In der Verhaltensökologie hat das Interesse an der Persönlichkeit von Tieren in den letzten zwei Jahrzehnten zugenommen und man geht davon aus, dass verschiedene Verhaltenstypen wichtige ökologische und evolutionäre Konsequenzen haben. Diese Konsequenzen sind besonders ausgeprägt, wenn eine konstante Verhaltensweise mit anderen Verhaltensweisen (sog. Verhaltenssyndrome) oder mit Life-History-Merkmalen (sog. Pace-of-life-Syndrome) korreliert. Bisher haben Studien zu Verhaltens- und Pace-of-life-Syndromen zu keinen schlüssigen Ergebnissen geführt – manche Hypothesen wurden bestätigt, andere jedoch nicht. Man nimmt an, dass Kovariation zwischen Merkmalen von der Umwelt geprägt ist. Da jedoch Tiere in Gefangenschaft in diesen Studien überrepräsentiert sind, kann dies ein möglicher Grund für die recht uneindeutigen Ergebnisse sein. Um diese Wissenslücke zu schließen, erforschte ich die Entstehung von Verhaltens- und Pace-of-life-Syndromen in einer Wildpopulation juveniler Haie, die relevanten ökologischen Bedingungen ausgesetzt waren. Ich untersuchte (1) ob ein Syndrom zwischen zwei konstanten Verhaltensweisen besteht und ob das Auftreten des Syndroms kontextabhängig ist, (2) ob ein Trade-off zwischen Wachstum und Mortalität durch Explorationsverhalten vermittelt wird, und (3) ob Persönlichkeit das Nahrungshabitat der Haie vorhersagen kann und ob dieser Zusammenhang kontextabhängig ist. Ich dokumentierte ein Verhaltenssyndrom das über Jahre und Standorte hinweg inkonsistent war und von inter-individueller Konkurrenz abhing. Ein Zusammenhang zwischen Explorationsverhalten und einem Wachstums-Mortalitäts-Trade-off war nur bei geringem Prädationsrisiko zu beobachten. Zudem sagte das Explorationsverhalten Nahrungshabitate nur bei geringem Prädationsrisiko vorher. Zusammengenommen deuten diese Ergebnisse darauf hin, dass die ökologischen Bedingungen eine entscheidende Rolle bei der Entstehung und Ausprägung von Persönlichkeits- und Pace-of-life-Syndromen spielen. / In behavioural ecology, interest in the study of animal personality (i.e. consistent individual differences in behaviour across time and/or context) has increased in the last two decades as it is believed to have important ecological and evolutionary consequences. These consequences are especially pronounced when a behaviour that is consistent covaries with other consistent behaviours (i.e. behavioural syndrome) or with life-history traits (i.e. pace-of-life syndrome). So far, studies of behavioural and pace-of-life-syndromes have produced ambiguous outcomes (e.g. hypotheses are sometimes verified and others not), and the prominence of studies on captive animals (i.e. as opposed to wild animals) in the literature may be a reason for inconclusive results as trait covariation has been hypothesized to be environmentally driven. To address this knowledge gap, I investigated the emergence of behavioural and pace-of-life-syndromes in a wild population of juvenile sharks subject to relevant ecological pressures (e.g. predation risk, inter-individual competition). I explored (1) whether a behavioural syndrome existed between two consistent traits (exploration and sociability) and whether the appearance of the syndrome was context dependent, (2) whether a growth-mortality trade-off was mediated by exploration personality and (3) whether personality could predict the foraging habitat of sharks and whether this link was context-dependent. First, I observed a behavioural syndrome between sociability and exploration personality which was inconsistent across years and locations and was dependent on inter-individual competition. Then, I found the association between exploration personality and a growth-mortality trade-off to only be observable in low predation risk. Similarly, I found that exploration personality only predicted wild foraging habitat when predation risk was low. Overall, these results suggest that ecological conditions play a crucial role in the emergence and the shaping of personality and trait association. This thesis offers a possible explanation for the ambiguous results of previous studies and highlights the importance of increasing the focus on wild study systems that are subject to relevant ecological pressures in future animal personality research.

Thesis advisor: Lisa Feldman Barrett / The aim of the present study was to examine whether individual differences in Time Insensitivity are related to subjective experiences of emotion and cognitive performance. Sixty-five undergraduates (52% female) completed self-report measures of cognitive flexibility and provided subjective self-reports of emotions following two time pressured cognitive tasks. As predicted, Time Insensitivity was related to self-reported cognitive flexibility, better cognitive performance during a time pressured task, as well as less negative subjective experience in response to these tasks. The results of the present study suggest that Time Insensitivity may have some beneficial outcomes. Limitations and implications for future directions are discussed. / Thesis (BA) — Boston College, 2004. / Submitted to: Boston College. College of Arts and Sciences. / Discipline: Psychology. / Discipline: College Honors Program.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第21603号 / 理博第4510号 / 新制||理||1647(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 森 哲, 教授 沼田 英治, 教授 中務 真人 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

Consistent between-individual differences in behaviour (termed “animal personality”) may be driven by adaptive differences in behavioural and physiological life-history traits. The Pace of Life Syndrome (POLS) hypothesis predicts a suit of correlations between those life-history traits along a fast-slow continuum. Therefore, according to the POLS, individuals that are fast-paced would be bolder, more explorative, show high growth-rates, lower immunity and a higher metabolic rate. A mechanistic link between such traits could also explain variation in cognitive traits, where bold individuals are faster at a given task but pay less attention to external cues and therefore make decisions less accurately. Here, I tested the POLS hypothesis focusing on between and within-individual variance in boldness, metabolic rate (MR), cognitive performance (as decision-making performance) and exploration in the hermit crab Pagurus bernhardus. In addition, I also investigated the potential role of anthropogenic disturbances (constant light exposure) as a driver of between and within-individual variation in boldness. Hermit crabs demonstrated consistent between-individual differences in boldness and exploration, providing evidence for the presence of animal personality. However, variation between individuals in boldness, exploration and cognitive performance were not underpinned by variation in MR. Although there were no between-individual correlations among MR and behaviour, MR did co-vary with within-individual variance in boldness. My results indicate that less predictable hermit crabs, on average, have a higher MR during startle responses compared with those that are relatively consistent in their behaviour. Boldness was positively correlated with exploration rate, indicating that more explorative were also bolder, as well as cognitive performance, as bold individuals had a better performance than shy. Finally, constant light exposure is likely to modify hermit crab personality and physiology. Hermit crabs kept under a constant light regime were less bold and had a higher metabolic rate, than when kept under standard light and dark regime, indicating possible effects light pollution in this species. These results only partially support the POLS hypothesis.

Animal personality is found in a wide range of taxa, yet our knowledge of what maintains consistent among-individual variation in behaviour is still incomplete. Many personality traits are associated with fitness, leading to the expectation that, under selection, genetic (and among-individual) variation will be eroded over time. Several adaptive models have been developed in order to explain this maintenance of variation. These include state-dependence, state-behaviour feedback loops, life-history and behavioural coadaptation and the Pace of Life syndrome. These models represent good starting points for thinking about what drives and maintains among-individual variation in behaviour, and while empirical support for these models is mixed, one thing they do have in common is the assumption of a significant genetic basis underpinning personality traits. Significant heritability is required for an evolutionary response to selection and for among-individual variation to be adaptive. The univariate estimates of heritability for personality traits that are growing in the literature, while useful, are likely insufficient to predict how personality traits will respond to selection. This thesis uses the Trinidadian guppy, Poecilia reticulata, and other species to explore patterns of among-individual and genetic variation in personality traits, advocating the benefits of using multivariate perspectives throughout. Firstly, the among-individual covariance structure between measures of boldness, growth and metabolic rate are estimated in a test of the Pace of Life syndrome. Secondly, an appraisal of the relative strength of maternal and genetic effects on offspring personality and how it changes over ontogeny. Next, a comprehensive treatment of sexual dimorphism in behaviour and size followed by analysis of genotype-by-sex interactions using both univariate and multivariate methods. Finally, a comparative analysis of personality in 7 species of small fish, identifying the main axis of among-individual variation from a single assay in each and quantifying the phylogenetic signal.

Avian life histories range along the axis from slow to fast, with slow life histories being characterized by greater investments in future reproduction, and fast by greater investments in current reproduction. The concept of pace-of-life syndromes refers to the coevolution of life strategies and related physiological, immunological and behavioral traits. Avian species from tropical areas are characterised by slower life histories (longer parental care, later maturation, smaller clutches, reduced metabolic rate) when compared to temperate zone species. Within this latitudinal gradient, investments in the total amount (weight) of body feathers have also been shown to be reduced in tropical birds. It remains unclear, however, whether feather growth itself follows this latitudinal pattern, and is slower in tropical species. Tropical birds have lower basal levels of corticosterone and testosterone, however available studies are based mainly on analyzes of hormone concentrations from plasma. The first aim of this diploma thesis was to evaluate differences between tropical (Cameroon) and temperate zone (the Czech Republic) passerine species in investments in tail feather growth by using methods of comparative ptilochronology. The second aim of the diploma thesis was to analyze concentrations of steroid...

Les sociologues s’accordent pour constater l’émergence de nouveaux rapports au temps, mais débattent des marges de manœuvre différenciées dont disposent les individus face à ces phénomènes. L’hypothèse que la vitesse de déplacement puisse être un levier est confortée par les études montrant que nos sociétés ont maîtrisé l’espace-temps grâce à l’amélioration des transports d’information et de marchandises. Cependant, peu de travaux empiriques abordent le sujet à l’échelle de l’individu. Cette thèse, qui repose sur une combinaison de méthodes qualitatives et quantitatives, a pour objectif de compléter ces recherches à travers l’étude des modes de transport utilisés et des pratiques spatiales quotidiennes en Île-de-France. Cette approche micro-sociale montre que l’augmentation du nombre et de la durée des activités hors du domicile n’est pas corrélée à une hausse des vitesses, mais à une diminution des distances parcourues pour se rendre à certaines activités. Ceci s’explique par les logiques de localisation des activités des individus. Pour celles auxquelles ils accordent de la valeur, ils maximisent la qualité de la destination, ce qui passe par l’utilisation de modes motorisés rapides. Pour les activités secondaires, ils minimisent leur temps de trajet en les localisant à proximité d’activités structurantes, ce qui induit l’usage de modes plus lents. Comme l’articulation entre ces deux logiques est plus aisée dans Paris qu’en banlieue, l’adoption d’un rythme de vie plus élevé et d’une planification plus souple y est facilitée. Plus que l’accès à la vitesse, ce sont les territoires de résidence qui induisent des disparités dans les marges de manœuvre face au temps / Times studies agree on the fact that our society is currently undergoing changes in terms of pace of life, but debates arise as to the uniformity of such changes within the population and individuals’ scope of action in the face of said phenomena. Given that society as a whole mastered time and space through the rise of transport speed, one is tempted to assume that it is also the case on the individual level. However, few empirical studies have taken it upon themselves to analyze the impact of speed in daily mobility on individuals’ pace of life. Using a combination of statistical analysis and in-depth interviews, this thesis aims at providing supplementary insight into the topic by analyzing transport modes and mobility patterns of inhabitants of the Greater Paris region. Through this micro-social analysis, we show that the more out-of-home activities people do, the lower their average speed in the course of a day is, whilst the average distance to a destination also decreases. This can be explained by the way they choose their destinations. For activities which are meaningful to them, they maximize the utility of their destination by using fast transport modes. For ancillary activities, they minimize travel time by locating them in the direct vicinity of their high-value activities, which prompts the use of slower modes. Finally, we show that the combination of these two rationales is easier in the center than in the suburbs, thus facilitating a high pace of life and more improvised schedules in the former. Hence, more than access to speed, it is residential location which induces disparities between people in their ability to deal with time demands

Animal behaviour is a state variable of the individual that deserves special attention given its determinant role in eco-evolutionary processes (Wolf et al. 2007 in Nature). The decomposition of the behavioural variation in between- and within-individual variability has revealed the existence of consistent between-individual differences referred to as personality or behavioural types (Dall et al. 2004 in Ecology Letters). Five axes of personality are usually recognized (exploration, aggressiveness, activity, sociability and boldness), and individual specificities along them tend to be correlated leading to what is known as behavioural syndromes. Recently, these patterns of covariation have been enlarged to accommodate movement behaviour within a personality-dependent spatial ecology theory (Spiegel et al. 2017 in Ecology Letters). Most animals tend to forage, reproduce and develop any activity within specific bounded space, which leads to the formation of home range (HR) areas (i.e., HR behaviour, Börger et al. 2008 in Ecology Letters). The increasing development of animal tracking technology is providing a huge amount of movement data revealing that HR behaviour is widespread among taxa and shows a large consistent variability, both at within- and between-individual level, which allows to define the existence of well-contrasted spatial behavioural types (SBTs). SBTs, as other personality traits, play an important role in selective processes as those impelled by harvesting activities. The Pace-of-Life-Syndrome (POLS) theory (Réale et al. 2010 in Philos. Trans. R. Soc. B Biol. Sci), hypothesises on how personality traits are expected to be correlated with life history (LH) traits along the fast-slow continuum (Stearns 1992 in Oxford Univ. Press) in the broadest sense. Accordingly, patterns of covariation between specific SBTs, physiology-related features and LHs would be expected to exist whenever they maximize the animal performance in a given environment. However, the way in which behavioural variation at the within-species level is translated to the wide range of LH traits remains a fundamental yet unresolved question, mainly due to the lack of a proper theoretical framework (Mathot & Frankenhuis, March 2018 in Behavioral Ecology and Sociobiology). Thus, unrevealing the mechanisms behind is certainly scientifically very exciting but also socially relevant. In such a context, this PhD thesis aimed to address from conceptual, empirical and theoretical perspectives cornerstone questions in behavioural ecology: what are the feasible mechanisms underpinning the establishment of HR areas and within-species variation, what are their consequences for animal functioning and performance (i.e., in. LH traits) at the individual and eco-evolutionary levels, or what are the implications for the assessment and conservation of wildlife of the existence of SBTs. The PhD thesis focusses in a fish heavily exploited by recreational fishers but it aims to provide general reasoning applicable to a wide range of wild animals. First, the PhD thesis proposes a mechanistic theory of personality-dependent movement behaviour based on dynamic energy budget models (i.e., a behavioural-bioenergetics theoretical model). Second, integrated in the field of animal personality (i.e., decomposition of behavioural variability into within- and between-individual’s components), it addresses empirically the study of behavioural variability in the main axis of personality for a marine fish species and looked for evidences of whether personality-mediated differences in energy acquisition may exist. Aiming to support empirically the possible connections between personality traits and space-use behaviour, the thesis provides some insights on the application of a novel-tracking algorithm to analyse the movement of individual fish submitted to different experimental conditions. Third, it provides two examples of how applying HR-related theoretical concepts may improve the management of natural resources: attending the properties of HR may facilitate the assessment of wildlife using fixed monitoring sampling stations, and considering SBTs may influence the assessment of the status of wild fish stocks. Finally, the adaptive value of the proposed behavioural-bioenergetics theory is explored by means of dynamic optimization to understand the eco-evolutionary consequences related with HR variability. In summary, this PhD thesis makes an important contribution to behavioural ecology by developing a unifying theory to test the generality and adaptive value of POLS based on dynamic energy budgets. This behavioural-bioenergetics model connects (1) personality traits (2) HR behaviour, (3) physiology and (4) LH traits through an interwoven of mass/energy fluxes, within which they interact and feedback with the ecological context. Overall, from an eco-evolutionary perspective, the proposed framework constitutes a powerful tool for exploring the ecological role of HR behaviour and predicting what combination of behavioural traits would be evolutionally favoured in a given ecological context. Moving forward to including managerial scenarios, this unifying theory provides scientifically founded knowledge that would promote to improve natural resource management by attending the behavioural component of animal populations.

Individuelle Verhaltensunterschiede (sog. „Animal personality“) werden oft als konsistent über die Zeit und situationsunabhängig angenommen. Vielfach werden solche Persönlichkeitsunterschiede zwischen Tieren einer Art durch individuelle Unterschiede im Energiehaushalt sowie Lebenszyklusvariablen (sog. ‚state variables‘, dt. Zustandsgrößen) erklärt. Dies ist in der „pace-of-life“ Hypothese zusammengefasst. In neueren Arbeiten wurde jedoch die Konsistenz von Persönlichkeitsmerkmalen über den Lebensverlauf und deren strikte Abhängigkeit von Zustandsgrößen in Frage gestellt. Die vorliegende Dissertation soll neue Einblicke in die Mechanismen, die zur Entstehung von Persönlichkeitsunterschieden im Verlauf des Lebens von Fischen, deren Situationsabhängigkeit sowie ihre Verbindung zu individuellen Unterschieden in Zustandsgrößen liefern. In einer Abfolge von fünf unabhängigen Studien untersuchte ich die genannten Annahmen und fand, dass (1) Persönlichkeitsunterschiede sich im Laufe des Lebens von Tieren vergrößern; (2) Persönlichkeitsabschätzungen bei jungen Tieren oft stärker vom experimentellen Aufbau beeinflusst werden als bei Erwachsenen; (3) der Energiehaushalt und Lebenszyklusvariablen Persönlichkeitsunterschiede sowohl unter Laborbedingungen als auch im Freiland nicht hinreichend erklären können; (4) Beziehungen zwischen Persönlichkeitsunterschieden und Unterschieden im Energiehaushalt und in Lebenszyklusvariablen fanden sich nur bei Fischen einer Population mit langsamer Lebenszyklusstrategie nicht jedoch in einer Population mit schnellem Lebenszyklus. Die vorliegende Arbeit suggeriert daher, dass sich erst im Verlauf des Lebens eines Tieres Persönlichkeitsunterschiede unvermeidbar entwickeln. Dies stellt Persönlichkeitsmessungen bei juvenilen Tieren grundsätzlich in Frage. Weiterhin scheinen Persönlichkeitsunterschiede und Zustandsgrößen unter bestimmten Umweltbedingungen und evolutiven Szenarien voneinander unabhängig zu sein. / Among-individual differences in behavior (i.e., animal personality) are assumed to be consistent over time and contexts. In theory, they are often explained by individual variations in energy costs of self-maintenance as well as life history among animals (i.e., state variables), commonly expressed as the pace-of-life syndrome hypothesis. Yet, recent theories have disputed the consistency of personality types over lifetime and their rigid state-dependency. This thesis aims to offer novel insights on the mechanisms behind the emergence and development of personality over lifetime of fishes, its context dependency, and its link to individual variation in state variables. In a sequence of five independent yet interconnected studies, I tested the assumptions above and observed that (1) personality differences increased during lifetime as a function of consistent declines in the behavioral plasticity with increasing age of animals; (2) personality estimates in young animals were weaker and thus more vulnerable to experimental biases compared to adults; (3) personality variation did not depend upon individual differences in energy costs of self-maintenance and life-history traits among individuals, under both laboratory and natural settings; and (4) the relationship between behavioral, metabolic, and life-history traits was manifested only in fish populations with slow rather than fast life-history strategies. This thesis suggests that personality variation in animals might be the inevitable outcome of development, raising questions about the reliability of personality estimates in juvenile individuals. Furthermore, individual variation in personality and “states” may act independently (i.e., phenotypes are uncorrelated) under environmental conditions and evolutionary contexts that mask or select against their trade-offs.

La sénescence de survie est un processus défini comme le déclin progressif des probabilités de survivre en fonction de l'âge, et caractérisé par deux paramètres clés : l'âge de début de sénescence et le taux de sénescence. En dépit de son caractère délétère sur la fitness, la sénescence est observée chez la plupart des êtres vivants et plusieurs théories évolutives de la sénescence ont été proposées pour expliquer ce paradoxe. Chez les mammifères, il existe une grande diversité de patrons de sénescence de survie entre les espèces, mais aussi entre les sexes. En général, les males sénescent plus tôt et plus vite que les femelles, et vivent donc moins longtemps. Un des objectifs de ma thèse fut d'identifier un (des) facteur(s), qui pourraient expliquer cette diversité aussi bien au niveau inter- que intra-spécifique chez les mammifères. Pour ce faire, j'ai utilisé une approche comparative qui corrige pour la phylogénie, et des données démographiques issues de populations captives (base de données Species360). Ma thèse confirme tout d'abord l'idée commune que les mammifères vivent généralement plus longtemps en captivité qu'en milieu sauvage, mais ce bénéfice est plus important pour les espèces ayant un cycle de vie rapide que pour celles ayant un cycle de vie très lent. De plus, l'intensité de la sélection sexuelle explique en partie les différences de patron de sénescence avec, chez les ruminants, les mâles des espèces polygynes commençant leur sénescence plus tôt et vivant moins longtemps que les mâles des espèces monogames. Cette thèse démontre également l'importance de définir précisément la forme et la force des relations allométriques pour éviter toute erreur d'interprétation. Enfin, mes résultats montrent que les pratiques managériales en zoo, plus que l'allocation à la reproduction, influencent le patron de sénescence de deux espèces de Varis. En conclusion, ma thèse a permis de mettre en évidence l'importance d'inclure les concepts de la sélection sexuelle dans le contexte des théories évolutives actuelles. Enfin, l'étude des facteurs ayant une influence sur le patron de sénescence de survie des espèces menacées, que cela soit en captivité ou dans leur milieu naturel, devrait permettre d'obtenir des informations clés pour aider à la conservation de ces espèces / The process of actuarial senescence is defined as the progressive decline of survival per time unit with increasing age and can be characterized by two keys metrics corresponding to the age at the onset of senescence and the rate of senescence. Despite its detrimental effect in terms of fitness, senescence is a nearly ubiquitous process across the tree of life and several evolutionary theories of senescence have been proposed to solve this apparent paradox. Across mammals, a large diversity of actuarial senescence patterns is observed among species, but also between sexes, with males often living shorter than females. One objective of my Ph.D. was to identify factor(s) influencing survival and actuarial senescence at inter- and intra-specific levels using a phylogenetic comparative approach based on demographic data obtained from captive populations (Species360 database). My thesis first confirms the common belief that mammals in zoos generally outlive their wild counterparts, but fast-living species benefit more of captive condition than slow-living species. In addition, I found that sexual selection intensity partly account for the diversity of actuarial senescence patterns with males of polygynous ruminant senescing earlier and living shorter than males of monogamous ruminants. Moreover, I highlight that evolutionary allometry of sexually selected traits need to be determined accurately to avoid any misinterpretation. Finally, my results confirm that management practices in zoos influence actuarial senescence pattern at intra-specific level in two species of Varecia but not the allocation in reproduction. In conclusion, my Ph.D. highlights the importance to include the concept of sexual selection within the evolutionary framework of current theories of senescence. Finally, the study of factors influencing the actuarial senescence patterns of threatened species is likely to provide key information in conservation projects of these species