En raison des forts coûts en temps et en énergie associés à chaque reproduction, les femelles ne sont généralement pas aussi disponibles que les mâles pour se reproduire. Les mâles entrent donc souvent en compétition pour accéder aux femelles disponibles. Ceci conduit à une forte sélection sexuelle chez les mâles. Un des exemples les plus frappants de compétition entre mâles peut être observé chez certaines espèces de crustacés chez qui les femelles ne sont sexuellement réceptives que pour un temps très limité. Les mâles ont donc évolué une stratégie de gardiennage précopulatoire grâce à laquelle ils monopolisent une femelle plusieurs jours avant qu’elle ne devienne réceptive. Ce comportement mâle est lui-même coûteux en temps et en énergie. En conséquence, il a été suggéré que les mâles devraient devenir sélectifs envers les femelles du fait du fort investissement que chaque reproduction représente pour eux. A l’aide d’un modèle mathématique, nous prédisons que les mâles effectuant de longs gardiennages précopulatoires devraient préférer s’apparier avec les grandes femelles plus fécondes. Toutefois, cette sélectivité devrait rester faible du fait de la forte compétition pour accéder aux femelles libres. Nous suggérons plutôt que les mâles devraient chercher à s’apparier avec des femelles de bonne qualité après s’être initialement apparié avec une femelle. Quand les mâles en couple rencontrent une femelle libre de meilleure qualité que leur propre femelle, ils devraient quitter leur femelle pour s’accoupler avec la nouvelle femelle. Contrairement à cette prédiction, nos expériences ont montré que les mâles en couple d’un crustacé amphipode Gammarus pulex ne changeaient pas systématiquement de femelle quand nous leurs proposions une femelle de meilleure qualité que leur propre femelle. Ils décidaient de changer de partenaire uniquement quand leur femelle était de mauvaise qualité, indépendamment de la qualité de la nouvelle femelle libre. D’autres expériences sont nécessaires pour comprendre le caractère adaptatif de ce comportement de changement de partenaire, seulement basé sur une partie de l’information disponible. Ces deux études soulignent la difficulté d’inférer des patrons de reproduction uniquement à partir des préférences individuelles. Dans la première étude, les mâles étaient contraints par la compétition pour accéder aux femelles libres. Dans la seconde, le processus de prise de décision des mâles conduisait à un comportement de choix apparemment sous-optimal. Ces contraintes n’ont que rarement été prises en compte malgré leur grande importance lorsqu’il s’agit de comprendre les causes comportementales d’un patron de reproduction très répandus chez les crustacés à gardiennage précopulatoire : l’homogamie pour la taille. Il a principalement été suggéré que ce patron de reproduction était issu d’une préférence mâle pour les grandes femelles associée à un avantage des grands mâles pour accéder aux femelles. Cette hypothèse n’a malgré tout reçu que peu de support empirique. A l’aide d’un modèle par simulation individu centrée, nous avons donc testé l’hypothèse selon laquelle une préférence mâle pour la distance à la mue des femelles serait à l’origine de l’homogamie pour la taille chez les crustacés à gardiennage précopulatoire. Quand les mâles préfèrent s’apparier avec des femelles qui sont strictement plus proches de la mue qu’eux, les couples formaient un patron d’homogamie pour la taille. Puisque plusieurs préférences différentes peuvent conduire à un même patron de reproduction, ce résultat souligne l’importance de considérer le processus complet de mise en couple pour étudier le lien entre les préférences individuelles et les patrons de reproduction. Les stratégies de femelles peuvent aussi jouer un rôle important dans les processus de mise en couple. Contrairement aux mâles, les femelles ont été décrites comme préférant les gardiennages courts du fait des coûts associés à la mise en couple. / Because of strong costs associated with each mating event, females are usually not as available for reproduction as males at any given time. Males are therefore in competition with each other for access to receptive females, hence leading to strong sexual selection. One textbook case of such a mating system occurs in moulting crustaceans where females can only be fertilized during a short period following their moult. This has favoured the evolution male strategies to monopolize females before their period of receptivity. Such a precopulatory mate guarding is widespread among many taxa and represents one of the most striking example of males’ competitive traits favoured by sexual selection. However, recent investigations have suggested that because males’ sexually selected traits often involve opportunity or mortality costs, males should become choosy towards females. Using a theoretical approach, we showed that males performing long lasting mate guarding should choose larger, more fecund females. However, under sequential encounter of potential mates, competition for female access decreases male choosiness before entering in precopula. We rather suggest that males should become choosy after initial pairing with a female. When encountering an unpaired female of better quality than their current female, paired males should switch partners. Contrary to our expectations, even under simultaneous encounters of two females, males did not seem to assess their relative quality. Instead they decided to change partner when their own female was of low absolute quality. This led to several cases where males forewent the possibility of increasing their fitness. Further investigations are needed to understand the adaptive significance of using only a subset of information in decision making. These two cases highlight the difficulty of inferring mating patterns from mating preferences only. In the first case, male preference was constrained by competition for access to females while in the second one, sampling processes led to apparent suboptimal mate choices. These potential constraints on decision making have rarely been acknowledge in precopulatory mate guarding crustaceans in spite of their major importance when inferring the causes of a well-known pairing pattern occurring in these species: size-assortative pairing. Size assortment among pairs has mainly been considered to come from a male directional preference for larger females associated with a large male advantage in getting access to preferred females. However, this hypothesis has received contrasted empirical support and little is known about the underlying pairing process causing size-assortative pairing. We investigated theoretically the possibility that a state-dependent male mating preference could account for size-assortative pairing. When males chose females which were exclusively closer to moult than them, assortative pairing by size arose under strong male-male competition. Because several preferences can account for a given pattern, this result emphasises the importance of considering the whole pairing process when studying the link between preferences and mate choice. Female strategies may also be of great importance during the pairing process. Contrary to males, females have been suggested to prefer short precopulatory mate guarding due to costs associated with pairing. Such a sexual conflict over guarding duration may have major effects on co-evolutionary dynamics between males and females traits. Proving its occurrence is yet challenging because empirical studies often lack a full economical survey of costs and benefits for females associated with male traits. Females benefits associated with long lasting precopulatory mate guarding have particularly been overlooked in previous studies. Here, we proposed several potential benefits for females and discuss their influence on sexual conflict over guarding duration.

En biologie évolutive, la variation phénotypique a longtemps été réduite au substrat sur lequel agissait la sélection naturelle. Toutefois, la constance intra-individuelle et le maintien intra-populationnel de certains comportements conduisirent au milieu des années 90 au développement du concept de personnalité animale, basé sur la caractérisation de traits tels que la néophobie, l’agressivité, les tendances exploratoires ou la prise de risque. De nombreuses études montrent que la personnalité est soumise à la sélection naturelle et est reliée à certaines stratégies biodémographiques, telles que la dispersion ou le comportement anti-prédateur.Les liens entre la personnalité animale et les stratégies d’appariement et d’approvisionnement, deux composantes fondamentales de la vie des organismes, ont pourtant été négligés jusqu’ici. Nous nous sommes donc attachés au cours de cette thèse à déterminer l’influence de la personnalité sur certains comportements sexuels et alimentaires à l’aide d’un organisme modèle en écologie comportementale : le Diamant mandarin (Taeniopygia guttata). Puis, à l’inverse, nous avons souligné le rôle des sélections naturelle et sexuelle dans le maintien des variations de personnalité.Nos principaux résultats indiquent que certains traits de personnalité mesurés sont inter-corrélés au niveau de notre population captive d’oiseaux, définissant un syndrome comportemental. De plus, la personnalité prédit de manière différentielle le succès d’approvisionnement entre les contextes de compétition par exploitation et par interférence. Ainsi, les individus proactifs sont dominants lors d’épisodes de compétition par interférence mais souffrent d’un plus faible succès d’approvisionnement lors de jeux producteur-chapardeur. Ces résultats soulèvent la possibilité que la sélection naturelle favorise différentes personnalités dans différents contextes, offrant un mécanisme d’explication du maintien des variations intra-populationnelles de personnalité. De plus, ils suggèrent que la personnalité contraint l’optimalité des comportements à travers les situations. Enfin, dans un contexte de choix du partenaire, nous avons montré que la personnalité des femelles utilisées dans des tests de préférence par association influence les mesures de la sélectivité, des scores de préférence et de leur répétabilité.L’étude conjointe de la personnalité animale et des stratégies d’appariement et d’approvisionnement constitue ainsi une voie prometteuse dans l’explication du maintien des variations de personnalité par sélections naturelle et sexuelle, ainsi que dans l’étude de l’influence de la personnalité sur les stratégies biodémographiques des organismes en contextes alimentaire et sexuel / In evolutionary biology, phenotypic variation has for a long time been considered as the raw material on which natural selection acts. However, research on the consistency of behaviour led to the development of the animal personality concept during the 1990s. This concept was based on the characterization of traits such as neophobia, aggressiveness, exploratory tendencies and risk-taking behaviour. Since then, several studies have shown that personality can evolve through natural selection and is related to many life-history traits, such as dispersal or anti-predator behaviour.Pairing strategies and foraging strategies are two fundamental components of an organism’s life, but their relationships with personality have so far been neglected. In this thesis, we determined the extent to which personality influences sexual and feeding behaviour, using the zebra finch (Taeniopygia guttata) as a model organism. Our work highlights the role of natural and sexual selection on the maintenance of personality variation.Some personality traits are related to each other within our sample, defining a behavioural syndrome. Moreover, personality predicted feeding success in competitive situations, but differently for scramble and interference competition. Proactive individuals were dominant in interference competition but had lower feeding success in producer-scrounger games. Interestingly, these results suggest that natural selection could favour different personalities depending on the context, perhaps explaining the maintenance of personality variation within populations. Moreover, personality could constrain behavioural optimality across situations. Finally, in a mate-choice context, we found that female personality influences selectivity, preference and its repeatability during spatial association tests.The joint study of personality and pairing and foraging strategies thus represents a promising avenue of research for understanding the maintenance of personality variation through natural and sexual selection. Moreover, personality can considerably influence some life-history traits in sexual and foraging contexts

De acordo com o paradigma Darwin-Bateman, fêmeas não ganham benefícios em copular com múltiplos machos (i.e., poliandria), dado que seu sucesso reprodutivo depende principalmente da qualidade dos seus parceiros sexuais. Porém, a poliandria é um comportamento amplamente encontrado nas espécies com reprodução sexuada. O objetivo central desta tese foi explorar as possíveis causas e consequências da poliandria. No Capítulo 1, utilizamos experimentos em laboratório para investigar como a experiência social antes e depois da maturação influencia o grau de poliandria. A espécie estudada foi o grilo australiano Teleogryllus commodus, para a qual o ambiente acústico antes da maturação determina o comportamento reprodutivo das fêmeas após a maturação. Nossos resultados mostram que o grau de poliandria não varia entre fêmeas criadas em dois ambientes acústicos distintos, um composto por machos de qualidade variada (QV) e outro composto por machos de alta qualidade (AQ). As fêmeas do grupo AQ foram menos responsivas aos machos de alta qualidade e aceitaram machos independentemente de sua qualidade, um padrão diferente daquele encontrado para as fêmeas do grupo QV. Quando a qualidade média dos machos encontrados pelas fêmeas do grupo QV foi baixa, o número total de machos aceitos foi baixo, porém quando a qualidade média dos machos foi alta, o número de machos aceitos foi alto. Portanto, há uma interação entre a experiência social antes e após a maturação na determinação do grau de poliandria. No Capítulo 2, utilizamos métodos comparativos para testar se o risco de competição espermática promovido pela poliandria determina a evolução de características masculinas relacionadas à monopolização de fêmeas. Nosso modelo de estudo foram rãs da subfamília Leptodactylinae, cujas espécies podem depositar ovos na água (onde o risco de competição espermática é alto) ou dentro de tocas na terra (onde o risco de competição espermática é baixo). Encontramos que há correlação entre o tipo de local de reprodução (aquático x terrestre) e quatro características masculinas relacionadas à monopolização de fêmeas. Nossos resultados provêem uma relação causal entre características dos machos e a reprodução terrestre, mas os padrões encontrados nem sempre seguiram as previsões propostas pela hipótese de que o risco de competição espermática é menor em espécies com reprodução terrestre. Tais resultados sugerem que as pressões seletivas que agem nas quatro características masculinas exploradas aqui não são as mesmas, provavelmente devido a diferenças nas funções que cada característica tem durante as interações inter- e intra-sexuais. Ao fim desta tese, fornecemos evidências de uma nova causa da poliandria, além de mostrarmos as consequências da poliandria sobre a evolução de características dos machos / According to the Darwin-Bateman paradigm, females derive no benefit from multiple mating (i.e., polyandry) and their reproductive success is mainly influenced by the quality of their partners. However, there is increasing evidence that polyandry is ubiquitous across many animal groups. The main goal of this thesis was to explore possible causes and consequences of polyandry. In Chapter 1, we conducted an experiment to investigate how the social experience pre- and post-maturity influences the degree of polyandry. We studied the Australian cricket Teleogryllus commodus, for which we know that the acoustic environment prior to maturity alters female reproductive behavior after maturity. We found that the degree of polyandry did not differ between females reared in two acoustic environments, one composed of varied-quality (VQ) males and other composed of only high-quality (HQ) males. However, whereas females from the HQ group were less responsive to high quality male calls, accepting males regardless of their quality, females from the VQ group showed a different response. When the mean male quality experienced by a female from the VQ group was low, the total number of accepted males was low, and when the mean male quality was high, the total number of accepted males was high. Thus, the social experience pre- and post-maturation interact to determine the degree of polyandry in females of the Australian cricket. In Chapter 2, we used comparative methods to test if the risk of sperm competition promoted by polyandry drives the evolution of male traits related to female monopolization. We studied frogs of the subfamily Leptodactylinae, which lay eggs either on the water (where sperm competition risk is high) or inside terrestrial burrows (where sperm competition risk is low). We found evidence of an evolutionary correlation between the type breeding site (aquatic x terrestrial) and four male traits related to female monopolization. Our study creates a causal link between male traits and terrestrial reproduction, but the patterns not always followed the hypothesis that sperm competition risk is lower in terrestrial breeders. This finding suggests that the selective pressure acting on male traits is not the same, probably because they have different functions during inter- and intrasexual interactions. In conclusion, we provide evidence of a new cause of polyandry and show that polyandry may have consequences for the evolution of weaponry traits in males

Em espécies que exibem cuidado paternal exclusivo, a qualidade esperada do comportamento paternal pode influenciar as decisões de acasalamento das fêmeas e determinar o sucesso de acasalamento dos machos. Nesta tese, investigamos a inter-relação entre o cuidado paternal e a seleção sexual utilizando o opilião Quindina limbata como organismo modelo. Os machos nesta espécie constroem ninhos na forma de taça que são visitados pelas fêmeas na busca de um sítio de oviposição. No primeiro capítulo, nós experimentalmente avaliamos a eficiência do atendimento aos ovos provido pelos machos e testamos se os ninhos desatendidos são adotamos por fêmeas e/ou por machos não relacionados com os ovos. As conclusões mais importantes deste capítulo são: (1) a proteção do macho é crucial para a sobrevivência dos ovos porque ninhos desatendidos são prontamente atacados por predadores; (2) a compensação do cuidado parental por fêmeas é rara, provavelmente porque elas estão associadas aos machos e não aos ninhos; (3) os machos adotam ovos não relacionados com eles e protegem eles tão eficientemente quanto os machos originais, provavelmente porque o cuidado dos ovos é um comportamento selecionada sexualmente. No segundo capítulo, nós testamos a existência de duas táticas alternativas de acasalamento nas fêmeas: residentes, na qual as fêmeas permanecem perto de um ninho, repelem fêmeas conespecíficas e copulam preferencialmente com um único macho dono de ninho, e visitantes, na qual as fêmeas não permanecem espacialmente associadas a ninhos, não repelem conespecíficas e copulam com vários machos donos de ninhos. Também investigamos se a monopolização de ninhos por fêmeas afeta o sucesso reprodutivo dos machos. As conclusões mais importantes deste capítulo são: (1) o comportamento das fêmeas parece ser uma tática reversível, na qual as fêmeas podem mudar de residentes a visitantes durante sua vida, provavelmente em resposta à condição corporal, e (2) a monogamia social imposta pelas fêmeas residentes pode afetar negativamente o grau da promiscuidade dos machos, mas não sua taça de acasalamento. Em conclusão, nós provemos evidência de que a preferência das fêmeas por machos que provem cuidado pode ter favorecido a manutenção do cuidado paternal. Além disso, demonstramos que a monopolização dos melhores machos ou ninhos pode ter favorecido a evolução das táticas alternativas de acasalamento nas fêmeas / In species exhibiting exclusive male care, the expected quality of paternal behavior can influence female mating decisions and thus determine male mating success. In this thesis, we investigated the interplay between paternal care and sexual selection using the harvestman Quindina limbata as model organism. Males of this species build cup-like mud nests that are visited by females in search of an oviposition site. In the first chapter, we experimentally evaluated the efficiency of egg-attendance provided by males and tested if unattended nests are adopted by females and/or unrelated males. The most important conclusions of this chapter are: (1) male protection is crucial for egg survival because unattended nests are promptly attacked by predators; (2) flexible compensation of parental care by females is rare, probably because they are associated to males and not to the nests; (3) males adopt unrelated eggs and protect them as efficiently as original owner males, probably because egg-attendance is a sexually-selected behavior. In the second chapter, we tested the existence of two reproductive tactics in females: resident, in which females remain close to a nest, repel conspecific females, and mate preferentially with a single nest-owner male, and wanderer, in which females are not spatially associated with nests, do not repel conspecific females, and mate with different nest-owner males. We also investigated whether nest monopolization by females may affect males\' reproductive success. The most important conclusions of this chapter are: (1) females\' behavior seems to be a reversible tactic, in which females can switch from resident to wanderer during their lifetime, probably in response to body condition, and (2) the social monogamy imposed by resident females may negatively affect the degree of male promiscuity, but not male\'s mating rate. In conclusion, we provide evidence that female preference for parental individuals may have favored the evolution and maintenance of paternal care. Moreover, we show that the monopolization of the best males or the best nests may have favored the evolution of alternative reproductive tactics in females

Physical properties of the environment may shape signalling traits by determining how effective signals are in influencing the behaviour of other individuals. Evidence abounds of signalling environment driving the evolution of colours and sounds, yet little is known about its influence upon gestural displays. Here, we performed a continent-wide phylogenetic comparative analysis to test the hypothesis that habitat structure drives the evolution of aerial sexual displays in passerine birds. We found that aerial displays are seven times more likely to evolve in open-habitat passerines than in forest ones, likely as a result of physical properties that allow aerial displays to transmit more broadly in open habitats. Our results provide an emblematic example of how environmental factors may help predict the direction of evolution of otherwise unpredictable sexual traits. The broader range of aerial displays in open habitats may also mean that females can sample more males, potentially leading to more intense sexual selection in open-habitat, aerial-displaying males / Propriedades físicas do ambiente podem influenciar a evolução de sinais ao determinar quão efetivos eles são em influenciar o comportamento de outro indivíduo. Diversos estudos mostram a influência do ambiente sobre a evolução de cores e sons. Entretanto, pouco se sabe de sua influência sobre sinais motores (i.e., displays). Nesse trabalho, conduzimos uma análise comparativa filogenética para testar a hipótese de que a estrutura do hábitat influencia a evolução de displays sexuais aéreos em aves Passeriformes. Descobrimos que display aéreos têm uma probabilidade sete vezes maior de evoluir em passeriformes de ambiente aberto do que nos florestais, provavelmente decorrente de propriedades físicas que permitem que displays aéreos sejam transmitidos mais amplamente em ambientes abertos. Nossos resultados são um exemplo emblemático de como fatores ambientais podem ajudar a prever a direção de evolução de caracteres sexuais, frequentemente tidos como imprevisíveis. O raio mais amplo de displays aéreos em ambientes abertos também pode permitir que fêmeas consigam amostrar mais machos da população, potencialmente intensificado a seleção sexual sobre machos de ambiente aberto que exibem displays aéreos

Dans un contexte de sélection sexuelle, les systèmes de communication permettant l’attraction et la stimulation du partenaire sexuel et la compétition entre individus du même sexe sont indispensables. Ceci est particulièrement vrai chez les espèces à système d’appariement polygyne de type lek, où les mâles rassemblés dans l’espace sont en compétition pour l’accès aux femelles. Chez notre sujet d’étude, l’outarde houbara nord-africaine les males réalisent des parades incluant des démonstrations visuelles et des vocalisations appelées booms, sur des sites espacés les uns des autres par des distances importantes dans un système qualifié de lek explosé. Notre objectif était d’étudier les systèmes de codage-décodage des informations exprimées durant la parade des mâles chez cette espèce. Nous avons démontré l’existence d’un codage de l’identité individuelle porté par des paramètres visuels et des paramètres acoustiques des vocalisations. Des associations significatives entre des paramètres des booms et des caractéristiques physiques et comportementales des mâles susceptibles de refléter leur qualité, ont également été mises en évidence. Ainsi, les mâles qui produisent les booms les plus graves avaient les masses les plus importantes et paradaient le plus intensément. Lorsque des interactions agonistiques étaient simulées expérimentalement en diffusant des leurres acoustiques, les mâles avaient des réponses comportementales différentes en fonction de la fréquence des booms diffusés. Ainsi le niveau fréquentiel des booms des autres mâles est effectivement perçu et décodé par les individus en compétition. Par ailleurs, plusieurs paramètres des booms sont génétiquement déterminés et héritables, et pourraient porter une information sur l’apparentement entre individus.Certains paramètres sont également liés à la consanguinité des individus. Nous avons montré également que les booms sont particulièrement bien adaptés à la communication à grande distance. En effet, les booms sont de très basse fréquence, se propagent à des distances supérieures à 640 m, et les paramètres supportant le codage-décodage de l’information sont résistants à la propagation et produits de manière redondante. Enfin, les signaux acoustiques et des signaux visuels et semblent agir en synergie (multimodalité) ce qui pourraient améliorer l’efficacité de la communication à grande distance. Ainsi notre étude a démontré que des informations complexes sont transmises durant la parade des mâles au travers de signaux acoustiques et visuels au sein d’un réseau de communication à grande distance. / Résumé en anglais ;In a context of sexual selection, the communication between individuals is essential to optimize both intra-sexual (competition between male) and inter-sexual (mate choice) processes. This is particularly true in species with lek mating systems, where both processes reach their full expression. The North African Houbara bustard, Chlamydotis undulata undulata, is a lekking bird where courting males aggregate in a so called ‘exploded-lek’, i.e. where aggregation is detectable only when mapped over a large area. During the breeding season, males perform a conspicuous and spectacular courtship including visual and acoustic (booms) signals. Our aim was to decipher the coding-decoding systems of the courtship in this species. We found that males can be individually discriminated on the basis of visual and acoustic parameters. The booms produced during the courtship are also related to males’ characteristics susceptible to reflect their quality. Individuals who produced the lowest frequency booms were those with the greatest weights and performed the highest intensity courtships. Playback experiment shows that the frequency content of the booms was perceived and decoded by males during agonistic interaction, leading to significantly different behavioural responses according to the frequency of the signals played back. Furthermore, several acoustic parameters of the booms are significantly heritable through genetic transmission and these parameters are also related to the level of inbreeding. Finally, we showed that the booms and the acoustic parameters supporting information are particularly well adapted to the transmission of information at long range as required in a long distance communication network constituted by an exploded lek. First, booms are of very low-frequency and propagate up to 640 m. Second, coding-decoding of information relies on sequentially redundant and propagation-resistant features. Third, acoustic signals are combined with visual signalling (multimodality) which may improve the long distance transmission of the information. This study brought evidence that complex information is conveyed by visual and acoustic signals during the courtship of males within a long distance network.

<p>Studies of hybrid zones offer important insights into the process of speciation. Much of the knowledge to be gained is dependent on an accurate estimation of the strength of pre- and post-zygotic isolation between hybridizing taxa. My results demonstrate that hybridization can variously affect different components of fitness. In Ficedula flycatchers, late-breeding females may directly benefit from pairing with a heterospecific male by gaining access to superior territories. The hybrid offspring possess an immune system that is as equally well functioning as in the parental species (the collared, F. albicollis, and pied flycatcher, F. hypoleuca). However, I found that a severe reduction in fertility persists for at least three generations after the actual hybridization event. Combining all information about the reproductive success of hybridizing individuals and their descendents revealed that postzygotic isolation between flycatchers is very strong; hybridizing individuals leave almost no descendents. This thesis presents one of few comprehensive summaries of the selection for/against assortative mating in a natural hybrid zone. These findings suggest a central role for intrinsic postzygotic isolation as a reproductive barrier separating newly evolved bird species, and contrast previous suggestions that postmating isolation is the slowest of the reproductive barriers to evolve in birds.</p><p>Despite this strong selection against hybridization, pre-mating isolation is incomplete. Hybridization often results from females lacking conspecific partners, but appears to be also caused by errors in species recognition. Much of this error probably reflects the short period of time that pied flycatchers on Gotland and Öland have been in sympatry. Compared to collared flycatchers, pied flycatchers are poorer able to discriminate between conspecific and heterospecific song, and male pied flycatchers more often falsely signal their own identity through heterospecific song copying. However, despite colonising the study site from other sympatric populations and having very little gene flow from allopatry, collared flycatchers also possess traits (e.g. delayed plumage maturation) that increase their hybridization risk. Once pre-mating isolation is strong, the rarity of hybridization probably inhibits further selection against traits promoting interspecific mating, especially when such traits may be beneficial in other contexts. This thesis highlights complex interactions between factors affecting hybridization rate that would not be detected if such a study were not field-based. Furthermore, it showcases likely examples in nature of a number of theoretical objections to the evolution of pre-mating barriers between populations living in sympatry.</p>

Studies of hybrid zones offer important insights into the process of speciation. Much of the knowledge to be gained is dependent on an accurate estimation of the strength of pre- and post-zygotic isolation between hybridizing taxa. My results demonstrate that hybridization can variously affect different components of fitness. In Ficedula flycatchers, late-breeding females may directly benefit from pairing with a heterospecific male by gaining access to superior territories. The hybrid offspring possess an immune system that is as equally well functioning as in the parental species (the collared, F. albicollis, and pied flycatcher, F. hypoleuca). However, I found that a severe reduction in fertility persists for at least three generations after the actual hybridization event. Combining all information about the reproductive success of hybridizing individuals and their descendents revealed that postzygotic isolation between flycatchers is very strong; hybridizing individuals leave almost no descendents. This thesis presents one of few comprehensive summaries of the selection for/against assortative mating in a natural hybrid zone. These findings suggest a central role for intrinsic postzygotic isolation as a reproductive barrier separating newly evolved bird species, and contrast previous suggestions that postmating isolation is the slowest of the reproductive barriers to evolve in birds. Despite this strong selection against hybridization, pre-mating isolation is incomplete. Hybridization often results from females lacking conspecific partners, but appears to be also caused by errors in species recognition. Much of this error probably reflects the short period of time that pied flycatchers on Gotland and Öland have been in sympatry. Compared to collared flycatchers, pied flycatchers are poorer able to discriminate between conspecific and heterospecific song, and male pied flycatchers more often falsely signal their own identity through heterospecific song copying. However, despite colonising the study site from other sympatric populations and having very little gene flow from allopatry, collared flycatchers also possess traits (e.g. delayed plumage maturation) that increase their hybridization risk. Once pre-mating isolation is strong, the rarity of hybridization probably inhibits further selection against traits promoting interspecific mating, especially when such traits may be beneficial in other contexts. This thesis highlights complex interactions between factors affecting hybridization rate that would not be detected if such a study were not field-based. Furthermore, it showcases likely examples in nature of a number of theoretical objections to the evolution of pre-mating barriers between populations living in sympatry.

Pygmy tarsiers (Tarsius pumilus) of Central Sulawesi, Indonesia are the only species of tarsier known to live exclusively at high altitudes. This study was the first to locate and observe multiple groups of this elusive primate. This research tested the hypothesis that variation in pygmy tarsier behavior and morphology correlates with measurable ecological differences that occur along an altitudinal gradient. As a response to decreased resources at higher altitudes and the associated effects on foraging competition and energy intake, pygmy tarsiers were predicted to exhibit lower population density, smaller group sizes, larger home ranges, and reduced sexually selected traits compared to lowland tarsiers. Six groups containing a total of 22 individuals were observed. Pygmy tarsiers were only found between 2000 and 2300 m, indicating allopatric separation from lowland tarsiers. As expected, the observed pygmy tarsiers lived at a lower density than lowland tarsier species, in association with decreased resources at higher altitudes. The estimated population density of pygmy tarsiers was 92 individuals per 100 ha, with 25 groups per 100 ha. However, contrary to expectation, home range sizes were not significantly larger than lowland tarsier home ranges, and average NPL was smaller than those of lowland tarsiers. The average home range size for the observed pygmy tarsiers was 2.0 ha, and the average nightly path length (NPL) was 365.36 m. Pygmy tarsiers exhibited a nonrandom, clumped distribution near forest edges. While insect abundance and biomass were found to decrease as altitude increased, insect abundance and biomass was higher along anthropogenic edges. Thus, tarsiers within the study area may mitigate the decreased availability of insects at high altitudes by remaining close to forest edges, which in turn may be related to smaller than expected home range sizes. Further, estimates of pygmy tarsier abundance may be inflated because of increased insect abundance along anthropogenic edges. Contrary to the prediction for smaller group sizes as a response to feeding competition, the observed pygmy tarsiers lived in relatively large groups with multiple adult males. However, in support of the prediction for energetic constraints on body proportions, the observed pygmy tarsiers did not exhibit sexually selected traits. The pygmy tarsiers exhibited low sexual dimorphism and small relative testes mass, a trend opposite from lowland tarsier species, which may indicate a constraint on the development of those traits. Considered together, these results suggest that the observed pygmy tarsiers have adapted to life in an environment with limited resources. Future studies should explore the possible contributing effects of seasonality and topography.

Contrairement à d’autres groupes animaux, chez les primates, la hiérarchie de dominance ne détermine pas systématiquement le succès reproductif des mâles. Afin de comprendre pourquoi, j’ai étudié les stratégies de reproduction des mâles et des femelles dans un groupe de macaques rhésus de la population semi-libre de Cayo Santiago (Porto Rico), collectant des données comportementales, hormonales et génétiques pendant deux saisons de reproduction. Les résultats se résument en cinq points. 1. Les nouveaux mâles qui ont immigré dans le groupe d’étude occupaient tous les rangs les plus subordonnés de la hiérarchie de dominance et ont monté en rang suite au départ de mâles plus dominants. Ainsi, l’acquisition d’un rang supérieur s’est faite passivement, en absence de conflits. Par conséquent, les mâles dominants étaient généralement d’âge mature et avaient résidé plus longtemps dans le groupe que les mâles subordonnés. 2. L’accès des mâles aux femelles est en accord avec le « modèle de la priorité d’accès » selon lequel le nombre de femelles simultanément en œstrus détermine le rang de dominance du mâle le plus subordonné qui peut avoir accès à une femelle (p. ex. le mâle de rang 4 s’il y a quatre femelles en œstrus). Bien que les mâles dominants aient eu plus de partenaires et aient monopolisé les femelles de qualité supérieure (dominance, parité, âge) pendant leur période ovulatoire (identifiée grâce au profil hormonal de la progestérone), le rang de dominance n’a pas déterminé le succès reproductif, les mâles intermédiaires ayant engendré significativement plus de rejetons que prédit. Il est possible que ces jeunes adultes aient produit un éjaculat de meilleure qualité que les mâles dominants d’âge mature, leur donnant un avantage au niveau de la compétition spermatique. 3. Les mâles dominants préféraient les femelles dominantes, mais cette préférence n’était pas réciproque, ces femelles coopérant plutôt avec les mâles intermédiaires, plus jeunes et moins familiers (c.-à-d. courte durée de résidence). Au contraire, les femelles subordonnées ont coopéré avec les mâles dominants. La préférence des femelles pour les mâles non familiers pourrait être liée à l’attrait pour un nouveau bagage génétique. 4. L’intensité de la couleur de la peau du visage des femelles pendant le cycle ovarien était corrélée au moment de la phase ovulatoire, une information susceptible d’être utilisée par les mâles pour maximiser leur probabilité de fécondation. 5. Les femelles retiraient des bénéfices directs de leurs liaisons sexuelles. En effet, les femelles en liaison sexuelle bénéficiaient d’un niveau de tolérance plus élevé de la part de leur partenaire mâle lorsqu’elles étaient à proximité d’une source de nourriture défendable, comparativement aux autres femelles. En somme, bien que les mâles dominants aient bénéficié d’une priorité d’accès aux femelles fertiles, cela s’est avéré insuffisant pour leur garantir la fécondation de ces femelles parce que celles-ci avaient plusieurs partenaires sexuels. Il semble que l’âge et la durée de résidence des mâles, corrélats de leur mode d’acquisition du rang, aient confondu l’effet du rang de dominance. / In contrast to most animal groups, dominance hierarchy does not systematically determine male reproductive success in primates. In order to investigate why, I studied male and female reproductive strategies in a group of free-ranging rhesus macaques on Cayo Santiago, Puerto Rico. I collected behavioural, genetic, and hormonal data during two consecutive mating seasons. My results are summarized below. 1. All new males who immigrated into the study group occupied the lowest-ranking position in the dominance hierarchy and rose in rank as the higher-ranking males left the group. Achieving a higher dominance rank occurred passively, without physical conflict. Thus, dominant males were mature individuals who resided longest in the group. 2. Male access to oestrus females followed the predictions of the ‘priority of access’ model, in which the number of females in oestrus determines the rank of the lowest-ranking male who can access a female (e.g. the fourth ranking male if four females are in oestrus). Even though dominant males obtained more mating partners and monopolised higher quality females (dominance, parity, age) during the ovulation window (as identified using progesterone profiles), dominance rank did not determine reproductive success, as intermediate-ranking males sired significantly more infants than predicted. It is likely that those young, intermediate-ranking adult males produced high quality ejaculate, giving them an advantage in sperm competition. 3. Dominant males preferred high-ranking females, but this preference was not reciprocal; high-ranking females cooperated with younger and less familiar intermediate-ranking males. Conversely, subordinate females cooperated with dominant males. Female preference for non-familiar males (i.e. short residency in the group) may be explained by an attraction to a novel genetic pool. 4. Female facial color intensity during the ovarian cycle was correlated with the timing of the ovulation window. This information may be used by males in order to maximize their fertilisation probability. 5. Consort females enjoyed a higher level of tolerance from their male partner when they were in proximity to a monopolisable food source, compared to other, non-consort females. This suggests that females obtained direct benefits from their sexual consorts. In conclusion, even though dominant males had priority access to ovulating females in the group, this was insufficient to guarantee fertilisation when females had several sexual partners. It appears that males’ age and length of residency, both correlates of their rank acquisition mode, may have been confounding factors in dominance rank.