The Silver Gull is a small gull (265 - 450g), which exhibits sexual dimorphism, with males larger than females. It has a protracted laying period of about 8 months over the winter on Penguin Island in Western Australia. The Silver Gull was studied on Penguin Island from 2000 to 2002. Completed clutches were removed from breeding pairs to induce repeat laying in order to determine the effect of increased reproductive effort on maternal body condition, egg production ability, offspring sex ratio and chick rearing capacity. Increased egg production had no significant effect on maternal body condition as measured by condition index, derived from mass divided by a measure of skeletal size. The seasonal period, divided into thirty-day intervals, had a significant impact on female condition index, with a decline in condition toward the end of the breeding season. While male condition also appeared to decline at the end of the season, this pattern was not significant. The initiation of laying varied between the three years of the study. The earliest occurred in 2000, which also experienced earlier rainfall than the later two years. Egg size and mass decreased throughout the breeding season although the number of eggs in a clutch did not decline. The size and mass of the eggs was significantly affected by the laying history of the parents, although this effect was dependent on the year in which the eggs were produced. The minimum interval required by Silver Gulls to replace a lost clutch is about 14 days. This interval increased from the start of the breeding season, but then declined toward the end, as summer was approaching. Laying interval increased significantly as the number of clutches produced by the parents also increased, up to 4 clutches in total. As more clutches were produced past this point, the laying interval became shorter. The probability of a replacement clutch being produced after clutch removal, declined as the clutch number increased and as the season progressed. Individuals that laid clutches with a larger mean mass were more likely to lay a replacement clutch. Increasing reproductive output caused a decline in the proportion of clutches that were replaced after clutch removal. The proportion of clutches that were replaced also varied between the years with the highest rates of replacement seen in 2000 which was also the year that experienced the earliest start in laying. The size of the original clutch in terms of its mean mass and volume was related to the size of the replacement clutch, but this relationship varied according to the timing of laying. During 2000 and 2001 male offspring predominated in the first two clutches produced by Silver Gulls. Further clutches that were produced demonstrated a sex ratio skewed toward females, the smaller sex in this species. Offspring sex ratio was close to equality in 2002 with very little effect caused by increased egg production. There was no effect of year, season or the laying history of the parents on hatching success. Growth rate in chicks was influenced by the year in which the chick hatched, the period during the season in which the chicks hatched, its sex and the laying history of the parents. The relationship between chick growth and the laying history, however, was complex with no consistent pattern emerging in terms of the performance of chicks from each treatment group. While the chicks from control groups generally grew faster than the chicks from manipulated parents, those individuals that were laid or raised by manipulated parents that had laid at least three clutches in total also performed well. Using the two main measures of reproductive success in the current study, egg production and chick rearing, those birds that were induced to lay multiple replacement clutches, were able to maintain a high level of condition and reproductive success. It is proposed that in the Silver Gull, only those individuals with a high level of condition continue to lay replacement clutches. If the female is unable to produce well provisioned eggs with a high chance of success, the breeding attempt is abandoned. Despite no loss of condition detected in female Silver Gulls with increasing clutch number, there was a significant shift in the offspring sex ratio toward females, indicating that strategies were in place to cope with the increased reproductive effort incurred as a consequence of repeat laying. Protracted laying in this species allows replacement of lost clutches only after maternal condition has been regained after laying.

Zooarchaeological mussel shell assemblages can be affected by an array of biases, one of which is cultural bias. Cultural biases may be exhibited in the transport of mussels from nonlocal mussel beds, and/or in preferential taste. There are a few methods used to help determine if cultural biases are at play (e.g., nestedness and detrended correspondence analysis). This thesis aims to test a new method, the life history approach, to determine if it is a viable method for assessing cultural bias in prehistoric mussel assemblages from the Tombigbee River drainage. Shell assemblages from the drainage previously have been demonstrated to not be culturally biased; therefore, these assemblages will act as a control against which to test the life history approach as a method for cultural bias assessment.

Saxicolous lichens on exposed bedrock are subjected to desiccation stress and intense light levels. Members of the genera Xanthoparmelia and Arctoparmelia are common foliose lichens on the Precambrian Shield, produce abundant sexual structures, and form part of the bedrock communities. The general goal of this thesis was to better understand the influence of community and underlying geology on three saxicolous lichens: Arctoparmelia centrifuga, Xanthoparmelia viriduloumbrina and X. cumberlandia. More specific goals were further examined in five chapters to investigate: 1) life history strategies of the three species, 2) a trade-off between fecundity and fungal secondary metabolite production; 3) an effect of substratum element composition on previously defined communities and lichen biology, 4) substratum preferences of Xanthoparmelia species, and 5) the photobiont guild hypothesis of the three species in a preliminary study. Field collections of lichens and environmental data were made in four locations on the Precambrian Shield in Manitoba and Ontario. Secondary metabolites were determined by digitally enhanced thin-layer chromatography. Fecundity was measured by number of apothecia, ascospores, and percent germination. Elements in rock samples were quantified by aqua-regia digest and inductively coupled plasma optical emission spectroscopy analysis and light microscopy was used to observe and quantify fungal germination and growth. The results showed eighty-one lichen species comprising three lichen communities; mossy rock, grassy rock, and treed rock communities. Lichen communities and fecundity were used to characterize life history strategies as competitive for Arctoparmelia centrifuga, stress tolerant for Xanthoparmelia viriduloumbrina, and ruderal generalist for Xanthoparmelia cumberlandia. A potential trade-off was reported for X. cumberlandia between sexual fecundity and a secondary metabolite. Substratum preferences were found at the genus level and element differences at the species level. Experimental evidence further supported geological preferences for the three species. Finally, the photobiont guild hypothesis could not be supported by this preliminary work. This research provides a broad overview of ecological and biological patterns found in Arctoparmelia and Xanthoparmelia species. The research forms a foundation for further studies in substratum preference and life history characterization in lichens. It can be further applied to habitat suitability modelling which may be valuable for phylogenetic context or in conservation biology of lichens. / October 2015

Between individual behavioral variation has been described in nearly every animal taxa where it has been measured. Often, these behavioral variations correlate across contexts, forming a behavioral syndrome. Despite a recent push to better understand the origins and consequences of behavioral syndromes, there still is no cohesive framework that describes this phenomenon. Here, I develop a social insect species into a model for measuring and testing behavioral syndromes at a new level of biological organization; the colony. This builds upon the rich literature describing between-colony variation in behavior and provides novel insights into the evolution of behavioral syndromes. In my first chapter I show that colonies do not vary from one another in foraging distance, nor is foraging distance directly associated with colony size. This was my first step in demonstrating that colony behavioral variation is not simply a byproduct of colony size. In chapter two, I expanded upon this finding by testing colonies both in the lab and in the field for a variety of ecologically relevant behaviors. Here, I found that there was a behavioral syndrome that reflected foraging distance, foraging effort to novel and familiar resources, response to threat and aggression. While there is a gradient of phenotypes, some colonies either travel farther to forage for food and respond more aggressively when confronted with a conspecific invader, but appear to invest less in each given incident or food source. I consider this to be more risk-tolerant; they increase their risk of external mortality for potentially larger pay-offs. On the other hand, risk-averse colonies deploy more foragers to exploit closer resources, increase their overall activity in the response to threat, but avoid travelling farther distances or aggressively engaging invaders. Additionally, there is between population variations in risk-taking phenotype. Across the western United States, colonies at more northern latitudes are more risk-tolerant than colonies at more southern latitudes. In chapter 3, I expand upon this latitudinal gradient in behavioral phenotype by investigating what ecological factors predict a colonies level of risk-tolerance. Specifically, I focused on ecological traits that reflected predation, competition, food resource availability and abiotic stress. I found that competition for nest sites and spatial clustering predicted behavioral type; colonies at high levels of nest site competition or spatial clustering were more risk-tolerant than colonies at lower levels of competition or were more spatially dispersed. In chapter 4, I used a common garden and brood transfer experiment to investigate if the relationship between the ecological environment and behavior was the result of phenotypic plasticity or local adaptation. I show that local adaptation is the most likely explanation, as colonies with more workers from the donor colony are more, behaviorally, like the donor colony than colonies with fewer donor workers. In chapter 5 I test if the risk-taking behavioral syndrome is the result of life history strategy variation. I test the growth rate and energy allocation towards either somatic effort or reproductive effort. I found that colonies which are risk-tolerant also grow faster and dedicate more energy towards reproductive effort, which is consistent with predictions built from life history theory. This body of work shows that behavioral syndromes can exist at a new level of organization, the colony, and that variation in behavioral type is the result of differential selection pressure between populations. This directly connects behavioral syndrome research to life history strategy research. As life history strategy theory is a well-understood field, this represents a true advancement in the field of behavioral syndromes.

The National Longitudinal Study of Adolescent Health was used to test predictions from life history theory. It was hypothesized that (1) in young adulthood an emerging life history strategy would exist as a common factor underlying many life history traits (e.g., health, relationship stability, and economic success), (2) both environmental harshness and unpredictability would account for unique variance in expression of adolescent and young adult life history strategies, and (3) adolescent life history traits would predict young adult life history strategy. These predictions were supported. The current findings suggest that the environmental parameters of harshness and unpredictability have concurrent effects on life history development in adolescence, as well as longitudinal effects into young adulthood. In addition, life history traits appear to be stable across developmental time from adolescence into young adulthood.

Doctor of Philosophy / Department of Psychological Sciences / Gary Brase / Although fertility decision-making has been the source of considerable theoretical and empirical investigation, the effect of several contextual variables on individuals’ fertility decision-making processes are not yet understood. For example, are individuals more strongly influenced by social forces that are informational or normative? Also, do individuals change their fertility intentions based on their current and developmental economic conditions? Further, how ‘shared’ are reproductive decisions within a couple, are males or females more likely to get what they want? This 3-study program of research used both experimental and exploratory qualitative methods to elucidate the nature of these unresolved issues within the domain of fertility decision-making. Study 1 (N = 344, M[subscript]age = 23, SD[subscript]age =6.41, 59.3% female) found that highly motivated individuals’ fertility intentions were more susceptible to informational, compared to normative messages (the opposite was true for unmotivated participants). Study 2 (N = 249, M[subscript]age = 24, SD[subscript]age =6.10, 61.4% female) found that exposure to mortality primes up-regulated fertility intentions for individuals with “fast” life history strategies, but facilitated the down-regulation of fertility intentions for individuals with “slow” life history strategies. Interestingly, resource scarcity primes were associated with the postponement of fertility plans in individuals’ with “fast” life history strategies. Study 3 (N = 120, M[subscript]age = 21, SD[subscript]age =4.96, 50% female) found that, contrary to predictions, the similarity of couples’ gender role attitudes, career-orientations, and education levels did not significantly predict the frequency of their use of statements coded as compromise and agreement or persuasion and disagreement in their discussions regarding their future reproductive plans. Findings do suggest that individuals with higher levels of education were more likely to use persuasion and disagreement statements in their child timing and number discussions with their romantic partner, indicative of greater decision-making power in that particular social exchange. Further, men and women in study 3 were equally likely to use statements coded as compromise and agreement, persuasion and disagreement, and concession when discussing both their future fertility plans as well as their future financial plans.

In addition to habitat loss and fragmentation, habitat degradation can have important consequences for biodiversity and population persistence, including effects on ecological and genetic processes beyond decreased demographic viability and the loss of genetic variation. Particularly interesting is the potential for evolutionary changes and adaptation to degraded habitats, that can affect population viability even in the short-term. Here, I explore how environmental changes after habitat degradation affect the evolutionary dynamics of populations of the rainforest cycad Zamia fairchildiana, specifically how habitat degradation affects gene dispersal, inbreeding, directional selection, and genotype-by-environment interactions, and the potential for genetic differentiation between populations. Colonies of Z. fairchildiana showed little genetic differentiation in neutral molecular markers across study sites, thus can be considered as subpopulations. Subpopulations in the disturbed habitat are experiencing different environmental conditions when compared to subpopulation in their native habitat. Disturbed-habitat subpopulations showed a faster life-history. This faster life history is associated with a weaker spatial genetic structure and higher levels of inbreeding in the disturbed-habitat subpopulations. In addition, higher light availability in the disturbed habitat seems to be a major agent of selection on traits like leaf production that have the potential to respond to selection in these subpopulations. Different traits were under selection in the native-habitat subpopulations, suggesting the potential for genetic differentiation between native and disturbed-habitat subpopulations. Genotype by environment interactions in seed germination and seedling survival, in response to light and water availability, further suggested that subpopulations can adaptively diverge between habitats, but the relative role of genetic and environmental factors, particularly maternal effects, on the magnitude and rate of genetic differentiation between subpopulations remains to be evaluated. These results suggest that habitat degradation can have important consequences for the evolutionary dynamics of populations of this cycad, not necessarily typical of habitat loss and fragmentation. This study identified factors and processes important for population persistence in degraded habitats, but population responses to habitat degradation are complex. Thus further studies and long-term experiments are required for better understanding the effects of habitat degradation on population viability.

Life history traits affect the timing and pattern of maturation, reproduction, and survival during an organism’s lifecycle and are the major components influencing Darwinian fitness. Co-evolved patterns of these traits are known as life history strategies (LHS) and variation occurs between individuals, populations, and species. The polymorphisms underlying LHS are important targets of natural selection, yet the underlying genes and physiological mechanisms remain largely unknown. Mapping the genetic basis of a LHS and subsequently unraveling the associated physiological mechanisms is a challenging task, as complex phenotypes are often polygenic. However, in several systems discrete LHS are maintained within the population and are inherited as a single locus with pleiotropic effects. These systems provide a promising starting point for investigation into LHS mechanisms and this thesis focuses on one such strategy - the Alba polymorphism in Colias butterflies. Alba is inherited as a single autosomal locus, expressed only in females, and simultaneously affects development rate, reproductive potential, and wing color. Alba females are white, while the alternative morph is yellow/orange. About 28 of 90 species exhibit polymorphic females, though whether the Alba mechanism and associated tradeoffs are conserved across the genus remains to be determined. In this thesis I primarily focus on the species Colias croceus and integrate results from lipidomics, transcriptomics, microscopy, and genomics to gain insights to the proximate mechanisms underlying Alba and Alba’s evolution within the genus. Lipidomics confirm that, consistent with findings in New World species, C. croceus Alba females have larger abdominal lipid stores than orange, an advantage which is temperature dependent and arises primarily due to mobilized lipids. Gene expression data suggests differences in resource allocation, with Alba females investing in reproduction rather than wing color, consistent with previous findings in other Colias species. Additionally, I identify a morphological basis for Alba’s white wing color. Alba females from C. croceus, an Old World species, and Colias eurytheme, a New World species both exhibit a significant reduction in pigment granules, the structures within the wing scale that contain pigment. This is a trait that seems to be unique to Colias as other white Pierid butterflies have an abundance of pigment granules, similar to orange females. I also map the genetic basis of Alba to a single genomic region containing an Alba specific, Jockey-like transposable element insertion. Interestingly this transposable element​ is located downstream of BarH-1, a gene known to affect pigment granule formation in Drosophila. Finally, I construct a phylogeny using a global distribution of 20 Colias species to facilitate investigations of Alba’s evolution within the genus. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 4: Manuscript.</p>

Pour comprendre l'origine des variations d'histoire de vie des organismes, il fautétudier et mettre en évidence les stratégies d'histoire de vie et les processus évolutifsqui les gouvernent. Ce travail de thèse a pour objectif d'étudier les stratégiesd'histoire de vie et leur conditionnement par les caractéristiques individuellesen conditions naturelles. Les stratégies d'histoire de vie sont vues comme un agencementde normes de réactions et de compromis évolutifs. Cependant, l'étude desprocessus évolutifs en milieu naturel se heurte à des problèmes d'ordre méthodologique.En effet, le suivi exhaustif au cours du temps d'individus d'une populationest difficilement réalisable, voire impossible en conditions naturelles. Les méthodesde capture-marquage-recapture permettent une observation partielle des histoires devie et des traits d'histoire de vie. Ce travail se base sur l'idée que nos observationsne sont que la partie visible de processus sous-jacents qu'il est nécessaire de prendreen compte pour ne pas biaiser nos inférences statistiques. J'utilise la modélisation àstructure cachée pour 1) séparer le processus d'observation du processus dynamiqued'intérêt, 2) modéliser les histoires de vie complètes des individus, 3) intégrer dansun cadre unique et cohérent les décisions d'histoire de vie et les compromis évolutifset 4) représenter explicitement les mécanismes sous-jacents qui génèrent nos observations.Dans ce cadre, on peut alors intégrer les théories et concepts de la biologieévolutive dans l'analyse statistique des données d'observations. J'illustre ce travailpar l'étude du conditionnement des stratégies d'histoire de vie dans une populationnaturelle de saumon Atlantique sur le Scorff (Morbihan) à partir de données de CMR.Mes résultats mettent en évidence des décisions d'histoire de vie statut-dépendanteset des compromis évolutifs qui n'auraient pas pu être mis en évidence hors du cadrede modélisation proposé. / Understanding the origin of life history variations of organisms requires studying life historystrategies and evolutionary processes that drive them. This thesis aims at studying life historystrategies under natural conditions and how they are conditioned by individual characteristics.Life history strategies are seen as a combination of reaction norms and evolutionarytrade-offs. The study of evolutionary processes in the wild faces to methodological issues.Indeed, the exhaustive monitoring of individuals over time is often impossible in the wild.Capture-mark-recapture methods allow a partial observation of life histories and life historytraits. This work was based on the idea that our observations are only the visible part ofunderlying processes that need to be accounted for to limit the risk of flawed statistical inferences.I resort to hidden structure modeling to 1) separate the observation process fromthe dynamic process of interest, 2) model the full life histories of individuals, 3) integratewithin a single and coherent framework life history decisions and evolutionary trade-offs and4) explicitly represent the underlying mechanisms that generate our observations. Withinthis framework, one can confront theories and concepts in evolutionary biology with observationaldata through appropriate statistical tools. Finally, I illustrate this work by studying theconditioning of life-history strategies in a natural population of Atlantic salmon on the Scorffriver (Morbihan) using CMR data. My results highlight status-dependent life history decisionsand evolutionary trade-offs that could not be identified without our proposed modelingframework.

Understanding the impacts of climate change is critical for improving the conservation and management of ecosystems worldwide. Ecosystems vary along a precipitation and temperature gradient, ranging from tropical jungles to arid deserts. The Great Basin is a semi-arid eco-region that is found within the western United States. Plant communities within the Great Basin range from sagebrush valleys to sub-alpine conifer forests found at high elevation areas. It is predicted that the Great Basin will experience prolonged periods of drought, more intense fires, and greater variability in average annual and monthly precipitation, all in response to changes in climate patterns. At the lower elevations, sagebrush communities are expected to experience less suitable habitat conditions, however, less is understood about vegetation response at upper elevations. Understanding forest composition and structure at these upper elevations within the Great Basin will help us better understand potential impacts from climate change. In chapter 1, we characterized Pinus longaeva (Great Basin bristlecone pine D.K. Bailey) forest structure and composition. We mapped this tree species distribution and characterized forest structure and composition using a sampling protocol that included both biophysical variables and individual tree characteristics. We collected data from 69 mixed and homogenous P. longaeva stands found within the Great Basin and Colorado Plateau. Results suggest that P. longaeva forest structure and composition exhibit high structural variability in tree characteristic measurements like density, basal area, growth rate, age, and in biophysical variables such as substrate type, slope, aspect, elevation, average monthly temperature and precipitation, latitude, and longitude. This study also found that variability in forest composition and structure in P. longaeva forests allows for greater flexibility in the breadth of life-history strategies and probable resiliency to climate change. In chapter 2 we used remote sensing images with high spatial resolution to identify 685 unique P. longaeva stands on 42 mountain ranges. Pinus longaeva was found on the White Mountains on the western edge of the Great Basin to the Colorado Plateau's Henry Mountain and West Tavaputs Plateau in the East, and from the Spring Mountains in the South to the Ruby and Spruce Mountains in the North. Stands covered 113,886 ha across the geographic distribution. A comparison between our maps and those produced by David Charlet found a total of 36% overlap of P. longaeva. We mapped 58 unique stands that the control dataset lacked and 11 stands that we did not include. We believe that this is the most comprehensive P. longaeva distribution map created to date.