The human voice may convey meaningful information about socially and evolutionarily relevant characteristics of the vocalizer. In turn, listeners may readily evaluate personal characteristics, such as body size, on the basis of nonverbal voice features. Research investigating vocal communication of physical size in humans has focused on two salient and largely independent voice features, fundamental frequency and/or corresponding harmonics (perceived as voice pitch) and formant frequencies (resonance frequencies of the supralaryngeal vocal tract). However, the degree to which fundamental and formant frequencies reliably predict variation in body size controlling for sex and age, and their relative role in the perception or accurate estimation of body size, has to date been unclear. In the current thesis, using meta-analysis, I establish that formants reliably predict variation in men’s and women’s heights and weights. In contrast, fundamental frequency only weakly predicts men’s heights and women’s weights. These findings corroborate work on many other mammals whose vocal production, like humans, follows the source-filter model. Despite the lack of a robust physical relationship between fundamental frequency and size within sexes, I further demonstrate that listeners utilize voice pitch to accurately gauge men’s relative height. My research suggests that voice pitch indirectly facilitates accurate size assessment by providing a carrier signal (i.e., dense harmonics) for formants. This is the first evidence that pitch does not confound accurate size estimation. Finally, I demonstrate that voices with lowered pitch, but not raised pitch, are perceived as larger when projected from a low than high spatial location. These results suggest that strong cross-modal perceptual biases linking low pitch to low elevation and large size may, in some contexts, cause errors in size estimation. Taken together, this thesis provides a detailed account of human vocal communication of body size, which can play a meaningful role in sexual and social contexts. / Dissertation / Doctor of Philosophy (PhD)

Sexual conflicts arise when there is a difference in how females and males of a species or population achieve their maximum reproductive fitness. In intralocus sexual conflicts, alleles at a given locus are exposed to conflicting, or antagonistic, selection pressures. Based on a Genome-wide association study on sexually antagonistic genes in Drosophila melanogaster (fruit flies), the aim of this study was to investigate whether the candidate gene CG3598 exhibits sexually antagonistic effects on fitness between the 2 identified alleles of the gene. The study was performed on females from a Canton-S population of D. melanogaster who, by genetic manipulation through CRISPR Cas9, carried one of the 2 alleles of the CG3598. 6 excision lines of females had allele 1 and 5 excision lines had allele 2. The females were mated with "wild type" males from a Canton-S population in mediums prepared with about 6 mg of live dry yeast, after which the females were moved to separate mediums to lay their eggs. After 12 days, the adult offspring were counted and statistical calculations were performed on the average number of offspring per female for each line and allele. An Independent sample t-test showed that the females’ average fecundity did not differ between alleles (p = 0.059) and a Nested ANOVA analysis indicated that the average fecundity for each line within each allele differed (p = 0.023). Due to the fact that similar studies have found the same result, it may be necessary to investigate and possibly change the experimental design of the method to enable competition between females with different genetic conditions in order to observe a difference in fertility based on the females' ability to compete. / Sexuella konflikter uppstår när det finns en skillnad i hur honor och hanar i en art eller population uppnår sin maximala reproduktiva fitness. Vid intralocus sexuella konflikter utsätts alleler vid ett givet locus för motstridiga, eller antagonistiska, selektionstryck. Baserat på en Genome-wide association study om sexuellt antagonistiska gener hos Drosophila melanogaster (bananflugor) syftade denna studie till att undersöka huruvida genkandidaten CG3598 uppvisar sexuellt antagonistiska effekter på fitness mellan de 2 identifierade allelerna av genen. Undersökningen utfördes på honor från en CantonS-population av D. melanogaster som genom genmodifiering av CRISPR Cas9 bar en av de olika alleler av genen CG3598. 6 linjer av honor bar allel 1 och 5 linjer bar allel 2. Honorna parades med ”wild type”-hanar från en CantonS-population i rör preparerade med ca 6 mg levande torrjäst, varpå honorna förflyttades till separata rör för att lägga sina ägg. Efter 12 dagar räknades de vuxna avkommorna och statistiska beräkningar utfördes på det genomsnittliga antalet avkommor per hona för respektive linje och allel. Ett oberoende t-test visade att honornas genomsnittliga fekunditet inte skiljde sig mellan alleler (p=0,059) och en Nested ANOVA-analys indikerade att genomsnittlig fekunditet för varje linje inom respektive allel skiljde sig (p=0,023). Då även liknande studier funnit samma resultat kan det vara nödvändig att studera och eventuellt förändra den experimentella designen av metoden för att möjliggöra konkurrens mellan honor med olika genetiska förutsättningar för att kunna observera en skillnad i fekunditet baserat på honornas förmåga att konkurrera.

In meiner Dissertation untersuche ich das Paarungs- und Fortpflanzungsverhalten des Europäischen Grasfrosches (Rana temporaria) in einem evolutionären Kontext. Mein Ziel ist es zu verstehen, welche Mechanismen zur Bildung von Paaren führen, ob die Partnerwahl die Paarungsmuster erklärt, die wir beobachten können, und ob es evolutive Vorteile gibt, die sich aus der Paarung mit einem bestimmten Partner ergeben. Die Suche nach und die Konkurrenz um Paarungspartner führt zur Entwicklung verschiedener Paarungssysteme, Strategien und Taktiken, um den Reproduktionserfolg während der gesamten Lebensdauer zu erhöhen. Das Paarungsverhalten wird durch natürliche und sexuelle Selektion beeinflusst, wobei beide in unterschiedliche Richtungen wirken können. Für die meisten Individuen ist das Überleben unerlässlich, um sich so oft wie möglich zu reproduzieren, und dadurch die reproduktive Gesamtfitness zu erhöhen. Andererseits könnte ein auffälliges Verhalten bei der Fortpflanzung das Prädationsrisiko erhöhen. Der Akt der Paarung selbst kann bereits mit Risiken verbunden sein, welche sich auf die Überlebensraten auswirken können. Durch sexuelle Selektion könnten bestimmte sekundäre Geschlechtsmerkmale begünstigt werden, entweder aufgrund von Vorteilen im Wettbewerb innerhalb eines Geschlechts (intrasexuell), oder aufgrund spezifischer Präferenzen zwischen den Geschlechtern (intersexuelle Selektion). Damit sich die Partnerwahl entwickeln kann, muss der gewählte Paarungspartner Vorteile aufweisen, von denen der wählende Partner profitiert, denn die Wahl ist mit energetischen Kosten und zeitlichem Aufwand verbunden. Als Frühlaicher muss der Europäische Grasfrosch mit einem eingeschränktem Paarungszeitraum umgehen. Die Männchen konkurrieren um den Zugang zu Weibchen und es wird angenommen, dass sich Weibchen während der Paarung und Reproduktion passiv verhalten, da der hohe "Männchen-Überschuss" keine Wahl zulassen würde. Aus evolutionärer Sicht sollten Weibchen jedoch das wählerische Geschlecht sein und entscheiden mit wem sie sich paaren, da sie mehr Energie in die Eierproduktion investieren. / In my thesis, I examine the mating and reproductive behaviour of the European Common Frog (Rana temporaria) in an evolutionary context. I aim to understand which mechanisms lead to the formation of pairs, if mate choice shapes the patterns of mating that we can observe and if there are benefits derived from pairing with a specific mate. The search and competition for mating partners lead to the evolution of various mating systems, strategies and tactics to increase lifetime reproductive success. The mating behaviour is influenced by natural and sexual selection, whereby both could act in different directions. For most individuals, survival is essential in order to reproduce as often as possible to increase lifetime reproductive fitness. On the other hand, reproduction could increase predation risk due to conspicuous behaviour and risks associated with mating itself. Sexual selection could favour specific secondary sexual traits, either due to advantages in intrasexual competition, or by specific preferences of the choosy sex (intersexual selection). For mate choice to evolve, there need to be benefits associated with the chosen mating partner, because choosiness involves costs in terms of energy and time constraints during mating. As an explosive breeder, the European Common Frog has to deal with time constraints during the short breeding season. The males are competing for the access to females and it is assumed that females are passive during breeding due to a high male-biased operational sex ratio. However, from an evolutionary perspective females should be the choosy sex and should decide with whom to mate, as they invest more energy into the production of eggs.

No description available.

Environmental factors can have profound effects on the strength and direction of selection and recent studies suggest that such environment-dependent selection can be sex-specific. Sexual selection theory predicts that male fitness is more condition dependent compared to female fitness, suggesting that male fitness is more sensitive to environmental stress. However, our knowledge about the effect of environmental factors on sex-specific reproductive performance and on sex differences in the opportunity for selection is still very limited. In the present study, we investigated the sex-specific effects of diet quality (yeast deprivation and flour type) in the red flour beetle Tribolium castaneum. Specifically, we manipulated yeast supplementation in wheat and whole-wheat flour in competition assays allowing us to test for sex-specific effects of food quality (i) on reproductive success and (ii) on the opportunity for selection. Our data show that yeast deprivation in wheat flour had significantly negative effects on body mass and reproductive success of both sexes, while high-quality flour (whole-wheat flour) was able to buffer the negative impact to a large extent. Importantly, our data suggest no sex-specific effect of dietary stress on reproductive success because the magnitude of the negative effect of yeast deprivation was similar for males and females. Moreover, our study demonstrates that low food quality inflated the opportunity for selection and did not differ between sexes neither under benign nor stressful dietary conditions. We discuss the implications of our findings for the adaptation to stressful environments.

S’approvisionner en nourriture est essentiel à la survie et au succès reproducteur. Lorsque les animaux font face à des changements environnementaux brutaux, ils doivent s’ajuster rapidement à leur nouvel environnement et parfois même innover dans leur façon de s’approvisionner. Des processus comportementaux et cognitifs, tels que l’innovation et l’apprentissage, permettent aux animaux d'intégrer de nouveaux comportements à leur répertoire comportemental afin de s'adapter de façon optimale. Les performances cognitives varient entre les individus d’une même population et bien que des études récentes se soient intéressées aux causes de ce phénomène, de convaincantes évidences sont manquantes afin d’expliquer pourquoi ces variations sont maintenues. Au cours de ce mémoire, les questions des pressions de sélection s'exerçant sur les performances d’alimentation par une tâche motrice nouvelle sont abordées afin de mieux comprendre l'évolution des capacités cognitives au sein d'une population captive de diamants mandarins (Taeniopygia guttata). Nous avons tout d'abord testé si les femelles diamants mandarins modifient leurs préférences d'accouplement après avoir observé la performance d'alimentation par une tâche motrice nouvelle des mâles. Afin de déterminer si les femelles sont capables de discriminer entre les mâles sur la base de leur capacité cognitive, nous avons également évalué les performances d’apprentissage de chacune d’elles. En effet, des études ont suggéré qu’il peut être coûteux, spécialement en terme de temps, de discriminer entre des partenaires potentiels sur cette base. La généralisation d’une préférence pour un mâle performant à d’autres mâles possédant le même phénotype permettrait la réduction de ces coûts. Nous avons donc finalement testé si les femelles diamants mandarins peuvent généraliser leur préférence après avoir observé les performances d’alimentation pour une tâche motrice nouvelle d’un mâle. Nos résultats suggèrent que les femelles diamants mandarins ne peuvent évaluer les capacités cognitives d’un mâle par l’intermédiaire de traits indicateurs. Toutefois, nous avons démontré qu’une observation directe des performances d’alimentation d’un mâle guide le choix d’appariement des femelles. Également, nous avons montré que les femelles peuvent généraliser l’apparence du mâle le plus performant et utiliser cette information lors de l’évaluation de nouveaux mâles. La relation entre les performances cognitives et le choix de partenaire pourraient s’expliquer par exemple par une meilleure exploitation de l’habitat, mais nécessite des études plus approfondies. / Successful foraging is essential for survival and reproductive success. When animals face rapidly change due to climate change or anthropogenic habitat destruction, they are force to quickly adjust their behaviour such as foraging. Innovation and learning, two processes related to cognitive functions, are know to allow animals to incorporate novel behaviours into their behavioural repertoires and thus to facilitate optimal responses to environmental change. Cognitive performance vary between and within individuals and although several studies have rencently addressed the causes, convicing evidences for why inter-individual variations in cognitive performance are maintained in a population are still lacking. During my Masters, I investigated different selective pressures acting on foraging performance on a novel motor task to better understand the evolution of cognitive abilities in a captive population of zebra finch (Taeniopygia guttata). Firstly, we investigated whether female zebra finches modify their mating preferences after having observed the foraging performance of males on a novel motor task. We also assessed each bird’s learning performance in a color associative task in order to check whether females could discriminate between the two males based on their learning performance. Discriminating among mates based on their cognitive ability might be very costly for females, especially in terms of time. Therefore, one way to reduce the cost of assessing mate would be to generalize their preferences to any male with the same phenotype as the most efficient observed individual. We finaly investigated whether female zebra finches can generalize their mating preferences after having observed a male’s foraging performance on a novel motor task. Our findings suggest that female zebra finches would be unable to assess male cognitive ability indirectly via morphological traits. However our results demonstrate that direct observation of the males’ performance on a foraging task can guide female mating preferences. We also demonstrates that female zebra finches can generalize the appearance of the male that is the most efficient at solving a motor task and then use this information to assessing new males. The relationship between cognitive performance and mating preference might be mediated throught habitat exploitation for example, but requires further investigation.

Evolution is a most fascinating feature of all living things in our world. The means of organismal evolution are diverse, comprising mainly of natural and sexual selection. Both of these phenomena may contribute to evolutionary adaptations in health and disease. The questions of immunity impact on the process of sexual selection and sexual selection for increased anti-parasite resistance are wide issues of the present research in natural sciences. Their clarification requires multidisciplinarily-based investigation combining a variety of partial results into a single united paradigm. My co-workers and I have chosen several specific issues to study, in order to fill some of the important gaps of the current knowledge. We proposed the Carotenoid maintenance handicap hypothesis to point out that optimisation instead of maximisation of the carotenoid intake and ornamental display may be physiologically convenient for the individual. In Scarlet rosefinch (Carpodacus erythrinus) we confirmed the role of the carotenoid-based ornamental colouration in the process of sexual selection and investigated several partial aspects of the association between immunogenetics, immune function, health and the ornament expression and mate choice. In Zebra finch (Taeniopygia guttata) we have shown that the present way of...

O presente trabalho se propõe a compreender a variabilidade cultural humana em relação à organização da família por meio das teorias darwinianas da seleção natural e da seleção sexual, e se encontra baseado em evidências vindas da comparação de dados etnográficos. Pretende-se demonstrar que existe uma alteração marcante no comportamento de todos os envolvidos nas relações familiares quando mudam as regras de herança. Essas alterações assim ocorrem porque as regras de herança direcionam o Investimento Parental Masculino (MPI), ora ao sobrinho, ora ao filho. A hipótese apresentada é a de que a variação do MPI modifica de maneira compreensível e previsível as relações familiares em organizações matrilineares e patrilineares. A prova da hipótese será feita por meio da aplicação do método intercultural com pesquisa etnográfica nos bancos de dados do HRAF, e utilizando-se da amostra PSF. Os resultados do presente trabalho indicarão que em sociedades em que o tio tem uma relação afetuosa com seu sobrinho, o marido espanca a mulher; e quando o pai se relaciona de maneira indulgente e amigável com seu filho, irmãos e irmãs se evitam, alinhando esses fatos de maneira que possam ser coerentemente entendidos com a teoria sociobiológica. A riqueza cultural, o sem número de costumes, as tradições, os tabus, as normas explícitas ou ocultas, os aparatos de coerção de toda sorte, as regras sobre herança, casamento, dote, preço da noiva, divisão de trabalho, enfim, todas as regras observadas em todos os agrupamentos humanos podem ser explicadas à luz das seguintes teorias: a) da “seleção de consangüíneos" (Hamilton, 1963), b) do “altruísmo recíproco" (Trivers, 1971), c) do “investimento parental e seleção sexual" (Trivers, 1972), e d) do “conflito entre prole e parentais" (Trivers, 1974). / The present work proposes to account for cultural variability in connection to human family organization, through Darwin’s theories of natural selection and sexual selection. It is also founded on evidence based on the comparison of ethnographic data of diverse human societies. It intends to demonstrate that there is a significant alteration in the behavior of family members when the rules of inheritance of wealth change. These alterations are triggered because the rules of inheritance dictate the MPI – Male Parental Investment – sometimes in the nephew and sometimes in the son. The hypothesis is that this variation in MPI (male investment in either the nephew or the son) modifies family relations in matrilineal and patrilineal organizations in a predictable manner. The role played by MPI in determining family tensions will be shown by applying cross cultural analysis to the data obtained from HRAF (Human Relations Area Files), using PSF (Probability Sample Files). The results of the present work show that societies in which the uncle’s relation with his nephew is affectionate, the husband beats the wife; on the other hand, when the father’s relation with his son is indulgent and friendly, brother and sister avoid each other. This variability of behavior can be coherently explained by sociobiological theory. Cultural wealth: the innumerable customs, the traditions, the taboos, the manifest and hidden norms, the apparatus of coercion, the rules of inheritance, marriage, dowry, bride price, the division of labor, in short, all the rules observed in all human groups can be explained in the light of the following theories: a) ‘kinship selection’ (Hamilton, 1963); b) ‘reciprocal altruism’ (Trivers, 1971; Axelrod, 1981); c) ‘parental investment and sexual selection’ (Trivers, 1972); and d) ‘parentoffspring conflict’ (Trivers, 1974).

No description available.

Postcopulatory sexual selection—sperm competition and cryptic female choice—has become a major area of research over the past 40 years. Within this field there are many outstanding questions at every level of analysis, from proximate to ultimate. The fitness consequences for both sexes in the period after copulation and before fertilization are considerable, but are obscured within the female reproductive tract. Our understanding of postcopulatory mechanisms is especially sparse in taxa other than birds and insects. Nearly nothing is known in reptiles except that multiple paternity is common and widespread, and often results from long-term sperm storage across breeding seasons. We present some of the very first data on the determinants of fertilization success in the context of sperm competition in reptiles, a group that accounts for 30% of terrestrial vertebrates. In the first chapter, "Asymmetric gametic isolation between two populations of red-sided garter snakes", we discuss the use of between-population crosses to reveal gametic isolation. The effect of population density and operational sex ratios on mating systems and the speciation process has fueled theoretical debate. We attempted to address these issues using two populations of red-sided garter snakes (Thamnophis sirtalis parietalis) from Manitoba, Canada. Our study populations differ markedly in their density mating aggregations, with a 10-fold difference between them. Using microsatellite markers for paternity analysis of litters produced from within and between population crosses. We found that the population with highest aggregation density, and presumably with the highest level of sexual conflict (i.e., when the evolutionary interests of the sexes differ) over mating, was also the population that exhibited homotypic sperm precedence. The less dense population showed a distinct postcopulatory male-size advantage. We also demonstrated that sperm stored within the female over hibernation can father 20-30% of offspring in a litter. In the second chapter, "Sperm competition and mate-order effects in red-sided garter snakes", we test whether females use mate-order effects to ensure that a larger (fitter) male will sire her offspring. Does that second male should have precedence in sperm competition? We tested for second-male precedence using singly-mated females that mated with a second male. Average proportion of paternity was shared equally among the first (P₁, i.e., proportion of offspring from a litter fathered by the first male to mate) and second males (P₂) to mate, and stored sperm (P[subscript ss]). This may be a case where last male precedence breaks down with more than two males. All females were spring virgins (they had not mated that spring, but may have stored sperm from fall matings); thus sperm stored presumably from fall matings is important in this system. As the interval between matings increased P₁ increased at the expense of P[subscript ss]. As the second male to mate's copulation duration increased, P₁ also increased at the expense of P₂. This last result may indicate female influence over sperm transfer during coerced matings. Copulatory plugs (CPs) are found in many taxa, but the functional significance is debated. Male garter snakes produce a gelatinous copulatory plug during mating that occludes the opening of the female reproductive tract for approximately two days. In chapter three, "Not just a chastity belt: the role of mating plugs in red-sided garter snakes revisited", we experimentally tested the role of the CPs. In snakes, sperm are produced in the testes and delivered through the ductus deferens, and the copulatory plug is thought to be produced by the sexual segment of the kidney and conveyed through the ureter. We manipulated the delivery of the two fluids separately by ligating the ducts. We confirmed that the CP is not formed in ureter-ligated males and that sperm leaks out immediately after copulation. The CP is analogous to a spermatophore. The protein matrix contains most of the sperm which are liberated as the plug dissolves within the female's vaginal pouch. One of the fundamental principles in sperm competition is that increased sperm numbers increase the odds of winning in competitions for fertilization success and males will adjust their ejaculate relative to competition and the quality of his mate. In chapter four, "Sperm depleted males and the unfortunate females who mate with them", we detect significant among-male variation in the number of sperm ejaculated, and that male mate-order reduces sperm numbers. Male sperm numbers drop significantly from one mating to the next, and this has implications for sperm competiveness, as Thamnophis sirtalis exhibits a disassociated reproductive tactic, in that sperm stores are produced outside the breeding season, and thus cannot be replenished after mating. Interestingly, however, the on average the mobility of the sperm increased for a male's second mating. Therefore, increased sperm quality may compensate for reduced numbers in a competitive context. Further, females increase their remating rate when mating with males that are unable to deliver sperm. In chapter five, "Sexual conflict during mating in red-sided garter snakes as evidenced by genital manipulation", we revisited the CP in the context of sexual conflict. Sex-differences in optimal copulation duration can be a source of conflict, as increased copulation duration may be advantageous for males as it delays female remating. Males of many species actively guard females to prevent them from remating, and in some cases males produce copulatory plugs to prevent remating. If precopulatory choice is limited at the time of her first mating, conflict may be especially onerous to a female. The size of the plug is influenced by the copulation duration. We experimentally tested the contribution of male and female control over copulation duration. We ablated the largest basal spine on the male's hemipene and found a reduction in copulation duration and an increase in the variation of plug mass. Further, we anesthetized the female's cloaca and found copulation duration increased, which suggests that males benefit from increased copulation duration while females actively try to reduce copulation duration. Therefore, sexual conflict is manifest in divergent copulation duration optima for males and females. / Graduation date: 2013