Toxicity and population structure of the Rough‐Skinned Newt (Taricha granulosa) outside the range of an arms race with resistant predators

Species interactions, and their fitness consequences, vary across the geographic range of a coevolutionary relationship. This spatial heterogeneity in reciprocal selection is predicted to generate a geographic mosaic of local adaptation, wherein coevolutionary traits are phenotypically variable from one location to the next. Under this framework, allopatric populations should lack variation in coevolutionary traits due to the absence of reciprocal selection. We examine phenotypic variation in tetrodotoxin (TTX) toxicity of the Rough‐Skinned Newt (Taricha granulosa) in regions of allopatry with its TTX‐resistant predator, the Common Garter Snake (Thamnophis sirtalis). In sympatry, geographic patterns of phenotypic exaggeration in toxicity and toxin‐resistance are closely correlated in prey and predator, implying that reciprocal selection drives phenotypic variation in coevolutionary traits. Therefore, in allopatry with TTX‐resistant predators, we expect to find uniformly low levels of newt toxicity. We characterized TTX toxicity in northwestern North America, including the Alaskan panhandle where Ta. granulosa occur in allopatry with Th. sirtalis. First, we used microsatellite markers to estimate population genetic structure and determine if any phenotypic variation in toxicity might be explained by historical divergence. We found northern populations of Ta. granulosa generally lacked population structure in a pattern consistent with northern range expansion after the Pleistocene. Next, we chose a cluster of sites in Alaska, which uniformly lacked genetic divergence, to test for phenotypic divergence in toxicity. As predicted, overall levels of newt toxicity were low; however, we also detected unexpected among‐ and within‐population variation in toxicity. Most notably, a small number of individuals contained large doses of TTX that rival means of toxic populations in sympatry with Th. sirtalis. Phenotypic variation in toxicity, despite limited neutral genetic divergence, suggests that factors other than reciprocal selection with Th. sirtalis likely contribute to geographic patterns of toxicity in Ta. granulosa.     

Behavioral constraints on local adaptation and counter‐gradient variation: Implications for climate change

Resource allocation to growth, reproduction, and body maintenance varies within species along latitudinal gradients. Two hypotheses explaining this variation are local adaptation and counter‐gradient variation. The local adaptation hypothesis proposes that populations are adapted to local environmental conditions and are therefore less adapted to environmental conditions at other locations. The counter‐gradient variation hypothesis proposes that one population out performs others across an environmental gradient because its source location has greater selective pressure than other locations. Our study had two goals. First, we tested the local adaptation and counter‐gradient variation hypotheses by measuring effects of environmental temperature on phenotypic expression of reproductive traits in the burying beetle, Nicrophorus orbicollis Say, from three populations along a latitudinal gradient in a common garden experimental design. Second, we compared patterns of variation to evaluate whether traits covary or whether local adaptation of traits precludes adaptive responses by others. Across a latitudinal range, N. orbicollis exhibits variation in initiating reproduction and brood sizes. Consistent with local adaptation: (a) beetles were less likely to initiate breeding at extreme temperatures, especially when that temperature represents their source range; (b) once beetles initiate reproduction, source populations produce relatively larger broods at temperatures consistent with their local environment. Consistent with counter‐gradient variation, lower latitude populations were more successful at producing offspring at lower temperatures. We found no evidence for adaptive variation in other adult or offspring performance traits. This suite of traits does not appear to coevolve along the latitudinal gradient. Rather, response to selection to breed within a narrow temperature range may preclude selection on other traits. Our study highlights that N. orbicollis uses temperature as an environmental cue to determine whether to initiate reproduction, providing insight into how behavior is modified to avoid costly reproductive attempts. Furthermore, our results suggest a temperature constraint that shapes reproductive behavior.     

Shell morphology and color of the subtidal whelk Buccinum undatum exhibit fine‐scaled spatial patterns

Geographical patterns in morphology can be the result of divergence among populations due to neutral or selective changes and/or phenotypic plasticity in response to different environments. Marine gastropods are ideal subjects on which to explore these patterns, by virtue of the remarkable intraspecific variation in life‐history traits and morphology often observed across relatively small spatial scales. The ubiquitous N‐Atlantic common whelk (Buccinum undatum) is well known for spatial variation in life‐history traits and morphology. Previous studies on genetic population structure have revealed that it exhibits significant differentiation across geographic distances. Within Breiðafjörður Bay, a large and shallow bay in W‐Iceland, genetic differentiation was demonstrated between whelks from sites separated by just 20 km. Here, we extended our previous studies on the common whelk in Breiðafjörður Bay by quantifying phenotypic variation in shell morphology and color throughout the Bay. We sought to test whether trait differentiation is dependent on geographic distance and/or environmental variability. Whelk in Breiðafjörður Bay displayed fine‐scale patterns of spatial variation in shape, thickness, and color diversity. Differentiation increased with increasing distance between populations, indicating that population connectivity is limited. Both shape and color varied along a gradient from the inner part of the bay in the east to the outer part in the west. Whelk shells in the innermost part of Breiðafjörður Bay were thick with an elongate shell, round aperture, and low color diversity, whereas in the outer part of the bay the shells were thinner, rounder, with a more elongate aperture and richer color diversity. Significant site‐specific difference in shell traits of the common whelk in correlation with environmental variables indicates the presence of local ecotypes and limited demographic connectivity.     

Adaptation in temporally variable environments: stickleback armor in periodically breaching bar‐built estuaries

The evolutionary consequences of temporal variation in selection remain hotly debated. We explored these consequences by studying threespine stickleback in a set of bar‐built estuaries along the central California coast. In most years, heavy rains induce water flow strong enough to break through isolating sand bars, connecting streams to the ocean. New sand bars typically re‐form within a few weeks or months, thereby re‐isolating populations within the estuaries. These breaching events cause severe and often extremely rapid changes in abiotic and biotic conditions, including shifts in predator abundance. We investigated whether this strong temporal environmental variation can maintain within‐population variation while eroding adaptive divergence among populations that would be caused by spatial variation in selection. We used neutral genetic markers to explore population structure and then analysed how stickleback armor traits, the associated genes Eda and Pitx1 and elemental composition (%P) varies within and among populations. Despite strong gene flow, we detected evidence for divergence in stickleback defensive traits and Eda genotypes associated with predation regime. However, this among‐population variation was lower than that observed among other stickleback populations exposed to divergent predator regimes. In addition, within‐population variation was very high as compared to populations from environmentally stable locations. Elemental composition was strongly associated with armor traits, Eda genotype and the presence of predators, thus suggesting that spatiotemporal variation in armor traits generates corresponding variation in elemental phenotypes. We conclude that gene flow, and especially temporal environmental variation, can maintain high levels of within‐population variation while reducing, but not eliminating, among‐population variation driven by spatial environmental variation.     

Genotypic differences in embryonic life history traits of Folsomia quadrioculata (Collembola: Isotomidae) across a wide geographical range

1. Life history traits play a central role in adaptation to specific environmental conditions. Egg development time, hatchling, and egg batch size in 10 populations of the soil‐dwelling collembolan species Folsomia quadrioculata (Tullberg, 1871) from diverse habitats across arctic and temperate regions, ranging in latitude from 43 to 81°N were studied. 2. For all traits, 15 °C was used as the reference temperature. Phenotypic plasticity was studied by changing temperature to 10 and 20 °C in hatchling size, and to 20 °C in egg development time. The effect of latitude, climatic zone, and summer temperature at their sites of sampling was tested to address the hypotheses that populations from cooler areas would have (i) a faster temperature‐dependent egg development, (ii) a larger hatchling size, (iii) smaller egg batches and (iv) higher phenotypic plasticity 3. The first and fourth hypothesis were not supported, whereas the second and third were partly supported when including the whole latitudinal gradient, but not within regions. Plasticity showed a complex pattern, including large differences between populations from similar macroclimates and considerable similarity between some populations from contrasting climates. Small effects of latitude and macroclimatic variables emphasised that local climate and microhabitat should be taken into account when evaluating thermal conditions as possible drivers of population‐specific differences in soil‐dwelling ectotherms. 4. There was a trade‐off between egg batch and hatchling sizes. A weak correlation between the population mean egg development time and the mean hatchling size suggested that the populations are, or have been, exposed to differential selection.     

Ten years of life in compost: temporal and spatial variation of North German Caenorhabditis elegans populations

The nematode Caenorhabditis elegans is a central laboratory model system in almost all biological disciplines, yet its natural life history and population biology are largely unexplored. Such information is essential for in‐depth understanding of the nematode's biology because its natural ecology provides the context, in which its traits and the underlying molecular mechanisms evolved. We characterized natural phenotypic and genetic variation among North German C. elegans isolates. We used the unique opportunity to compare samples collected 10 years apart from the same compost heap and additionally included recent samples for this and a second site, collected across a 1.5‐year period. Our analysis revealed significant population genetic differentiation between locations, across the 10‐year time period, but for only one location a trend across the shorter time frame. Significant variation was similarly found for phenotypic traits of likely importance in nature, such as choice behavior and population growth in the presence of pathogens or naturally associated bacteria. Phenotypic variation was significantly influenced by C. elegans genotype, time of isolation, and sampling site. The here studied C. elegans isolates may provide a valuable, genetically variable resource for future dissection of naturally relevant gene functions.     

Parasite‐mediated selection in a natural metapopulation of Daphnia magna

Parasite‐mediated selection varying across time and space in metapopulations is expected to result in host local adaptation and the maintenance of genetic diversity in disease‐related traits. However, nonadaptive processes like migration and extinction‐(re)colonization dynamics might interfere with adaptive evolution. Understanding how adaptive and nonadaptive processes interact to shape genetic variability in life‐history and disease‐related traits can provide important insights into their evolution in subdivided populations. Here we investigate signatures of spatially fluctuating, parasite‐mediated selection in a natural metapopulation of Daphnia magna. Host genotypes from infected and uninfected populations were genotyped at microsatellite markers, and phenotyped for life‐history and disease traits in common garden experiments. Combining phenotypic and genotypic data a Q_ST–F_ST‐like analysis was conducted to test for signatures of parasite mediated selection. We observed high variation within and among populations for phenotypic traits, but neither an indication of host local adaptation nor a cost of resistance. Infected populations have a higher gene diversity (Hs) than uninfected populations and Hs is strongly positively correlated with fitness. These results suggest a strong parasite effect on reducing population level inbreeding. We discuss how stochastic processes related to frequent extinction‐(re)colonization dynamics as well as host and parasite migration impede the evolution of resistance in the infected populations. We suggest that the genetic and phenotypic patterns of variation are a product of dynamic changes in the host gene pool caused by the interaction of colonization bottlenecks, inbreeding, immigration, hybrid vigor, rare host genotype advantage and parasitism. Our study highlights the effect of the parasite in ameliorating the negative fitness consequences caused by the high drift load in this metapopulation.     

Geographic variation in lizard phenotypes: importance of the incubation environment

Geographic variation in phenotypes can result from proximate environmental effects as well as from underlying genetic factors. Reciprocal transplant experiments, in which organisms are moved from one area to another, offer a powerful technique to partition the effects of these two factors. However, many studies that have utilized this technique have focused on the post-hatching organism only and ignored potential effects of environmental influences acting during embryonic development. We examined the phenotypic responses of hatchling scincid lizards ( Lampropholis guichenoti ) incubated in the laboratory under thermal regimes characteristic of natural nests in two study areas in southeastern Australia. Although the sites were less than 120 km apart, lizards from these two areas differed in thermal regimes of natural nests, and in hatchling phenotypes (morphology, locomotor performance). We incubated eggs from each area under the thermal regimes typical of both sites. Some of the traits we measured (e.g. hatchling mass and snout-vent length) showed little or no phenotypic plasticity in response to differences in incubation conditions, whereas other traits (e.g. incubation period, tail length, inter-limb length, body shape, locomotor performance) were strongly influenced by the thermal regime experienced by the embryo. Thus, a significant proportion of the geographic variation in morphology and locomotor performance of hatchling lizards may be directly induced by differences in nest temperatures rather than by genetic divergence. We suggest that future studies using the reciprocal transplant design should consider environmental influences on all stages of the life-history, including embryonic development as well as post-hatching life.     

Genome‐wide differentiation in closely related populations: the roles of selection and geographic isolation

Population divergence in geographic isolation is due to a combination of factors. Natural and sexual selection may be important in shaping patterns of population differentiation, a pattern referred to as ‘isolation by adaptation’ (IBA). IBA can be complementary to the well‐known pattern of ‘isolation by distance’ (IBD), in which the divergence of closely related populations (via any evolutionary process) is associated with geographic isolation. The barn swallow Hirundo rustica complex comprises six closely related subspecies, where divergent sexual selection is associated with phenotypic differentiation among allopatric populations. To investigate the relative contributions of selection and geographic distance to genome‐wide differentiation, we compared genotypic and phenotypic variation from 350 barn swallows sampled across eight populations (28 pairwise comparisons) from four different subspecies. We report a draft whole‐genome sequence for H. rustica, to which we aligned a set of 9493 single nucleotide polymorphisms (SNPs). Using statistical approaches to control for spatial autocorrelation of phenotypic variables and geographic distance, we find that divergence in traits related to migratory behaviour and sexual signalling, as well as geographic distance, together explain over 70% of genome‐wide divergence among populations. Controlling for IBD, we find 42% of genomewide divergence is attributable to IBA through pairwise differences in traits related to migratory behaviour and sexual signalling alone. By (i) combining these results with prior studies of how selection shapes morphological differentiation and (ii) accounting for spatial autocorrelation, we infer that morphological adaptation plays a large role in shaping population‐level differentiation in this group of closely related populations.     

Geographic variation in body size, sexual size dimorphism and fitness components of a seed beetle: local adaptation versus phenotypic plasticity

Variation in body size, growth and life history traits of ectotherms along latitudinal and altitudinal clines is generally assumed to represent adaptation to local environmental conditions, especially adaptation to temperature. However, the degree to which variation along these clines is due to adaptation vs plasticity remains poorly understood. In addition, geographic patterns often differ between females and males – e.g. sexual dimorphism varies along latitudinal clines, but the extent to which these sex differences are due to genetic differences between sexes vs sex differences in plasticity is poorly understood. We use common garden experiments (beetles reared at 24, 30 and 36°C) to quantify the relative contribution of genetically‐based differentiation among populations vs phenotypic plasticity to variation in body size and other traits among six populations of the seed‐feeding beetle Stator limbatus collected from various altitudes in Arizona, USA. We found that temperature induces substantial plasticity in survivorship, body size and female lifetime fecundity, indicating that developmental temperature significantly affects growth and life history traits of S. limbatus. We also detected genetic differences among populations for body size and fecundity, and genetic differences among populations in thermal reaction norms, but the altitude of origin (and hence mean temperature) does not appear to explain these genetic differences. This and other recent studies suggest that temperature is not the major environmental factor that generates geographic variation in traits of this species. In addition, though there was no overall difference in plasticity of body size between males and females (when averaged across populations), we did find that the degree to which dimorphism changed with temperature varied among populations. Consequently, future studies should be extremely cautious when using only a few study populations to examine environmental effects on sexual dimorphism.     

Subspecies delineation amid phenotypic,geographic and genetic discordance in a songbird

Understanding the processes that drive divergence within and among species is a long‐standing goal in evolutionary biology. Traditional approaches to assessing differentiation rely on phenotypes to identify intra‐ and interspecific variation, but many species express subtle morphological gradients in which boundaries among forms are unclear. This intraspecific variation may be driven by differential adaptation to local conditions and may thereby reflect the evolutionary potential within a species. Here, we combine genetic and morphological data to evaluate intraspecific variation within the Nelson's (Ammodramus nelsoni) and salt marsh (Ammodramus caudacutus) sparrow complex, a group with populations that span considerable geographic distributions and a habitat gradient. We evaluated genetic structure among and within five putative subspecies of A. nelsoni and A. caudacutus using a reduced‐representation sequencing approach to generate a panel of 1929 SNPs among 69 individuals. Although we detected morphological differences among some groups, individuals sorted along a continuous phenotypic gradient. In contrast, the genetic data identified three distinct clusters corresponding to populations that inhabit coastal salt marsh, interior freshwater marsh and coastal brackish–water marsh habitats. These patterns support the current species‐level recognition but do not match the subspecies‐level taxonomy within each species—a finding which may have important conservation implications. We identified loci exhibiting patterns of elevated divergence among and within these species, indicating a role for local selective pressures in driving patterns of differentiation across the complex. We conclude that this evidence for adaptive variation among subspecies warrants the consideration of evolutionary potential and genetic novelty when identifying conservation units for this group.     

Phenotypic variation in an oviparous montane lizard (Bassiana duperreyi): the effects of thermal and hydric incubation environments

Recent studies have shown that incubation temperatures can profoundly affect the phenotypes of hatchling lizards, but the effects of hydric incubation environments remain controversial. We examined incubation-induced phenotypic variation in Bassiana duperreyi (Gray, 1938; Sauria: Scincidae), an oviparous montane lizard from south-eastern Australia. We incubated eggs from this species in four laboratory treatments, mimicking cool and moist, cool and dry, warm and moist, and warm and dry natural nest-sites, and assessed several morphological and behavioural traits of lizards after hatching. Incubation temperature influenced a lizard's hatching success, incubation period, tail length and antipredator behaviour, whereas variation in hydric conditions did not engender significant phenotypic variation for most traits. However, moisture affected incubation period slightly differently in males and females, and for a given snout-vent length moisture interacted weakly with temperature to affect lizard body mass. Although incubation conditions can substantially affect phenotypic variation among hatchling lizards, the absence of strong hydric effects suggests that hatchling lizards react less plastically to variation in moisture levels than they do to thermal conditions. Thus, our data do not support the generalization that water availability during embryogenesis is more important than temperature in determining the phenotypes of hatchling reptiles.     

Ecological,evolutionary, and conservation implications of incubation temperature‐dependent phenotypes in birds

Incubation is a vital component of reproduction and parental care in birds. Maintaining temperatures within a narrow range is necessary for embryonic development and hatching of young, and exposure to both high and low temperatures can be lethal to embryos. Although it is widely recognized that temperature is important for hatching success, little is known about how variation in incubation temperature influences the post‐hatching phenotypes of avian offspring. However, among reptiles it is well known that incubation temperature affects many phenotypic traits of offspring with implications for their future survival and reproduction. Although most birds, unlike reptiles, physically incubate their eggs, and thus behaviourally control nest temperatures, variation in temperature that influences embryonic development still occurs among nests within a population. Recent research in birds has primarily been limited to populations of megapodes and waterfowl; in each group, incubation temperature has substantial effects on hatchling phenotypic traits important for future development, survival, and reproduction. Such observations suggest that incubation temperature (and incubation behaviours of parents) is an important but underappreciated parental effect in birds and may represent a selective force instrumental in shaping avian reproductive ecology and life‐history traits. However, much more research is needed to understand how pervasive phenotypic effects of incubation temperature are among birds, the sources of variation in incubation temperature, and how effects on phenotype arise. Such insights will not only provide foundational information regarding avian evolution and ecology, but also contribute to avian conservation.     

Spatial and environmental correlates of intraspecific morphological variation in three species of passerine birds from the Purus–Madeira interfluvium,Central Amazonia

Biogeographic studies in Amazonia typically describe biodiversity across interfluvia, rarely within them, where geographic variability in morphological traits might be observed. We tested for intraspecific phenotypic variation in three bird species within the Purus–Madeira interfluvium (Central Amazon) and whether phenotypes were correlated with environmental heterogeneity or geographic distance among sites. We compared coloration indexes derived from reflectance spectra and morphometrics of up to five adult individuals of each sex among 11 sites within the interfluvium and contrasted them with proxies for geographic distance and environmental variation (tree basal area and bird community). Environmental heterogeneity was minimally spatially autocorrelated, and there were no obvious geographical barriers to dispersal in the study region. The null hypothesis was that we would see either no phenotypic variation or random variation that was not explained by the tested variables. Half of the cases analyzed showed intraspecific morphological variation. Coloration varied more frequently than morphometrics, and color was better explained by environmental heterogeneity, particularly in males, whereas brightness also varied with geographic distance. Geographic distance explained the only case of variation in morphometrics. Our results indicate that coloration, particularly plumage brightness, is more labile than morphometric traits and that plumage color might be under stronger effects of local adaptation than brightness, which also seems to be under effects of neutral drift and gene flow among populations. Higher frequencies of association between male coloration and the environment suggest a role of non-arbitrary mechanisms of sexual selection on the expression of male phenotypes, whereas arbitrary intersexual selection might explain the randomly distributed variation that is not explained by environmental heterogeneity or geographic distance. We revealed intraspecific phenotypic variation in a spatial extent usually not considered in biogeographic studies in the Amazon and demonstrate that both local adaptation and neutral drift are important to explain intraspecific trait diversification at this geographical scale.     

Geographic variation in hatchling size in an oviparous skink: effects of maternal investment and incubation thermal environment

Geographic variation in offspring size can be viewed as an adaptive response to local environmental conditions, but the causes of such variation remain unclear. Here, we compared the size and composition of eggs laid by female Chinese skinks (Plestiodon chinensis) from six geographically distinct populations in southeastern China to evaluate geographic variation in hatchling size. We also incubated eggs from these six populations at three constant temperatures (24, 28 and 32 °C) to evaluate the combined effects of incubation temperature and population source on hatchling size. Egg mass and composition varied among populations, and interpopulation differences in yolk dry mass and energy content were still evident after accounting for egg mass. Population mean egg mass and thus hatchling mass were greater in the colder localities. Females from three northern populations increased offspring size by laying larger eggs relative to their own size. Females from an inland population in Rongjiang could increase offspring size by investing relatively more dry materials and thus more energy into individual eggs without enlarging the size of their eggs. The degree of embryonic development at oviposition was almost the same across the six populations, so was the rate of embryonic development and thus incubation length at any given temperature. Both incubation temperature and population source affected hatchling traits examined, but the relative importance of these two factors varied between traits. Our data show that in P. chinensis hatchling traits reflecting overall body size (body mass, snout‐vent length and tail length) are more profoundly affected by population source. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 283–296.     

Geographic variation of flower traits in Narcissus papyraceus (Amaryllidaceae): do pollinators matter?

Aim The geographic clinal variation of traits in organisms can indicate the possible causes of phenotypic evolution. We studied the correlates of flower trait variation in populations of a style‐dimorphic plant, Narcissus papyraceus Ker‐Gawl., within a region of high biogeographical significance, the Strait of Gibraltar. This species shows a geographic gradient in the style‐morph ratio, suggested to be driven by pollinator shifts. We tested whether parallel geographic variation of perianth traits also exists, concomitant with vegetative trait variation or genetic similarity of plant populations. Location The Strait of Gibraltar region (SG hereafter, including both south‐western Iberian Peninsula and north‐western Morocco). Methods We used univariate and multivariate analyses of flower and vegetative traits in 23 populations. We applied Mantel tests and partial Mantel correlations on vegetative and flower traits and geographic locations of populations to test for spatial effects. We used Moran’s autocorrelation analyses to explore the spatial structure within the range, and performed the analyses with and without the Moroccan samples to test for the effects of the SG on spatial patterns. Amplified fragment length polymorphism data were used to estimate the genetic distance between populations and to ascertain its relationship with morphometric distance. Results There was high variation between and within populations in both flower and vegetative traits. Mantel correlations between geographic and morphometric distances were not significant, but the exclusion of Moroccan populations revealed some distance effect. Partial Mantel correlation did not detect a significant correlation between flower and vegetative morphometric distances after controlling for geographic distance. There were opposite trends in spatial autocorrelograms of flower and vegetative traits. The genetic distance between pairs of populations was directly correlated with geographic distance; however, flower morphometric and genetic distances were not significantly correlated. Main conclusions The SG had some influence on phenotypes, although the causes remain to be determined. The opposite trend of variation in flower and vegetative traits, and the lack of correlation between genetic distance and dissimilarity of flower phenotypes favour the hypothesis of pollinator‐mediated selection on flower morphology, although this may affect only particular traits and populations rather than overall phenotypes. Although stochastic population processes may have a small effect, other factors may account for the high flower variation within and between populations.     

The effects of cultural drift on geographic variation in echolocation calls of the Chinese rufous horseshoe bat (Rhinolophus sinicus)

Drift, selection, or their combined effects commonly drive geographic variation in traits. Clarifying the relative roles of each process is a long‐standing research goal in evolutionary biology. Acoustic signals of bats are a phenotypic characteristic that plays an important role in social organization and species recognition. We extensively sampled the Chinese rufous horseshoe bat (Rhinolophus sinicus) throughout China and Vietnam and reconstructed a species phylogeny to better understand the patterns and causes of the geographic variation of acoustic signals. Our results showed that the resting frequency (RF) of calls varied with latitude, sex, and distance among different colony locations. RF differences were not correlated with genetic distance (based on only one nuclear locus and the mitochondrial locus), climatic factors (mean annual temperature and mean annual precipitable water), or body size, although differences in calls increased with distance among various populations. This suggests that cultural drift may play more important roles than genetic drift and acoustic adaptation in shaping acoustic differences within regions in R. sinicus.     

The contributions of evolutionary divergence and phenotypic plasticity to geographic variation in the western fence lizard,Sceloporus occidentalis

Local genetic adaptation and phenotypic plasticity are two mechanisms that can have marked effects on the morphology, performance, and behaviour of animals, producing geographic variation among populations. However, few studies have examined how these mechanisms interact during ontogeny to shape organismal phenotypes. We incubated eggs of the western fence lizard, Sceloporus occidentalis, from four populations (representing two latitudes and altitudes) in either a warm or cool environment in the laboratory. We then raised the hatchlings under common laboratory conditions, measured morphological and performance traits until 5 weeks of age, and compared juvenile morphology with that of field‐caught adults from each population. The results obtained indicate that some phenotypic traits that contribute to performance (body size, hindlimb length, head shape) were relatively canalized in juveniles and differed among populations in a way that was consistent with adults from their population of origin. However, other traits (forelimb length, inter‐limb length, mass, tail length), showed significant effects of incubation temperature, and this environmentally induced variation persisted throughout the experiment. Although selection pressure may be stronger for traits that are integral to survival, developmental effects might still have a lasting impact on traits less important to organismal fitness. We discuss the results obtained in the present study the context of the life history of these animals. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 84–98.     

Selection on QTL and complex traits in complex environments

Understanding genetic variation for complex traits in heterogeneous environments is a fundamental problem in biology. In this issue of Molecular Ecology, Fournier‐Level et al. ( 2013 ) analyse quantitative trait loci (QTL) influencing ecologically important phenotypes in mapping populations of Arabidopsis thaliana grown in four habitats across its native European range. They used causal modelling to quantify the selective consequences of life history and morphological traits and QTL on components of fitness. They found phenology QTL colocalizing with known flowering time genes as well as novel loci. Most QTL influenced fitness via life history and size traits, rather than QTL having direct effects on fitness. Comparison of phenotypes among environments found no evidence for genetic trade‐offs for phenology or growth traits, but genetic trade‐offs for fitness resulted because flowering time had opposite fitness effects in different environments. These changes in QTL effects and selective consequences may maintain genetic variation among populations.     

Root architecture,plant size and soil nutrient variation in natural populations of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Phenotypic variation in ecologically important traits may vary at large and small geographic scales, and may be shaped by natural selection. Here our explicit aim is to evaluate phenotypic differentiation among local populations and examine its relationship with ecological edaphic and climatic features that could lead to local adaptation. We characterized six populations of the model plant Arabidopsis thaliana over 3 years in the field in its native range. At each site, we measured edaphic conditions and aboveground and belowground phenotypes. In addition, we grew plants from the six characterized populations in a common greenhouse along with an additional fifteen populations from the Iberian Peninsula to examine evolutionary and ecological differentiation among populations, and relationships between geographic and ecological distance to phenotypic differences among populations. Significant differences in aboveground and belowground traits, population density, and micro- and macro-nutrient soil concentrations were found among the field populations. In particular, root architectural traits differed significantly among field populations. Complex patterns of ecological differences among population and plant phenotypes emerged when examining edaphic conditions in the Extremadura region, and geographic and climate variables at a broader scale of the Iberian Peninsula. We report levels of phenotypic variation at the local scale comparable to those found at broad geographic scales and report that local edaphic conditions contribute to population-level phenotypic variation in root and shoot traits. To our knowledge, these are the first reports of among population root architectural variation from natural field populations for this model organism. We demonstrate how ecological features, such as soil nutrients, can be associated with the phenotypic variation of A. thaliana measured in natural populations and may contribute to adaptive differentiation at a local scale.