Clines In Traits Research Articles

Within- and between-population comparisons suggest independently acting selection maintaining parallel clines in Scots pine (Pinus sylvestris).

Parallel clines in traits related to adaptation in a species can be due to independent selection on a pair of traits, or due to selection in one trait resulting in a parallel cline in a correlated trait. To distinguish between the mechanisms giving rise to parallel adaptive population divergence of multiple traits along an environmental gradient we need to study variation, correlations, and selective forces within individual populations along the gradient. In many tree species, budset timing (BST) forms a latitudinal cline, and parallel clinal variation is also found in other seedling traits, such as first-year height (FYH) and fall frost injury (FFI). In this study, we set up a common garden experiment with open pollinated progeny from natural populations of Scots pine (Pinus sylvestris), with one large sample from single population (500 families) and smaller samples from across a latitudinal gradient. BST, FYH and induced FFI were first measured in a greenhouse. The seedlings were then planted in the field, where survival and height were measured at the age of 9 years as fitness proxies. We compared between- and within-population variation and genetic correlations of these three seedling traits, and estimated selection gradients at the family level in our main population, taking into account the potential effects of seed weight. Between-population genetic correlations between seedling traits were high (0.76-0.95). Within-population genetic correlations in the main population were lower (0.14-0.35), as in other populations (0.10-0.39). Within population, extensive adaptive variation persists in the seedling traits, in line with rather weak selection gradients, yet maintaining the clines. Although our sampling does not cover the whole cline equally, the results suggest that the individual clines in these traits are maintained by largely independently acting selection, which results in fewer constraints in adaptation under changing climate.

The paradox of adaptive trait clines with nonclinal patterns in the underlying genes

Multivariate climate change presents an urgent need to understand how species adapt to complex environments. Population genetic theory predicts that loci under selection will form monotonic allele frequency clines with their selective environment, which has led to the wide use of genotype-environment associations (GEAs). This study used a set of simulations to elucidate the conditions under which allele frequency clines are more or less likely to evolve as multiple quantitative traits adapt to multivariate environments. Phenotypic clines evolved with nonmonotonic (i.e., nonclinal) patterns in allele frequencies under conditions that promoted unique combinations of mutations to achieve the multivariate optimum in different parts of the landscape. Such conditions resulted from interactions among landscape, demography, pleiotropy, and genetic architecture. GEA methods failed to accurately infer the genetic basis of adaptation under a range of scenarios due to first principles (clinal patterns did not evolve) or statistical issues (clinal patterns evolved but were not detected due to overcorrection for structure). Despite the limitations of GEAs, this study shows that a back-transformation of multivariate ordination can accurately predict individual multivariate traits from genotype and environmental data regardless of whether inference from GEAs was accurate. In addition, frameworks are introduced that can be used by empiricists to quantify the importance of clinal alleles in adaptation. This research highlights that multivariate trait prediction from genotype and environmental data can lead to accurate inference regardless of whether the underlying loci display clinal or nonmonotonic patterns.

Deep ecomorphological and genetic divergence in Steller's Jays (Cyanocitta stelleri, Aves: Corvidae).

The relationship between ecology and morphology is a cornerstone of evolutionary biology, and quantifying variation across environments can shed light on processes that give rise to biodiversity. Three morphotypes of the Steller's Jay (Cyanocitta stelleri) occupy different ecoregions in western North America, which vary in climate and landcover. These morphotypes (Coastal, Interior, Rocky Mountain) differ in size, plumage coloration, and head pattern. We sampled 1080 Steller's Jays from 68 populations (plus 11 outgroups) to address three main questions using data on morphology, plumage, genetics (mtDNA, microsatellites), and ecological niches: (1) How do phenotypic and genetic traits vary within and among populations, morphotypes, and ecoregions? (2) How do population-level differences in Steller's Jays compare with other sister species pairs of North American birds? (3) What can we infer about the population history of Steller's Jays in relation to past climates, paleoecology, and niche evolution? We found substantial morphological, genetic, and ecological differentiation among morphotypes. The greatest genetic divergence separated Coastal and Interior morphotypes from the Rocky Mountain morphotype, which was associated with warmer, drier, and more open habitats. Microsatellites revealed additional structure between Coastal and Interior groups. The deep mtDNA split between Coastal/Interior and Rocky Mountain lineages of Steller's Jay (ND2 ~ 7.8%) is older than most North American avian sister species and dates to approximately 4.3mya. Interior and Rocky Mountain morphotypes contact across a narrow zone with steep clines in traits and reduced gene flow. The distribution of the three morphotypes coincides with divergent varieties of ponderosa pine and Douglas fir. Species distribution models support multiple glacial refugia for Steller's Jays. Our integrative dataset combined with extensive geographic sampling provides compelling evidence for recognizing at least two species of Steller's Jay.

Evolutionary Rescue as a Mechanism Allowing a Clonal Grass to Adapt to Novel Climates.

Filing gaps in our understanding of species' abilities to adapt to novel climates is a key challenge for predicting future range shifts and biodiversity loss. Key knowledge gaps are related to the potential for evolutionary rescue in response to climate, especially in long-lived clonally reproducing species. We illustrate a novel approach to assess the potential for evolutionary rescue using a combination of reciprocal transplant experiment in the field to assess performance under a changing climate and independent growth chamber assays to assess growth- and physiology-related plant trait maxima and plasticities of the same clones. We use a clonal grass, Festuca rubra, as a model species. We propagated individual clones and used them in a transplant experiment across broad-scale temperature and precipitation gradients, simulating the projected direction of climate change in the region. Independent information on trait maxima and plasticities of the same clones was obtained by cultivating them in four growth chambers representing climate extremes. Plant survival was affected by interaction between plant traits and climate change, with both trait plasticities and maxima being important for adaptation to novel climates. Key traits include plasticity in extravaginal ramets, aboveground biomass, and osmotic potential. The direction of selection in response to a given climatic change detected in this study mostly contradicted the natural trait clines indicating that short-term selection pressure as identified here does not match long-term selection outcomes. Long-lived clonal species exposed to different climatic changes are subjected to consistent selection pressures on key traits, a necessary condition for adaptation to novel conditions. This points to evolutionary rescue as an important mechanism for dealing with climate change in these species. Our experimental approach may be applied also in other model systems broadening our understanding of evolutionary rescue. Such knowledge cannot be easily deduced from observing the existing field clines.

Range-wide variations in common milkweed traits and their effect on monarch larvae.

Leaf economic spectrum (LES) theory has historically been employed to inform vegetation models of ecosystem processes, but largely neglects intraspecific variation and biotic interactions. We attempt to integrate across environment-plant trait-herbivore interactions within a species at a range-wide scale. We measured traits in 53 populations spanning the range of common milkweed (Asclepias syriaca) and used a common garden to determine the role of environment in driving patterns of intraspecific variation. We used a feeding trial to determine the role of plant traits in monarch (Danaus plexippus) larval development. Trait-trait relationships largely followed interspecific patterns in LES theory and persisted in a common garden when individual traits change. Common milkweed showed intraspecific variation and biogeographic clines in traits. Clines did not persist in a common garden. Larvae ate more and grew larger when fed plants with more nitrogen. A longitudinal environmental gradient in precipitation corresponded to a resource gradient in plant nitrogen, which produces a gradient in larval performance. Biogeographic patterns in common milkweed traits can sometimes be predicted from LES, are largely driven by environmental conditions, and have consequences for monarch larval performance. Changes to nutrient dynamics of landscapes with common milkweed could potentially influence monarch population dynamics. We show how biogeographic trends in intraspecific variation can influence key ecological interactions, especially in common species with large distributions.

Rapid evolution of latitudinal clines in growth and defence of an invasive weed.

Re-establishment of heritable latitudinal clines in growth-related traits has been recognised as evidence for adaptive evolution in invasive plants. However, less information is known about latitudinal clines in defence and joint clinal evolution of growth and defence in invasive plants. We planted 14 native Argentinean populations and 14 introduced Chinese populations of Alternanthera philoxeroides in replicate common gardens in China. We investigated the latitudinal clines of traits related to growth and defence, and plasticity of these traits in relation to experiment site and soil nitrogen. We found that chemical defence decreased with latitude in introduced populations but increased with latitude in native populations. For growth rate, latitudinal clines were positive in introduced populations but nonexistent in native populations. There were also parallel positive latitudinal clines in total/shoot biomass and specific leaf area. Experiment site affected the occurrence or magnitude of latitudinal clines in growth rate, branch intensity and triterpenoid saponins concentration. Introduced populations were more plastic to experiment site and soil nitrogen than native populations. We provide evidence for rapid evolution of clines in growth and defence in an invasive plant. Altered herbivory gradients and trade-off between growth and defence may explain nonparallel clines between the native and introduced ranges.

Müllerian mimicry of a quantitative trait despite contrasting levels of genomic divergence and selection.

Hybrid zones, where distinct populations meet and interbreed, give insight into how differences between populations are maintained despite gene flow. Studying clines in genetic loci and adaptive traits across hybrid zones is a powerful method for understanding how selection drives differentiation within a single species, but can also be used to compare parallel divergence in different species responding to a common selective pressure. Here, we study parallel divergence of wing colouration in the butterflies Heliconius erato and H.melpomene, which are distantly related Müllerian mimics which show parallel geographic variation in both discrete variation in pigmentation, and quantitative variation in structural colour. Using geographic cline analysis, we show that clines in these traits are positioned in roughly the same geographic region for both species, which is consistent with direct selection for mimicry. However, the width of the clines varies markedly between species. This difference is explained in part by variation in the strength of selection acting on colour traits within each species, but may also be influenced by differences in the dispersal rate and total strength of selection against hybrids between the species. Genotyping-by-sequencing also revealed weaker population structure in H.melpomene, suggesting the hybrid zones may have evolved differently in each species, which may also contribute to the patterns of phenotypic divergence in this system. Overall, we conclude that multiple factors are needed to explain patterns of clinal variation within and between these species, although mimicry has probably played a central role.

Contrasting plant adaptation strategies to latitude in the native and invasive range of Spartina alterniflora.

Biological invasions offer model systems of contemporary evolution. We examined trait differences and evolution across geographic clines among continents of the intertidal grass Spartina alterniflora within its invasive and native ranges. We sampled vegetative and reproductive traits in the field at 20 sites over 20° latitude in China (invasive range) and 28 sites over 17° in the US (native range). We grew both Chinese and US plants in a glasshouse common garden for 3yr. Chinese plants were c.15% taller, c. 10% denser, and set up to four times more seed than US plants in both the field and common garden. The common garden experiments showed a striking genetic cline of seven-fold greater seed set at higher latitudes in the introduced but not the native range. By contrast, there was a slight genetic cline in some vegetative traits in the native but not the introduced range. Our results are consistent with others showing that introduced plants can evolve rapidly in the new range. S.alterniflora has evolved different trait clines in the native and introduced ranges, showing the importance of phenotypic plasticity and genetic control of change during the invasion process.

Spatial Population Structure Determines Extinction Risk in Climate-Induced Range Shifts.

Climate change is an escalating threat facing populations around the globe, necessitating a robust understanding of the ecological and evolutionary mechanisms dictating population responses. However, populations respond to climate change not in isolation but rather in the context of their existing ranges. In particular, spatial population structure within a range (e.g., trait clines, starkness of range edges, etc.) likely interacts with other ecological and evolutionary processes during climate-induced range shifts. Here, we use an individual-based model to explore the interacting roles of several such factors in range shift dynamics. We show that increased spatial population structure (driven primarily by a steeper environmental gradient) severely increases a population's extinction risk. Further, we show that while evolution of heightened dispersal during range shifts can aid populations in tracking changing conditions, it can also interact negatively with adaptation to the environmental gradient, leading to reduced fitness and contributing to the increased extinction risk observed in populations structured along steep environmental gradients. Our results demonstrate that the effect of dispersal evolution on range-shifting populations is dependent on environmental context and that spatial population structure can substantially increase extinction risk in range shifts.

Interannual variation in season length is linked to strong co‐gradient plasticity of phenology in a montane annual plant

Species are commonly distributed along latitudinal and elevational gradients of growing season length to which they might respond via phenotypic plasticity and/or adaptive genetic differentiation. However, the relative contribution of these processes and whether plasticity, if it occurs, facilitates expansion along season-length gradients remain unclear, but are important for predicting species fates during anthropogenic change. We quantified phenological trait variation in the montane annual Rhinanthus minor for three generations at 12 sites across 900m of elevation in the Canadian Rocky Mountains and conducted a reciprocal transplant experiment for two generations among nine sites. We compared clines and interannual variation of phenological traits between natural and transplanted individuals. Season length declined by c.37% along our elevational gradient and, as expected, plants emerged, reached first flower and made their first seed in c.41% fewer growing degree days under shorter growing seasons. Although reciprocal transplants revealed modest genetic differentiation across elevation, trait clines primarily were due to striking co-gradient plasticity that paralleled genetic differentiation. Co-gradient plasticity likely evolved in response to considerable interannual variation in season length across our elevational transect, and should prepare R.minor to make adaptive changes to phenology in response to ongoing climate change predicted for montane environments.

Rapid growth and defence evolution following multiple introductions.

Rapid adaptation can aid invasive populations in their competitive success. Resource allocation trade‐off hypotheses predict higher resource availability or the lack of natural enemies in introduced ranges allow for increased growth and reproduction, thus contributing to invasive success. Evidence for such hypotheses is however equivocal and tests among multiple ranges over productivity gradients are required to provide a better understanding of the general applicability of these theories.Using common gardens, we investigated the adaptive divergence of various constitutive and inducible defence‐related traits between the native North American and introduced European and Australian ranges, while controlling for divergence due to latitudinal trait clines, individual resource budgets, and population differentiation, using >11,000 SNPs.Rapid, repeated clinal adaptation in defence‐related traits was apparent despite distinct demographic histories. We also identified divergence among ranges in some defence‐related traits, although differences in energy budgets among ranges may explain some, but not all, defence‐related trait divergence. We do not identify a general reduction in defence in concert with an increase in growth among the multiple introduced ranges as predicted trade‐off hypotheses. Synthesis: The rapid spread of invasive species is affected by a multitude of factors, likely including adaptation to climate and escape from natural enemies. Unravelling the mechanisms underlying invasives' success enhances understanding of eco‐evolutionary theory and is essential to inform management strategies in the face of ongoing climate change. OPEN RESEARCH BADGES This article has been awarded Open Materials, Open Data, Preregistered Research Designs Badges. All materials and data are publicly accessible via the Open Science Framework at https://doi.org/10.6084/m9.figshare.8028875.v1, https://github.com/lotteanna/defence_adaptation,https://doi.org/10.1101/435271.

Biogeography of a plant invasion: genetic variation and plasticity in latitudinal clines for traits related to herbivory

AbstractThe juxtaposition of plant‐species invasions with latitudinal gradients in herbivore pressure is an important yet mostly unexplored issue in invasion biology. Latitudinal clines in defense and palatability to herbivores are expected to exist in native plant species but the evolution of these clines may lag behind for invasive plant species resulting in non‐parallel latitudinal clines that may impact invasion success. Our study focused on a native and European invasive lineages of the common reed Phragmites australis in North America. Using native and invasive genotypes of P. australis collected across a 17° latitudinal range, we performed experiments in replicate northern and southern common gardens to investigate whether these two lineages exhibited different genetically based latitudinal clines in defenses, nutritional condition, and palatability to their herbivores, the aphid Hyalopterus pruni and the fall armyworm Spodoptera frugiperda. We also tested whether invasive genotypes are more phenotypically plastic than native genotypes and whether plasticity varies with latitude. Although invasive genotypes did not exhibit higher defense levels (leaf toughness, phenolics, percent carbon), they were considerably less palatable to their herbivores than native genotypes. Genetically based latitudinal clines were evident for both native and invasive P. australis and for all defenses, nutrients, and at least one palatability trait for each herbivore. In 36% of the cases where clines were evident, they were non‐parallel between the two lineages. These data suggest that clines in the invasive genotypes of P. australis evolved within the past ~100 years. Moreover, our study showed that the occurrence and direction of latitudinal clines in plant traits were commonly dependent on where the study was conducted (north or south), indicating strong phenotypic plasticity in these genetic‐based clines. Finally, traits for invasive genotypes of P. australis were 2.5 times more plastic than traits for native genotypes. Interestingly, plasticity for native but not invasive genotypes was strongly dependent on latitude of origin. Such spatial heterogeneity within and between the native and invasive lineages of P. australis with respect to their interactions with herbivores can generate substantial spatial variability in biotic resistance that can have important implications for the establishment and spread of invasive genotypes and species.

Contemporary evolution during invasion: evidence for differentiation, natural selection, and local adaptation.

Biological invasions are 'natural' experiments that can improve our understanding of contemporary evolution. We evaluate evidence for population differentiation, natural selection and adaptive evolution of invading plants and animals at two nested spatial scales: (i) among introduced populations (ii) between native and introduced genotypes. Evolution during invasion is frequently inferred, but rarely confirmed as adaptive. In common garden studies, quantitative trait differentiation is only marginally lower (~3.5%) among introduced relative to native populations, despite genetic bottlenecks and shorter timescales (i.e. millennia vs. decades). However, differentiation between genotypes from the native vs. introduced range is less clear and confounded by nonrandom geographic sampling; simulations suggest this causes a high false-positive discovery rate (>50%) in geographically structured populations. Selection differentials (¦s¦) are stronger in introduced than in native species, although selection gradients (¦β¦) are not, consistent with introduced species experiencing weaker genetic constraints. This could facilitate rapid adaptation, but evidence is limited. For example, rapid phenotypic evolution often manifests as geographical clines, but simulations demonstrate that nonadaptive trait clines can evolve frequently during colonization (~two-thirds of simulations). Additionally, QST-FST studies may often misrepresent the strength and form of natural selection acting during invasion. Instead, classic approaches in evolutionary ecology (e.g. selection analysis, reciprocal transplant, artificial selection) are necessary to determine the frequency of adaptive evolution during invasion and its influence on establishment, spread and impact of invasive species. These studies are rare but crucial for managing biological invasions in the context of global change.

Parallel invasions produce heterogenous patterns of life history adaptation: rapid divergence in an invasive insect

Biotic invasions provide a natural experiment in evolution: when invasive species colonize new ranges, they may evolve new clines in traits in response to environmental gradients. Yet it is not clear how rapidly such patterns can evolve and whether they are consistent between regions. We compare four populations of the invasive cabbage white butterfly (Pieris rapae) from North America and Japan, independently colonized by P. rapae 150 years ago and 300 years ago, respectively. On each continent, we employed a northern and southern population to compare the effects of latitude on body mass, development rate and immune function. For each population, we used a split-sibling family design in which siblings were reared at either warm (26.7 °C) or cool (20 °C) temperatures to determine reaction norms for each trait. Latitudinal patterns in development time were similar between the two continents. In contrast, there were strong geographical differences in reaction norms for body size, but no consistent effects of latitude; there were no detectable effects of latitude or continent on immune function. These results imply that some life history traits respond consistently to selection along climatic gradients, whereas other traits may respond to local environmental factors, or not at all.

Adaptive geographical clines in the growth and defense of a native plant

Broad‐scale geographical gradients in the abiotic environment and interspecific interactions should select for clinal adaptation. How trait clines evolve has recently received increased attention because of anticipated climate change and the importance of rapid evolution in invasive species. This issue is particularly relevant for clines in growth and defense of plants, because both sets of traits are closely tied to fitness and because such sessile organisms experience strong local selection. Yet despite widespread recognition that growth and defense traits are intertwined, the general issue of their joint clinal evolution is not well resolved.To address heritable clinal variation and adaptation of growth and defense traits of common milkweed (Asclepias syriaca), we planted seed from 22 populations encompassing the species' latitudinal range in common gardens near the range center (New York) and toward the range edges (New Brunswick and North Carolina). Populations were differentiated in 13 traits, and six traits showed genetically based latitudinal clines. Higher‐latitude populations had earlier phenology, lower shoot biomass, more root buds and clonal growth, higher root‐to‐shoot ratio, and greater latex production. The cline in shoot biomass was consistent in all three locations. Selection on phenology was reversed in New Brunswick and North Carolina, with early genotypes favored in the north but not the south. We found no clines in foliar trichomes or toxic cardenolides. Annual precipitation of source populations explained variation in phenology, clonal growth, root‐to‐shoot ratio, and latex. Across four traits measured in New Brunswick and North Carolina, we found garden‐by‐latitude (and garden‐by‐precipitation) interactions, indicating plasticity in genetically based trait clines.In the two gardens with substantial herbivory (New York and North Carolina), northern populations showed higher resistance to insects. Resistance to aphids was driven by trichomes and water content, while resistance to monarch caterpillars was driven by latex. However, surveys of natural populations indicated that leaf damage and insect diversity on milkweed are low at the geographical extremes (New Brunswick and North Carolina) and higher toward the range center. We speculate that milkweed plants evolved clines in growth traits in response to climate, and that this set the template for tolerance to herbivory, which subsequently shaped the evolution of defensive traits.

Polymorphism in the couch potato gene clines in eastern Australia but is not associated with ovarian dormancy in Drosophila melanogaster

Natural selection can generate parallel latitudinal clines in traits and gene frequencies across continents, but these have rarely been linked. An amino acid (isoleucine to lysine, or I462K) polymorphism of the couch potato (cpo) gene in Drosophila melanogaster is thought to control female reproductive diapause cline in North America (Schmidt et al. 2008, Proc Natl Acad Sci USA, 105, 16207-16211). Here, we show that under standard diapause-inducing conditions (12 °C and short photoperiod) (Saunders et al. 1989, Proc Natl Acad Sci USA, 86, 3748-3752), egg maturation in Australian flies is delayed, but not arrested at previtellogenic stages. At 12 °C, the phenotypic distribution in egg development was bimodal at stages 8 and 14 and showed a strong nonlinear pattern on the east coast of Australia, with incidence of egg maturation delay (ovarian dormancy) increasing both toward tropical and temperate climates. Furthermore, we found no evidence for an association between the cpo I462K polymorphism and ovarian dormancy at either 12 or 10 °C (when egg maturation was often delayed at stage 7). Owing to strong linkage disequilibrium, the latitudinal cline in cpo allele frequencies was no longer evident once variation in the In(3R)P inversion polymorphism was taken into account. Our results suggest that the standard diapause-inducing conditions (12 °C and short photoperiod) were not sufficient to cause the typical previtellogenic developmental arrest in Australian flies and that the cpo I462K polymorphism does not explain the observed delay in egg development. In conclusion, ovarian dormancy does not show a simple latitudinal cline, and the lack of cpo-dormancy association suggests a different genetic basis to reproductive dormancy in North America and Australia.

Large scale geographic clines of parasite damage to Populus tremula L.

According to the geographic mosaic theory of coevolution (GMTC), clines of traits reflecting local co‐adaptation (including resistance genes) should be common between a host and its parasite and should persist across time. To test the GMTC‐assumption of persistent clinal patterns we compared the natural prevalence of two parasites on aspen Populus tremula trees: mining moths of the genus Phyllocnistis and leaf rust Melampsora spp. Damage data were collated from the Swedish National Forest Damage Inventory (2004–2006). In addition, occurrence of the parasites was scored in field conditions in two common gardens in the north and south of Sweden over five growing seasons (2004–2008), then related to biomass (stem height and diameter) and to concentrations of eleven leaf phenolics. Phyllocnistis mainly occurred in the northern garden, a distribution range which was confirmed by the countrywide inventory, although Phyllocnistis was more abundant on southern clones, providing evidence for possible local maladaptation. Melampsora occurred all over the country and in both gardens, but built up more quickly on northern clones, which suggests a centre of local clone maladaptation in the north. Stem growth also followed a clinal pattern as did the concentration of three phenolic compounds: benzoic acid, catechin and cinnamic acid. However, only benzoic acid was related to parasite presence: negatively to Phyllocnistis and positively to Melampsora and it could thus be a potential trait under selection.In conclusion, clines of Phyllocnistis were stronger and more persistent compared to Melampsora, which showed contrasting clines of varying strength. Our data thus support the assumption of the GMTC model that clines exist in the border between hot and cold spots and that they may be less persistent for parasites with an elevated gene flow, and/or for parasites which cover relatively larger hot spots surrounded by fewer cold spots.

Adaptation and colonization history affect the evolution of clines in two introduced species

Phenotypic and genetic clines have long been synonymous with adaptive evolution. However, other processes (for example, migration, range expansion, invasion) may generate clines in traits or loci across geographical and environmental gradients. It is therefore important to distinguish between clines that represent adaptive evolution and those that result from selectively neutral demographic or genetic processes. We tested for the differentiation of phenotypic traits along environmental gradients using two species in the genus Silene, whilst statistically controlling for colonization history and founder effects. We sampled seed families from across the native and introduced ranges, genotyped individuals and estimated phenotypic differentiation in replicated common gardens. The results suggest that post-glacial expansion of S. vulgaris and S. latifolia involved both neutral and adaptive genetic differentiation (clines) of life history traits along major axes of environmental variation in Europe and North America. Phenotypic clines generally persisted when tested against the neutral expectation, although some clines disappeared (and one cline emerged) when the effects of genetic ancestry were statistically removed. Colonization history, estimated using genetic markers, is a useful null model for tests of adaptive trait divergence, especially during range expansion and invasion when selection and gene flow may not have reached equilibrium.

Genetic, morphological, and feather isotope variation of migratory willow warblers show gradual divergence in a ring

The circular distribution of the willow warbler Phylloscopus trochilus around the Baltic Sea shares many features with the classic examples of ring species; however, the system is much younger. It has previously been shown that a secondary contact zone is located in central Scandinavia, where there are narrow clines for several morphological traits coincident with a migratory divide. Here we analyse multiple traits and genes from > 1700 males captured on breeding territories at 77 sites spread around the Baltic Sea to test the following hypothesis. If the secondary contact zone in Scandinavia is a result of divergence in two allopatric refuge populations during the last glaciation, we expect to find a similar secondary contact zone somewhere else around the circular distribution. Our results show that the trait clines were wider and displaced from each other along the eastern side of the Baltic Sea. Analyses of 12 microsatellite loci confirmed that the genome is very similar between the terminal forms (F(ST) = 0). Two AFLP-derived markers filtered out from a genomic scan instead appear to be maintained by selection. These markers exhibited steep clines at the secondary contact zone in Scandinavia, but as for the phenotypic traits, had vastly different cline centres east of the Baltic Sea. The trait clines along the ring distribution outside the Scandinavian secondary contact zone thus seem to have been shaped by independent action of selection or drift during the process of postglacial colonization.

Establishment of parallel altitudinal clines in traits of native and introduced forbs

Due to altered ecological and evolutionary contexts, we might expect the responses of alien plants to environmental gradients, as revealed through patterns of trait variation, to differ from those of the same species in their native range. In particular, the spread of alien plant species along such gradients might be limited by their ability to establish clinal patterns of trait variation. We investigated trends in growth and reproductive traits in natural populations of eight invasive Asteraceae forbs along altitudinal gradients in their native and introduced ranges (Valais, Switzerland, and Wallowa Mountains, Oregon, USA). Plants showed similar responses to altitude in both ranges, being generally smaller and having fewer inflorescences but larger seeds at higher altitudes. However, these trends were modified by region-specific effects that were independent of species status (native or introduced), suggesting that any differential performance of alien species in the introduced range cannot be interpreted without a fully reciprocal approach to test the basis of these differences. Furthermore, we found differences in patterns of resource allocation to capitula among species in the native and the introduced areas. These suggest that the mechanisms underlying trait variation, for example, increasing seed size with altitude, might differ between ranges. The rapid establishment of clinal patterns of trait variation in the new range indicates that the need to respond to altitudinal gradients, possibly by local adaptation, has not limited the ability of these species to invade mountain regions. Studies are now needed to test the underlying mechanisms of altitudinal clines in traits of alien species.