<p> Many floral characters have can affect plant fitness by altering the frequency or efficiency of pollinator visits. More frequent pollinator visitation may lead to larger pollen loads and greater outcrossing rates, which in tern may lead to greater numbers of offspring, with higher genetic variability and less inbreeding depression. Mating systems may moderate the strength of natural selection on traits that attract pollinators through its effects on inbreeding depression, and the ability to self-pollinate autogamously. Natural selection is expected to be stronger in largely outcrossing systems where inbreeding depression is high, but reduced in largely selfing systems where deleterious alleles have been purged.</p> <p> Here I investigate the role of floral display height in attracting pollinators, and its consequences for plant fitness in two closely related monocarpic herbs (Hesperis matronalis and Alliaria petiolata) with different mating systems. Though others have looked at height as a potential trait attracting pollinators, no one has yet clearly separated the effects of floral display height from that of plant size. I experimentally altered plant height independently of plant size by placing potted plants into either elevated or non-elevated treatments and observed pollinator foraging behaviour and reproduction.</p> <p> Pollinators preferred taller floral displays in the predominant outcrosser Hesperis matronalis. Elevated plants produced more seeds than control plants although their seeds were smaller, reflecting a strong seed size-number trade-off. Path analysis showed seed number was affected directly by plant height, presumably though greater pollinator visitation, and indirectly by plant mass through increased fruit number.</p> <p> In Alliaria petiolata, a predominant self-pollinating species, pollinators visited taller displays in field observations and experimentally elevated plants had significantly greater seed per fruit. Experimentally outcrossed and self-pollinated plants showed no difference in seed production, but offspring quality was significantly higher in outcrossed plants for early traits. Measures of maternal fitness increased significantly with both maternal and seed mass. For individual seeds, seed mass was the most important factor in seed fitness, where larger seeds germinated sooner, more frequently, and had higher survivorship than smaller seeds, regardless of pollination treatment. Competition strongly affected all seedling traits, with increased density having a significant negative effect on seedling mass, total leaf area, and maximum leaf length; and, a positive effect on hypocotyl length and petiole length. Seedling mass at harvest was not significantly affected by elevation treatment, seed mass, or maternal mass, however, this may have been due to lower competition brought about by low germination and survival; thus, competition did not accentuate the effect of maternal size or elevation treatment on seedling. Taken together, the results of these two studies suggest the importance of height, in attracting pollinators, independent of size in both species. The consequences of pollinator attraction were highly significant in the predominant outcrosser Hesperis matronalis, while maternal size and seed size played more important roles in the fitness of the predominant selfer Alliaria petiolata.</p> / Thesis / Master of Science (MSc)

Patterns of heterospecific matings are important to understand in order to learn about speciation. I proposed a classical signal detection model as a representation of heterospecific mating patterns in both sexes. From the model I proposed that males would be able to achieve successive heterospecific mating success, as those males were more similar to conspecifics. I also proposed that restrictive females that rejected heterospecific matings would also be more likely to reject conspecific matings. I used the sibling species Drosophila pseudoobscura and D. persimilis to examine heterospecific mating patterns in males and females. I found that males previously accepted as heterospecific mates were more likely to achieve heterospecific matings than males that had previously been rejected as heterospecific mates. This was most likely due to decreased courtship by rejected males. Males rejected heterospecifically but mated conspecifically also were less likely to achieve heterospecific matings than males that had initially been accepted as heterospecific mates. This indicated that the same males were able to achieve successive heterospecific matings, despite rejected males also having had initial mating success. This was again likely influenced by decreased courtship in rejected males. Furthermore, I found that females that rejected heterospecific males were also more likely to reject conspecific males than naive females. There was also a non-significant trend of females that previously accepted heterospecific males were more likely to again accept heterospecific males than females that had previously rejected heterospecific males. / Thesis / Master of Science (MSc)

The “risk allocation hypothesis” can predict how prey species will respond to predation risk, balancing vigilance with potentially risky behaviours. In order to maximize fitness, an optimal behavioural repertoire can evolve to respond to predation threat to allow for both survival and future fitness gains. High locomotor activity and time spent engaging in mating behaviours are expected to put Drosophila melanogaster at a greater risk to predation. With direct predator exposure, Drosophila are predicted to reduce activity and mating, which over years of exposure, will be reflected in evolved behavioural traits and evolved changes in the genome. Predation as a selective force shows alterations in flies genomes of experimentally evolved populations. Locomotor activity was found to be reduced in the presence of zebra jumping spiders (Salticus scenicus), presumably due to these spiders as visual hunters, using movement to detect prey. This behaviour is reflected in populations of Drosophila that have been constantly under selection by predators. Flies evolved with spider predators or mantid predators (Tenodera aridifolia sinensis) showed reduced locomotor activity when no predators are present. Interestingly, while alterations are seen for locomotory activity, the presumed risky behaviours of courtship and mating did not show an evolved response. Wild caught populations under threat from spiders, as well as the evolved populations when no predators were present showed no alterations in courtship or copulation behaviour. It appears that although there may be potential risks associated with mating behaviours, the benefits to future fitness when mating outweigh the potential costs from predation risks in Drosophila. / Thesis / Master of Science (MSc) / Predation is a profound selective force, with many anti-predator adaptations seen throughout the diversity of life. Antipredatory behavioural adaptations must balance immediate and future fitness effects, to maximize overall fitness. In Drosophila melanogaster, research into natural ecology and role of predation is generally lacking for behavioural & evolutionary studies. I will discuss research on the influence of predator exposure on Drosophila behaviours, as well as the evolution of Drosophila behaviours and genomes through experimentally evolved populations of Drosophila. While predation risk has resulted in changes in Drosophila locomotory activity, predation has not altered mating behaviours.

Despite decades of effort in developing evidence-based treatments, drug addiction remains one of the most problematic and enduring public health crises. Developing a new generation of theoretically-derived interventions constitutes an important clinical and scientific gap that, if addressed, may open innovative treatment opportunities. Based on the Reinforcer Pathology theory, altering the temporal window over which reinforcers are integrated (i.e., measured by delay discounting) would alter drug valuation and consumption. The first investigation—in 2 separate studies— test the Reinforcer Pathology theory by examining the effect of expanding and constricting the temporal window of integration using two mating narratives (long-term and short-term relationships, respectively) on cigarette valuation among cigarette smokers. The second investigation, test the Reinforcer Pathology theory by assessing the effect of remotely delivered Episodic Future Thinking (EFT) narratives (expands the temporal window) on real-world alcohol consumption among individuals with alcohol use disorder (AUD). Together, these investigations supported the Reinforcer Pathology theory and demonstrated its relevance for understanding and intervening in addiction. The current findings provide scientific justification to further investigate Reinforcer Pathology based interventions that expand the temporal window to change drug valuation and consumption. The construction of multi-component treatments that incorporate Reinforcer Pathology based interventions to systematically alter the temporal window may provide a novel intervention to reduce alcohol consumption. / Doctor of Philosophy / The following studies provide evidence that altering the temporal widow (how far in the future one can imagine and integrate into the present) would alter drug valuation. In the following studies, we used narratives describing long-term or short-term mating relationships (Study 1) and Episodic Future Thinking (EFT; represents one's capability to pre-experience the future; Study 2) to alter the valuation of cigarettes and alcohol, respectively. In the first study, cigarette smokers who read and vividly imagined long-term romance relationship narrative (expands the temporal window) valued cigarettes less than control (imagined looking for a lost key). In contrast, those who read and vividly imagined a short-term sexual encounter (shortens the temporal window) valued cigarettes more than controls. The second study employed EFT (expands the temporal window) as a strategy to reduce alcohol consumption, in real-world settings, over two weeks in individuals with alcohol use disorder. The study found that expanding the temporal window using EFT reduced alcohol consumption. Together, these two studies provide support to employing interventions that extend the temporal window to change drug valuation and consumption. The construction of multi-component treatments that incorporate interventions expanding the temporal window may reduce drug consumption.

Gravimetric analysis was used to determine the release rates and longevities of several designs of pheromone dispensers for mating disruption of leafrollers and codling moth, Cydia pomonella (Linneaus). Release rates were described by linear equations for at least four months, but by the end of the season release rates tended to become erratic. Biocontrol’s red-brown codling moth dispenser lasted for up to four months, and one application of the dispenser in early May should control codling moth for the entire season in Virginia apple orchards. Ecogen and Hercon leafroller dispensers lasted for a shorter time than the codling moth dispensers and would require two applications per season to provide the best control of their target pests. In a commercial northern Virginia apple orchard, the effects of pheromone trap height and pheromone dispenser height on captures of the tufted apple bud moth, Platynota idaeusalis (Walker), were studied. Low traps (2.0 m) were more sensitive than high traps (4.5 m) for monitoring tufted apple bud moth. The number of moths caught in 1994 and 1995 in a two-hectare mating disruption plot with pheromone dispensers placed in the upper third of the tree was not significantly different from the number caught in a plot with pheromone dispensers placed at head height. Fruit damage was very high in both pheromone plots in 1994, but by 1995 it appeared that mating disruption was able to reduce fruit damage due to leafroller larvae. / Master of Science

Considering the high reproductive investment of the social male and the cost to the female of losing this benefit by soliciting copulations outside the social pair bond, it is expected that most raptor populations would exhibit low to no occurrence of extra-pair paternity (EPP). This holds true for the majority of raptor species studied to date with only one exception of an urban Cooper's hawk (Accipiter cooperii) study which reported an unexpectedly high extra-pair young frequency of 19.29%. In our study we examined the frequency of EPP within a red-shouldered hawk (Buteo lineatus) population residing in the suburban/urban matrix of southwest Ohio. During the breeding seasons of 2018 and 2019, 181 breeding age and nestling individuals were color-banded and sampled for genetic analysis using nine microsatellite loci. After genotyping a total of 40 broods (with at least two nestlings per brood) and both presumptive parents of each brood, no clear evidence of EPP was detected. However, at one nest site, the entire brood of four chicks was not sired by the adult male observed during the courtship period, nor another adult male observed tending the chicks later in the season. We suspect that this particular nest represented two instances of rapid mate replacement rather than extra-pair fertilization by a third unsampled male, because none of the chicks were sired by either of the two adult males observed at the nest. We also reviewed potential factors contributing to our finding of overall genetic monogamy in our study population in comparison to other raptor taxa EPP studies. Our results suggested that factors other than habitat composition alone play an important role in determining the type of breeding strategy exhibited by different raptor populations.

The drivers of fidelity and divorce of pair-bonded individuals, along with their fitness consequences, are of great interest as they influence mating systems, population structure and productivity, and gene flow. Socially monogamous birds offer an ideal opportunity to study divorce since they show great variability in the extent to which pair bonds are maintained. However, there has been little consensus as to whether divorce is a behavioural adaptation to improve a mating situation, or a consequence of other processes. Moreover, the biological and ecological correlates of fidelity are difficult to address because previous work has been based on indirect and potentially biased methods. Finally, in terms of process, the link between the process of mate choice and subsequent mating decisions has been largely inaccessible to study. My doctoral thesis addressed these significant gaps in our understanding of cause, process and consequence in the formation and dissolution of pair bonds in socially monogamous birds. I accomplished this in three principal ways. First, I conducted a robust phylogenetic meta-analysis on 84 studies across 64 species to assess the existing empirical evidence that divorce in socially monogamous birds is adaptive (in terms of breeding success). This analysis revealed that divorce is, in general, adaptive as it is both triggered by relatively low breeding success and leads to improvement in success. Next, I developed a novel probabilistic multievent capture–mark–recapture framework that provides joint estimates of survival and fidelity while explicitly accounting for imperfect detection, capture heterogeneity, and uncertainty in pair status. By applying this model to breeding data on a wild great tit population I showed that birds that remain faithful to their partner exhibit higher survival rates and are more likely to remain faithful in the next breeding season than do birds that change partners. Subsequently, I confirmed the generality of a survival benefit by applying the model to breeding data on other tit populations. Then, by applying the model to data from a population of mute swans, I showed that fidelity decreases the likelihood of skipping breeding and mortality in this long-lived species, and that these effects depended on age, individual quality, and immigration status. Finally, I investigated how the timing of pair formation influences breeding success and divorce probability using five years of data on the over-winter social behaviour of great tits. I showed that early pair formation had a positive effect on fitness components, influencing the likelihood of divorce only indirectly, through breeding success. Further, my work revealed that males, but not females, with higher numbers of the female associates in winter, and males whose future breeding partners were ranked low amongst these, divorced more often. My research makes a significant contribution to our understanding of divorce and fidelity, and generates a number of important implications for future studies. First, my work establishes that divorce is adaptive for breeding success. Second, my results highlight that survival is an important (and likely, widespread) fitness consequence of pairing decisions. Third, I provide a novel statistically rigorous modelling framework for estimating fidelity-rates and testing hypothesis about fidelity that overcomes many of the inherent biases in traditional estimates. Fourth, it provides the first evidence for a selective advantage of early pair formation in wild, thus highlighting that there are benefits to pair familiarity that manifest via social associations of individuals prior to breeding. Finally, my work reveals the selective pressures operating via the social environment can ultimately influence the mating strategies individuals adopt.

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.

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

Mating behaviour of cockroaches is spectacular phenomenon with three different mating patterns traditionally distinguished. The most common and probably ancestral is type A - female climbing. Then it is type B - male climbing and type C - no climbing. Mating type was described in around 30 Blaberidae species. As it is the only cockroach family where all three types are present, it is an interesting model group for studying evolutionary questions. The main goal of the thesis was detail exploration and description of mating sequences in 21 Blaberidae species, 17 of which haven't been examined in this aspect before. An intrageneric diversity in mating patterns (genus Pycnoscelus) was described for the first time. Moreover, in case of P. tenebrigera, it was not possible to assign its mating pattern to any of the three traditional types. Multidimensional statistical analysis and biological interpretation of qualitative behavioural elements revealed two well separated clusters of mating sequensces. Consequently, we propose to distinguish only two types of mating pattern - ancestral type (identical with type A) and derived type. We also discussed the relationship between wing reduction and occurence of derived mating types. Evaluation of quantitative behavioural elements revealed positive correlation between...