What stops populations expanding into new territory beyond the edge of a range margin? Recent models addressing this problem have brought together population genetics and population ecology, and some ...have included interactions among species at range edges. Here, we review these models of adaptation at environmental or parapatric margins, and discuss the contrasting effects of migration in either swamping local adaptation, or supplying the genetic variation that is necessary for adaptation to continue. We illustrate how studying adaptation at range margins (both with and without hybridization) can provide insight into the genetic and ecological factors that limit evolution more generally, especially in response to current rates of environmental change.
Climate change is altering life at multiple scales, from genes to ecosystems. Predicting the vulnerability of populations to climate change is crucial to mitigate negative impacts. We suggest that ...regional patterns of spatial and temporal climatic variation scaled to the traits of an organism can predict where and why populations are most vulnerable to climate change. Specifically, historical climatic variation affects the sensitivity and response capacity of populations to climate change by shaping traits and the genetic variation in those traits. Present and future climatic variation can affect both climate change exposure and population responses. We provide seven predictions for how climatic variation might affect the vulnerability of populations to climate change and suggest key directions for future research.
Predicting the vulnerability of populations to climate change is crucial.
Spatial and temporal climatic variation strongly influence vulnerability.
Historical climatic variation can shape traits that affect vulnerability.
Present and future climatic variation affect exposure and population responses.
Maps of climatic variability can predict where populations are most vulnerable.
Ecology Letters (2012) 15: 378–392
Forest trees are the dominant species in many parts of the world and predicting how they might respond to climate change is a vital global concern. Trees are ...capable of long‐distance gene flow, which can promote adaptive evolution in novel environments by increasing genetic variation for fitness. It is unclear, however, if this can compensate for maladaptive effects of gene flow and for the long‐generation times of trees. We critically review data on the extent of long‐distance gene flow and summarise theory that allows us to predict evolutionary responses of trees to climate change. Estimates of long‐distance gene flow based both on direct observations and on genetic methods provide evidence that genes can move over spatial scales larger than habitat shifts predicted under climate change within one generation. Both theoretical and empirical data suggest that the positive effects of gene flow on adaptation may dominate in many instances. The balance of positive to negative consequences of gene flow may, however, differ for leading edge, core and rear sections of forest distributions. We propose future experimental and theoretical research that would better integrate dispersal biology with evolutionary quantitative genetics and improve predictions of tree responses to climate change.
The changes in species' geographical distribution demanded by climate change are often critically limited by the availability of key interacting species. In such cases, species' persistence will ...depend on the rapid evolution of biotic interactions. Understanding evolutionary limits to such adaptation is therefore crucial for predicting biological responses to environmental change. The recent poleward range expansion of the UK brown argus butterfly has been associated with a shift in female preference from its main host plant, rockrose (Cistaceae), onto Geraniaceae host plants throughout its new distribution. Using reciprocal transplants onto natural host plants across the UK range, we demonstrate reduced fitness of females from recently colonised Geraniaceae‐dominated habitat when moved to ancestral rockrose habitats. By contrast, individuals from ancestral rockrose habitats show no reduction in fitness on Geraniaceae. Climate‐driven range expansion in this species is therefore associated with the rapid evolution of biotic interactions and a significant loss of adaptive variation.
Advances in phenology (the annual timing of species' life-cycles) in response to climate change are generally viewed as bioindicators of climate change, but have not been considered as predictors of ...range expansions. Here, we show that phenology advances combine with the number of reproductive cycles per year (voltinism) to shape abundance and distribution trends in 130 species of British Lepidoptera, in response to ~0.5 °C spring-temperature warming between 1995 and 2014. Early adult emergence in warm years resulted in increased within- and between-year population growth for species with multiple reproductive cycles per year (n = 39 multivoltine species). By contrast, early emergence had neutral or negative consequences for species with a single annual reproductive cycle (n = 91 univoltine species), depending on habitat specialisation. We conclude that phenology advances facilitate polewards range expansions in species exhibiting plasticity for both phenology and voltinism, but may inhibit expansion by less flexible species.
Two decades of intensive research have provided compelling evidence for a link between biodiversity and ecosystem functioning (B-EF). Whereas early B-EF research concentrated on species richness and ...single processes, recent studies have investigated different measures of both biodiversity and ecosystem functioning, such as functional diversity and joint metrics of multiple processes. There is also a shift from viewing assemblages in terms of their contribution to particular processes toward placing them within a wider food web context. We review how the responses and predictors in B-EF experiments are quantified and how biodiversity effects are shaped by multitrophic interactions. Further, we discuss how B-EF metrics and food web relations could be addressed simultaneously. We conclude that addressing traits, multiple processes and food web interactions is needed to capture the mechanisms that underlie B-EF relations in natural assemblages.
Accurately predicting biological impacts of climate change is necessary to guide policy. However, the resolution of climate data could be affecting the accuracy of climate change impact assessments. ...Here, we review the spatial and temporal resolution of climate data used in impact assessments and demonstrate that these resolutions are often too coarse relative to biologically relevant scales. We then develop a framework that partitions climate into three important components: trend, variance, and autocorrelation. We apply this framework to map different global climate regimes and identify where coarse climate data is most and least likely to reduce the accuracy of impact assessments. We show that impact assessments for many large mammals and birds use climate data with a spatial resolution similar to the biologically relevant area encompassing population dynamics. Conversely, impact assessments for many small mammals, herpetofauna, and plants use climate data with a spatial resolution that is orders of magnitude larger than the area encompassing population dynamics. Most impact assessments also use climate data with a coarse temporal resolution. We suggest that climate data with a coarse spatial resolution is likely to reduce the accuracy of impact assessments the most in climates with high spatial trend and variance (e.g., much of western North and South America) and the least in climates with low spatial trend and variance (e.g., the Great Plains of the USA). Climate data with a coarse temporal resolution is likely to reduce the accuracy of impact assessments the most in the northern half of the northern hemisphere where temporal climatic variance is high. Our framework provides one way to identify where improving the resolution of climate data will have the largest impact on the accuracy of biological predictions under climate change.
• As climate change transforms seasonal patterns of temperature and precipitation, germination success at marginal temperatures will become critical for the long-term persistence of many plant ...species and communities. If populations vary in their environmental sensitivity to marginal temperatures across a species’ geographical range, populations that respond better to future environmental extremes are likely to be critical for maintaining ecological resilience of the species.
• Using seeds from two to six populations for each of nine species of Mediterranean plants, we characterized patterns of among-population variation in environmental sensitivity by quantifying genotype-by-environment interactions (G × E) for germination success at temperature extremes, and under two light regimes representing conditions below and above the soil surface.
• For eight of nine species tested at hot and cold marginal temperatures, we observed substantial among-population variation in environmental sensitivity for germination success, and this often depended on the light treatment. Importantly, different populations often performed best at different environmental extremes.
• Our results demonstrate that ongoing changes in temperature regime will affect the phenology, fitness, and demography of different populations within the same species differently. We show that quantifying patterns of G × E for multiple populations, and understanding how such patterns arise, can test mechanisms that promote ecological resilience.
Ecology Letters (2010) 13: 485-494 All species are restricted in their distribution. Currently, ecological models can only explain such limits if patches vary in quality, leading to asymmetrical ...dispersal, or if genetic variation is too low at the margins for adaptation. However, population genetic models suggest that the increase in genetic variance resulting from dispersal should allow adaptation to almost any ecological gradient. Clearly therefore, these models miss something that prevents evolution in natural populations. We developed an individual-based simulation to explore stochastic effects in these models. At high carrying capacities, our simulations largely agree with deterministic predictions. However, when carrying capacity is low, the population fails to establish for a wide range of parameter values where adaptation was expected from previous models. Stochastic or transient effects appear critical around the boundaries in parameter space between simulation behaviours. Dispersal, gradient steepness, and population density emerge as key factors determining adaptation on an ecological gradient.
Population genetic models of evolution along linear environmental gradients cannot explain why adaptation stops at ecological margins. This is because, unless models impose reductions in carrying ...capacity at species’ edges, the dominant effect of gene flow is to increase genetic variance and adaptive potential rather than swamping local adaptation. This allows the population to match even very steep changes in trait optima. We extend our previous simulations to explore two nonlinear models of ecological gradients: (a) a sigmoid (steepening) gradient and (b) a linear gradient with a flat centre of variable width. We compare the parameter conditions that allow local adaptation and range expansion from the centre, with those that permit the persistence of a perfectly adapted population distributed across the entire range. Along nonlinear gradients, colonization is easier, and extinction rarer, than along a linear gradient. This is because the shallow environmental gradient near the range centre does not cause gene flow to increase genetic variation, and so does not result in reduced population density. However, as gradient steepness increases, gene flow inflates genetic variance and reduces local population density sufficiently that genetic drift overcomes local selection, creating a finite range margin. When a flat centre is superimposed on a linear gradient, gene flow increases genetic variation dramatically at its edges, leading to an abrupt reduction in density that prevents niche expansion. Remarkably local interruptions in a linear ecological gradient (of a width much less than the mean dispersal distance) can prevent local adaptation beyond this flat centre. In contrast to other situations, this effect is stronger and more consistent where carrying capacity is high. Practically speaking, this means that habitat improvement at patch margins will make evolutionary rescue more likely. By contrast, even small improvements in habitat at patch centres may confine populations to limited areas of ecological space.
PDF
