Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Climate change and amphibian declines: is there a link?

Abstract. Global climates have been changing, sometimes rapidly and dramatically, throughout the evolutionary history of amphibians. Therefore, existing amphibian species have been derived from those that have survived major climatic disturbances. Although recent global climate change has resulted in warming in many regions, temperatures in some areas to date have not changed measurably, or have even cooled. Declines of some amphibian populations have been correlated with climate events, but demonstrations of direct causal relationships need further research. Data are available indicating some indirect effect of climate change on the initiation of breeding activities of some amphibians that occur earlier than in previous springs, but the costs and benefits of these changes are just beginning to be investigated. Climate may also play an indirect role in facilitating epidemics of infectious disease. Regardless of the role that climate changes may have played in past and current amphibian declines, future shifts in climate, should they prove as dramatic as predicted, will certainly pose challenges for surviving amphibian populations and for successful recovery efforts of species that have suffered declines.     

Contrasting mechanisms underlie short‐ and longer‐term soil respiration responses to experimental warming in a dryland ecosystem

Soil carbon losses to the atmosphere through soil respiration are expected to rise with ongoing temperature increases, but available evidence from mesic biomes suggests that such response disappears after a few years of experimental warming. However, there is lack of empirical basis for these temporal dynamics in soil respiration responses, and for the mechanisms underlying them, in drylands, which collectively form the largest biome on Earth and store 32% of the global soil organic carbon pool. We coupled data from a 10 year warming experiment in a biocrust‐dominated dryland ecosystem with laboratory incubations to confront 0–2 years (short‐term hereafter) versus 8–10 years (longer‐term hereafter) soil respiration responses to warming. Our results showed that increased soil respiration rates with short‐term warming observed in areas with high biocrust cover returned to control levels in the longer‐term. Warming‐induced increases in soil temperature were the main drivers of the short‐term soil respiration responses, whereas longer‐term soil respiration responses to warming were primarily driven by thermal acclimation and warming‐induced reductions in biocrust cover. Our results highlight the importance of evaluating short‐ and longer‐term soil respiration responses to warming as a mean to reduce the uncertainty in predicting the soil carbon–climate feedback in drylands.     

Experimental warming drives a seasonal shift in the timing of host‐parasite dynamics with consequences for disease risk

Multi‐species experiments are critical for identifying the mechanisms through which climate change influences population dynamics and community interactions within ecological systems, including infectious diseases. Using a host–parasite system involving freshwater snails, amphibians and trematode parasites, we conducted a year‐long, outdoor experiment to evaluate how warming affected net parasite production, the timing of infection and the resultant pathology. Warming of 3 °C caused snail intermediate hosts to release parasites 9 months earlier and increased infected snail mortality by fourfold, leading to decreased overlap between amphibians and parasites. As a result, warming halved amphibian infection loads and reduced pathology by 67%, despite comparable total parasite production across temperature treatments. These results demonstrate that climate–disease theory should be expanded to account for predicted changes in host and parasite phenology, which may often be more important than changes in total parasite output for predicting climate‐driven changes in disease risk.     

Temporal dynamics of marginal steppic vegetation over a 26‐year period of substantial environmental change

Questions: (1) Is climate a strong driver of vegetation dynamics, including interannual variation, in a range margin steppic community? (2) Are there long‐term trends in cover and species richness in this community, and are these consistent across species groups and species within groups? (3) Can long‐term trends in plant community data be related to variation in local climate over the last three decades? Location: A range margin steppic grassland community in central Germany. Methods: Cover, number and size of all individuals of all plant species present in three permanent 1‐m² plots were recorded in spring for 26 years (1980–2005). Climatic data for the study area were used to determine the best climatic predictor for each plant community, functional group and species variable (annual data and interannual variation) using best subsets regression. Results: April and autumn temperature showed the highest correlation with total cover and species richness and with interannual variations of cover and richness. However, key climate drivers differed between the five most abundant species. Similarly, total cover and number and cover of perennials significantly decreased over time, while no trend was found for the cover and number of annuals. However, within functional groups there were also contrasting species‐specific responses. Long‐term temperature increases and high interannual variability in both temperature and precipitation were strongly related to long‐term trends and interannual variations in plant community data. Conclusions: Temporal trends in vegetation were strongly associated with temporal trends in climate at the study site, with key roles for autumn and spring temperature and precipitation. Dynamics of functional groups and species within groups and their relationships to changes in temperature and precipitation reveal complex long‐term and interannual patterns that cannot be inferred from short‐term studies with only one or a few individual species. Our results also highlight that responses detected at the functional group level may mask contrasting responses within functional groups. We discuss the implications of these findings for attempts to predict the future response of biodiversity to climate change.     

Species and community responses to short‐term climate manipulation: Microarthropods in the sub‐Antarctic

Abstract: Both species and community‐level investigations are important for understanding the biotic impacts of climate change, because current evidence suggests that individual species responses are idiosyncratic. However, few studies of climate change impacts have been conducted on entire terrestrial arthropod communities living in the same habitat in the southern Hemisphere, and the effects of precipitation changes on them are particularly poorly understood. Here we investigate the species‐ and community‐level responses of microarthropods inhabiting a keystone plant species, on sub‐Antarctic Marion Island, to experimental reduction in precipitation, warming and shading. These climate manipulations were chosen based on observed climate trends and predicted indirect climate change impacts on this system. The dry‐warm and shade inducing treatments that were imposed effected significant species‐ and community‐level responses after a single year. Although the strongest community‐level trends included a dramatic decline in springtail abundance and total biomass under the dry‐warm and shade treatments, species responses were generally individualistic, that is, springtails responded differently to mites, and particular mite and springtail species responded differently to each other. Our results therefore provide additional support for the dynamic rather than static model for community responses to climate change, in the first such experiment in the sub‐Antarctic. In conclusion, these results show that an ongoing decline in precipitation and increase in temperature is likely to have dramatic direct and indirect effects on this microarthropod community. Moreover, they indicate that while at a broad scale it may be possible to make generalizations regarding species responses to climate change, these generalizations are unlikely to translate into predictable effects at the community level.     

Convergent ecosystem responses to 23‐year ambient and manipulated warming link advancing snowmelt and shrub encroachment to transient and long‐term climate–soil carbon feedback

Ecosystem responses to climate change can exert positive or negative feedbacks on climate, mediated in part by slow‐moving factors such as shifts in vegetation community composition. Long‐term experimental manipulations can be used to examine such ecosystem responses, but they also present another opportunity: inferring the extent to which contemporary climate change is responsible for slow changes in ecosystems under ambient conditions. Here, using 23 years of data, we document a shift from nonwoody to woody vegetation and a loss of soil carbon in ambient plots and show that these changes track previously shown similar but faster changes under experimental warming. This allows us to infer that climate change is the cause of the observed shifts in ambient vegetation and soil carbon and that the vegetation responses mediate the observed changes in soil carbon. Our findings demonstrate the realism of an experimental manipulation, allow attribution of a climate cause to observed ambient ecosystem changes, and demonstrate how a combination of long‐term study of ambient and experimental responses to warming can identify mechanistic drivers needed for realistic predictions of the conditions under which ecosystems are likely to become carbon sources or sinks over varying timescales.     

Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.     

Moving forward in global‐change ecology: capitalizing on natural variability

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global‐ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short‐term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate‐change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short‐term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.     

Population‐level thermal performance of a cold‐water ectotherm is linked to ontogeny and local environmental heterogeneity

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Predicting impacts of climate change on biodiversity: a role for semi‐mechanistic community‐level modelling

Aim Robust and reliable predictions of the effects of climate change on biodiversity are required in formulating conservation and management strategies that best retain biodiversity into the future. Significant challenges in modelling climate change impacts arise from limitations in our current knowledge of biodiversity. Community‐level modelling can complement species‐level approaches in overcoming these limitations and predicting climate change impacts on biodiversity as a whole. However, the community‐level approaches applied to date have been largely correlative, ignoring the key processes that influence change in biodiversity over space and time. Here, we suggest that the development of new ‘semi‐mechanistic’ community‐level models would substantially increase our capacity to predict climate change impacts on biodiversity. Location Global. Methods Drawing on an expansive review of biodiversity modelling approaches and recent advances in semi‐mechanistic modelling at the species level, we outline the main elements of a new semi‐mechanistic community‐level modelling approach. Results Our quantitative review revealed a sharp divide between mechanistic and non‐mechanistic biodiversity modelling approaches, with very few semi‐mechanistic models developed to date. Main conclusions We suggest that the conceptual framework presented here for combining mechanistic and non‐mechanistic community‐level approaches offers a promising means of incorporating key processes into predictions of climate change impacts on biodiversity whilst working within the limits of our current knowledge.     

Contemporary climate change in the Sonoran Desert favors cold‐adapted species

Impacts of long‐term climate shifts on the dynamics of intact communities within species ranges are not well understood. Here, we show that warming and drying of the Southwestern United States over the last 25 years has corresponded to a shift in the species composition of Sonoran Desert winter annuals, paradoxically favoring species that germinate and grow best in cold temperatures. Winter rains have been arriving later in the season, during December rather than October, leading to the unexpected result that plants are germinating under colder temperatures, shifting community composition to favor slow growing, water‐use efficient, cold‐adapted species. Our results demonstrate how detailed ecophysiological knowledge of individual species, combined with long‐term demographic data, can reveal complex and sometimes unexpected shifts in community composition in response to climate change. Further, these results highlight the potentially overwhelming impact of changes in phenology on the response of biota to a changing climate.     

Phenological responses of Icelandic subarctic grasslands to short‐term and long‐term natural soil warming

The phenology of vegetation, particularly the length of the growing season (LOS; i.e., the period from greenup to senescence), is highly sensitive to climate change, which could imply potent feedbacks to the climate system, for example, by altering the ecosystem carbon (C) balance. In recent decades, the largest extensions of LOS have been reported at high northern latitudes, but further warming‐induced LOS extensions may be constrained by too short photoperiod or unfulfilled chilling requirements. Here, we studied subarctic grasslands, which cover a vast area and contain large C stocks, but for which LOS changes under further warming are highly uncertain. We measured LOS extensions of Icelandic subarctic grasslands along natural geothermal soil warming gradients of different age (short term, where the measurements started after 5 years of warming and long term, i.e., warmed since ≥50 years) using ground‐level measurements of normalized difference vegetation index. We found that LOS linearly extended with on average 2.1 days per °C soil warming up to the highest soil warming levels (ca. +10°C) and that LOS had the potential to extend at least 1 month. This indicates that the warming impact on LOS in these subarctic grasslands will likely not saturate in the near future. A similar response to short‐ and long‐term warming indicated a strong physiological control of the phenological response of the subarctic grasslands to warming and suggested that genetic adaptations and community changes were likely of minor importance. We conclude that the warming‐driven extension of the LOSs of these subarctic grasslands did not saturate up to +10°C warming, and hence that growing seasons of high‐latitude grasslands are likely to continue lengthening with future warming (unless genetic adaptations or species shifts do occur). This persistence of the warming‐induced extension of LOS has important implications for the C‐sink potential of subarctic grasslands under climate change.     

Continent‐scale global change attribution in European birds ‐ combining annual and decadal time scales

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes‐based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long‐term population growth rates documents simultaneous responses that can be attributed to both climate change and land‐use change, including long‐term increases in populations of hot‐dwelling species and declines in long‐distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long‐term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high‐resolution time series are available in large‐scale biodiversity surveys.     

Experimental whole‐stream warming alters community size structure

How ecological communities respond to predicted increases in temperature will determine the extent to which Earth's biodiversity and ecosystem functioning can be maintained into a warmer future. Warming is predicted to alter the structure of natural communities, but robust tests of such predictions require appropriate large‐scale manipulations of intact, natural habitat that is open to dispersal processes via exchange with regional species pools. Here, we report results of a two‐year whole‐stream warming experiment that shifted invertebrate assemblage structure via unanticipated mechanisms, while still conforming to community‐level metabolic theory. While warming by 3.8 °C decreased invertebrate abundance in the experimental stream by 60% relative to a reference stream, total invertebrate biomass was unchanged. Associated shifts in invertebrate assemblage structure were driven by the arrival of new taxa and a higher proportion of large, warm‐adapted species (i.e., snails and predatory dipterans) relative to small‐bodied, cold‐adapted taxa (e.g., chironomids and oligochaetes). Experimental warming consequently shifted assemblage size spectra in ways that were unexpected, but consistent with thermal optima of taxa in the regional species pool. Higher temperatures increased community‐level energy demand, which was presumably satisfied by higher primary production after warming. Our experiment demonstrates how warming reassembles communities within the constraints of energy supply via regional exchange of species that differ in thermal physiological traits. Similar responses will likely mediate impacts of anthropogenic warming on biodiversity and ecosystem function across all ecological communities.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

Can amphibians take the heat? Vulnerability to climate warming in subtropical and temperate larval amphibian communities

Predicting the biodiversity impacts of global warming implies that we know where and with what magnitude these impacts will be encountered. Amphibians are currently the most threatened vertebrates, mainly due to habitat loss and to emerging infectious diseases. Global warming may further exacerbate their decline in the near future, although the impact might vary geographically. We predicted that subtropical amphibians should be relatively susceptible to warming‐induced extinctions because their upper critical thermal limits (CT_max) might be only slightly higher than maximum pond temperatures (T_max). We tested this prediction by measuring CT_max and T_max for 47 larval amphibian species from two thermally distinct subtropical communities (the warm community of the Gran Chaco and the cool community of Atlantic Forest, northern Argentina), as well as from one European temperate community. Upper thermal tolerances of tadpoles were positively correlated (controlling for phylogeny) with maximum pond temperatures, although the slope was steeper in subtropical than in temperate species. CT_max values were lowest in temperate species and highest in the subtropical warm community, which paradoxically, had very low warming tolerance (CT_max–T_max) and therefore may be prone to future local extinction from acute thermal stress if rising pond T_max soon exceeds their CT_max. Canopy‐protected subtropical cool species have larger warming tolerance and thus should be less impacted by peak temperatures. Temperate species are relatively secure to warming impacts, except for late breeders with low thermal tolerance, which may be exposed to physiological thermal stress in the coming years.     

Experimental evidence of gradual size‐dependent shifts in body size and growth of fish in response to warming

A challenge facing ecologists trying to predict responses to climate change is the few recent analogous conditions to use for comparison. For example, negative relationships between ectotherm body size and temperature are common both across natural thermal gradients and in small‐scale experiments. However, it is unknown if short‐term body size responses are representative of long‐term responses. Moreover, to understand population responses to warming, we must recognize that individual responses to temperature may vary over ontogeny. To enable predictions of how climate warming may affect natural populations, we therefore ask how body size and growth may shift in response to increased temperature over life history, and whether short‐ and long‐term growth responses differ. We addressed these questions using a unique setup with multidecadal artificial heating of an enclosed coastal bay in the Baltic Sea and an adjacent reference area (both with unexploited populations), using before‐after control‐impact paired time‐series analyses. We assembled individual growth trajectories of ~13,000 unique individuals of Eurasian perch and found that body growth increased substantially after warming, but the extent depended on body size: Only among small‐bodied perch did growth increase with temperature. Moreover, the strength of this response gradually increased over the 24 year warming period. Our study offers a unique example of how warming can affect fish populations over multiple generations, resulting in gradual changes in body growth, varying as organisms develop. Although increased juvenile growth rates are in line with predictions of the temperature–size rule, the fact that a larger body size at age was maintained over life history contrasts to that same rule. Because the artificially heated area is a contemporary system mimicking a warmer sea, our findings can aid predictions of fish responses to further warming, taking into account that growth responses may vary both over an individual's life history and over time.     

Long‐term mammal and nocturnal bird trends are influenced by vegetation type,weather and climate in temperate woodlands

Many studies have documented the individual effects of variables such as vegetation, long‐term climate and short‐term weather on biodiversity. Few, however, have explicitly explored how interactions among these major drivers can influence species abundance. We used data from a 15‐year study (2002–2017) in the endangered temperate woodlands of south‐eastern Australia to test hypotheses associated with the effects of vegetation type, long‐term climate and short‐term weather on population trajectories of seven species of (largely) nocturnal mammals and birds. Despite prolonged drought conditions, there was a significant increase in the abundance of some species over time (e.g. the Eastern Grey Kangaroo). It is possible that destocking of domestic livestock may have reduced competition with Kangaroos, thereby facilitating increases in abundance. The Common Brushtail Possum and Common Ringtail Possum were significantly less likely to occur in replanted woodlands, possibly because of the paucity of nesting sites. We found no evidence that replanted woodlands are refuges for exotic pest species like the European Rabbit and Red Fox. Short‐ and long‐term rainfall and vegetation type had important independent and combined effects on animal abundance. That is, responses to periods of high short‐term rainfall were dependent on vegetation type and whether sites occurred in long‐term climatically wet versus climatically dry locations. For example, the Red Fox responded positively to high levels of short‐term rainfall, but only at climatically dry sites. Our results highlight the complementary value of different vegetation types across the landscape and the context‐specific responses of animals to short‐term fluctuations in moisture availability. They also underscore the value of long‐term monitoring at a landscape scale for examining how multiple interacting factors influence trends in animal abundance.     

Both life‐history plasticity and local adaptation will shape range‐wide responses to climate warming in the tundra plant Silene acaulis

Many predictions of how climate change will impact biodiversity have focused on range shifts using species‐wide climate tolerances, an approach that ignores the demographic mechanisms that enable species to attain broad geographic distributions. But these mechanisms matter, as responses to climate change could fundamentally differ depending on the contributions of life‐history plasticity vs. local adaptation to species‐wide climate tolerances. In particular, if local adaptation to climate is strong, populations across a species’ range—not only those at the trailing range edge—could decline sharply with global climate change. Indeed, faster rates of climate change in many high latitude regions could combine with local adaptation to generate sharper declines well away from trailing edges. Combining 15 years of demographic data from field populations across North America with growth chamber warming experiments, we show that growth and survival in a widespread tundra plant show compensatory responses to warming throughout the species’ latitudinal range, buffering overall performance across a range of temperatures. However, populations also differ in their temperature responses, consistent with adaptation to local climate, especially growing season temperature. In particular, warming begins to negatively impact plant growth at cooler temperatures for plants from colder, northern populations than for those from warmer, southern populations, both in the field and in growth chambers. Furthermore, the individuals and maternal families with the fastest growth also have the lowest water use efficiency at all temperatures, suggesting that a trade‐off between growth and water use efficiency could further constrain responses to forecasted warming and drying. Taken together, these results suggest that populations throughout species’ ranges could be at risk of decline with continued climate change, and that the focus on trailing edge populations risks overlooking the largest potential impacts of climate change on species’ abundance and distribution.