A plant's perspective of extremes: terrestrial plant responses to changing climatic variability

We review observational, experimental, and model results on how plants respond to extreme climatic conditions induced by changing climatic variability. Distinguishing between impacts of changing mean climatic conditions and changing climatic variability on terrestrial ecosystems is generally underrated in current studies. The goals of our review are thus (1) to identify plant processes that are vulnerable to changes in the variability of climatic variables rather than to changes in their mean, and (2) to depict/evaluate available study designs to quantify responses of plants to changing climatic variability. We find that phenology is largely affected by changing mean climate but also that impacts of climatic variability are much less studied, although potentially damaging. We note that plant water relations seem to be very vulnerable to extremes driven by changes in temperature and precipitation and that heatwaves and flooding have stronger impacts on physiological processes than changing mean climate. Moreover, interacting phenological and physiological processes are likely to further complicate plant responses to changing climatic variability. Phenological and physiological processes and their interactions culminate in even more sophisticated responses to changing mean climate and climatic variability at the species and community level. Generally, observational studies are well suited to study plant responses to changing mean climate, but less suitable to gain a mechanistic understanding of plant responses to climatic variability. Experiments seem best suited to simulate extreme events. In models, temporal resolution and model structure are crucial to capture plant responses to changing climatic variability. We highlight that a combination of experimental, observational, and/or modeling studies have the potential to overcome important caveats of the respective individual approaches.     

Climate variability and vulnerability to climate change: a review

The focus of the great majority of climate change impact studies is on changes in mean climate. In terms of climate model output, these changes are more robust than changes in climate variability. By concentrating on changes in climate means, the full impacts of climate change on biological and human systems are probably being seriously underestimated. Here, we briefly review the possible impacts of changes in climate variability and the frequency of extreme events on biological and food systems, with a focus on the developing world. We present new analysis that tentatively links increases in climate variability with increasing food insecurity in the future. We consider the ways in which people deal with climate variability and extremes and how they may adapt in the future. Key knowledge and data gaps are highlighted. These include the timing and interactions of different climatic stresses on plant growth and development, particularly at higher temperatures, and the impacts on crops, livestock and farming systems of changes in climate variability and extreme events on pest‐weed‐disease complexes. We highlight the need to reframe research questions in such a way that they can provide decision makers throughout the food system with actionable answers, and the need for investment in climate and environmental monitoring. Improved understanding of the full range of impacts of climate change on biological and food systems is a critical step in being able to address effectively the effects of climate variability and extreme events on human vulnerability and food security, particularly in agriculturally based developing countries facing the challenge of having to feed rapidly growing populations in the coming decades.     

Designing ecological climate change impact assessments to reflect key climatic drivers

Identifying the climatic drivers of an ecological system is a key step in assessing its vulnerability to climate change. The climatic dimensions to which a species or system is most sensitive – such as means or extremes – can guide methodological decisions for projections of ecological impacts and vulnerabilities. However, scientific workflows for combining climate projections with ecological models have received little explicit attention. We review Global Climate Model (GCM) performance along different dimensions of change and compare frameworks for integrating GCM output into ecological models. In systems sensitive to climatological means, it is straightforward to base ecological impact assessments on mean projected changes from several GCMs. Ecological systems sensitive to climatic extremes may benefit from what we term the ‘model space’ approach: a comparison of ecological projections based on simulated climate from historical and future time periods. This approach leverages the experimental framework used in climate modeling, in which historical climate simulations serve as controls for future projections. Moreover, it can capture projected changes in the intensity and frequency of climatic extremes, rather than assuming that future means will determine future extremes. Given the recent emphasis on the ecological impacts of climatic extremes, the strategies we describe will be applicable across species and systems. We also highlight practical considerations for the selection of climate models and data products, emphasizing that the spatial resolution of the climate change signal is generally coarser than the grid cell size of downscaled climate model output. Our review illustrates how an understanding of how climate model outputs are derived and downscaled can improve the selection and application of climatic data used in ecological modeling.     

Means and extremes: building variability into community‐level climate change experiments

Experimental studies assessing climatic effects on ecological communities have typically applied static warming treatments. Although these studies have been informative, they have usually failed to incorporate either current or predicted future, patterns of variability. Future climates are likely to include extreme events which have greater impacts on ecological systems than changes in means alone. Here, we review the studies which have used experiments to assess impacts of temperature on marine, freshwater and terrestrial communities, and classify them into a set of ‘generations’ based on how they incorporate variability. The majority of studies have failed to incorporate extreme events. In terrestrial ecosystems in particular, experimental treatments have reduced temperature variability, when most climate models predict increased variability. Marine studies have tended to not concentrate on changes in variability, likely in part because the thermal mass of oceans will moderate variation. In freshwaters, climate change experiments have a much shorter history than in the other ecosystems, and have tended to take a relatively simple approach. We propose a new ‘generation’ of climate change experiments using down‐scaled climate models which incorporate predicted changes in climatic variability, and describe a process for generating data which can be applied as experimental climate change treatments.     

Projected timing of perceivable changes in climate extremes for terrestrial and marine ecosystems

Human and natural systems have adapted to and evolved within historical climatic conditions. Anthropogenic climate change has the potential to alter these conditions such that onset of unprecedented climatic extremes will outpace evolutionary and adaptive capabilities. To assess whether and when future climate extremes exceed their historical windows of variability within impact‐relevant socioeconomic, geopolitical, and ecological domains, we investigate the timing of perceivable changes (time of emergence; TOE) for 18 magnitude‐, frequency‐, and severity‐based extreme temperature (10) and precipitation (8) indices using both multimodel and single‐model multirealization ensembles. Under a high‐emission scenario, we find that the signal of frequency‐ and severity‐based temperature extremes is projected to rise above historical noise earliest in midlatitudes, whereas magnitude‐based temperature extremes emerge first in low and high latitudes. Precipitation extremes demonstrate different emergence patterns, with severity‐based indices first emerging over midlatitudes, and magnitude‐ and frequency‐based indices emerging earliest in low and high latitudes. Applied to impact‐relevant domains, simulated TOE patterns suggest (a) unprecedented consecutive dry day occurrence in >50% of 14 terrestrial biomes and 12 marine realms prior to 2100, (b) earlier perceivable changes in climate extremes in countries with lower per capita GDP, and (c) emergence of severe and frequent heat extremes well‐before 2030 for the 590 most populous urban centers. Elucidating extreme‐metric and domain‐type TOE heterogeneities highlights the challenges adaptation planners face in confronting the consequences of elevated twenty‐first century radiative forcing.     

Effects of climate extremes on the terrestrial carbon cycle: concepts,processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance‐induced mechanisms and processes to also operate in an extreme context. The paucity of well‐defined studies currently renders a quantitative meta‐analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land‐cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground‐based observational case studies reveals that many regions in the (sub‐)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.     

Riparian Ecosystems in the 21st Century: Hotspots for Climate Change Adaptation?

Riparian ecosystems in the 21st century are likely to play a critical role in determining the vulnerability of natural and human systems to climate change, and in influencing the capacity of these systems to adapt. Some authors have suggested that riparian ecosystems are particularly vulnerable to climate change impacts due to their high levels of exposure and sensitivity to climatic stimuli, and their history of degradation. Others have highlighted the probable resilience of riparian ecosystems to climate change as a result of their evolution under high levels of climatic and environmental variability. We synthesize current knowledge of the vulnerability of riparian ecosystems to climate change by assessing the potential exposure, sensitivity, and adaptive capacity of their key components and processes, as well as ecosystem functions, goods and services, to projected global climatic changes. We review key pathways for ecological and human adaptation for the maintenance, restoration and enhancement of riparian ecosystem functions, goods and services and present emerging principles for planned adaptation. Our synthesis suggests that, in the absence of adaptation, riparian ecosystems are likely to be highly vulnerable to climate change impacts. However, given the critical role of riparian ecosystem functions in landscapes, as well as the strong links between riparian ecosystems and human well-being, considerable means, motives and opportunities for strategically planned adaptation to climate change also exist. The need for planned adaptation of and for riparian ecosystems is likely to be strengthened as the importance of many riparian ecosystem functions, goods and services will grow under a changing climate. Consequently, riparian ecosystems are likely to become adaptation ‘hotspots’ as the century unfolds.     

Beyond realism in climate change experiments: gradient approaches identify thresholds and tipping points

Experimental evidence for impacts of increased climatic variability and extremes on ecosystems is urgently needed. The constraint in our knowledge, however, is not caused by the uncertainty in the applied climate scenarios. We need mechanistic understanding from experiments challenging ecological thresholds coupled with ecosystem models to allow for meaningful up‐scaling.     

Cascading effects of climate extremes on vertebrate fauna through changes to low‐latitude tree flowering and fruiting phenology

Forest vertebrate fauna provide critical services, such as pollination and seed dispersal, which underpin functional and resilient ecosystems. In turn, many of these fauna are dependent on the flowering phenology of the plant species of such ecosystems. The impact of changes in climate, including climate extremes, on the interaction between these fauna and flora has not been identified or elucidated, yet influences on flowering phenology are already evident. These changes are well documented in the mid to high latitudes. However, there is emerging evidence that the flowering phenology, nectar/pollen production, and fruit production of long‐lived trees in tropical and subtropical forests are also being impacted by changes in the frequency and severity of climate extremes. Here, we examine the implications of these changes for vertebrate fauna dependent on these resources. We review the literature to establish evidence for links between climate extremes and flowering phenology, elucidating the nature of relationships between different vertebrate taxa and flowering regimes. We combine this information with climate change projections to postulate about the likely impacts on nectar, pollen and fruit resource availability and the consequences for dependent vertebrate fauna. The most recent climate projections show that the frequency and intensity of climate extremes will increase during the 21st century. These changes are likely to significantly alter mass flowering and fruiting events in the tropics and subtropics, which are frequently cued by climate extremes, such as intensive rainfall events or rapid temperature shifts. We find that in these systems the abundance and duration of resource availability for vertebrate fauna is likely to fluctuate, and the time intervals between episodes of high resource availability to increase. The combined impact of these changes has the potential to result in cascading effects on ecosystems through changes in pollinator and seed dispersal ecology, and demands a focused research effort.     

Impacts of changes in climate variability on regional vegetation in China: NDVI-based analysis from 1982 to 2000

Three methods were used to distinguish the characteristics of changes in climate variability and normalized difference vegetation index (NDVI) during the period from 1982 to 2000 in China. Great changes in climate variability and an increased trend in NDVI were observed. The changes in precipitation variability were greater than the changes in temperature variability in each month, which is attributed to changes in the monsoon system in East Asia. The abrupt changes in climate and NDVI were more significant in 1983 than in the other years due to the impacts of El Niño/Southern Oscillation (ENSO). Using these results, the influences of changes in climate variability on vegetation were studied in the whole nation, and eight regions were defined according to the vegetation division map of China. The results show that abrupt climate changes at a small scale cannot cause abrupt NDVI changes directly. At a nationwide level, over a longer time scale the persistence of above/below average temperature determines the changes in NDVI; at a shorter time scale, changes in the magnitude of precipitation influence NDVI significantly. Such regional climate variability affects vegetation in different ways owing to the diversity of vegetation types, climatic conditions and topography of the land.     

Vulnerability of sea turtle nesting grounds to climate change

Given the potential vulnerability of sea turtles to climate change, a growing number of studies are predicting how various climatic processes will affect their nesting grounds. However, these studies are limited by scale, because they predict how a single climatic process will affect sea turtles but processes are likely to occur simultaneously and cause cumulative effects. This study addresses the need for a structured approach to investigate how multiple climatic processes may affect a turtle population. Here, we use a vulnerability assessment framework to assess the cumulative impact of various climatic processes on the nesting grounds used by the northern Great Barrier Reef (nGBR) green turtle population. Further, we manipulate the variables from this framework to allow users to investigate how mitigating different climatic processes individually or simultaneously can influence the vulnerability of the nesting grounds. Our assessment indicates that nesting grounds closer to the equator, such as Bramble Cay and Milman Island, are the most vulnerable to climate change. In the short‐term (by 2030), sea level rise will cause the most impact on the nesting grounds used by the nGBR green turtle population. However, in the longer term, by 2070 sand temperatures will reach levels above the upper transient range and the upper thermal threshold and cause relatively more impact on the nGBR green turtle population. Thus, in the long term, a reduction of impacts from sea‐level rise may not be sufficient, as rookeries will start to experience high vulnerability values from increased temperature. Thus, in the long term, reducing the threats from increased temperature may provide a greater return in conservation investment than mitigating the impacts from other climatic processes. Indeed, our results indicate that if the impacts from increased temperature are mitigated, the vulnerability values of almost all rookeries will be reduced to low levels.     

Climatic seasonality, resource bottlenecks, and abundance of rainforest birds: implications for global climate change

We demonstrate that within-year climatic variability, particularly rainfall seasonality, is the most significant variable explaining spatial patterns of bird abundance in Australian tropical rainforest. The likely mechanism causing this pattern is a resource bottleneck (insects, nectar, and fruit) during the dry season that limits the population size of many species. The patterns support both the diversity–climatic–stability hypothesis and the species–energy hypothesis but clearly show that seasonality in energy availability may be a more significant factor than annual totals or means. An index of dry season severity is proposed that quantifies the combined effect of the degree of dryness and the duration of the dry season. We suggest that the predicted increases in seasonality due to global climate change could produce significant declines in bird abundance, further exacerbating the impacts of decreased range size, increased fragmentation, and decreased population size likely to occur as a result of increasing temperature. We suggest that increasing climatic seasonality due to global climate change has the potential to have significant negative impacts on tropical biodiversity.     

Integrating mechanistic and empirical model projections to assess climate impacts on tree species distributions in northwestern North America

Empirical and mechanistic models have both been used to assess the potential impacts of climate change on species distributions, and each modeling approach has its strengths and weaknesses. Here, we demonstrate an approach to projecting climate‐driven changes in species distributions that draws on both empirical and mechanistic models. We combined projections from a dynamic global vegetation model (DGVM) that simulates the distributions of biomes based on basic plant functional types with projections from empirical climatic niche models for six tree species in northwestern North America. These integrated model outputs incorporate important biological processes, such as competition, physiological responses of plants to changes in atmospheric CO₂ concentrations, and fire, as well as what are likely to be species‐specific climatic constraints. We compared the integrated projections to projections from the empirical climatic niche models alone. Overall, our integrated model outputs projected a greater climate‐driven loss of potentially suitable environmental space than did the empirical climatic niche model outputs alone for the majority of modeled species. Our results also show that refining species distributions with DGVM outputs had large effects on the geographic locations of suitable habitat. We demonstrate one approach to integrating the outputs of mechanistic and empirical niche models to produce bioclimatic projections. But perhaps more importantly, our study reveals the potential for empirical climatic niche models to over‐predict suitable environmental space under future climatic conditions.     

Climate drives temporal replacement and nested‐resultant richness patterns of Scottish coastal vegetation

Beta diversity quantifies spatial and/or temporal variation in species composition. It is comprised of two distinct components, species replacement and nestedness, which derive from opposing ecological processes. Using Scotland as a case study and a β‐diversity partitioning framework, we investigate temporal replacement and nestedness patterns of coastal grassland species over a 34‐yr time period. We aim to 1) understand the influence of two potentially pivotal processes (climate and land‐use changes) on landscape‐scale (5 × 5 km) temporal replacement and nestedness patterns, and 2) investigate whether patterns from one β‐diversity component can mask observable patterns in the other. We summarised key aspects of climate driven macro‐ecological variation as measures of variance, long‐term trends, between‐year similarity and extremes, for three important climatic predictors (minimum temperature, water‐balance and growing degree‐days). Shifts in landscape‐scale heterogeneity, a proxy of land‐use change, was summarised as a spatial multiple‐site dissimilarity measure. Together, these climatic and spatial predictors were used in a multi‐model inference framework to gauge the relative contribution of each on temporal replacement and nestedness patterns. Temporal β‐diversity patterns were reasonably well explained by climate change but weakly explained by changes in landscape‐scale heterogeneity. Climate was shown to have a greater influence on temporal nestedness than replacement patterns over our study period, linking nestedness patterns, as a result of imbalanced gains and losses, to climatic warming and extremes respectively. Important climatic predictors (i.e. growing degree‐days) of temporal β‐diversity were also identified, and contrasting patterns between the two β‐diversity components revealed. Results suggest climate influences plant species recruitment and establishment processes of Scotland's coastal grasslands, and while species extinctions take time, they are likely to be facilitated by climatic perturbations. Our findings also highlight the importance of distinguishing between different components of β‐diversity, disentangling contrasting patterns than can mask one another.     

Plasticity in Dendroclimatic Response across the Distribution Range of Aleppo Pine (Pinus halepensis)

We investigated the variability of the climate-growth relationship of Aleppo pine across its distribution range in the Mediterranean Basin. We constructed a network of tree-ring index chronologies from 63 sites across the region. Correlation function analysis identified the relationships of tree-ring index to climate factors for each site. We also estimated the dominant climatic gradients of the region using principal component analysis of monthly, seasonal, and annual mean temperature and total precipitation from 1,068 climatic gridpoints. Variation in ring width index was primarily related to precipitation and secondarily to temperature. However, we found that the dendroclimatic relationship depended on the position of the site along the climatic gradient. In the southern part of the distribution range, where temperature was generally higher and precipitation lower than the regional average, reduced growth was also associated with warm and dry conditions. In the northern part, where the average temperature was lower and the precipitation more abundant than the regional average, reduced growth was associated with cool conditions. Thus, our study highlights the substantial plasticity of Aleppo pine in response to different climatic conditions. These results do not resolve the source of response variability as being due to either genetic variation in provenance, to phenotypic plasticity, or a combination of factors. However, as current growth responses to inter-annual climate variability vary spatially across existing climate gradients, future climate-growth relationships will also likely be determined by differential adaptation and/or acclimation responses to spatial climatic variation. The contribution of local adaptation and/or phenotypic plasticity across populations to the persistence of species under global warming could be decisive for prediction of climate change impacts across populations. In this sense, a more complex forest dynamics modeling approach that includes the contribution of genetic variation and phenotypic plasticity can improve the reliability of the ecological inferences derived from the climate-growth relationships.     

The impacts of climate change in coastal marine systems

Anthropogenically induced global climate change has profound implications for marine ecosystems and the economic and social systems that depend upon them. The relationship between temperature and individual performance is reasonably well understood, and much climate-related research has focused on potential shifts in distribution and abundance driven directly by temperature. However, recent work has revealed that both abiotic changes and biological responses in the ocean will be substantially more complex. For example, changes in ocean chemistry may be more important than changes in temperature for the performance and survival of many organisms. Ocean circulation, which drives larval transport, will also change, with important consequences for population dynamics. Furthermore, climatic impacts on one or a few 'leverage species' may result in sweeping community-level changes. Finally, synergistic effects between climate and other anthropogenic variables, particularly fishing pressure, will likely exacerbate climate-induced changes. Efforts to manage and conserve living marine systems in the face of climate change will require improvements to the existing predictive framework. Key directions for future research include identifying key demographic transitions that influence population dynamics, predicting changes in the community-level impacts of ecologically dominant species, incorporating populations' ability to evolve (adapt), and understanding the scales over which climate will change and living systems will respond.     

Climate change accelerates local disease extinction rates in a long‐term wild host–pathogen association

Pathogens are a significant component of all plant communities. In recent years, the potential for existing and emerging pathogens of agricultural crops to cause increased yield losses as a consequence of changing climatic patterns has raised considerable concern. In contrast, the response of naturally occurring, endemic pathogens to a warming climate has received little attention. Here, we report on the impact of a signature variable of global climate change – increasing temperature – on the long‐term epidemiology of a natural host–pathogen association involving the rust pathogen Triphragmium ulmariae and its host plant Filipendula ulmaria. In a host–pathogen metapopulation involving approximately 230 host populations growing on an archipelago of islands in the Gulf of Bothnia we assessed changes in host population size and pathogen epidemiological measures over a 25‐year period. We show how the incidence of disease and its severity declines over that period and most importantly demonstrate a positive association between a long‐term trend of increasing extinction rates in individual pathogen populations of the metapopulation and increasing temperature. Our results are highly suggestive that changing climatic patterns, particularly mean monthly growing season (April‐November) temperature, are markedly influencing the epidemiology of plant disease in this host–pathogen association. Given the important role plant pathogens have in shaping the structure of communities, changes in the epidemiology of pathogens have potentially far‐reaching impacts on ecological and evolutionary processes. For these reasons, it is essential to increase understanding of pathogen epidemiology, its response to warming, and to invoke these responses in forecasts for the future.     

Converging Climate Sensitivities of European Forests Between Observed Radial Tree Growth and Vegetation Models

The impacts of climate variability and trends on European forests are unevenly distributed across different bioclimatic zones and species. Extreme climate events are also becoming more frequent and it is unknown how they will affect feedbacks of CO₂ between forest ecosystems and the atmosphere. An improved understanding of species differences at the regional scale of the response of forest productivity to climate variation and extremes is thus important for forecasting forest dynamics. In this study, we evaluate the climate sensitivity of aboveground net primary production (NPP) simulated by two dynamic global vegetation models (DGVM; ORCHIDEE and LPJ-wsl) against tree ring width (TRW) observations from about 1000 sites distributed across Europe. In both the model simulations and the TRW observations, forests in northern Europe and the Alps respond positively to warmer spring and summer temperature, and their overall temperature sensitivity is larger than that of the soil-moisture-limited forests in central Europe and Mediterranean regions. Compared with TRW observations, simulated NPP from ORCHIDEE and LPJ-wsl appear to be overly-sensitive to climatic factors. Our results indicate that the models lack biological processes that control time lags, such as carbohydrate storage and remobilization, that delay the effects of radial growth dynamics to climate. Our study highlights the need for re-evaluating the physiological controls on the climate sensitivity of NPP simulated by DGVMs. In particular, DGVMs could be further enhanced by a more detailed representation of carbon reserves and allocation that control year-to-year variation in plant growth.     

Climatic thresholds shape northern high‐latitude fire regimes and imply vulnerability to future climate change

Boreal forests and arctic tundra cover 33% of global land area and store an estimated 50% of total soil carbon. Because wildfire is a key driver of terrestrial carbon cycling, increasing fire activity in these ecosystems would likely have global implications. To anticipate potential spatiotemporal variability in fire‐regime shifts, we modeled the spatially explicit 30‐yr probability of fire occurrence as a function of climate and landscape features (i.e. vegetation and topography) across Alaska. Boosted regression tree (BRT) models captured the spatial distribution of fire across boreal forest and tundra ecoregions (AUC from 0.63–0.78 and Pearson correlations between predicted and observed data from 0.54–0.71), highlighting summer temperature and annual moisture availability as the most influential controls of historical fire regimes. Modeled fire–climate relationships revealed distinct thresholds to fire occurrence, with a nonlinear increase in the probability of fire above an average July temperature of 13.4°C and below an annual moisture availability (i.e. P‐PET) of approximately 150 mm. To anticipate potential fire‐regime responses to 21st‐century climate change, we informed our BRTs with Coupled Model Intercomparison Project Phase 5 climate projections under the RCP 6.0 scenario. Based on these projected climatic changes alone (i.e. not accounting for potential changes in vegetation), our results suggest an increasing probability of wildfire in Alaskan boreal forest and tundra ecosystems, but of varying magnitude across space and throughout the 21st century. Regions with historically low flammability, including tundra and the forest–tundra boundary, are particularly vulnerable to climatically induced changes in fire activity, with up to a fourfold increase in the 30‐yr probability of fire occurrence by 2100. Our results underscore the climatic potential for novel fire regimes to develop in these ecosystems, relative to the past 6000–35 000 yr, and spatial variability in the vulnerability of wildfire regimes and associated ecological processes to 21st‐century climate change.     

Evolutionary responses to global change: lessons from invasive species

Biologists have recently devoted increasing attention to the role of rapid evolution in species' responses to environmental change. However, it is still unclear what evolutionary responses should be expected, at what rates, and whether evolution will save populations at risk of extinction. The potential of biological invasions to provide useful insights has barely been realised, despite the close analogies to species responding to global change, particularly climate change; in both cases, populations encounter novel climatic and biotic selection pressures, with expected evolutionary responses occurring over similar timescales. However, the analogy is not perfect, and invasive species are perhaps best used as an upper bound on expected change. In this article, we review what invasive species can and cannot teach us about likely evolutionary responses to global change and the constraints on those responses. We also discuss the limitations of invasive species as a model and outline directions for future research.