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.     

Conserving biodiversity under climate change: the rear edge matters

Modern climate change is producing poleward range shifts of numerous taxa, communities and ecosystems worldwide. The response of species to changing environments is likely to be determined largely by population responses at range margins. In contrast to the expanding edge, the low-latitude limit (rear edge) of species ranges remains understudied, and the critical importance of rear edge populations as long-term stores of species' genetic diversity and foci of speciation has been little acknowledged. We review recent findings from the fossil record, phylogeography and ecology to illustrate that rear edge populations are often disproportionately important for the survival and evolution of biota. Their ecological features, dynamics and conservation requirements differ from those of populations in other parts of the range, and some commonly recommended conservation practices might therefore be of little use or even counterproductive for rear edge populations.     

Expanding population edges: theories,traits, and trade‐offs

Recent patterns of global change have highlighted the importance of understanding the dynamics and mechanisms of species range shifts and expansions. Unique demographic features, spatial processes, and selective pressures can result in the accumulation and evolution of distinctive phenotypic traits at the leading edges of expansions. We review the characteristics of expanding range margins and highlight possible mechanisms for the appearance of phenotypic differences between individuals at the leading edge and core of the range. The development of life history traits that increase dispersal or reproductive ability is predicted by theory and supported with extensive empirical evidence. Many examples of rapid phenotypic change are associated with trade‐offs that may influence the persistence of the trait once expansion ends. Accounting for the effects of edge phenotypes and related trade‐offs could be critical for predicting the spread of invasive species and population responses to climate change.     

Developmental trap or demographic bonanza? Opposing consequences of earlier phenology in a changing climate for a multivoltine butterfly

A rapidly changing climate has the potential to interfere with the timing of environmental cues that ectothermic organisms rely on to initiate and regulate life history events. Short‐lived ectotherms that exhibit plasticity in their life history could increase the number of generations per year under warming climate. If many individuals successfully complete an additional generation, the population experiences an additional opportunity to grow, and a warming climate could lead to a demographic bonanza. However, these plastic responses could become maladaptive in temperate regions, where a warmer climate could trigger a developmental pathway that cannot be completed within the growing season, referred to as a developmental trap. Here we incorporated detailed demography into commonly used photothermal models to evaluate these demographic consequences of phenological shifts due to a warming climate on the formerly widespread, multivoltine butterfly (Pieris oleracea). Using species‐specific temperature‐ and photoperiod‐sensitive vital rates, we estimated the number of generations per year and population growth rate over the set of climate conditions experienced during the past 38 years. We predicted that populations in the southern portion of its range have added a fourth generation in recent years, resulting in higher annual population growth rates (demographic bonanzas). We predicted that populations in the Northeast United States have experienced developmental traps, where increases in the thermal window initially caused mortality of the final generation and reduced growth rates. These populations may recover if more growing degree days are added to the year. Our framework for incorporating detailed demography into commonly used photothermal models demonstrates the importance of using both demography and phenology to predict consequences of phenological shifts.     

Losing your edge: climate change and the conservation value of range‐edge populations

Populations occurring at species' range edges can be locally adapted to unique environmental conditions. From a species' perspective, range‐edge environments generally have higher severity and frequency of extreme climatic events relative to the range core. Under future climates, extreme climatic events are predicted to become increasingly important in defining species' distributions. Therefore, range‐edge genotypes that are better adapted to extreme climates relative to core populations may be essential to species' persistence during periods of rapid climate change. We use relatively simple conceptual models to highlight the importance of locally adapted range‐edge populations (leading and trailing edges) for determining the ability of species to persist under future climates. Using trees as an example, we show how locally adapted populations at species' range edges may expand under future climate change and become more common relative to range‐core populations. We also highlight how large‐scale habitat destruction occurring in some geographic areas where many species range edge converge, such as biome boundaries and ecotones (e.g., the arc of deforestation along the rainforest‐cerrado ecotone in the southern Amazonia), can have major implications for global biodiversity. As climate changes, range‐edge populations will play key roles in helping species to maintain or expand their geographic distributions. The loss of these locally adapted range‐edge populations through anthropogenic disturbance is therefore hypothesized to reduce the ability of species to persist in the face of rapid future climate change.     

Differential effects of past climate warming on mountain and flatland species distributions: a multispecies North American mammal assessment

Aim The magnitude of predicted range shifts during climate change is likely to be different for species living in mountainous environments compared with those living in flatland environments. The southern edges of ranges in mountain species may not shift northwards during warming as populations instead migrate up available elevational gradients; overall latitudinal range appears therefore to expand. In contrast, flatland species should shift range centroids northwards but not expand or contract their latitudinal range extent. These hypotheses were tested utilizing Late Pleistocene and modern occurrence data. Location North America. Methods The location and elevation of modern and Late Pleistocene species occurrences were collected from data bases for 26 species living in mountain or flatland environments. Regressions of elevation change over latitude, and southern and northern range edges were calculated for each species for modern and fossil data sets. A combination of regressions and anova s were used to test whether flatland species shift range edges and latitudinal extents more than mountain species do. Results Flatland species had significantly larger northward shifts at southern range edges than did mountain‐dwelling species from the Late Pleistocene to the present. There was also a significant negative correlation between the amount of change in the latitude of the southern edge of the range and the amount of elevational shifting from the Late Pleistocene to the present. Although significant, only c. 25% of the variance could be explained by this relationship. In addition, there was a weak indication that overall range expansion was less in flatland‐dwelling than in mountain‐dwelling species. Main conclusions The approach used here was to examine past species’ range responses to warming that occurred after the last ice ages as a means to better predict potential future responses to continued warming. The results confirm predictions of differential southern edge and overall range shifts for species occupying mountain and flatland regions in North America. The findings may be broadly applicable in other regions, thus allowing better modelling of future range and distribution related responses.     

Forecasted range shifts of arid-land fishes in response to climate change

Climate change is poised to alter the distributional limits, center, and size of many species. Traits may influence different aspects of range shifts, with trophic generality facilitating shifts at the leading edge, and greater thermal tolerance limiting contractions at the trailing edge. The generality of relationships between traits and range shifts remains ambiguous however, especially for imperiled fishes residing in xeric riverscapes. Our objectives were to quantify contemporary fish distributions in the Lower Colorado River Basin, forecast climate change by 2085 using two general circulation models, and quantify shifts in the limits, center, and size of fish elevational ranges according to fish traits. We examined relationships among traits and range shift metrics either singly using univariate linear modeling or combined with multivariate redundancy analysis. We found that trophic and dispersal traits were associated with shifts at the leading and trailing edges, respectively, although projected range shifts were largely unexplained by traits. As expected, piscivores and omnivores with broader diets shifted upslope most at the leading edge while more specialized invertivores exhibited minimal changes. Fishes that were more mobile shifted upslope most at the trailing edge, defying predictions. No traits explained changes in range center or size. Finally, current preference explained multivariate range shifts, as fishes with faster current preferences exhibited smaller multivariate changes. Although range shifts were largely unexplained by traits, more specialized invertivorous fishes with lower dispersal propensity or greater current preference may require the greatest conservation efforts because of their limited capacity to shift ranges under climate change.     

A review and meta‐analysis of the effects of climate change on Holarctic mountain and upland bird populations

Mountain regions are globally important areas for biodiversity but are subject to multiple human‐induced threats, including climate change, which has been more severe at higher elevations. We reviewed evidence for impacts of climate change on Holarctic mountain bird populations in terms of physiology, phenology, trophic interactions, demography and observed and projected distribution shifts, including effects of other factors that interact with climate change. We developed an objective classification of high‐elevation, mountain specialist and generalist species, based on the proportion of their breeding range occurring in mountain regions. Our review found evidence of responses of mountain bird populations to climate (extreme weather events, temperature, rainfall and snow) and environmental (i.e. land use) change, but we know little about either the underlying mechanisms or the synergistic effects of climate and land use. Long‐term studies assessing reproductive success or survival of mountain birds in relation to climate change were rare. Few studies have considered shifts in elevational distribution over time and a meta‐analysis did not find a consistent direction in elevation change. A meta‐analysis carried out on future projections of distribution shifts suggested that birds whose breeding distributions are largely restricted to mountains are likely to be more negatively impacted than other species. Adaptation responses to climate change rely mostly on managing and extending current protected areas for both species already present, and for expected colonizing species that are losing habitat and climate space at lower elevation. However, developing effective management actions requires an improvement in the current knowledge of mountain species ecology, in the quality of climate data and in understanding the role of interacting factors. Furthermore, the evidence was mostly based on widespread species rather than mountain specialists. Scientists should provide valuable tools to assess the status of mountain birds, for example through the development of a mountain bird population index, and policy‐makers should influence legislation to develop efficient agri‐environment schemes and forestry practices for mountain birds, as well as to regulate leisure activities at higher elevations.     

Local adaptation at range edges: comparing elevation and latitudinal gradients

Local adaptation at range edges influences species’ distributions and how they respond to environmental change. However, the factors that affect adaptation, including gene flow and local selection pressures, are likely to vary across different types of range edge. We performed a reciprocal transplant experiment to investigate local adaptation in populations of Plantago lanceolata and P. major from central locations in their European range and from their latitudinal and elevation range edges (in northern Scandinavia and Swiss Alps, respectively). We also characterized patterns of genetic diversity and differentiation in populations using molecular markers. Range‐centre plants of P. major were adapted to conditions at the range centre, but performed similarly to range‐edge plants when grown at the range edges. There was no evidence for local adaptation when comparing central and edge populations of P. lanceolata. However, plants of both species from high elevation were locally adapted when compared with plants from high latitude, although the reverse was not true. This asymmetry was associated with greater genetic diversity and less genetic differentiation over the elevation gradient than over the latitudinal gradient. Our results suggest that adaptation in some range‐edge populations could increase their performance following climate change. However, responses are likely to differ along elevation and latitudinal gradients, with adaptation more likely at high‐elevation. Furthermore, based upon these results, we suggest that gene flow is unlikely to constrain adaptation in range‐edge populations of these species.     

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.     

Rapid evolution of phenology during range expansion with recent climate change

Although climate warming is expected to make habitat beyond species’ current cold range edge suitable for future colonization, this new habitat may present an array of biotic or abiotic conditions not experienced within the current range. Species’ ability to shift their range with climate change may therefore depend on how populations evolve in response to such novel environmental conditions. However, due to the recent nature of thus far observed range expansions, the role of rapid adaptation during climate change migration is only beginning to be understood. Here, we evaluated evolution during the recent native range expansion of the annual plant Dittrichia graveolens, which is spreading northward in Europe from the Mediterranean region. We examined genetically based differentiation between core and edge populations in their phenology, a trait that is likely under selection with shorter growing seasons and greater seasonality at northern latitudes. In parallel common garden experiments at range edges in Switzerland and the Netherlands, we grew plants from Dutch, Swiss, and central and southern French populations. Population genetic analysis following RAD‐sequencing of these populations supported the hypothesized central France origins of the Swiss and Dutch range edge populations. We found that in both common gardens, northern plants flowered up to 4 weeks earlier than southern plants. This differentiation in phenology extended from the core of the range to the Netherlands, a region only reached from central France over approximately the last 50 years. Fitness decreased as plants flowered later, supporting the hypothesized benefits of earlier flowering at the range edge. Our results suggest that native range expanding populations can rapidly adapt to novel environmental conditions in the expanded range, potentially promoting their ability to spread.     

Temperature tracking by North Sea benthic invertebrates in response to climate change

Climate change is a major threat to biodiversity and distributions shifts are one of the most significant threats to global warming, but the extent to which these shifts keep pace with a changing climate is yet uncertain. Understanding the factors governing range shifts is crucial for conservation management to anticipate patterns of biodiversity distribution under future anthropogenic climate change. Soft‐sediment invertebrates are a key faunal group because of their role in marine biogeochemistry and as a food source for commercial fish species. However, little information exists on their response to climate change. Here, we evaluate changes in the distribution of 65 North Sea benthic invertebrate species between 1986 and 2000 by examining their geographic, bathymetric and thermal niche shifts and test whether species are tracking their thermal niche as defined by minimum, mean or maximum sea bottom (SBT) and surface (SST) temperatures. Temperatures increased in the whole North Sea with many benthic invertebrates showing north‐westerly range shifts (leading/trailing edges as well as distribution centroids) and deepening. Nevertheless, distribution shifts for most species (3.8–7.3 km yr^?1 interquantile range) lagged behind shifts in both SBT and SST (mean 8.1 km yr^?1), resulting in many species experiencing increasing temperatures. The velocity of climate change (VoCC) of mean SST accurately predicted both the direction and magnitude of distribution centroid shifts, while maximum SST did the same for contraction of the trailing edge. The VoCC of SBT was not a good predictor of range shifts. No good predictor of expansions of the leading edge was found. Our results show that invertebrates need to shift at different rates and directions to track the climate velocities of different temperature measures, and are therefore lagging behind most temperature measures. If these species cannot withstand a change in thermal habitat, this could ultimately lead to a drop in benthic biodiversity.     

Integrated modeling predicts shifts in waterbird population dynamics under climate change

Climate change has been identified as one of the most important drivers of wildlife population dynamics. The in‐depth knowledge of the complex relationships between climate and population sizes through density dependent demographic processes is important for understanding and predicting population shifts under climate change, which requires integrated population models (IPMs) that unify the analyses of demography and abundance data. In this study we developed an IPM based on Gaussian approximation to dynamic N‐mixture models for large scale population data. We then analyzed four decades (1972–2013) of mallard Anas platyrhynchos breeding population survey, band‐recovery and climate data covering a large spatial extent from North American prairies through boreal habitat to Alaska. We aimed to test the hypothesis that climate change will cause shifts in population dynamics if climatic effects on demographic parameters that have substantial contribution to population growth vary spatially. More specifically, we examined the spatial variation of climatic effects on density dependent population demography, identified the key demographic parameters that are influential to population growth, and forecasted population responses to climate change. Our results revealed that recruitment, which explained more variance of population growth than survival, was sensitive to the temporal variation of precipitation in the southern portion of the study area but not in the north. Survival, by contrast, was insensitive to climatic variation. We then forecasted a decrease in mallard breeding population density in the south and an increase in the northwestern portion of the study area, indicating potential shifts in population dynamics under future climate change. Our results implied that different strategies need to be considered across regions to conserve waterfowl populations in the face of climate change. Our modelling approach can be adapted for other species and thus has wide application to understanding and predicting population dynamics in the presence of global change.     

A resurrection study reveals limited evolution of phenology in response to recent climate change across the geographic range of the scarlet monkeyflower

Premise of the studyAs global climate change alters drought regimes, rapid evolution of traits that facilitate adaptation to drought can rescue populations in decline. The evolution of phenological advancement can allow plant populations to escape drought, but evolutionary responses in phenology can vary across a species'' range due to differences in drought intensity and standing genetic variation.Methods Mimulus cardinalis, a perennial herb spanning a broad climatic gradient, recently experienced a period of record drought. Here, we used a resurrection study comparing flowering time and stem height at first flower of pre‐drought ancestors and post‐drought descendants from northern‐edge, central, and southern‐edge populations in a common environment to examine the evolution of drought escape across the latitudinal range.Key resultsContrary to the hypothesis of the evolution of advanced phenology in response to recent drought, flowering time did not advance between ancestors and descendants in any population, though storage condition and maternal effects could have impacted these results. Stem height was positively correlated with flowering time, such that plants that flowered earlier were shorter at first flower. This correlation could constrain the evolution of earlier flowering time if selection favors flowering early at a large size.ConclusionsThese findings suggest that rapid evolution of phenology will not rescue these populations from recent climate change. Future work is needed to examine the potential for the evolution of alternative drought strategies and phenotypic plasticity to buffer M. cardinalis populations from changing climate.     

Cold range edges of marine fishes track climate change better than warm edges

Species around the world are shifting their ranges in response to climate change. To make robust predictions about climate‐related colonizations and extinctions, it is vital to understand the dynamics of range edges. This study is among the first to examine annual dynamics of cold and warm range edges, as most global change studies average observational data over space or over time. We analyzed annual range edge dynamics of marine fishes—both at the individual species level and pooled into cold‐ and warm‐edge assemblages—in a multi‐decade time‐series of trawl surveys conducted on the Northeast US Shelf during a period of rapid warming. We tested whether cold edges show stronger evidence of climate tracking than warm edges (due to non‐climate processes or time lags at the warm edge; the biogeography hypothesis or extinction debt hypothesis), or whether they tracked temperature change equally (due to the influence of habitat suitability; the ecophysiology hypothesis). In addition to exploring correlations with regional temperature change, we calculated species‐ and assemblage‐specific sea bottom and sea surface temperature isotherms and used them to predict range edge position. Cold edges shifted further and tracked sea surface and bottom temperature isotherms to a greater degree than warm edges. Mixed‐effects models revealed that for a one‐degree latitude shift in isotherm position, cold edges shifted 0.47 degrees of latitude, and warm edges shifted only 0.28 degrees. Our results suggest that cold range edges are tracking climate change better than warm range edges, invalidating the ecophysiology hypothesis. We also found that even among highly mobile marine ectotherms in a global warming hotspot, few species are fully keeping pace with climate.     

The shaping of genetic variation in edge‐of‐range populations under past and future climate change

With rates of climate change exceeding the rate at which many species are able to shift their range or adapt, it is important to understand how future changes are likely to affect biodiversity at all levels of organisation. Understanding past responses and extent of niche conservatism in climatic tolerance can help predict future consequences. We use an integrated approach to determine the genetic consequences of past and future climate changes on a bat species, Plecotus austriacus. Glacial refugia predicted by palaeo‐modelling match those identified from analyses of extant genetic diversity and model‐based inference of demographic history. Former refugial populations currently contain disproportionately high genetic diversity, but niche conservatism, shifts in suitable areas and barriers to migration mean that these hotspots of genetic diversity are under threat from future climate change. Evidence of population decline despite recent northward migration highlights the need to conserve leading‐edge populations for spearheading future range shifts.     

Genes on the edge: A framework to detect genetic diversity imperiled by climate change

Ongoing global warming is disrupting several ecological and evolutionary processes, spanning different levels of biological organization. Species are expected to shift their ranges as a response to climate change, with relevant implications to peripheral populations at the trailing and leading edges. Several studies have analyzed the exposure of species to climate change but few have explored exposure at the intraspecific level. We introduce a framework to forecast exposure to climate change at the intraspecific level. We build on existing methods by combining correlative species distribution models, a model of species range dynamics, and a model of phylogeographic interpolation. We demonstrate the framework by applying it to 20 Iberian amphibian and reptile species. Our aims were to: (a) identify which species and intraspecific lineages will be most exposed to future climate change; (b) test if nucleotide diversity at the edges of species ranges are significantly higher or lower than on the overall range; and (c) analyze if areas of higher species gain, loss, and turnover coincide with those predicted for lineages richness and nucleotide diversity. We found that about 80% of the studied species are predicted to contract their range. Within each species, some lineages were predicted to contract their range, while others were predicted to maintain or expand it. Therefore, estimating the impacts of climate change at the species level only can underestimate losses at the intraspecific level. Some species had significant high amount of nucleotide at the trailing or leading edge, or both, but we did not find a consistent pattern across species. Spatial patterns of species richness, gain, loss, and turnover were fairly concurrent with lineages richness and nucleotide diversity. Our results support the need for increased attention to intraspecific diversity regarding monitoring and conservation strategies under climate change.     

Dynamics of range margins for metapopulations under climate change

We link spatially explicit climate change predictions to a dynamic metapopulation model. Predictions of species'' responses to climate change, incorporating metapopulation dynamics and elements of dispersal, allow us to explore the range margin dynamics for two lagomorphs of conservation concern. Although the lagomorphs have very different distribution patterns, shifts at the edge of the range were more pronounced than shifts in the overall metapopulation. For Romerolagus diazi (volcano rabbit), the lower elevation range limit shifted upslope by approximately 700 m. This reduced the area occupied by the metapopulation, as the mountain peak currently lacks suitable vegetation. For Lepus timidus (European mountain hare), we modelled the British metapopulation. Increasing the dispersive estimate caused the metapopulation to shift faster on the northern range margin (leading edge). By contrast, it caused the metapopulation to respond to climate change slower, rather than faster, on the southern range margin (trailing edge). The differential responses of the leading and trailing range margins and the relative sensitivity of range limits to climate change compared with that of the metapopulation centroid have important implications for where conservation monitoring should be targeted. Our study demonstrates the importance and possibility of moving from simple bioclimatic envelope models to second-generation models that incorporate both dynamic climate change and metapopulation dynamics.     

Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae)

Environmental variation often induces shifts in functional traits, yet we know little about whether plasticity will reduce extinction risks under climate change. As climate change proceeds, phenotypic plasticity could enable species with limited dispersal capacity to persist in situ, and migrating populations of other species to establish in new sites at higher elevations or latitudes. Alternatively, climate change could induce maladaptive plasticity, reducing fitness, and potentially stalling adaptation and migration. Here, we quantified plasticity in life history, foliar morphology, and ecophysiology in Boechera stricta (Brassicaceae), a perennial forb native to the Rocky Mountains. In this region, warming winters are reducing snowpack and warming springs are advancing the timing of snow melt. We hypothesized that traits that were historically advantageous in hot and dry, low‐elevation locations will be favored at higher elevation sites due to climate change. To test this hypothesis, we quantified trait variation in natural populations across an elevational gradient. We then estimated plasticity and genetic variation in common gardens at two elevations. Finally, we tested whether climatic manipulations induce plasticity, with the prediction that plants exposed to early snow removal would resemble individuals from lower elevation populations. In natural populations, foliar morphology and ecophysiology varied with elevation in the predicted directions. In the common gardens, trait plasticity was generally concordant with phenotypic clines from the natural populations. Experimental snow removal advanced flowering phenology by 7 days, which is similar in magnitude to flowering time shifts over 2–3 decades of climate change. Therefore, snow manipulations in this system can be used to predict eco‐evolutionary responses to global change. Snow removal also altered foliar morphology, but in unexpected ways. Extensive plasticity could buffer against immediate fitness declines due to changing climates.     

Tree demography suggests multiple directions and drivers for species range shifts in mountains of Northeastern United States

Climate change is expected to lead to upslope shifts in tree species distributions, but the evidence is mixed partly due to land‐use effects and individualistic species responses to climate. We examined how individual tree species demography varies along elevational climatic gradients across four states in the northeastern United States to determine whether species elevational distributions and their potential upslope (or downslope) shifts were controlled by climate, land‐use legacies (past logging), or soils. We characterized tree demography, microclimate, land‐use legacies, and soils at 83 sites stratified by elevation (~500 to ~1200 m above sea level) across 12 mountains containing the transition from northern hardwood to spruce‐fir forests. We modeled elevational distributions of tree species saplings and adults using logistic regression to test whether sapling distributions suggest ongoing species range expansion upslope (or contraction downslope) relative to adults, and we used linear mixed models to determine the extent to which climate, land use, and soil variables explain these distributions. Tree demography varied with elevation by species, suggesting a potential upslope shift only for American beech, downslope shifts for red spruce (more so in cool regions) and sugar maple, and no change with elevation for balsam fir. While soils had relatively minor effects, climate was the dominant predictor for most species and more so for saplings than adults of red spruce, sugar maple, yellow birch, cordate birch, and striped maple. On the other hand, logging legacies were positively associated with American beech, sugar maple, and yellow birch, and negatively with red spruce and balsam fir – generally more so for adults than saplings. All species exhibited individualistic rather than synchronous demographic responses to climate and land use, and the return of red spruce to lower elevations where past logging originally benefited northern hardwood species indicates that land use may mask species range shifts caused by changing climate.