A review on trade-offs at the warm and cold ends of geographical distributions.

Species' range limits are ubiquitous. This suggests that the evolution of the ecological niche is constrained in general and at the edges of distributions in particular. While there may be many ecological and genetic reasons for this phenomenon, here we focus on the potential role of trade-offs.

European common frog () recolonized Switzerland from multiple glacial refugia in northern Italy via trans- and circum-Alpine routes.

The high mountain ranges of Western Europe had a profound effect on the biotic recolonization of Europe from glacial refugia. The Alps present a particularly interesting case because they form an absolute barrier to dispersal for most taxa, obstructing recolonization from multiple refugia in northern Italy. Here, we investigate the effect of the European Alps on the phylogeographic history of the European common frog .

Ecological causes of fluctuating natural selection on habitat choice in an amphibian.

We estimated natural selection targeting three traits related to habitat choice in a frog (Pseudacris maculata) breeding in pools on the rocky shores of Isle Royale, Michigan, over 16 years. Our aim was to identify the form and ecological causes of annual variation in directional and correlational selection as expressed in the survival and growth of tadpoles. We found directional selection favoring early breeding, but pool choice was under weak stabilizing selection.

Demographic Processes Linked to Genetic Diversity and Positive Selection across a Species' Range.

Demography determines the strength of genetic drift, which generally reduces genetic variation and the efficacy of selection. Here, we disentangled the importance of demographic processes at a local scale (census size and mating system) and at a species-range scale (old split between population clusters, recolonization after the last glaciation cycle, and admixture) in determining within-population genomic diversity and genomic signatures of positive selection. Analyses were based on re-sequence data from 52 populations of North American collected across its entire distribution.

Adaptation to elevation but limited local adaptation in an amphibian.

We performed a reciprocal transplant experiment to estimate "parallel" adaptation to elevation and "unique" adaptation to local sites at the same elevation, using the frog Rana temporaria in the Swiss Alps. It is important to distinguish these two processes because they have different implications for population structure and ecological specialization. Larvae were reared from hatching to metamorphosis within enclosures installed in their pond of origin, in three foreign ponds at the same elevation, and in four ponds at different elevation (1500-2000 m higher or lower).

Relative importance of isolation-by-environment and other determinants of gene flow in an alpine amphibian.

A pattern of population structure called isolation-by-environment (IBE) evolves when gene flow connecting populations in different habitats is lower than expected. Although IBE is widespread, there is limited information on its magnitude compared with other factors influencing gene flow. We estimated the relative importance of IBE in the frog Rana temporaria in the Swiss Alps, a geographic context in which IBE should be relatively pronounced.

Gene Flow Limits Adaptation along Steep Environmental Gradients.

When environmental variation is spatially continuous, dispersing individuals move among nearby sites with similar habitat conditions. But as an environmental gradient becomes steeper, gene flow may connect more divergent habitats, and this is predicted to reduce the slope of the adaptive cline that evolves. We compared quantitative genetic divergence of frog populations along a 2,000-m elevational gradient in eastern Switzerland (new experimental results) with divergence along a 1,550-km latitudinal gradient in Fennoscandia (previously published results).

Predator-induced changes in the chemical defence of a vertebrate.

1. Inducible defences are ubiquitous in the animal kingdom, but little is known about facultative changes in chemical defences in response to predators, especially so in vertebrates. 2.

A Practical Guide to the Study of Distribution Limits.

Factors that limit the geographic distribution of species are broadly important in ecology and evolutionary biology, and understanding distribution limits is imperative for predicting how species will respond to environmental change. Good data indicate that factors such as dispersal limitation, small effective population size, and isolation are sometimes important. But empirical research highlights no single factor that explains the ubiquity of distribution limits.

Accumulation of Mutational Load at the Edges of a Species Range.

Why species have geographically restricted distributions is an unresolved question in ecology and evolutionary biology. Here, we test a new explanation that mutation accumulation due to small population size or a history of range expansion can contribute to restricting distributions by reducing population growth rate at the edge. We examined genomic diversity and mutational load across the entire geographic range of the North American plant Arabidopsis lyrata, including old, isolated populations predominantly at the southern edge and regions of postglacial range expansion at the northern and southern edges.

Is bigger really better? Relative and absolute body size influence individual growth rate under competition.

Models suggest that the mechanism of competition can influence the growth advantage associated with being large (in absolute body size or relative to other individuals in the population). Large size is advantageous under interference, but disadvantageous under exploitative competition. We addressed this prediction in a laboratory experiment on tadpoles competing for limited food.

Spatially heterogeneous selection in nature favors phenotypic plasticity in anuran larvae.

Theory holds that adaptive phenotypic plasticity evolves under spatial or temporal variation in natural selection. I tested this prediction in a classic system of predator-induced plasticity: frog tadpoles (Rana temporaria) reacting to predaceous aquatic insects. An outdoor mesocosm experiment manipulating exposure to Aeshna dragonfly larvae revealed plasticity in most characters: growth, development, behavior, and external morphology.

Quantitative Genetic Architecture at Latitudinal Range Boundaries: Reduced Variation but Higher Trait Independence.

Species distribution limits are hypothesized to be caused by small population size and limited genetic variation in ecologically relevant traits, but earlier studies have not evaluated genetic variation in multivariate phenotypes. We asked whether populations at the latitudinal edges of the distribution have altered quantitative genetic architecture of ecologically relevant traits compared with midlatitude populations. We calculated measures of evolutionary potential in nine Arabidopsis lyrata populations spanning the latitudinal range of the species in eastern and midwestern North America.

The relative importance of prey-borne and predator-borne chemical cues for inducible antipredator responses in tadpoles.

Chemical cues that evoke anti-predator developmental changes have received considerable attention, but it is not known to what extent prey use information from the smell of predators and from cues released through digestion. We conducted an experiment to determine the importance of various types of cues for the adjustment of anti-predator defences. We exposed tadpoles (common frog, Rana temporaria) to water originating from predators (caged dragonfly larvae, Aeshna cyanea) that were fed different types and quantities of prey outside of tadpole-rearing containers.

Isocline analysis of competition predicts stable coexistence of two amphibians.

We investigated the interaction between larvae of two anuran amphibian species (Rana temporaria and Bufo bufo) to test models of two-species competition. The study had a response surface experimental design with four replicates, each consisting of 24 density combinations. Larval performance-and, by assumption, change in population size-was defined by a linear combination of survival, growth, and development.

Permeability of the landscape matrix between amphibian breeding sites.

For organisms that reproduce in discrete habitat patches, land cover between patches (known as the matrix) is important for dispersal among breeding sites. Models of patchy populations often incorporate information on the permeability of the matrix to dispersal, sometimes based on expert opinion. I estimated the relative resistance to gene flow of land cover types and barriers using F(ST) calculated from microsatellite markers in two amphibians, within an 800-km(2) area in northern Switzerland.

Phenotypic plasticity alone cannot explain climate-induced change in avian migration timing.

Recent climate change has been linked to shifts in the timing of life-cycle events in many organisms, but there is debate over the degree to which phenological changes are caused by evolved genetic responses of populations or by phenotypic plasticity of individuals. We estimated plasticity of spring arrival date in 27 species of bird that breed in the vicinity of an observatory in eastern North America. For 2441 individuals detected in multiple years, arrival occurred earlier during warm years, especially in species that migrate short distances.

Non-interactive multiple predator effects on tadpole survival.

Interactions among and within three species of predators were estimated in terms of their effects on prey survival using short-term predation experiments. The prey were tadpoles (Rana temporaria), and the predators were dragonfly larvae (Anax imperator), newts (Triturus alpestris), and backswimmers (Notonecta glauca). Mortality rate per predator imposed by Triturus and Notonecta did not decline with predator density, whereas the predation rate of Anax was strongly reduced when the number of predator individuals increased.

Predator-induced changes in metabolism cannot explain the growth/predation risk tradeoff.

Defence against predators is usually accompanied by declining rates of growth or development. The classical growth/predation risk tradeoff assumes reduced activity as the cause of these declines. However, in many cases these costs cannot be explained by reduced foraging effort or enhanced allocation to defensive structures under predation risk.

A comparative test of the adaptive plasticity hypothesis: relationships between habitat and phenotype in anuran larvae.

The hypothesis that phenotypic plasticity is maintained by divergent natural selection acting across different environments predicts that populations and species exposed to highly variable environments will express high levels of plasticity. I tested this prediction by measuring the behavioral and morphological responses to aeshnid dragonfly larvae of 16 tadpole species and asking whether predator-induced plasticity is greater in species that experience more variable densities of predators in nature. Tadpole phenotypes were measured in a series of similar experiments in outdoor artificial ponds carried out over a 9-yr period.

Body size, competitive interactions, and the local distribution of Triturus newts.

1. Pairs of European Triturus newt species of similar size tend not to co-occur syntopically, suggesting that similarity in body size is associated with competitive interactions that prevent coexistence. I tested this hypothesis with an experiment involving larvae of four species in 675-L artificial ponds.

The change in quantitative genetic variation with inbreeding.

Inbreeding is known to reduce heterozygosity of neutral genetic markers, but its impact on quantitative genetic variation is debated. Theory predicts a linear decline in additive genetic variance (V(A)) with increasing inbreeding coefficient (F) when loci underlying the trait act additively, but a nonlinear hump-shaped relationship when dominance and epistasis are important. Predictions for heritability (h2) are similar, although the exact shape depends on the value of h2 in the absence of inbreeding.

Genomic compatibility occurs over a wide range of parental genetic similarity in an outcrossing plant.

The theory of inbreeding and outbreeding suggests that there is a hump-shaped relationship between the genetic similarity of sexually reproducing parents and the performance of their offspring. Inbreeding depression occurs when genetic similarity is high, whereas hybrid breakdown is expected when genetic similarity is low. Between these extremes, the effect of genetic similarity on fitness is unclear.

A threefold genetic allee effect: population size affects cross-compatibility, inbreeding depression and drift load in the self-incompatible Ranunculus reptans.

A decline in population size can lead to the loss of allelic variation, increased inbreeding, and the accumulation of genetic load through drift. We estimated the fitness consequences of these processes in offspring of controlled within-population crosses from 13 populations of the self-incompatible, clonal plant Ranunculus reptans. We used allozyme allelic richness as a proxy for long-term population size, which was positively correlated with current population size.

NATURAL SELECTION FOR ENVIRONMENTALLY INDUCED PHENOTYPES IN TADPOLES.

Models suggest that phenotypic plasticity is maintained in situations where the optimal phenotype differs through time or space, so that selection acts in different directions in different environments. Some empirical work supports the general premise of this prediction because phenotypes induced by a particular environment sometimes perform better than other phenotypes when tested in that environment. We have extended these results by estimating the targets of selection in Pseudacris triseriata tadpoles in environments without predators and with larval Anax dragonflies.