Bergmann's rule is maintained during a rapid range expansion in a damselfly

Climate‐induced range shifts result in the movement of a sample of genotypes from source populations to new regions. The phenotypic consequences of those shifts depend upon the sample characteristics of the dispersive genotypes, which may act to either constrain or promote phenotypic divergence, and the degree to which plasticity influences the genotype–environment interaction. We sampled populations of the damselfly Erythromma viridulum from northern Europe to quantify the phenotypic (latitude–body size relationship based on seven morphological traits) and genetic (variation at microsatellite loci) patterns that occur during a range expansion itself. We find a weak spatial genetic structure that is indicative of high gene flow during a rapid range expansion. Despite the potentially homogenizing effect of high gene flow, however, there is extensive phenotypic variation among samples along the invasion route that manifests as a strong, positive correlation between latitude and body size consistent with Bergmann's rule. This positive correlation cannot be explained by variation in the length of larval development (voltinism). While the adaptive significance of latitudinal variation in body size remains obscure, geographical patterns in body size in odonates are apparently underpinned by phenotypic plasticity and this permits a response to one or more environmental correlates of latitude during a range expansion.     

Latitudinal and temperature-dependent variation in embryonic development and growth in Rana temporaria

Variation in seasonal time constraints and temperature along latitudinal gradients are expected to select for life history trait differentiation, but information about the relative importance of these factors in shaping patterns of divergence in embryonic traits remains sparse. We studied embryonic survival, growth and development rates in the common frog (Rana temporaria) along a 1,400-km latitudinal gradient across Sweden by raising embryos from four populations in the laboratory at seven temperatures (9 degrees C, 12 degrees C, 15 degrees C, 18 degrees C, 21 degrees C, 24 degrees C, 27 degrees C). We found significant differences in mean values of all traits between the populations and temperature treatments, but this variation was not latitudinally ordered. In general, embryonic survival decreased at the two highest temperatures in all populations, but less so in the southernmost as compared to the other populations. The northernmost population developed slowest at the lowest temperature, while the two mid-latitude populations were slowest at the other temperatures. Hatchling size increased with increasing temperature especially in the two northern populations, whereas the two southern populations showed peak hatchling size at 15 degrees C. Analyses of within-population genetic variation with a half-sib design revealed that there was significant additive genetic variation in all traits, and egg size-related maternal effects were important in the case of hatchling size. Overall, our results indicate that unlike larval growth and development, variation in embryonic development and growth in R. temporaria cannot be explained in terms of a latitudinal gradient in season length. While adaptation to a latitudinal variation in temperature might have contributed to the observed differentiation in embryonic performance, the effects of other, perhaps more local environmental factors, seem to have overridden them in importance.     

Morphological and genetic divergence of intralacustrine stickleback morphs in Iceland: a case for selective differentiation?

The evolutionary processes involved in population divergence and local adaptation are poorly understood. Theory predicts that divergence of adjacent populations is possible but depends on several factors including gene flow, divergent selection, population size and the number of genes involved in divergence and their distribution on the genome. We analyse variation in neutral markers, markers linked to putative quantitative trait loci and morphological traits in a recent (<10000 years) zone of primary divergence between stickleback morphs in Lake Thingvallavatn, Iceland. Environmental factors, especially predation, are clearly implicated in reducing gene flow between morphs. There is continuous morphological and genetic variation between habitats with a zone centre similar to secondary contact zones. Individual microsatellite loci are implicated as being linked to adaptive variation by direct tests as well as by differences in cline shape. Patterns of linkage disequilibria indicate that the morphs have diverged at several loci. This divergence shows parallels and differences with the well-studied limnetic-benthic stickleback morphs, both in phenotypic divergence and at the genomic level.     

Allometric influence on phenotypic variation in the Song Sparrow (Melospiza melodia)

Song Sparrow ( Melospiza melodia ) populations found along the Pacific Coast of North America, from Baja California to the islands off the coast of Alaska, exhibit extensive morphological variation. With a multivariate analysis of size and shape, I describe a portion of this pattern and examine how it could be maintained despite gene flow among the populations. Because shape differences fall along geographic barriers, I suggest that similarities among Song Sparrow populations in multivariate shape reflect their pattern of genetic relatedness. A general pattern of Song Sparrow post-nestling growth allometry has been discovered: bill characteristics are positively allometric and all other characteristics are negatively allometric. In contrast to shape, multivariate patterns of body size variation do not correspond to geographic relationships. In combination with evidence of Song Sparrow phenotypic plasticity, it is proposed that multivariate body size is an environmentally plastic trait and that specific traits exhibit levels of phenotypic plasticity in proportion to their rate of growth with respect to body size. In this way local environmental factors which alter body size may change an entire suite of allometrically related traits and thus create striking patterns of morphological variation.     

Signals of local adaptation across an environmental gradient among highly connected populations of the Dead Sea Sparrow Passer moabiticus in Israel

Populations found at the edge of a species range often have decreased genetic diversity, which together with high gene flow may reduce the ability of a species to adapt to local environmental conditions. The Dead Sea Sparrow Passer moabiticus occupies a disjointed range, where the Israeli populations are considered peripheral and fragmented. The species is also thought to have undergone a recent range expansion. We aimed to describe the genetic and morphological variation of the Israeli populations and to determine the extent of gene flow among them. We expected that because of the small latitudinal gradient across Israel and the recent range expansion of the species that Dead Sea Sparrow populations would show no significant morphological adaptation to local environmental conditions, and that considerable gene flow would be taking place among populations. Our findings indicate the existence of gene flow, suggesting high connectivity among populations, but recovered no support for a recent range expansion, possibly due to insufficient time since expansion for mutations to have accumulated. However, despite recurrent gene flow among populations, latitudinal variation in wing length (male and female) and body mass (male) was indicative of local adaptation across Israel, in accordance with Bergmann's rule.     

Testing limits to adaptation along altitudinal gradients in rainforest Drosophila

Given that evolution can generate rapid and dramatic shifts in the ecological tolerance of a species, what prevents populations adapting to expand into new habitat at the edge of their distributions? Recent population genetic models have focused on the relative costs and benefits of migration between populations. On the one hand, migration may limit adaptive divergence by preventing local populations from matching their local selective optima. On the other hand, migration may also contribute to the genetic variance necessary to allow populations to track these changing optima. Empirical evidence for these contrasting effects of gene flow in natural situations are lacking, largely because it remains difficult to acquire. Here, we develop a way to explore theoretical models by estimating genetic divergence in traits that confer stress resistance along similar ecological gradients in rainforest Drosophila. This approach allows testing for the coupling of clinal divergence with local density, and the effects of genetic variance and the rate of change of the optimum on the response to selection. In support of a swamping effect of migration on phenotypic divergence, our data show no evidence for a cline in stress-related traits where the altitudinal gradient is steep, but significant clinal divergence where it is shallow. However, where clinal divergence is detected, sites showing trait means closer to the presumed local optimum have more genetic variation than sites with trait means distant from their local optimum. This pattern suggests that gene flow also aids a sustained response to selection.     

Latitudinal trait variation and responses to drought in Arabidopsis lyrata

Species may respond in three ways to environmental change: adapt, migrate, or go extinct. Studies of latitudinal clines can provide information on whether species have adapted to abiotic stress such as temperature and drought in the past and what the traits underlying adaptation are. We investigated latitudinal trait variation and response to drought in North American populations of Arabidopsis lyrata. Plants from nine populations collected over 13° latitude were grown under well-watered and dry conditions. A total of 1,620 seedlings were raised and 12 phenological, physiological, morphological, and life history traits were measured. Two traits, asymptotic rosette size and the propensity to flower, were significantly associated with latitude: plants from northern locations grew to a larger size and were more likely to flower in the first season. Most traits displayed a plastic response to drought, but plasticity was never related linearly with latitude nor was it enhanced in populations from extreme latitudes with reduced water availability. Populations responded to drought by adopting mixed strategies of resistance, tolerance, and escape. The study shows that latitudinal adaptation in A. lyrata involves the classic life history traits, size at and timing of reproduction. Contrary to recent theoretical predictions, adaptation to margins is based on fixed trait differences and not on phenotypic plasticity, at least with respect to drought.     

Weak impact of fine‐scale landscape heterogeneity on evolutionary potential in Arabidopsis lyrata

Microhabitat heterogeneity can lead to fine‐scale local adaptation when gene flow is restricted, which may be important for the maintenance of genetic variation within populations. This study tested whether microhabitat heterogeneity was associated with trait differences in a population of Arabidopsis lyrata and studied its impact on the genetic variance–covariance (G) matrix. Maternal seed families were collected from dune tops and bottoms, two microhabitats known to vary significantly in water availability. In a common garden experiment, replicate individuals per family were raised under wet and dry conditions, and physiological, morphological and life‐history traits were assessed. Plants from the two microenvironments differed in their response to treatment in two performance components, in stomata density and most strongly in flowering time. Under wet conditions, plants originating from dune bottoms flowered 4 weeks earlier than those from dune tops. Only one of three G‐matrix comparisons revealed that habitat heterogeneity and evolutionary potential were positively linked. The number of independent trait dimensions was larger in the entire population than within subpopulations separated by microhabitat under wet conditions. However, the size of the G‐matrix was no larger in the entire population than within subpopulations separated by microhabitat, and trait correlation structure between microhabitats and treatments was not significantly different. These results indicate that fine‐scale habitat heterogeneity likely led to local adaptation, which weakly affected levels of across‐trait genetic variation.     

Patterns of genetic diversity vary among shoot and root functional traits in Norway spruce Picea abies along a latitudinal gradient

Roots constitute a major segment of plant biomass, and variation in belowground traits in situ correlates with environmental gradients at large spatial scales. Local adaptation of populations maintains intraspecific genetic variation in various shoot traits, but the contribution of genetic factors to adaptation to soil heterogeneity remains poorly known. I established a common-garden experiment with three Norway spruce Picea abies populations sampled between 60° and 67° N in Finland, each represented by 13 or 15 maternal families, to determine whether belowground traits are as genetically differentiated among populations as those in the shoot along a collective latitudinal gradient of temperature and soil heterogeneity. Two growing season simulations enabled testing for among-population differences in phenotypic plasticity. I phenotyped 777 first-year seedlings from shoot to root to capture functional traits that may influence survival in the wild: autumn phenology, shoot growth, root system size, root architecture, root morphology and growth allocation. All traits exhibited within-population genetic diversity, but among-population differentiation ranged from strong in shoot traits to nonexistent in root system architecture and morphology that are scaled to root system size. However, latitudinal trends characterised root-to-shoot ratio and root tip-to-shoot ratio that account for among-population differences in aboveground growth. Overall trait variability was multidimensional with variable among- versus within-population trends: for example, phenology and shoot growth covaried across populations, but their association within individual populations was variable. Shoot growth correlated positively with root system size, but not with root architecture or morphology. Finally, the two higher-latitude populations exhibited greater phenotypic plasticity in shoot traits and growth allocation. The results demonstrate varying patterns of genetic variation in functional traits of Norway spruce in the boreal zone, suggesting simultaneous adaptation to multiple environmental factors. Functional traits that exhibit phenotypic plasticity, genetic diversity and little covariation will promote long-term survival of populations in fluctuating environments.     

Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Latitudinal clines in the grasshopper Dichroplus elongatus: Coevolution of the A genome and B chromosomes?

Argentine populations of Dichroplus elongatus (Orthoptera: Acrididae) are polymorphic for B chromosomes. Previous studies showed that B chromosomes affect body size and some fitness components in Northwestern populations. We studied phenotype and B′s variation patterns along a latitudinal cline as well as the relationship between karyotype and body size related traits in 17 populations from East. Body size related traits showed a ‘saw tooth’ pattern of variation being small at low and high latitudes and large at intermediate latitudes in most of the analysed populations. Analyses of variance and principal components demonstrated that in most analysed populations B carrier males are associated with a decrease in body size related traits with respect to individuals with standard karyotype. Accordingly with the relationship between karyotype and body size, an opposite pattern of latitudinal variation in the frequencies of B′s with respect to body size variation was observed in this area. i.e. smaller individuals tend to have a higher frequency of B chromosomes. The comparison of the differentiation of both karyotype and body size traits with molecular neutral markers demonstrated the relative importance of selection moulding chromosome and phenotype variation. The observed pattern of phenotypic variation is likely to be the result of local adaptation to season length along the latitudinal gradient. The observed contrary pattern of B′s clinal variation may reflect the population ability to maintain this chromosome in relation to the local adaptation. The available evidence indicates that the distribution of B chromosome frequency was shaped by selective factors.     

Latitudinal clines in body size,but not in thermal tolerance or heat‐shock cognate 70 (HSC70), in the highly‐dispersing intertidal gastropod Littorina keenae (Gastropoda: Littorinidae)

Natural populations of widely‐distributed animals often exhibit clinal variation in phenotypic traits or in allele frequencies of a particular gene over their geographical range. A planktotrophic intertidal snail, Littorina keenae is broadly distributed along the north‐eastern Pacific coast through a large latitudinal range (24°50′N–43°18′N). We tested for latitudinal clines in two complex phenotypic traits – thermal tolerance and body size – and one single locus trait – heat shock cognate 70 (HSC70) – in L. keenae along almost its entire geographical range. We found only weak evidence for a latitudinal cline in the thermal tolerance and no evidence for a cline in allele frequencies at HSC70. However, as predicted by Bergmann's rule, we detected a strong latitudinal cline that accounted for 60% of the variance in body size (R² = 0.598; P < 0.001). In contrast, body size did not significantly affect thermal tolerance. HSC70 showed no genetic differentiation among the populations, supporting our previous mitochondrial gene‐based estimate of high gene flow during this snail's free‐swimming larval stage. Given that L. keenae experiences panmixia along its species range, the observed size cline may be partially or entirely caused by a phenotypically plastic response to local thermal environments rather than by genetic divergence in body size among populations in response to locally optimizing natural selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 494–505.     

LOCAL ADAPTATION MAINTAINS CLINAL VARIATION IN MELANIN‐BASED COLORATION OF EUROPEAN BARN OWLS (TYTO ALBA)

Ecological parameters vary in space, and the resulting heterogeneity of selective forces can drive adaptive population divergence. Clinal variation represents a classical model to study the interplay of gene flow and selection in the dynamics of this local adaptation process. Although geographic variation in phenotypic traits in discrete populations could be remainders of past adaptation, maintenance of adaptive clinal variation requires recurrent selection. Clinal variation in genetically determined traits is generally attributed to adaptation of different genotypes to local conditions along an environmental gradient, although it can as well arise from neutral processes. Here, we investigated whether selection accounts for the strong clinal variation observed in a highly heritable pheomelanin‐based color trait in the European barn owl by comparing spatial differentiation of color and of neutral genes among populations. Barn owl's coloration varies continuously from white in southwestern Europe to reddish‐brown in northeastern Europe. A very low differentiation at neutral genetic markers suggests that substantial gene flow occurs among populations. The persistence of pronounced color differentiation despite this strong gene flow is consistent with the hypothesis that selection is the primary force maintaining color variation among European populations. Therefore, the color cline is most likely the result of local adaptation.     

The effect of urbanisation on avian morphology and latitudinal gradients in body size

Urban areas occupy a large and growing proportion of the earth. Such sites exhibit distinctive characteristics relative to adjacent rural habitats, and many species have colonised and now successfully exploit urban habitats. The change in selection pressures as a result of urbanisation has led to trait divergence in some urban populations relative to their rural counterparts, but studies have generally been local in scale and the generality of differentiation thus remains unknown. The European blackbird Turdus merula is one of the commonest urban bird species in the Western Palearctic, but populations vary substantially in the length of time they have been urbanised. Here we investigate patterns of morphological variation in European blackbirds occupying 11 paired urban and rural habitats across much of the urbanised range of this species and spanning 25° of latitude. First, we assessed the extent to which urban and rural blackbirds are differentiated morphologically and the consistency of any differentiation across the range. Paired urban and rural Blackbird populations frequently exhibited significant morphological differences, but the magnitude and direction of differentiation was site dependent. We then investigated whether the nature of latitudinal gradients in body‐size differed between urban and rural populations, as predicted by differences in the climatic regimes of urban and rural areas. Blackbird body‐size exhibited strong latitudinal gradients, but their form did not differ significantly between urban and rural habitats. The latitudinal gradient in body size may be a consequence of Seebohm's rule, that more migratory populations occurring at high latitudes have longer wings. We conclude that while there can be substantial morphological variation between adjacent urban and rural bird populations, such differentiation may not apply across a species’ range. Locality specific differentiation of urban and rural blackbirds may arise if the selection pressures acting on blackbird morphology vary in an inconsistent manner between urban and rural habitats. Alternatively, phenotypic divergence could arise in a stochastic manner depending on the morphological traits of colonists, through founder effects.     

Molecular and morphometric variation in chromosomally differentiated populations of the grasshopper Sinipta dalmani (Orthopthera: Acrididae)

Sinipta dalmani is an Argentine grasshopper whose chromosome polymorphisms have been widely studied through cytogenetic, morphometric, and fitness component analyses. The present work analysed molecular and morphometric variation in seven chromosomally differentiated populations from Entre Rios and Buenos Aires provinces to analyse population structure. Molecular studies were performed studying RAPD loci and morphometric analyses were carried out measuring five morphometric traits. Genetic variability was high in all studied populations and was characterized by a decrease in H as a function of latitude and temperature. Both conventional F(ST) analysis and Bayesian approach for dominant marker showed that there were significant genetic differences among all populations, between provinces, and among populations within provinces. Entre Rios populations showed higher mean numbers of migrants per generation as well as low genetic differentiation and high gene flow with almost all populations whereas Buenos Aires populations may be considered as a result of a more recently colonization. There is considerable morphometric variation between populations and this variation correlates with latitude and temperature. Our results suggest that selection contributes to phenotypic differentiation among populations by moulding the differences in trait means whereas genetic drift is responsible for differences in the matrix of variance-covariance. The gene flow detected is insufficient to prevent phenotypic and chromosome divergences.     

Wing trait–inversion associations in Drosophila subobscura can be generalized within continents,but may change through time

Clinal variation is one of the most emblematic examples of the action of natural selection at a wide geographical range. In Drosophila subobscura, parallel clines in body size and inversions, but not in wing shape, were found in Europe and South and North America. Previous work has shown that a bottleneck effect might be largely responsible for differences in wing trait–inversion association between one European and one South American population. One question still unaddressed is whether the associations found before are present across other populations of the European and South American clines. Another open question is whether evolutionary dynamics in a new environment can lead to relevant changes in wing traits–inversion association. To analyse geographical variation in these associations, we characterized three recently laboratory founded D. subobscura populations from both the European and South American latitudinal clines. To address temporal variation, we also characterized the association at a later generation in the European populations. We found that wing size and shape associations can be generalized across populations of the same continent, but may change through time for wing size. The observed temporal changes are probably due to changes in the genetic content of inversions, derived from adaptation to the new, laboratory environment. Finally, we show that it is not possible to predict clinal variation from intrapopulation associations. All in all this suggests that, at least in the present, wing traits–inversion associations are not responsible for the maintenance of the latitudinal clines in wing shape and size.     

The evolution of novel herbicide tolerance in a noxious weed: the geographic mosaic of selection

Understanding how genetic variation is organized over geography has long been of interest to evolutionary biologists given that traits can vary within and among populations, across regions, and at continental or global scales. The pattern of regional variation can have an important impact on trait evolution at the local or population level. Using a common garden, we asked whether a geographically variable mosaic of tolerance to the widely applied herbicide RoundUp^® existed in two closely related co-occurring species of morning glory, Ipomoea purpurea and I. hederacea. We assayed RoundUp tolerance in over 1,700 plants representing 290 families from 29 populations in the southeastern United States. Our findings suggest that the two species of morning glory partition their respective levels of genetic variation for tolerance to glyphosate differently. Variation for tolerance in I. purpurea appears to exist among maternal lines and regions, whereas in I. hederacea, variation in tolerance existed only among populations. In addition, we find a significant hotspot of tolerance or positive spatial aggregation of this trait on a local scale in I. purpurea populations from the Coastal Plain. This suggests that either similar regimes of selection or gene flow between populations can produce a geographic mosaic of tolerance. These results highlight the fact that the genetic variation underlying an adaptive trait can exist at many different scales, whether it be within- or among-populations, among geographical ‘hotspots,’ or among distinct ecological regions. Given these results, the partitioning of genetic variation should be considered before making predictions about an adaptive trait’s evolutionary trajectory.     

Genetic covariances promote climatic adaptation in Australian Drosophila*

Evolutionary potential for adaptation hinges upon the orientation of genetic variation for traits under selection, captured by the additive genetic variance-covariance matrix (G), as well as the evolutionary stability of G. Yet studies that assess both the stability of G and its alignment with selection are extraordinarily rare. We evaluated the stability of G in three Drosophila melanogaster populations that have adapted to local climatic conditions along a latitudinal cline. We estimated population- and sex-specific G matrices for wing size and three climatic stress-resistance traits that diverge adaptively along the cline. To determine how G affects evolutionary potential within these populations, we used simulations to quantify how well G aligns with the direction of trait divergence along the cline (as a proxy for the direction of local selection) and how genetic covariances between traits and sexes influence this alignment. We found that G was stable across the cline, showing no significant divergence overall, or in sex-specific subcomponents, among populations. G also aligned well with the direction of clinal divergence, with genetic covariances strongly elevating evolutionary potential for adaptation to climatic extremes. These results suggest that genetic covariances between both traits and sexes should significantly boost evolutionary responses to environmental change.