•
Nora (& Selim) (plus que des collègues de labo, vous êtes devenus des Amis ! Merci !),
Jeannot & Laure (pour votre amitié et votre chaleureuse et rassurante présence à mes côtés
durant ma thèse, pour votre formidable accueil dans votre havre de paix en Ariège quand
j’avais besoin de changer d’air), Pingu & Marina (pour m’avoir accueillie et aidée à mon
arrivée à Toulouse et pour votre chaleureux soutien), Natalia (merci d’avoir partagé ces
Thank you to all my friends from Toulouse or elsewhere: Aurélie, Mélanie (& Virgil),
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années thèse avec moi ! Bon vent à toi !), Sophie (pour ton soutien et notre longue amitié),
Anne & Nacho, Sarah, Laurette, Christian & Jeannine Bravard (mes profs de bio du lycée
grâce à qui j’ai pris goût à la recherche en biologie et qui m’ont toujours soutenue dans mes
études depuis le lycée), Popy (pour m’avoir hébergée à plusieurs reprises lors de mes venues à
Lyon pour voir JMG), Zitoune (merci pour les photos de chevreuil !) et tous les autres potes
de Ker 51, Maryline P. et Cécile M. (de m’avoir fait découvrir le radio-tracking chevreuil à
Chizé), Annaïg & Arnaud, Stéph, Larbi, Abdel, Emilien.
•
m’avoir initiée à la nature, à la science et à la recherche, d’avoir toujours cru en moi et de
m’avoir toujours soutenue et encouragée dans mes études, mes stages naturalistes et mes
expatriations au bout du monde. Merci aussi d’être venus assister à ma soutenance de thèse et
d’avoir organisé ma fête de soutenance. Merci à la joyeuse tribu de la famille Hibert pour
leur soutien, leur chaleureux accueil dans la famille et d’avoir contribué par leur marque de
soutien ou leur présence à la fête de ma soutenance de thèse.
I would like also to thank my parents and grand-parents for their support. Merci de
•
et tes encouragements du premier au dernier jour de ma thèse, ton aide sur R et sur la
sélection d’habitat, tes suggestions (quelle couleur de fond d’écran vais-je mettre ?), d’avoir
supporté mes sautes d’humeur, de m’avoir remonté le moral quand je me décourageais,
d’avoir partagé mes moments de joie, d’avoir pris soin de moi, d’avoir été si patient et si
compréhensif, de m’avoir fait découvrir le W et le Niger et d’être venu découvrir Bogesund et
la Suède, d’avoir sacrifié beaucoup de ton encadrement de thèse pour que nous puissions
vivre ensemble à Toulouse, d’avoir été toujours à mes côtés même séparés par plusieurs
milliers de kilomètres et plusieurs dizaines de degrés (toi au Niger moi en Suède avec 50°C de
différence !)…
Finally, I would like to warmly and affectionately thank Fabrice. Merci pour ton soutien
My research and salary have been supported by the French Ministry of Research (MRT
and cotutelle grants), FORMAS-INRA and SLU.
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TABLE OF CONTENTS
LIST OF TABLES………………………………………………………………………..…13
LIST OF FIGURES…………………………………………………………………………15
ABSTRACT…………………………………………………………………………………19
INTRODUCTION...................................................................................................... 21
I.GENERAL INTRODUCTION: CONTEXT OF THE STUDY.......................................................... 23
II.MATING SYSTEMS AND SEXUAL SELECTION: WHAT THE THEORY TELLS US.............. 27
II.1. DIVERSITY AND EVOLUTION OF MATING SYSTEMS........................................................... 27
II.1.1.Parental care and party size...................................................................................................... 27
II.1.2. Strategy of monopolization of mates and evolution of male mating systems........................... 28
II.2. THE THEORY OF SEXUAL SELECTION..................................................................................... 32
II.2.1. Origin of the competition over mates....................................................................................... 32
II.2.2.Darwin’s theory of sexual selection ......................................................................................... 32
II.2.3.The various forms of intra-sexual selection.............................................................................. 34
II.2.4. Inter-sexual selection and evolution of female mate choice..................................................... 34
II.3.
VIEW…………………………………………………………………………………………………………..37
II.3.1. How to measure sexual selection?............................................................................................ 37
II.3.2. Variation in the opportunity for sexual selection...................................................................... 39
II.3.3.Variation in sexually selected traits and mechanisms of competition over mates.................... 40
VARIATION IN SEXUAL SELECTION AMONG SPECIES: THE THEORETICAL POINT OF
III.
STUDIES.............................................................................................................................................................. 43
MATING SYSTEMS AND SEXUAL SELECTION: NEW INSIGHTS FROM EMPIRICAL
III.1.
MODEL………………………………………………………………………………………………………..43
THE IMPORTANCE OF MAMMALS, AND ESPECIALLY UNGULATES, AS A STUDY
III.2. A NEW POWERFUL APPROACH BASED ON MOLECULAR TECHNIQUES.......................... 46
III.3.
SELECTION: A COMPLEX LINK................................................................................................................. 47
PARTY SIZE, SEXUAL DIMORPHISM AND THE OPPORTUNITY FOR SEXUAL
III.4.
COMPETITION OVER MATES: THE CASE OF THE UNGULATES........................................................ 49
III.4.1. Body size and mass .................................................................................................................. 50
III.4.2.Antler and horn size ................................................................................................................. 51
III.4.3.Antler and horn asymmetry...................................................................................................... 53
III.4.4. Territory size, location, and habitat quality.............................................................................. 55
III.4.5. Chemical signals....................................................................................................................... 56
III.4.6.Acoustic signals........................................................................................................................ 57
III.4.7.Visual signals ........................................................................................................................... 58
MATING SYSTEMS, SEXUALLY SELECTED TRAITS AND MECHANISMS OF
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III.5.
AND/OR TERRITORIAL MATING SYSTEM IN UNGULATES................................................................ 59
CONCLUSION: LACK OF DATA IN SPECIES WITH LOW SEXUAL SIZE DIMORPHISM
IV.
AND SEXUAL SELECTION IN ROE DEER..................................................................................................... 61
THE STUDY: MALE REPRODUCTIVE SUCCESS, THE EVOLUTION OF TERRITORIALITY
IV.1.
SEXUAL SIZE DIMORPHISM...................................................................................................................... 61
THE STUDY SPECIES: THE EUROPEAN ROE DEER, A TERRITORIAL SPECIES WITH LOW
IV.2.SUBJECT OF THE STUDY.............................................................................................................. 63
IV.3.STRUCTURE OF THE STUDY....................................................................................................... 65
MATERIALS AND METHODS................................................................................. 71
I. STUDY POPULATIONS..................................................................................................................... 73
II.
STUDIES.............................................................................................................................................................. 79
THE METHODS: A COMBINATION OF MOLECULAR TOOLS AND FIELD ECOLOGY
II.1. LONGITUDINAL DATA AND POPULATION SAMPLING (all 3 populations) .......................... 79
II.2.MOLECULAR ANALYSES (Bogesund and Trois Fontaines)......................................................... 81
II.2.1.Considerations on the lifetime breeding success as proxy of fitness........................................ 81
II.2.2.Tissue sampling........................................................................................................................ 84
II.2.3.Microsatellite genotyping......................................................................................................... 85
II.2.4.Test of the suitability of microsatellite markers....................................................................... 87
II.2.5. Limitation, identification and quantification of genotyping errors........................................... 89
II.2.6.Paternity analyses..................................................................................................................... 91
II.2.7.Estimation of male breeding success and the opportunity for sexual selection........................ 94
II.3.FIELD ECOLOGY STUDIES (Bogesund)....................................................................................... 95
II.3.1.Fawn capture............................................................................................................................ 95
II.3.2.Pellet-group counts and estimation of a local population abundance....................................... 96
II.3.3.Capture-Mark-Resighting (Petersen method)........................................................................... 97
II.3.4.Radio-tracking and estimation of the size of males’ territory................................................... 98
II.3.5. Vegetation sampling and map of habitat quality.................................................................... 100
PART 1: VARIATION IN MALE BREEDING SUCCESS AND OPPORTUNITY FOR
SEXUAL SELECTION ........................................................................................... 103
PAPER 1: MATING SYSTEM, SEXUAL DIMORPHISM AND THE OPPORTUNITY
FOR SEXUAL SELECTION IN A TERRITORIAL UNGULATE............................... 105
PART 2: AGE-RELATED VARIATION IN MALE BREEDING SUCCESS ............ 131
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PAPER 2: AGE-SPECIFIC VARIATION IN MALE BREEDING SUCCESS AND
MATING TACTICS OF ROE DEER........................................................................ 133
PART 3: PHENOTYPIC TRAITS AS SEXUALLY SELECTED TRAITS................ 155
PAPER 3: ANTLER SIZE PROVIDES AN HONEST SIGNAL OF MALE
PHENOTYPIC QUALITY IN ROE DEER................................................................ 157
PAPER 4: IS BIGGER BETTER IN A LOW SEXUAL SIZE DIMORPHIC
UNGULATE? CONTINUED SEXUAL SELECTION ON MALE BODY MASS AND
ANTLER SIZE IN ROE DEER................................................................................ 177
PART 4: ROLE OF THE TERRITORY IN MALE BREEDING SUCCESS AND
EVOLUTION OF TERRITORIALITY...................................................................... 193
PAPER 5: ACCESS TO MATES IN ROE DEER IS DETERMINED BY THE SIZE OF
A MALE’S TERRITORY, BUT NOT ITS HABITAT QUALITY................................. 195
PART 5: A FIRST STEP TOWARDS THE FEMALE POINT OF VIEW: FEMALE
MULTIPLE MATING .............................................................................................. 221
PAPER 6: MULTIPLE PATERNITY IS LIKELY BUT INFREQUENT IN THE
POLYTOCOUS EUROPEAN ROE DEER.............................................................. 223
DISCUSSION......................................................................................................... 235
I. SYNTHESIS OF MAIN RESULTS.................................................................................................. 237
II. DISCUSSION AND PERSPECTIVES............................................................................................. 239
II.1.
SELECTION IN ROE DEER......................................................................................................................... 239
II.1.1.First data on male lifetime breeding success in roe deer based on molecular analyses.......... 239
II.1.2.An attempt to compare estimates of the variance in male lifetime reproductive success between
different ungulate species.......................................................................................................................... 242
II.1.3. New insights on the pattern of variation in the opportunity for sexual selection among ungulate
species……………………………………………………………………………………………………..243
VARIANCE IN MALE BREEDING SUCCESS AND THE OPPORTUNITY FOR SEXUAL
II.2.
SELECTED TRAITS..................................................................................................................................... 246
II.2.1.Age......................................................................................................................................... 247
DETERMINANTS OF THE VARIANCE IN MALE BREEDING SUCCESS AND SEXUALLY
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II.2.2.
II.2.3.
II.2.4.
II.2.5.
II.2.6.
Body mass and antler size ...................................................................................................... 248
The abundance of females in territories.................................................................................. 249
Territory size and habitat quality within the male’s territory................................................. 251
Other potential sexually selected traits................................................................................... 253
New insights into the adaptive function and evolution of male territoriality ......................... 254
II.3.
IN THE TWO SEXES.................................................................................................................................... 257
II.3.1.Comparison of the distribution of lifetime breeding success.................................................. 257
II.3.2.Comparison of the effects of age on breeding success and survival....................................... 259
II.3.3.Comparison of the ultimate factors that affect breeding success............................................ 261
II.3.4.New insights into the evolution of the low sexual size dimorphism in roe deer .................... 263
COMPARISON OF THE EXTENT AND CAUSES OF VARIATION IN BREEDING SUCCESS
II.4. ALTERNATIVE MATING TACTICS IN YOUNG AND OLD MALES...................................... 264
II.5. THE ROLE OF FEMALES IN ROE DEER SEXUAL SELECTION ............................................ 268
II.6.
SEXUAL SELECTION TO APPLIED ISSUES IN CONSERVATION AND POPULATION
MANAGEMENT........................................................................................................................................... 271
LET’S GO FURTHER: FROM THEORETICAL KNOWLEDGE ON MATING SYSTEMS AND
II.7.CONCLUSION................................................................................................................................ 275
II.8. PERSPECTIVES............................................................................................................................. 276
REFERENCES....................................................................................................... 281
APPENDIX…………………………………………………………………………………303
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LIST OF TABLES
INTRODUCTION
Table 1. Taxonomic organization of the world’s ruminants (from Grubb 2005). 45
Table 2. Main long-term monitored populations of ungulates in the world. 65
MATERIALS AND METHODS
Table 3. Descriptive statistics for the microsatellite loci used to genotype Bogesund (A) and
Trois Fontaines (B) DNA samples. 86
Table 4. Sample size for paternity analyses at Bogesund (A) and Trois Fontaines (B). 93
DISCUSSION
Table 5. Summary of the main reproductive parameters in male and female roe deer at
Bogesund. 259
Note that tables from papers are not listed here.
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LIST OF FIGURES
INTRODUCTION
Figure 1. The present distribution of European roe deer in western Europe. In addition to the
areas marked on the map, roe deer extend further in Russia to the Ural mountains and are
found in scattered populations in Turkey. From Linnell et al. (1998a). 23
Figure 2. Variation of the number of roe deer allowed to be shot (in light) and actually shot
(in dark) in France since 1973. Data ONCFS. 24
Figure 3. The two-step process influencing mating systems in cases where males do not
provide parental care. Males thus have two main options to maximize the number of matings:
they could either compete for females directly (A) or indirectly (B). Figure adapted from
Krebs & Davies (1993). 28
Figure 4. Schematic figure illustrating the various polygynous mating systems in mammals.
Big ♂ signs represent dominant or territorial males. Small ♂ signs represent subordinates or
non-territorial males. Dotted circles represent female home ranges. Solid circles represent
male territories. 30
Figure 5. Method to estimate the intensity of sexual selection on a phenotypic trait. The slope
of the multiple regression line of fitness on trait represents the selection gradient and is an
indication of directional selection. 38
Figure 6. Illustration of the continuum of opportunity for sexual selection from monogamous
and monomorphic species on the left-hand side to highly polygynous and highly dimorphic
species on the right-hand side. 40
Figure 7. Relationship between the continuum of mating systems and the potential for the
two modes of sexual selection. Adapted from the figure of Min (1997). 42
Figure 8. Reproductive cycle of male (not in italic) and female (in italic) roe deer. 63
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