Marine mammals comprise a paraphyletic group of species whose current abundance and distribution has been greatly shaped by past environmental changes and anthropogenic impacts. This thesis describes molecular ecological approaches to answer questions regarding habitat requirements, genetic differentiation, and life-history trade-offs in three species of marine mammals.  The annual sea-ice dynamics of the Arctic may have large effects on the abundance and distribution of Arctic species such as the pagophilic ringed seal (Pusa hispida). Paper I describes and applies a simple molecular method for isolating and characterizing a relatively large set of single nucleotide polymorphisms (SNPs) in the ringed seal. These SNPs have been genotyped in a yet-to-be-analysed dataset which will form the basis in an assessment of the micro-evolutionary effects of annual sea-ice dynamics on ringed seal.  Current management efforts directed towards the North Atlantic fin whale (Balaenoptera physalus) are hampered by an unclear understanding of population structure. Paper II investigates the DNA basis for the high levels of genetic differentiation that have been reported in allozyme studies of the North Atlantic fin whale. We find that additional processes (at the organismal level) may have contributed to shaping the phenotype of the underlying allozyme variation. Telomeres may potentially serve as markers for determining the chronological and biological age of animals where other means of inference is difficult. Paper III describes the application and evaluation of four qPCR assays for telomere length estimation in humpback whales (Megaptera novaeangliae), finding that reliable telomere length estimates require extensive quality control. Paper IV applies the best performing qPCR assay to test whether telomeres may provide a method for genetic determination of chronological age in whales and concludes that the biological and experimental variation in telomere length estimates is too large to determine age with sufficient resolution. Finally, because telomere length and rate of telomere loss also may be affected by other cellular and organismal processes, such as resource allocation among self-maintenance mechanisms, growth and reproduction, Paper V describes the correlations between individual telomere length and rate of telomere loss, and sex, maturity status and female reproductive output. We found that the costs of reproduction in terms of telomere loss are higher in mature humpback whales than in juveniles; that reproductive costs are higher in males than females; and that differences among females tend to correlate with reproductive output. / At the time of doctoral defence the following papers were unpublished and had a status as follows: Paper 2: Submitted; Paper 3:Submitted; Paper 4: Manuscript; Paper 5:Manuscript

Phenotypic plasticity is the ability of a genotype to express different phenotypes in different environments. Despite its common occurrence, few have investigated differences in plasticity between populations, the selection pressures responsible for it, and costs and constraints associated with it. In this thesis, I investigated this by studying local adaptation and phenotypic plasticity in populations of the common frog Rana temporaria, inhibiting islands with different pool types (temporary, permanent or both). The tadpoles develop in these pools, and have to finish metamorphosis before the pool dries out. I found that the tadpoles were locally adapted both in development time and in phenotypic plasticity of development time. Tadpoles from islands with temporary pools had a genetically shorter development time than tadpoles from islands with permanent pools. The population differentiation in development time, estimated as QST, was larger than the population differentiation in neutral molecular markers (FST), which suggest that divergent selection among the populations is responsible for the differentiation. Moreover, tadpoles from islands with more variation in pool drying regimes had higher phenotypic plasticity in development time than tadpoles from islands with only one pool type present. Interestingly, increased migration among populations did not select for increased plasticity, rather it was the local environmental variation that was important. This adaptation has occurred over a short time scale, as the islands are less than 300 generations old. In temporary pools, it is adaptive to finish development before the pool dries out. This could be achieved by entering the metamorphosis at a smaller size, as a smaller size takes shorter time to reach. However, I found that there is a minimum threshold size below which tadpoles’ cannot enter metamorphosis, and that there had been no evolution of this threshold size in populations living in temporary environments. That suggests that this developmental threshold is tightly linked to physiological constraints in the developmental process. Despite their expected importance as constrains on the evolution of plasticity, costs of plasticity are often not found in nature.  However, theories of why they are absent have not been tested empirically. In this thesis, I show that fitness costs of phenotypic plasticity are only found in populations with genotypes expressing high levels of phenotypic plasticity, while in populations with low-plastic genotypes, I find costs of not being plastic. This suggests that costs of plasticity increase with increased level of plasticity in the population, and that might be a reason why costs of plasticity are hard to detect.

This thesis deals with the evolution of individuals within a species adapted to utilize specific resources, i.e. resource polymorphism. Although a well-known phenomenon, the understanding of the mechanisms behind is not complete. Considering the ruling theories, resource polymorphism is suggested to depend on severe competition for resources, the presence of open niches to be occupied leading to a reduction in competition, and disruptive selection where generalist are out-competed due trade-offs in foraging efficiency for different prey. In order to study resource polymorphism, I have used fish as the animal group in focus and the methods I have used range over laboratory experiments, field experiments, literature surveys and theoretical modelling. In my work, I have showed that different resource use induces different body shapes and that the rate of change is dependent of the encounter rate of different resources. The induced body changes partly led to increased foraging efficiency but surprisingly I did not find any trade-offs due to specialization. However, when studying predation risk in relation to resource polymorphism, my studies point towards that resource use and predation risk may act as balancing factors in such a way that disruptive selection can take place. My work also shows that population feedbacks have to be explored when considering the evolution of resource polymorphism. In pond and field experiments, I found that changes in resource densities affected the actual resource use despite previous adaptations to certain resources. By performing a literature survey, I found that cannibalism indirectly by its effect on population dynamics seems to facilitate the evolution of resource polymorphism. Modelling a size-structured population, I found that resource dynamics were stabilized, and the relative availability of different resources was levelled out due to cannibalism. Taken together, my studies strongly suggest that to understand the development of resource polymorphism in consumer populations, future studies have to include the effect of a dynamic environment both with respect to resources and predators.

Individuals within a single population can vary widely in their phenotype e.g. in their body shape. These differences are an important source of biodiversity and they can precede evolutionary divergence within a population. In this thesis we use the biological invasion of the zebra mussels into Swedish lakes to investigate which processes create or maintain phenotypic diversity within populations of the two native fish species perch and roach and the mussel itself. Both fishes have specially adapted body shapes that depend on whether they feed in the near-shore or open-water habitat of lakes. This habitat-specific divergence was more pronounced in lakes with zebra mussels, probably because resources in both habitats were in higher supply due to the mussels’ effects on the lakes. Divergence in perch body shapes between habitats was also higher in lakes with a higher water clarity, suggesting that visual conditions can affect the resource use and thus also the expression of a habitat-specific body shape. When investigating the diversity of body shapes in the mussel itself we found that mussels from one lake changed their shell shape when exposed to different predators: fish predators induced a more elongated shell shape while crayfish predators induced a rounder shell. These specific shell shapes probably serve as two alternative predator defenses protecting the mussel from predation. We conclude that the availability and use of distinct resources is an important source of diversity within populations. Abiotic conditions can play a previously underappreciated role by promoting or impairing the use of the distinct resources thus affecting the divergence. The diversity of shell shapes we found in the zebra mussels complements our study by demonstrating that not only consumer responses to resources but also resources’ responses to predators can generate phenotypic diversity.

<p>Koi and goldfish have been released into rivers in South Africa since the 1800&rsquo / s for food and sport fish and have since spread extensively. These fish are present in most of the river systems in South Africa and pose an additional threat the indigenous cyprinids in the Western Cape. Monogenean parasites of the genus Gyrodactylus are of particular concern, as their unique biology renders them a possible threat. Gyrodactylus kherulensis and G. kobayashii were identified from koi and goldfish respectively imported from Asia, Europe and locally bred fish. Morphometrics and the use of statistical classifiers, which includes univariate (ANOVA and Kruskal-Wallis), bivariate (Pearson&rsquo / s correlation) and multivariate (Principal Component Analysis) placed the two species within their respective groups. There was some intraspecific variation among the different populations collected from the various locations, especially in the hamulus and ventral bar features, but the marginal hooklets, however, remained static for both helminth species.</p>

A central goal of evolutionary biology is to understand the processes responsible for morphological, genetic and behavioural differentiation between sexes and among geographically distinct populations. Perhaps the most significant processes are genetic drift, natural selection, phenotypic plasticity and sexual selection. The main aim of this thesis was to investigate differentiation among individuals and populations of the sex-role reversed pipefish (Syngnathus typhle) and, consequently, determine which processes may be responsible for emerging patterns. This unique species is characterised by males predominately choosing amongst displaying females. In this thesis I revealed, on a microgeographic scale, morphological differentiation without genetic divergence among populations. Interestingly, females differed in size whereas the males did not. For females in this sex-role reversed species, the costs of expressing a plastic phenotype may be outweighed by the potential gains from greater survivorship, higher fecundity or increased mating success. Thus, females gain the ability to make themselves as conspicuous and attractive to males as possible in the specific environment they are living. Moreover, behavioural experiments, which focussed on describing “personalities”, reproductive investment strategies, and mate-sampling tactics, also indicated that males as well as females had the behavioural plasticity required to adjust to the environment in which they live. To this end, using video playbacks as experimental stimuli may be especially rewarding in this species. Overall, the studies in this thesis acknowledge the ability of species to fine-tune their phenotype to maximise fitness and, therefore, highlight the importance of considering patterns of differentiation in an environment-specific context.

Temperature variation affects most biological parameters from the molecular level to community structure and dynamics. Current studies on thermal biology assess how populations vary in response to environmental temperature, which can help determine how populations differentially respond to climate change. To date, temperature fluctuation effects on endothermic poikilotherms such as the common eastern bumblebee (Bombus impatiens) are unknown even though bumblebees are the most important natural pollinators in North America. A cold-acclimation experiment with B. impatiens colonies revealed individuals acclimated to 5°C or 10°C at night did not differ in resting metabolic rate, flight metabolic rate, wingbeat frequency, or morphological measurements, compared to the control group. Moreover, an infrared camera showed that all colonies maintained maximum nest temperature consistently above 36.8°C. A latitudinal sampling of flight metabolic rate and morphological measurements of B. impatiens from four locations spanning Ontario (N 45°; W 75°) to North Carolina (N 34°; W 77°) indicated no latitudinal trend in the measured variables. This study shows that bumblebees are well equipped to face a wide range of environmental temperatures, both in the short term and long term, and can use a combination of behavioural and physiological mechanisms to regulate body and nest temperatures. These results are reassuring on the direct effects of climate change on bumblebee ecology, but further studies on the indirect effect of temperature variation on North American bumblebees are required to predict future ecosystem dynamics.

Cells switch phenotypes or behaviors to adapt to various environmental stimuli. Often there are multiple alternative phenotypes, hence a cell chooses one phenotype among them, a process which we term a ``decision switch'. At the cellular level, decision switches are governed by gene regulation, hence they are intrinsically stochastic. Here we investigate two aspects of decision switches: how copy number of genetic components facilitates multiple phenotypes and how temporal dynamics of gene regulation with stochastic fluctuations affect switching a cell fate. First, we demonstrate that gene expression can be sensitive to changes in the copy number of genes and promoters, and alternative phenotypes may arise due to bistability within gene regulatory networks. Our analysis in phage-lambda-infected E. coli cells exhibit drastic change in gene expression by changing the copy number of viral genes, suggesting phages can determine their fates collectively via sharing gene products. Second, we examine decision switches mediated by temporal dynamics of gene regulation. We consider a case when temporal gene expression triggers a corresponding cell fate, and apply it to the lysis-lysogeny decision switch by phage lambda. Our analysis recapitulates the systematic bias between lysis and lysogeny by the viral gene copy number. We also present a quantitative measure of cell fate predictability based on temporal gene expression. Analyses using our framework suggest that the future fate of a cell can be highly correlated with temporal gene expression, and predicted if the current gene expression is known.

Thesis (Ph. D.)--Ohio State University, 2009. / Title from first page of PDF file. Includes bibliographical references (p. 151-168).

All organisms live in environments that are variable across space and time. Variation in selection across these environments may lead to the evolution of generalist genotypes that express phenotypic plasticity, in which one genotype can alter their phenotype (e.g., morphology, behavior, physiology) to match changes in environmental conditions, so that they may survive across a range of environments. In many egg-laying organisms that lack parental care, choosing an oviposition site is critical. The egg is an immobile stage of an animal's life cycle and mothers must balance a complex set of risks in deciding where to place their eggs. Because many biotic and abiotic factors are sources of selection on offspring survival, there is an advantage for females to evolve strategies in oviposition site selection to improve survival. This dissertation focuses on phenotypic plasticity in an offspring protection strategy that is triggered by natural enemies. In the seed beetle Mimosestes amicus (Coleoptera: Chrysomelidae: Bruchinae), females lay eggs on the outside of seed pods of legume trees and beetle larvae bore into and develop in the limited and discrete tissue of the seed. While most eggs are laid singly, I documented that beetle females superimpose eggs atop each other ("egg stacking") in response to the presence of egg parasitoids or parasitized eggs. In my first chapter, I investigated whether egg stacking is a strategy for protecting eggs from parasitism. In my second chapter, I examined female responses to variation in the number and dispersion of parasitized eggs on seed pods. Lastly, I investigated whether the intensity of stacking was affected by egg limitation (the risk of depleting her eggs before utilizing all hosts) or time limitation (losing reproductive ability or dying before laying all of her eggs). This study is unique in that it extends life history theory on egg and time costs to explain variation in egg protection behavior. The insights gained from this dissertation provide a foundation upon which we can examine how interactions among trophic levels impact the behavioral decisions made by insects that allow them to increase offspring survival.