The evolution of species' distributions: reciprocal transplants across the elevation ranges of Mimulus cardinalis and M. lewisii

Every species occupies a limited geographic area, but it remains unclear why traits that limit distribution do not evolve to allow range expansion. Hypotheses for the evolutionary stability of geographic ranges assume that species are maladapted at the range boundary and unfit beyond the current range, but this assumption has rarely been tested. To examine how fitness varies across species' ranges, we reciprocally transplanted two species of monkeyflowers, Mimulus cardinalis and M. lewisii, within and beyond their present elevation ranges. We used individuals of known parentage from populations collected across the elevation ranges of both species to examine whether populations are adapted to position within the range. For both species we found the greatest average fitness at elevations central within the range, reduced fitness at the range margin, and zero or near-zero fitness when transplanted beyond their present elevation range limits. However, the underlying causes of fitness variation differed between the species. At high elevations beyond its range, M. cardinalis displayed reduced growth and fecundity, whereas at low elevations M. lewisii experienced high mortality. Weak differences in performance were observed among populations within each species and these were not related to elevation of origin. Low fitness of both species at their range margin and weak differentiation among populations within each species suggest that adaptation to the environment at and beyond the range margin is hindered, illustrating that range margins provide an interesting system in which to study limits to adaptation.     

Interactions between soil habitat and geographic range location affect plant fitness

Populations are often found on different habitats at different geographic locations. This habitat shift may be due to biased dispersal, physiological tolerances or biotic interactions. To explore how fitness of the native plant Chamaecrista fasciculata depends on habitat within, at and beyond its range edge, we planted seeds from five populations in two soil substrates at these geographic locations. We found that with reduced competition, lifetime fitness was always greater or equivalent in one habitat type, loam soils, though early-season survival was greater on sand soils. At the range edge, natural populations are typically found on sand soil habitats, which are also less competitive environments. Early-season survival and fitness differed among source populations, and when transplanted beyond the range edge, range edge populations had greater fitness than interior populations. Our results indicate that even when the optimal soil substrate for a species does not change with geographic range location, the realized niche of a species may be restricted to sub-optimal habitats at the range edge because of the combined effects of differences in abiotic and biotic effects (e.g. competitors) between substrates.     

Environmental marginality and geographic range limits: a case study with Arabidopsis lyrata ssp. lyrata

Understanding the factors that govern the distribution of species is a central goal of evolutionary ecology. It is commonly assumed that geographic range limits reflect ecological niche limits and that species experience increasingly marginal conditions towards the edge of their ranges. Using spatial data and ecological niche models we tested these hypotheses in Arabidopsis lyrata. Specifically, we asked whether range limits coincide with predicted niche limits in this system and whether the suitability of sites declines towards the edge of the species’ range in North America. We further explored patterns of environmental change towards the edge of the range and asked whether genome‐wide patterns of genetic diversity decline with increasing peripherality and environmental marginality. Our results suggest that latitudinal range limits coincide with niche limits. Populations experienced increasingly marginal environments towards these limits – though patterns of environmental change were more complex than most theoretical models for range limits assume. Genomic diversity declined towards the edge of the species’ range and with increasing distance from the estimated centre of the species’ niche in environmental space, but not with the suitability of sites based on niche model predictions. Thus while latitudinal range limits in this system are broadly associated with niche limits, the link between environmental conditions and genetic diversity (and thus the adaptive potential of populations) is less clear.     

Maladaptation beyond a geographic range limit driven by antagonistic and mutualistic biotic interactions across an abiotic gradient

Species’ geographic range limits often result from maladaptation to the novel environments beyond the range margin. However, we rarely know which aspects of the n‐dimensional environment are driving this maladaptation. Especially of interest is the influence of abiotic versus biotic factors in delimiting species’ distributions. We conducted a 2‐year reciprocal transplant experiment involving manipulations of the biotic environment to explore how spatiotemporal gradients in precipitation, fatal mammalian herbivory, and pollination affected lifetime fitness within and beyond the range of the California annual plant, Clarkia xantiana ssp. xantiana. In the first, drier year of the experiment, fitness outside the range edge was limited mainly by low precipitation, and there was some evidence for local adaptation within the range. In the second, wetter year, we did not observe abiotic limitations to plant fitness outside the range; instead biotic interactions, especially herbivory, limited fitness outside the range. Together, protection from herbivory and supplementation of pollen resulted in three‐ to sevenfold increases in lifetime fitness outside the range margin in the abiotically benign year. Overall, our work demonstrates the importance of biotic interactions, particularly as they interact with the abiotic environment, in determining fitness beyond geographic range boundaries.     

Evolutionarily stable range limits set by interspecific competition

A combination of abiotic and biotic factors probably restricts the range of many species. Recent evolutionary models and tests of those models have asked how a gradual change in environmental conditions can set the range limit, with a prominent idea being that gene flow disrupts local adaptation. We investigate how biotic factors, explicitly competition for limited resources, result in evolutionarily stable range limits even in the absence of the disruptive effect of gene flow. We model two competing species occupying different segments of the resource spectrum. If one segment of the resource spectrum declines across space, a species that specializes on that segment can be driven to extinction, even though in the absence of competition it would evolve to exploit other abundant resources and so be saved. The result is that a species range limit is set in both evolutionary and ecological time, as the resources associated with its niche decline. Factors promoting this outcome include: (i) inherent gaps in the resource distribution, (ii) relatively high fitness of the species when in its own niche, and low fitness in the alternative niche, even when resource abundances are similar in each niche, (iii) strong interspecific competition, and (iv) asymmetric interspecific competition. We suggest that these features are likely to be common in multispecies communities, thereby setting evolutionarily stable range limits.     

Mixed population genomics support for the central marginal hypothesis across the invasive range of the cane toad (Rhinella marina) in Australia

Understanding factors that cause species' geographic range limits is a major focus in ecology and evolution. The central marginal hypothesis (CMH) predicts that species cannot adapt to conditions beyond current geographic range edges because genetic diversity decreases from core to edge due to smaller, more isolated edge populations. We employed a population genomics framework using 24 235–33 112 SNP loci to test major predictions of the CMH in the ongoing invasion of the cane toad (Rhinella marina) in Australia. Cane toad tissue samples were collected along broad‐scale, core‐to‐edge transects across their invasive range. Geographic and ecological core areas were identified using GIS and habitat suitability indices from ecological niche modelling. Bayesian clustering analyses revealed three genetic clusters, in the northwest invasion‐front region, northeast precipitation‐limited region and southeast cold temperature‐limited region. Core‐to‐edge patterns of genetic diversity and differentiation were consistent with the CMH in the southeast, but were not supported in the northeast and showed mixed support in the northwest. Results suggest cold temperatures are a likely contributor to southeastern range limits, consistent with CMH predictions. In the northeast and northwest, ecological processes consisting of a steep physiological barrier and ongoing invasion dynamics, respectively, are more likely explanations for population genomic patterns than the CMH.     

A synthesis of transplant experiments and ecological niche models suggests that range limits are often niche limits

Global change has made it important to understand the factors that shape species’ distributions. Central to this area of research is the question of whether species’ range limits primarily reflect the distribution of suitable habitat (i.e. niche limits) or arise as a result of dispersal limitation. Over‐the‐edge transplant experiments and ecological niche models are commonly used to address this question, yet few studies have taken advantage of a combined approach for inferring the causes of range limits. Here, we synthesise results from existing transplant experiments with new information on the predicted suitability of sites based on niche models. We found that individual performance and habitat suitability independently decline beyond range limits across multiple species. Furthermore, inferences from transplant experiments and niche models were generally concordant within species, with 31 out of 40 cases fully supporting the hypothesis that range limits are niche limits. These results suggest that range limits are often niche limits and that the factors constraining species’ ranges operate at scales detectable by both transplant experiments and niche models. In light of these findings, we outline an integrative framework for addressing the causes of range limits in individual species.     

Local adaptation primes cold‐edge populations for range expansion but not warming‐induced range shifts

According to theory, edge populations may be poised to expand species’ ranges if they are locally adapted to extreme conditions, or ill‐suited to colonise beyond‐range habitat if their offspring are genetically and competitively inferior. We tested these contrasting predictions by transplanting low‐, mid‐, and high‐elevation (edge) populations of an annual plant throughout and above its elevational distribution. Seed from poor‐quality edge habitat (one of two transects) had inferior emergence, but edge seeds also had adaptive phenology (both transects). High‐elevation plants flowered earlier, required less heat accumulation to mature seed, and so achieved higher lifetime fitness at and above the range edge. Experimental warming improved fitness above the range, but eliminated the advantage of local cold‐edge populations, supporting recent models in which cold‐adapted edge populations do not facilitate warming‐induced range shifts. The highest above‐range fitness was achieved by a ‘super edge phenotype’ from a neighbouring mountain, suggesting key adaptations exist regionally even if absent from local edge populations.     

Range edges in heterogeneous landscapes: Integrating geographic scale and climate complexity into range dynamics

The impacts of climate change have re‐energized interest in understanding the role of climate in setting species geographic range edges. Despite the strong focus on species' distributions in ecology and evolution, defining a species range edge is theoretically and empirically difficult. The challenge of determining a range edge and its relationship to climate is in part driven by the nested nature of geography and the multidimensionality of climate, which together generate complex patterns of both climate and biotic distributions across landscapes. Because range‐limiting processes occur in both geographic and climate space, the relationship between these two spaces plays a critical role in setting range limits. With both conceptual and empirical support, we argue that three factors—climate heterogeneity, collinearity among climate variables, and spatial scale—interact to shape the spatial structure of range edges along climate gradients, and we discuss several ways that these factors influence the stability of species range edges with a changing climate. We demonstrate that geographic and climate edges are often not concordant across species ranges. Furthermore, high climate heterogeneity and low climate collinearity across landscapes increase the spectrum of possible relationships between geographic and climatic space, suggesting that geographic range edges and climatic niche limits correspond less frequently than we may expect. More empirical explorations of how the complexity of real landscapes shapes the ecological and evolutionary processes that determine species range edges will advance the development of range limit theory and its applications to biodiversity conservation in the context of changing climate.     

Energy and spatial order in niche and community

The concept of a varying evolutionary tempo that is regulated by energy was first suggested in the 1950s. It was based on the observation that energy‐rich habitats appear to be the centres of evolutionary change, producing novel characters more frequently and having greater speciation rates. Subsequently, a number of studies have found positive relationships between evolutionary rate and energy. Gradients of energy occur across a range of scales and these have been invoked to explain, for example, higher tropical species richness. Precipitation has also been found to influence evolutionary rate, suggesting that biologically available energy and productivity are the important variables in the relationship rather than solar energy alone. Here, we take the theoretical step of investigating at smaller scales the implications of faster evolutionary tempo where productivity is greater: first at the level of the species population and, subsequently, for the same‐guild co‐habitants in a community. To facilitate, this we begin by applying the concept of gradients of available energy, from more productive to less productive sectors within the species niche, in the context of source‐sink theory. We then propose that a species population will have its highest rate of evolution, for changes that confer a positive selection coefficient throughout the niche, in that sector where it has best adaptive fit and greatest per capita energy flux. We also posit that, where it conferred a shift in fitness, an evolutionary change at niche optimum could subsequently affect conspecific populations occupying lower energy niche sectors via the selection‐mediated dispersal of the apomorphy into the more marginal components of the niche. We then infer that this type of change in a species with a generally higher per capita energy flux might negatively affect adjacent more slowly evolving species (living in less productive peripheral niches) in situations where the increased fitness conferred by a particular apomorphy was relevant to conditions occurring beyond the limits of the progenitor niche hypervolume. We therefore suggest a directional component to evolution at the compressed scales of niche and community level microevolution whereby populations and species occupying more productive conditions have greater tempos of change with attendant enhancements of both their competitive influence and their evolutionary potential. In this manner, previously recorded macro‐evolutionary patterns indicating faster evolution with increased energy at larger scales are interpreted in the context of proposed micro‐evolutionary relationships at intraspecific and locally interspecific scales. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 696–714.     

Ecological niche differentiation in peripheral populations: a comparative analysis of eleven Mediterranean plant species

The ‘central‐peripheral’ hypothesis has provided a baseline for many studies of population dynamics and genetic variability at species distribution limits. Although peripheral populations are often assumed to occur in ecologically marginal conditions, little is known about whether they effectively occur in a distinct ecological niche. A cross‐taxa analysis of 11 Mediterranean vascular plants were studied. We quantified variation in the ecological niche between populations at the northern range limits of species in Mediterranean France and those in the central part of the distribution in continental Spain or Italy in 2013–2014. We analyzed both the macro‐ecological niche where populations occur in terms of broad habitat and altitudinal range and the micro‐ecological niche where individual plants grow in terms of soil and structural biotic and abiotic characteristics. Most species occur in a single broad habitat type common to central and peripheral populations and have a narrower altitudinal range in the latter. In contrast, for the micro‐ecological niche we detected marked variation in several niche parameters among central and peripheral populations. Although many differences are species‐specific some are common to several species. We found a trend towards narrower micro‐niche breadth in peripheral populations. Our results illustrate the importance of studying the precise ecological characteristics where plants grow and the pertinence of a multi‐species approach to correctly assess niche variation. The ecological originality of peripheral populations underlines their evolutionary potential and conservation significance.     

Connecting genomic patterns of local adaptation and niche suitability in teosintes

The central abundance hypothesis predicts that local adaptation is a function of the distance to the centre of a species’ geographic range. To test this hypothesis, we gathered genomic diversity data from 49 populations, 646 individuals and 33,464 SNPs of two wild relatives of maize, the teosintes Zea mays ssp. parviglumis and Zea. mays. ssp. mexicana. We examined the association between the distance to their climatic and geographic centroids and the enrichment of SNPs bearing signals of adaptation. We identified candidate adaptive SNPs in each population by combining neutrality tests and cline analyses. By applying linear regression models, we found that the number of candidate SNPs is positively associated with niche suitability, while genetic diversity is reduced at the limits of the geographic distribution. Our results suggest that overall, populations located at the limit of the species’ niches are adapting locally. We argue that local adaptation to this limit could initiate ecological speciation processes and facilitate adaptation to global change.     

Disentangling the effects of geographic peripherality and habitat suitability on neutral and adaptive genetic variation in Swiss stone pine

It is generally accepted that the spatial distribution of neutral genetic diversity within a species’ native range mostly depends on effective population size, demographic history, and geographic position. However, it is unclear how genetic diversity at adaptive loci correlates with geographic peripherality or with habitat suitability within the ecological niche. Using exome‐wide genomic data and distribution maps of the Alpine range, we first tested whether geographic peripherality correlates with four measures of population genetic diversity at > 17,000 SNP loci in 24 Alpine populations (480 individuals) of Swiss stone pine (Pinus cembra) from Switzerland. To distinguish between neutral and adaptive SNP sets, we used four approaches (two gene diversity estimates, F_ST outlier test, and environmental association analysis) that search for signatures of selection. Second, we established ecological niche models for P. cembra in the study range and investigated how habitat suitability correlates with genetic diversity at neutral and adaptive loci. All estimates of neutral genetic diversity decreased with geographic peripherality, but were uncorrelated with habitat suitability. However, heterozygosity (H_e) at adaptive loci based on Tajima's D declined significantly with increasingly suitable conditions. No other diversity estimates at adaptive loci were correlated with habitat suitability. Our findings suggest that populations at the edge of a species' geographic distribution harbour limited neutral genetic diversity due to demographic properties. Moreover, we argue that populations from suitable habitats went through strong selection processes, are thus well adapted to local conditions, and therefore exhibit reduced genetic diversity at adaptive loci compared to populations at niche margins.     

Broad-scale latitudinal variation in female reproductive success contributes to the maintenance of a geographic range boundary in bagworms (Lepidoptera: Psychidae)

Background

Geographic range limits and the factors structuring them are of great interest to biologists, in part because of concerns about how global change may shift range boundaries. However, scientists lack strong mechanistic understanding of the factors that set geographic range limits in empirical systems, especially in animals.

Methodology/Principal Findings

Across dozens of populations spread over six degrees of latitude in the American Midwest, female mating success of the evergreen bagworm Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae) declines from ∼100% to ∼0% near the edge of the species range. When coupled with additional latitudinal declines in fecundity and in egg and pupal survivorship, a spatial gradient of bagworm reproductive success emerges. This gradient is associated with a progressive decline in local abundance and an increased risk of local population extinction, up to a latitudinal threshold where extremely low female fitness meshes spatially with the species'' geographic range boundary.

Conclusions/Significance

The reduction in fitness of female bagworms near the geographic range limit, which concords with the abundant centre hypothesis from biogeography, provides a concrete, empirical example of how an Allee effect (increased pre-reproductive mortality of females in sparsely populated areas) may interact with other demographic factors to induce a geographic range limit.     

Habitat suitability and the constraints of migration in New World warblers

Identifying the factors that influence geographic range limits can illustrate the various ecological, physiological, and evolutionary constraints imposed on a species. The range limits of migratory birds are particularly challenging to study as they occur in disjunct regions at different times of the year, which can impose different constraints. Travel between breeding and wintering regions poses a significant energetic and navigational challenge to birds, although it is not clear how these movements influence breeding dispersal and range expansion. Here I ask whether the possible costs of migration limit the breeding ranges of wood warblers, a group of birds with an extensive history of ecological and evolutionary studies. I used occurrence records for multiple wood warbler species, breeding primarily in the boreal forest of North America, to generate environmental niche models. I tested for over‐prediction of habitat suitability into the western boreal forest, where most these species do not have occurrence records but where there is presumably suitable habitat. I found that some of these vagile taxa, primarily found east of the Rocky Mountains, also have predicted habitat suitability that extends into the north and west, where they have little to no occurrence records. I discuss several possible explanations for this discordance. In particular, the patterns are consistent with the costs of a long‐distance migration limiting the benefits of range expansion, as migration may become too onerous as the distance between breeding and wintering areas increases. These results speak to the process of niche filling more generally and call attention to an under‐appreciated explanation for why migratory species may not fully occupy their fundamental niche.     

The role of climate for the range limits of parapatric European land salamanders

Abrupt range limits of parapatric species may serve as a model system to understand the factors that determine species’ range borders. Theory suggests that parapatric range limits can be caused by abiotic conditions along environmental gradients, biotic interactions or a combination of both. Geographic ranges of the parapatric salamanders, Salamandra salamandra and S. atra, meet in small contact zones in the European Alps and to date, the cause of parapatry and the restricted range of S. atra remain elusive. We combine multivariate approaches and climatic data analysis to explore niche differentiation among the two salamanders with respect to the available climatic environment at their contact zones. Our purpose is to evaluate whether climatic conditions explain the species’ sharp range limits or if biotic interactions may play a role for range delimitation. Analyses were carried out in three contact zones in Switzerland to assess possible geographic variation. Our results indicate that both species occur at localities with different climatic conditions as well as the presence of a strong climatic gradient across the species’ range limits. Although the species’ climatic niches differ moderately (with a wider niche breadth for S. atra), interspecific niche overlap is found. Comparisons among the contact zones confirm geographic variation in the species’ climatic niches as well as in the conditions within the geographically available space. Our results suggest that the change in climatic conditions along the recognized gradient represents a determining factor for species’ range limits within contact zones. However, our analyses of geographic variation in climatic conditions reveal that both salamander species can occur in a much wider range of conditions than observed within contact zones. This finding and the interspecific climatic niche overlap within each contact zone provides indirect evidence that biotic interactions (likely competition) between the two species may also determine their range limits.     

Thermal Carrying Capacity for a Thermally-Sensitive Species at the Warmest Edge of Its Range

Anthropogenic environmental change is causing unprecedented rates of population extirpation and altering the setting of range limits for many species. Significant population declines may occur however before any reduction in range is observed. Determining and modelling the factors driving population size and trends is consequently critical to predict trajectories of change and future extinction risk. We tracked during 12 years 51 populations of a cold-water fish species (brown trout Salmo trutta) living along a temperature gradient at the warmest thermal edge of its range. We developed a carrying capacity model in which maximum population size is limited by physical habitat conditions and regulated through territoriality. We first tested whether population numbers were driven by carrying capacity dynamics and then targeted on establishing (1) the temperature thresholds beyond which population numbers switch from being physical habitat- to temperature-limited; and (2) the rate at which carrying capacity declines with temperature within limiting thermal ranges. Carrying capacity along with emergent density-dependent responses explained up to 76% of spatio-temporal density variability of juveniles and adults but only 50% of young-of-the-year''s. By contrast, young-of-the-year trout were highly sensitive to thermal conditions, their performance declining with temperature at a higher rate than older life stages, and disruptions being triggered at lower temperature thresholds. Results suggest that limiting temperature effects were progressively stronger with increasing anthropogenic disturbance. There was however a critical threshold, matching the incipient thermal limit for survival, beyond which realized density was always below potential numbers irrespective of disturbance intensity. We additionally found a lower threshold, matching the thermal limit for feeding, beyond which even unaltered populations declined. We predict that most of our study populations may become extinct by 2100, depicting the gloomy fate of thermally-sensitive species occurring at thermal range margins under limited potential for adaptation and dispersal.     

A macroecological approach to evolutionary rescue and adaptation to climate change

Despite the widespread use of ecological niche models (ENMs) for predicting the responses of species to climate change, these models do not explicitly incorporate any population‐level mechanism. On the other hand, mechanistic models adding population processes (e.g. biotic interactions, dispersal and adaptive potential to abiotic conditions) are much more complex and difficult to parameterize, especially if the goal is to predict range shifts for many species simultaneously. In particular, the adaptive potential (based on genetic adaptations, phenotypic plasticity and behavioral adjustments for physiological responses) of local populations has been a less studied mechanism affecting species’ responses to climatic change so far. Here, we discuss and apply an alternative macroecological framework to evaluate the potential role of evolutionary rescue under climate change based on ENMs. We begin by reviewing eco‐evolutionary models that evaluate the maximum sustainable evolutionary rate under a scenario of environmental change, showing how they can be used to understand the impact of temperature change on a Neotropical anuran species, the Schneider's toad Rhinella diptycha. Then we show how to evaluate spatial patterns of species’ geographic range shift using such models, by estimating evolutionary rates at the trailing edge of species distribution estimated by ENMs and by recalculating the relative amount of total range loss under climate change. We show how different models can reduce the expected range loss predicted for the studied species by potential ecophysiological adaptations in some regions of the trailing edge predicted by ENMs. For general applications, we believe that parameters for large numbers of species and populations can be obtained from macroecological generalizations (e.g. allometric equations and ecogeographical rules), so our framework coupling ENMs with eco‐evolutionary models can be applied to achieve a more accurate picture of potential impacts from climate change and other threats to biodiversity.     

Range limits and parasite prevalence in a freshwater snail

Geographical range limits are thought to be set by species' physiological or ecological adaptation to abiotic factors, but the importance of biotic factors such as parasitism in determining range limits has not been well explored. In this study the prevalence of trematode parasitism in populations of a freshwater gastropod snail, Lymnaea stagnalis, increased sharply as this species approached its western UK range limit. The likelihood of trematode infection increased with snail size, but high prevalence at the range edge was not a result of interpopulation variation in snail size. Changes in population growth rates resulting from high rates of parasitism at the range edge could contribute to range limitation. The mechanism driving high rates of parasitism at the range edge is not clear, but changes in abiotic factors towards the range limit may influence snail life history and immune response to trematode infection, indirectly altering the prevalence of parasites in marginal host populations.     

Fitness declines towards range limits and local adaptation to climate affect dispersal evolution during climate‐induced range shifts

Dispersal ability will largely determine whether species track their climatic niches during climate change, a process especially important for populations at contracting (low‐latitude/low‐elevation) range limits that otherwise risk extinction. We investigate whether dispersal evolution at contracting range limits is facilitated by two processes that potentially enable edge populations to experience and adjust to the effects of climate deterioration before they cause extinction: (i) climate‐induced fitness declines towards range limits and (ii) local adaptation to a shifting climate gradient. We simulate a species distributed continuously along a temperature gradient using a spatially explicit, individual‐based model. We compare range‐wide dispersal evolution during climate stability vs. directional climate change, with uniform fitness vs. fitness that declines towards range limits (RLs), and for a single climate genotype vs. multiple genotypes locally adapted to temperature. During climate stability, dispersal decreased towards RLs when fitness was uniform, but increased when fitness declined towards RLs, due to highly dispersive genotypes maintaining sink populations at RLs, increased kin selection in smaller populations, and an emergent fitness asymmetry that favoured dispersal in low‐quality habitat. However, this initial dispersal advantage at low‐fitness RLs did not facilitate climate tracking, as it was outweighed by an increased probability of extinction. Locally adapted genotypes benefited from staying close to their climate optima; this selected against dispersal under stable climates but for increased dispersal throughout shifting ranges, compared to cases without local adaptation. Dispersal increased at expanding RLs in most scenarios, but only increased at the range centre and contracting RLs given local adaptation to climate.