Reef-building corals (Order: Scleractinia) are undergoing rapid taxonomic revision after molecular systematics disputed the relationships at all taxonomic levels within traditional classification. New morphological characters are being used to produce evolutionary relationships supported by molecular phylogenetics. While these characters are providing more congruent taxonomic relationships, their variation has not been fully explored. Additionally, phenotypic plasticity (changes in morphology resulting from environmental factors influencing the expressed phenotype despite a shared genotype) is prevalent amongst Scleractinia. In order to better understand the nature of these characters and explore their variation, I created a series of aquaria-based experiments designed to test the stability of these new morphological characters in response to differing environmental conditions. Light intensity and temperature were chosen as the environmental factors varied in these experiments on the basis of being a known trigger for environmentally-driven plasticity and their importance in calcification rate. In addition to aquaria-based phenotypic plasticity experiments I also examined a group (Family: Euphylliidae) within Scleractinia that had been divided by molecular phylogeny into two disparate groups. My research focused on morphological features viewed at magnifications observable by scanning electron microscopy (SEM) called micromorphology. Although variation in the skeletal micromorphology is observable, the new morphological characters that are used in taxonomy display only small amounts of variation caused by changing environmental conditions and were found to be stable for use in taxonomic studies. Additionally, I found a few micromorphological features distinguishing the two groups previously assigned to Euphylliidae including the shape of the septal margins and the fine-scale skeletal texture.

Mainstream evolutionary biology lacks a mature theory of phenotype. Following from the Modern Synthesis, researchers tend to assume an unrealistically simple mapping of genotype to phenotype, or else trust that the complexities of developmental architecture can be adequately captured by measuring trait variances and covariances. In contrast, the growing field of evolutionary developmental biology (evo-devo) explicitly examines the relationship between developmental architecture and evolutionary change, but lacks a rigorous quantitative and predictive framework. In my dissertation, I strive to integrate quantitative genetics and evo-devo, using both theoretical and empirical studies of plasticity. My first paper explores the effect of realistic development on the evolution of phenotypic plasticity when there is migration between two discrete environments. The model I use reveals that nonadditive developmental interactions can constrain the evolution of phenotypic plasticity in the presence of stabilizing selection. In my second paper, I examine the manner in which the genetically controlled responsiveness of traits to each other is shaped by selection and can in turn shape the phenotypic response to selection. Here, results indicate that developmental entanglement through plasticity can facilitate rapid multivariate adaptation in response to a novel selective pressure. In my final paper, I examine patterns of gene expression underlying ancestral plasticity and adaptive loss of melanin in Daphnia melanica. My results indicate that the developmental mechanism underlying ancestral plasticity has been co-opted to facilitate rapid adaptation to an introduced predator.

<p>Many prey species are able to develop different morphological structures as defence against</p><p>for example predators. Some of these structures are induced only by individuals exposed to a</p><p>predator. This phenomenon is called phenotypic plasticity. In this paper we examine whether</p><p>cuticle thickness in Leucorrhinia dubia (Odonata) larvae differed between specimens caught</p><p>in fish containing lakes and fish-free lakes respectively. We measured the thickness of the</p><p>cuticle from four different parts of the larvae; profemur, pronotum, ninth segment sternite and</p><p>ninth segment tergite. Our results showed a significantly thicker exocuticle on profemur in</p><p>larvae with a head width bigger than 4.5 mm caught in lakes with fish. The smaller larvae</p><p>showed a tendency to have thinner exocuticle on profemur in presence of fish. We discuss the</p><p>probability that the differences in exocuticle thickness on profemur could be some kind of</p><p>trade-off situation. The results also showed a tendency among the large larvae; the large</p><p>individuals from lakes containing fish had a slightly thicker exocuticle on pronotum than the</p><p>bigger individuals from fish-free lakes.</p>

Many prey species are able to develop different morphological structures as defence against for example predators. Some of these structures are induced only by individuals exposed to a predator. This phenomenon is called phenotypic plasticity. In this paper we examine whether cuticle thickness in Leucorrhinia dubia (Odonata) larvae differed between specimens caught in fish containing lakes and fish-free lakes respectively. We measured the thickness of the cuticle from four different parts of the larvae; profemur, pronotum, ninth segment sternite and ninth segment tergite. Our results showed a significantly thicker exocuticle on profemur in larvae with a head width bigger than 4.5 mm caught in lakes with fish. The smaller larvae showed a tendency to have thinner exocuticle on profemur in presence of fish. We discuss the probability that the differences in exocuticle thickness on profemur could be some kind of trade-off situation. The results also showed a tendency among the large larvae; the large individuals from lakes containing fish had a slightly thicker exocuticle on pronotum than the bigger individuals from fish-free lakes.

Phenotypic plasticity allows some animals to change their behavioural, morphological, performance, and life history traits in response to changes in environmental conditions such as the presence of predators. These changes can enhance survival, but come at a cost. Some of these phenotypic changes are predator and diet specific. I examined the effects of predator diet on the performance, life-history, and morphology of developing Northern Leopard Frog (Lithobates pipiens) tadpoles. Tadpoles were either exposed to cues from fish free water, cues from Brown Bullhead (Ameiurus nebulosus) fed a diet of trout pellets, or cues from A. nebulosus fed a L. pipiens tadpoles diet. Tadpoles exposed to predatory fish cues had smaller bodies, deeper tail fins, slower growth and development rates, and better rotational performance than tadpoles that were not exposed to predatory fish cues. Moreover, tadpoles appeared to differentiate between predatory fish diet and produced diet-specific responses in tail morphology and activity, although the latter effect was only marginally significant. Hatching, metamorphosis rates, and linear performance were not affected by the treatments. These results suggest that A. nebulosus can induce phenotypic changes in L. pipiens tadpoles, with some of these changes being diet specific.

Growth rates are highly variable, both within and among genotypes and populations. The resolution of the trade-off between size and age at maturity has been the study of extensive research by life historians. The fitness advantages of large body size and rapid development time are well supported, leading to two predictions. First, realized growth rates should be maximized. Second, growth rate will be subject to strong stabilizing or directional selection, and consequently, low genetic variability. In real populations, despite the advantages of rapid growth, animals often, in fact, grow at rates lower than the maximum rate that is physiologically possible, even in the absence of external constraints on growth rate (e.g. resource restriction or risk of predation while foraging). This implies that growth may have direct fitness consequences that are independent of the size and age of maturity, thereby lowering the optimal rate of growth. In addition to inducing plastic declines in growth rate, such costs may also select for lower intrinsic rates of growth. Despite the strong fitness effects arising from attaining a large body size quickly, variation in growth rate persists at both the phenotypic and genetic levels. The evolutionary and ecological factors contributing to this variation in growth rate are the focus of this thesis. Growth rate variation in insect model species was produced by the manipulation of resource levels during development. By comparing fitness-associated traits and body composition of adults from different treatment groups, I identify direct costs of rapid growth that could explain why animals benefit from growth at submaximal rates. In the second part of the thesis, the relationship between environmental variation and genetic variance in growth rate is investigated by quantitative genetic analysis of body size at different ages and in different growth environments. The results of this analysis suggest that environmental stress can lead to increased genetic variance via decanalization. This has consequences for the evolvability of growth rates in changing environments.

Growth rates are highly variable, both within and among genotypes and populations. The resolution of the trade-off between size and age at maturity has been the study of extensive research by life historians. The fitness advantages of large body size and rapid development time are well supported, leading to two predictions. First, realized growth rates should be maximized. Second, growth rate will be subject to strong stabilizing or directional selection, and consequently, low genetic variability. In real populations, despite the advantages of rapid growth, animals often, in fact, grow at rates lower than the maximum rate that is physiologically possible, even in the absence of external constraints on growth rate (e.g. resource restriction or risk of predation while foraging). This implies that growth may have direct fitness consequences that are independent of the size and age of maturity, thereby lowering the optimal rate of growth. In addition to inducing plastic declines in growth rate, such costs may also select for lower intrinsic rates of growth. Despite the strong fitness effects arising from attaining a large body size quickly, variation in growth rate persists at both the phenotypic and genetic levels. The evolutionary and ecological factors contributing to this variation in growth rate are the focus of this thesis. Growth rate variation in insect model species was produced by the manipulation of resource levels during development. By comparing fitness-associated traits and body composition of adults from different treatment groups, I identify direct costs of rapid growth that could explain why animals benefit from growth at submaximal rates. In the second part of the thesis, the relationship between environmental variation and genetic variance in growth rate is investigated by quantitative genetic analysis of body size at different ages and in different growth environments. The results of this analysis suggest that environmental stress can lead to increased genetic variance via decanalization. This has consequences for the evolvability of growth rates in changing environments.

Phenotypic plasticity is defined as an organism’s ability to express differentphenotypes depending on the environment. Predation is one of the key forces inecology and can indirectly cause a change of the phenotype in fish populations.Pigmentation change in order to match the background is one type of camouflage usedin fish and other organisms. Moreover, pigmentation might depend on environmentalconditions such as turbidity and water colour that affect the light spectrum and thusthe visibility in the water. The phenotypic variation in body pigmentation of perch(Perca fluviatilis L.) has rarely been studied to this date. In this study, I examined ifbody pigmentation of perch varied between different environments and betweenstructurally different habitats (littoral/pelagic). I tested long-term (phenotypicplasticity) and short-term (physiological-behavioural) changes in pigmentation byusing long-term pre-treatments and short-term aquarium experiments. Differences instructurally-diverse habitats were investigated in an extensive field study.Furthermore, experimental results were compared to data from the field. The resultsshow that pigmentation is determined by environmental factors, such as water colouror turbidity, and by structural complexity. Since fishes adapted their pigmentation totheir visual environment, pigmentation is likely used as predator avoidancemechanism in perch. Moreover, it was demonstrated that the environmentally-inducedpigmentation pattern determines the magnitude of short-term pigmentation in perch.

Marine consumers and their prey display plasticity that affects the outcomes of their dynamic interactions as well as community structure and ecosystem function. Aquatic chemical signals induced plasticity in consumers and prey from a broad range of taxonomy (phytoplankton to fishes), sizes (microscopic to macroscopic), and habitats (pelagic to benthic), and this complex plasticity strongly affected consumer-prey interactions. Two fishes,

Phenotypic plasticity, the ability of a genotype to express different phenotypes in different environments, allows organisms to cope with variation in resources and invade novel environments. Biologists have long been fascinated with the costs and tradeoffs that generate and maintain variation in plasticity, such as possible increases in brain size and delays in reproduction associated with the evolution of learning. However, the costs of plasticity vary: many studies have failed to find costs of plasticity, the degree of costs often vary with the system or environments considered, and many costs of plasticity are variable even within the lifetime of an individual. This research adopts a developmental perspective to predict the degree and incidence of costs of plasticity, using host learning in butterflies as a case study. Learning, a mechanism of plasticity that develops through a trial-and-error sampling process, should result in developmental costs and allocation of energy towards development (at the expense of reproduction). Furthermore, costs of learning should be less pronounced in environments for which organisms have innate biases and for learned traits underlain by short-term memory, relative to long-term memory (which requires more developmental re-structuring). This research found support for all three predictions across three levels of costs: behavioral costs, tissue costs, and fecundity trade-offs. Butterflies exhibited genetic variation in their ability to learn to recognize different colored hosts. Genotypes with higher proxies for long-term memory emerged with relatively larger neural investment and smaller reproductive investment. In contrast to these costs of long-term learning, proxies of short-term learning were only correlated with increased exploration of a range of possible resources (types of non-hosts) early in the host-learning process. Family-level costs of plasticity emerged from the ability to learn to locate a red host, for which butterflies do not have an innate bias. Costs of learning were also induced by learning itself: following exposure to novel (red) host environments, individual butterflies, regardless of genetic background, increased exploratory behavior, increased neural investment, and re-allocated energy away from reproduction towards other functions (e.g., flight). Considering developmental mechanisms helps to predict how costs will influence the evolution of learning and plasticity.