For evolution to occur over time, it is necessary for animals and plants to show phenotypic variation. If the individuals within populations of a species do not show observable differences among themselves, there will be a lack of driving force for natural selection to act on and decide which characteristic gets inherited from one generation to the next. It is hence important to study phenotypic variation, especially against environmental gradients such as latitude and altitude, which gives us an insight into the pattern of change according to essential factors such as temperature and length of seasons. The latter would impose time constraints on growing populations, leading to periods of unfavourable conditions limiting their growth and development. In many cases, such organisms would tend to compensate for the period of slow growth and catch up to the others that did not have to endure the same situation and grow to the same size as them. This study aims to find differences in three key larval life-history traits of the moor frog (Rana arvalis), namely metamorphic mass, larval period, and growth rate, and find what kind of pattern is observed in case of these phenotypic variations. It also aims to find differences in the strength of compensatory response between populations from lower and higher latitudes. A common garden experiment was conducted with populations originating from both sides of the Baltic Sea, ranging from southern Sweden and Latvia to central Finland and northern Sweden. It was expected that the northern populations would grow faster and show a countergradient variation pattern since they are faced with more strict time constraints at higher latitudes, but in most cases, we observed a co-gradient pattern, wherein the environmental effect amplifies the individual’s genetic predisposition instead of opposing it. We also observed a stronger compensatory response in the northern populations as compared to their southern counterparts. Effects of climate change and subsequent rise in temperatures making the environment unpredictable over time could be used to speculate about the reason behind the results obtained. Epigenetics could also be used as an approach to study long lasting changes in an organism’s gene expression to make it adapt better to changing conditions and hence show different patterns of variation from studies in the past. Studying such changes, expected or not, is important to keep up with the needs of the species that require conservation, and will help conservation biologists to formulate strategies that would be effective even in the face of constant change in the world.

<p>The main aim of this work was to identify local adaptation processes in amphibian populations, thereby improving the general understanding of genetics and mechanisms behind the evolution and maintenance of biological diversity. Phenotypic and genetic variation in life-history traits was studied within and between populations common frog (<i>Rana temporaria</i>) populations along a 1600 km transect from southern Sweden to northern Finland.</p><p>Embryonic and larval development and growth was investigated both under field and laboratory conditions. The results suggest ample genetic diversity in larval life-history traits among Fennoscandian common frog populations. Larval developmental rate along the gradient has evolved a countergradient variation pattern of genotypes and phenotypes as indicated by the positive relationship between developmental rate and latitude under laboratory conditions and the lack of such a relationship in the field. The data suggest that this pattern has evolved because of time constraints due to decreasing length of growth season with latitude. Neither field-caught adults nor laboratory raised larvae displayed a linear latitudinal size cline as expected from the so called Bergmanns rule. Rather, size increased towards the mid-latitude populations and decreased thereafter, indicating that body size is a product of direct environmental induction or a trade-off with other life-history characters. Age and size at hatching showed no consistent latitudinal pattern, indicating that the embryonic stage is not as time constrained as the larval stage.</p><p>A large part of the variation in age and size at metamorphosis among populations was due to additive genetic effects. However, small, but significant maternal effects, mostly due to variation in egg size and non-additive genetic effects also contributed to among population variation. A comparison of divergence in presumably neutral molecular genetic markers (F_ST) and quantitative characters (Q_ST) revealed that although both estimates of divergence were relatively high, estimates of Q_ST was generally higher than those of F_ST, indicating that the genetic variation observed in larval traits is primarily a result of natural selection rather than genetic drift. Hence, our results reinforce the conclusion that intraspecific genetic heterogeneity in the young northern European ecosystems may be more widespread than previously anticipated</p>

The main aim of this work was to identify local adaptation processes in amphibian populations, thereby improving the general understanding of genetics and mechanisms behind the evolution and maintenance of biological diversity. Phenotypic and genetic variation in life-history traits was studied within and between populations common frog (Rana temporaria) populations along a 1600 km transect from southern Sweden to northern Finland. Embryonic and larval development and growth was investigated both under field and laboratory conditions. The results suggest ample genetic diversity in larval life-history traits among Fennoscandian common frog populations. Larval developmental rate along the gradient has evolved a countergradient variation pattern of genotypes and phenotypes as indicated by the positive relationship between developmental rate and latitude under laboratory conditions and the lack of such a relationship in the field. The data suggest that this pattern has evolved because of time constraints due to decreasing length of growth season with latitude. Neither field-caught adults nor laboratory raised larvae displayed a linear latitudinal size cline as expected from the so called Bergmanns rule. Rather, size increased towards the mid-latitude populations and decreased thereafter, indicating that body size is a product of direct environmental induction or a trade-off with other life-history characters. Age and size at hatching showed no consistent latitudinal pattern, indicating that the embryonic stage is not as time constrained as the larval stage. A large part of the variation in age and size at metamorphosis among populations was due to additive genetic effects. However, small, but significant maternal effects, mostly due to variation in egg size and non-additive genetic effects also contributed to among population variation. A comparison of divergence in presumably neutral molecular genetic markers (FST) and quantitative characters (QST) revealed that although both estimates of divergence were relatively high, estimates of QST was generally higher than those of FST, indicating that the genetic variation observed in larval traits is primarily a result of natural selection rather than genetic drift. Hence, our results reinforce the conclusion that intraspecific genetic heterogeneity in the young northern European ecosystems may be more widespread than previously anticipated

Altitudinal gradients offer attractive opportunities for studies of population differentiation in response to environmental heterogeneity. In this thesis, I examined population differentiation along altitudinal gradients by combining common-garden experiments with field studies and experiments in alpine, subalpine and boreal populations of the perennial herb Solidago virgaurea. More specifically, I determined whether leaf physiology in terms of nitrogen concentration and resorption, flowering phenology, flower production and reproductive effort vary along altitudinal gradients. Nitrogen concentration in green leaves were higher in alpine than in subalpine and boreal populations. These differences persisted when plants were grown from seeds in a common-garden experiment at two sites, suggesting that the differences have a genetic component. There was mixed support for a trade-off between maximized carbon gain through the maintenance of high nitrogen concentration, and minimized nitrogen loss through high resorption. In their natural habitats alpine populations began flowering later than subalpine populations, but this difference was reversed when plants were grown in a common environment. This suggests that genetic differences among populations counteract environmental effects and reduce phenotypic variation in flowering time among populations. Flowering time thus shows countergradient genetic variation in S. virgaurea. In a common-garden experiment, boreal populations produced more flowers and had a higher reproductive effort than subalpine and alpine populations indicating habitat-specific genetic differences in reproductive allocation. In a field study, which included three populations, seed set was close to zero in the alpine population, intermediate in the subalpine population, and high in the boreal population. Experimental flower removal showed that seed production was associated with a considerable cost in terms of reduced flowering propensity the following year, but did not support the hypothesis that a large floral display is important for pollination success.

As the earth’s climate changes due to anthropogenic emissions, it has increasingly become an imperative within the ecological community to understand existing species adaptations to climate change. Much focus has been paid to how a species might react to climate change, but the role of locally adapted traits and responsible environmental mechanisms have received less attention. Quantifying how sublethal (e.g. growth rates) and lethal (e.g. thermal tolerance) trait performance vary between populations can thus improve our understanding of how populations, and the entire species, will react to climate change. Here, I quantified the spatial patterns of performance of several traits in populations of the predatory marine snail Urosalpinx cinerea from across two thermal gradients on the Pacific and Atlantic coasts of North America. In chapter 2, I quantified local adaptation and plasticity of thermal tolerance, warming tolerance, and developmental traits of Urosalpinx. I found that while low latitude populations have evolved higher thermal tolerance than their low latitude counterparts, they also demonstrate negative plasticity in response to higher acclimation temperatures. This is likely a result of low latitude population adaptation to cooler developmental conditions. Further, low latitude populations live in environments much closer to their thermal maxima than high latitude counterparts, resulting in higher climate sensitivity in low latitudes. In chapter 3, I quantified growth and consumption rates of Urosalpinx via a common garden experiment. I found evidence for a novel pattern of trait adaptation, wherein high latitude populations tended to have higher trait performance at higher thermal optima than low latitude counterparts. This can be attributed to the maximizing of growth rate during short growing seasons at high latitudes. Together, these results demonstrate that local adaptation in endemic across two traits in Urosalpinx. I demonstrate that these traits tend to be adapted to aspects of the environment directly related to aspects of Urosalpinx phenology, and not to environmental means as is commonly assumed. These insights suggest that models of organismal performance under climate change must consider not only the potential for local adaptation in populations, but also the aspects of the environment to which these populations are evolved.

<p>  </p> <p>Evolution is the foundation for all of biology. However, our approaches and understanding of evolution—simply, the change of allele frequencies from one generation to the next—have themselves evolved over time. In this dissertation I explore multiple approaches to understand evolution and the consequences of evolution across variable scales and study organisms. First, I use meta-analytic techniques and Bayesian hierarchical models to investigate the phenotypic consequences of two forms of cryptic local adaptation, co- and countergradient variation, by leveraging a decades-old quantitative genetics approach (Chapter 1). I find large effects for both co- and countergradient variation, however they are obscured in natural settings by concurrent large environmental effects. I also show that these large effects are ubiquitous across phenotypic traits, organisms, and environmental gradients, suggesting that while similar phenotypes may be the evolutionary end point, the mechanisms to achieve those phenotypes likely vary. In the following chapter I explore the rapid evolution of a unique and understudied species introduction, pink salmon (<em>Oncorhynchus gorbuscha</em>) in the Great Lakes. Pink salmon were introduced into Lake Superior in a single introduction event and have broken two obligate life histories, anadromy (though they treat the Great Lakes like surrogate oceans) and their fixed two-year life cycle, making them ripe subjects for contemporary evolution. Using whole-genome sequence data, I first investigate the effects of a genetic drift in the form of a bottleneck at introduction and characterize the subsequent loss of genetic diversity (Chapter 2). I show that despite a large loss of genetic diversity, pink salmon also rapidly adapted to their novel environment based on signals of putative selection across numerous regions of the genome, particularly in a period gene associated with their daily circadian clock (<em>per2</em>). Next, I explore how genome structure likely aided adaptation by pink salmon to the Great Lakes, providing evidence that a supergene (~29 Mbp) containing an inversion on chromosome 10 swept to near fixation in the Great Lakes (Chapter 3) and likely aided in osmoregulatory adaptation to this novel environment. Finally, I end with a short perspective chapter (Chapter 4) where I highlight potential future research directions for each of the previous chapters. Together, this research investigates the drivers and consequences of evolution across multiple scales and shows the powerful effect of genetic drift and genetic adaptation in shaping the genomic and phenotypic attributes of populations.</p>