Individual-scale inference to anticipate climate-change vulnerability of biodiversity

Anticipating how biodiversity will respond to climate change is challenged by the fact that climate variables affect individuals in competition with others, but interest lies at the scale of species and landscapes. By omitting the individual scale, models cannot accommodate the processes that determine future biodiversity. We demonstrate how individual-scale inference can be applied to the problem of anticipating vulnerability of species to climate. The approach places climate vulnerability in the context of competition for light and soil moisture. Sensitivities to climate and competition interactions aggregated from the individual tree scale provide estimates of which species are vulnerable to which variables in different habitats. Vulnerability is explored in terms of specific demographic responses (growth, fecundity and survival) and in terms of the synthetic response (the combination of demographic rates), termed climate tracking. These indices quantify risks for individuals in the context of their competitive environments. However, by aggregating in specific ways (over individuals, years, and other input variables), we provide ways to summarize and rank species in terms of their risks from climate change.     

Structural overshoot of tree growth with climate variability and the global spectrum of drought‐induced forest dieback

Ongoing climate change poses significant threats to plant function and distribution. Increased temperatures and altered precipitation regimes amplify drought frequency and intensity, elevating plant stress and mortality. Large‐scale forest mortality events will have far‐reaching impacts on carbon and hydrological cycling, biodiversity, and ecosystem services. However, biogeographical theory and global vegetation models poorly represent recent forest die‐off patterns. Furthermore, as trees are sessile and long‐lived, their responses to climate extremes are substantially dependent on historical factors. We show that periods of favourable climatic and management conditions that facilitate abundant tree growth can lead to structural overshoot of aboveground tree biomass due to a subsequent temporal mismatch between water demand and availability. When environmental favourability declines, increases in water and temperature stress that are protracted, rapid, or both, drive a gradient of tree structural responses that can modify forest self‐thinning relationships. Responses ranging from premature leaf senescence and partial canopy dieback to whole‐tree mortality reduce canopy leaf area during the stress period and for a lagged recovery window thereafter. Such temporal mismatches of water requirements from availability can occur at local to regional scales throughout a species geographical range. As climate change projections predict large future fluctuations in both wet and dry conditions, we expect forests to become increasingly structurally mismatched to water availability and thus overbuilt during more stressful episodes. By accounting for the historical context of biomass development, our approach can explain previously problematic aspects of large‐scale forest mortality, such as why it can occur throughout the range of a species and yet still be locally highly variable, and why some events seem readily attributable to an ongoing drought while others do not. This refined understanding can facilitate better projections of structural overshoot responses, enabling improved prediction of changes in forest distribution and function from regional to global scales.     

Influences of forest structure, climate and species composition on tree mortality across the eastern US

Few studies have quantified regional variation in tree mortality, or explored whether species compositional changes or within-species variation are responsible for regional patterns, despite the fact that mortality has direct effects on the dynamics of woody biomass, species composition, stand structure, wood production and forest response to climate change. Using bayesian analysis of over 430,000 tree records from a large eastern US forest database we characterised tree mortality as a function of climate, soils, species and size (stem diameter). We found (1) mortality is U-shaped vs. stem diameter for all 21 species examined; (2) mortality is hump-shaped vs. plot basal area for most species; (3) geographical variation in mortality is substantial, and correlated with several environmental factors; and (4) individual species vary substantially from the combined average in the nature and magnitude of their mortality responses to environmental variation. Regional variation in mortality is therefore the product of variation in species composition combined with highly varied mortality-environment correlations within species. The results imply that variation in mortality is a crucial part of variation in the forest carbon cycle, such that including this variation in models of the global carbon cycle could significantly narrow uncertainty in climate change predictions.     

Sources of Variability in Net Primary Production Predictions at a Regional Scale: A Comparison Using PnET-II and TEM 4.0 in Northeastern US Forests

Because model predictions at continental and global scales are necessarily based on broad characterizations of vegetation, soils, and climate, estimates of carbon stocks and fluxes made by global terrestrial biosphere models may not be accurate for every region. At the regional scale, we suggest that attention can be focused more clearly on understanding the relative strengths of predicted net primary productivity (NPP) limitation by energy, water, and nutrients. We evaluate the sources of variability among model predictions of NPP with a regional-scale comparison between estimates made by PnET-II (a forest ecosystem process model previously applied to the northeastern region) and TEM 4.0 (a terrestrial biosphere model typically applied to the globe) for the northeastern US. When the same climate, vegetation, and soil data sets were used to drive both models, regional average NPP predictions made by PnET-II and TEM were remarkably similar, and at the biome level, model predictions agreed fairly well with NPP estimates developed from field measurements. However, TEM 4.0 predictions were more sensitive to regional variations in temperature as a result of feedbacks between temperature and belowground N availability. In PnET-II, the direct link between transpiration and photosynthesis caused substantial water stress in hardwood and pine forest types with increases in solar radiation; predicted water stress was relieved substantially when soil water holding capacity (WHC) was increased. Increasing soil WHC had little effect on TEM 4.0 predictions because soil water storage was already sufficient to meet plant demand with baseline WHC values, and because predicted N availability under baseline conditions in this region was not limited by water. Because NPP predictions were closely keyed to forest cover type, the relative coverage of low- versus high-productivity forests at both fine and coarse resolutions was an important determinant of regional NPP predictions. Therefore, changes in grid cell size and differences in the methods used to aggregate from fine to coarse resolution were important to NPP predictions insofar as they changed the relative proportions of forest cover. We suggest that because the small patches of high-elevation spruce-fir forest in this region are substantially less productive than forests in the remainder of the region, more accurate NPP predictions will result if models applied to this region use land cover input data sets that retain as much fine-resolution forest type variability as possible. The differences among model responses to variations in climate and soil WHC data sets suggest that the models will respond quite differently to scenarios of future climate. A better understanding of the dynamic interactions between water stress, N availability, and forest productivity in this region will enable models to make more accurate predictions of future carbon stocks and fluxes. Received 19 June 1998; accepted 25 June 1999.     

Climate change,woodpeckers, and forests: Current trends and future modeling needs

The structure and composition of forest ecosystems are expected to shift with climate‐induced changes in precipitation, temperature, fire, carbon mitigation strategies, and biological disturbance. These factors are likely to have biodiversity implications. However, climate‐driven forest ecosystem models used to predict changes to forest structure and composition are not coupled to models used to predict changes to biodiversity. We proposed integrating woodpecker response (biodiversity indicator) with forest ecosystem models. Woodpeckers are a good indicator species of forest ecosystem dynamics, because they are ecologically constrained by landscape‐scale forest components, such as composition, structure, disturbance regimes, and management activities. In addition, they are correlated with forest avifauna community diversity. In this study, we explore integrating woodpecker and forest ecosystem climate models. We review climate–woodpecker models and compare the predicted responses to observed climate‐induced changes. We identify inconsistencies between observed and predicted responses, explore the modeling causes, and identify the models pertinent to integration that address the inconsistencies. We found that predictions in the short term are not in agreement with observed trends for 7 of 15 evaluated species. Because niche constraints associated with woodpeckers are a result of complex interactions between climate, vegetation, and disturbance, we hypothesize that the lack of adequate representation of these processes in the current broad‐scale climate–woodpecker models results in model–data mismatch. As a first step toward improvement, we suggest a conceptual model of climate–woodpecker–forest modeling for integration. The integration model provides climate‐driven forest ecosystem modeling with a measure of biodiversity while retaining the feedback between climate and vegetation in woodpecker climate change modeling.     

Environmental heterogeneity and neighbourhood interference modulate the individual response of Juniperus thurifera tree-ring growth to climate

Individual variation of tree-ring growth response to climate and heterogeneity of the local environment are usually neglected in dendrochronological research. Even if there is evidence showing that individual responsiveness to climate may depend on intrinsic traits such as tree age, size or sex, its modulation by the local heterogeneity of extrinsic factors has been less studied. Using an extensive, strictly regular sampling scheme across a 3300 ha woodland, we assessed the individual variation of tree-ring growth responses to climate in 100 Juniperus thurifera L. trees. The climatic response was evaluated by bootstrapped correlations of both population- and individual-based tree-ring chronologies with monthly records of precipitation, cloudiness, minimum and maximum temperatures. We studied also the influence of extrinsic abiotic (elevation, slope, heat load, tree location) and biotic (competition from neighbouring trees) factors on the individual growth variation and its climatic response. At a population level, growth was controlled by February–March precipitation, April minimum temperature, and June water stress. A significant proportion of individuals did not respond to those variables, but were sensitive to others not relevant at the population level. Inter-annual growth variation was strongly modulated by competition, whereas trees under lower competition levels, in eastern and warmer areas, were the most responsive to climate. The individual climatic response was, at least partially, modulated by the local heterogeneity of extrinsic factors. By considering environmental heterogeneity and neighbourhood interference we can identify the spectrum of site-dependent climatic responses in a population, which in turn will enable more realistic predictions of tree responses to ongoing climate change.     

Altered dynamics of forest recovery under a changing climate

Forest regeneration following disturbance is a key ecological process, influencing forest structure and function, species assemblages, and ecosystem–climate interactions. Climate change may alter forest recovery dynamics or even prevent recovery, triggering feedbacks to the climate system, altering regional biodiversity, and affecting the ecosystem services provided by forests. Multiple lines of evidence – including global‐scale patterns in forest recovery dynamics; forest responses to experimental manipulation of CO₂, temperature, and precipitation; forest responses to the climate change that has already occurred; ecological theory; and ecosystem and earth system models – all indicate that the dynamics of forest recovery are sensitive to climate. However, synthetic understanding of how atmospheric CO₂ and climate shape trajectories of forest recovery is lacking. Here, we review these separate lines of evidence, which together demonstrate that the dynamics of forest recovery are being impacted by increasing atmospheric CO₂ and changing climate. Rates of forest recovery generally increase with CO₂, temperature, and water availability. Drought reduces growth and live biomass in forests of all ages, having a particularly strong effect on seedling recruitment and survival. Responses of individual trees and whole‐forest ecosystems to CO₂ and climate manipulations often vary by age, implying that forests of different ages will respond differently to climate change. Furthermore, species within a community typically exhibit differential responses to CO₂ and climate, and altered community dynamics can have important consequences for ecosystem function. Age‐ and species‐dependent responses provide a mechanism by which climate change may push some forests past critical thresholds such that they fail to recover to their previous state following disturbance. Altered dynamics of forest recovery will result in positive and negative feedbacks to climate change. Future research on this topic and corresponding improvements to earth system models will be a key to understanding the future of forests and their feedbacks to the climate system.     

The impacts of increasing drought on forest dynamics,structure, and biodiversity in the United States

We synthesize insights from current understanding of drought impacts at stand‐to‐biogeographic scales, including management options, and we identify challenges to be addressed with new research. Large stand‐level shifts underway in western forests already are showing the importance of interactions involving drought, insects, and fire. Diebacks, changes in composition and structure, and shifting range limits are widely observed. In the eastern US, the effects of increasing drought are becoming better understood at the level of individual trees, but this knowledge cannot yet be confidently translated to predictions of changing structure and diversity of forest stands. While eastern forests have not experienced the types of changes seen in western forests in recent decades, they too are vulnerable to drought and could experience significant changes with increased severity, frequency, or duration in drought. Throughout the continental United States, the combination of projected large climate‐induced shifts in suitable habitat from modeling studies and limited potential for the rapid migration of tree populations suggests that changing tree and forest biogeography could substantially lag habitat shifts already underway. Forest management practices can partially ameliorate drought impacts through reductions in stand density, selection of drought‐tolerant species and genotypes, artificial regeneration, and the development of multistructured stands. However, silvicultural treatments also could exacerbate drought impacts unless implemented with careful attention to site and stand characteristics. Gaps in our understanding should motivate new research on the effects of interactions involving climate and other species at the stand scale and how interactions and multiple responses are represented in models. This assessment indicates that, without a stronger empirical basis for drought impacts at the stand scale, more complex models may provide limited guidance.     

Climate-driven,but dynamic and complex? A reconciliation of competing hypotheses for species’ distributions

Estimates of the percentage of species “committed to extinction” by climate change range from 15% to 37%. The question is whether factors other than climate need to be included in models predicting species’ range change. We created demographic range models that include climate vs. climate-plus-competition, evaluating their influence on the geographic distribution of Pinus edulis, a pine endemic to the semiarid southwestern U.S. Analyses of data on 23,426 trees in 1941 forest inventory plots support the inclusion of competition in range models. However, climate and competition together only partially explain this species’ distribution. Instead, the evidence suggests that climate affects other range-limiting processes, including landscape-scale, spatial processes such as disturbances and antagonistic biotic interactions. Complex effects of climate on species distributions—through indirect effects, interactions, and feedbacks—are likely to cause sudden changes in abundance and distribution that are not predictable from a climate-only perspective.     

Climate variability drives recent tree mortality in Europe

Tree mortality is an important process in forest ecosystems, frequently hypothesized to be highly climate sensitive. Yet, tree death remains one of the least understood processes of forest dynamics. Recently, changes in tree mortality have been observed in forests around the globe, which could profoundly affect ecosystem functioning and services provisioning to society. We describe continental‐scale patterns of recent tree mortality from the only consistent pan‐European forest monitoring network, identifying recent mortality hotspots in southern and northern Europe. Analyzing 925,462 annual observations of 235,895 trees between 2000 and 2012, we determine the influence of climate variability and tree age on interannual variation in tree mortality using Cox proportional hazard models. Warm summers as well as high seasonal variability in precipitation increased the likelihood of tree death. However, our data also suggest that reduced cold‐induced mortality could compensate increased mortality related to peak temperatures in a warming climate. Besides climate variability, age was an important driver of tree mortality, with individual mortality probability decreasing with age over the first century of a trees life. A considerable portion of the observed variation in tree mortality could be explained by satellite‐derived net primary productivity, suggesting that widely available remote sensing products can be used as an early warning indicator of widespread tree mortality. Our findings advance the understanding of patterns of large‐scale tree mortality by demonstrating the influence of seasonal and diurnal climate variation, and highlight the potential of state‐of‐the‐art remote sensing to anticipate an increased likelihood of tree mortality in space and time.     

Simulating forest ecosystem response to climate warming incorporating spatial effects in north-eastern China

Aim Predictions of ecosystem responses to climate warming are often made using gap models, which are among the most effective tools for assessing the effects of climate change on forest composition and structure. Gap models do not generally account for broad‐scale effects such as the spatial configuration of the simulated forest ecosystems, disturbance, and seed dispersal, which extend beyond the simulation plots and are important under changing climates. In this study we incorporate the broad‐scale spatial effects (spatial configurations of the simulated forest ecosystems, seed dispersal and fire disturbance) in simulating forest responses to climate warming. We chose the Changbai Natural Reserve in China as our study area. Our aim is to reveal the spatial effects in simulating forest responses to climate warming and make new predictions by incorporating these effects in the Changbai Natural Reserve. Location Changbai Natural Reserve, north‐eastern China. Method We used a coupled modelling approach that links a gap model with a spatially explicit landscape model. In our approach, the responses (establishment) of individual species to climate warming are simulated using a gap model (linkages ) that has been utilized previously for making predictions in this region; and the spatial effects are simulated using a landscape model (LANDIS) that incorporates spatial configurations of the simulated forest ecosystems, seed dispersal and fire disturbance. We used the recent predictions of the Canadian Global Coupled Model (CGCM2) for the Changbai Mountain area (4.6 °C average annual temperature increase and little precipitation change). For the area encompassed by the simulation, we examined four major ecosystems distributed continuously from low to high elevations along the northern slope: hardwood forest, mixed Korean pine hardwood forest, spruce‐fir forest, and sub‐alpine forest. Results The dominant effects of climate warming were evident on forest ecosystems in the low and high elevation areas, but not in the mid‐elevation areas. This suggests that the forest ecosystems near the southern and northern ranges of their distributions will have the strongest response to climate warming. In the mid‐elevation areas, environmental controls exerted the dominant influence on the dynamics of these forests (e.g. spruce‐fir) and their resilience to climate warming was suggested by the fact that the fluctuations of species trajectories for these forests under the warming scenario paralleled those under the current climate scenario. Main conclusions With the spatial effects incorporated, the disappearance of tree species in this region due to the climate warming would not be expected within the 300‐year period covered by the simulation. Neither Korean pine nor spruce‐fir was completely replaced by broadleaf species during the simulation period. Even for the sub‐alpine forest, mountain birch did not become extinct under the climate warming scenario, although its occurrence was greatly reduced. However, the decreasing trends characterizing Korean pine, spruce, and fir indicate that in simulations beyond 300 years these species could eventually be replaced by broadleaf tree species. A complete forest transition would take much longer than the time periods predicted by the gap models.     

A GIS-based simulation program to predict multi-species size-structure dynamics for natural forests in Hokkaido, northern Japan

A simulation program that runs on a geographic information system (GIS) was developed to predict the multi-species size-structure dynamics of forest stands. Because important characteristics of a forest stand, including woody biomass accumulation, carbon storage, commercial value of timber, and functions for environmental conservation, can be inferred from the size structures of the component populations, management plans can be made from the predictions of the size-structure dynamics. For example, the simulation can incorporate various forms of thinning; forest managers can then try several thinning plans in simulated forest stands and choose the appropriate plan that achieves the best results. Using GIS to predict the size-structure dynamics of forest stands is of practical importance, because GIS has been used widely in forest management and can easily handle spatial distributions of environmental information (e.g., climate, geology, soils) that may influence tree performance. To predict size-structure dynamics, the program numerically solves a continuum equation that describes size-structure dynamics based on growth and mortality rates of individual trees. When predicting size-structure dynamics of a forest stand, the program obtains the environmental information of the stand from a database stored in the GIS and calculates environmental factors such as warmth index and potential evapotranspiration/precipitation ratio that influence growth and mortality rates. The simulation program calculates growth and mortality rates using published growth and mortality models that incorporate the effects of size of the individual, competition between trees, and abiotic environmental factors. To demonstrate the effects of abiotic environmental factors on the multi-species size-structure dynamics, sensitivity analyses were conducted. The size-structure dynamics varied in a way that was predictable from the responses of the growth and mortality rates to variations in the abiotic environmental factors. To demonstrate the size-structure dynamics in different locations, five test runs of the simulation program were also performed using the same initial size-structure and five different sets of abiotic environmental conditions from five locations. At the end of the simulation, the predicted size structures differed because the growth and mortality rates differed among the five locations. Finally, the response of the size structure to thinning was clarified. The result showed how the size structure of a component species in a forest stand is dependent on the presence of other species.     

Patterns and Drivers of Tree Mortality in Iberian Forests: Climatic Effects Are Modified by Competition

Tree mortality is a key process underlying forest dynamics and community assembly. Understanding how tree mortality is driven by simultaneous drivers is needed to evaluate potential effects of climate change on forest composition. Using repeat-measure information from c. 400,000 trees from the Spanish Forest Inventory, we quantified the relative importance of tree size, competition, climate and edaphic conditions on tree mortality of 11 species, and explored the combined effect of climate and competition. Tree mortality was affected by all of these multiple drivers, especially tree size and asymmetric competition, and strong interactions between climate and competition were found. All species showed L-shaped mortality patterns (i.e. showed decreasing mortality with tree size), but pines were more sensitive to asymmetric competition than broadleaved species. Among climatic variables, the negative effect of temperature on tree mortality was much larger than the effect of precipitation. Moreover, the effect of climate (mean annual temperature and annual precipitation) on tree mortality was aggravated at high competition levels for all species, but especially for broadleaved species. The significant interaction between climate and competition on tree mortality indicated that global change in Mediterranean regions, causing hotter and drier conditions and denser stands, could lead to profound effects on forest structure and composition. Therefore, to evaluate the potential effects of climatic change on tree mortality, forest structure must be considered, since two systems of similar composition but different structure could radically differ in their response to climatic conditions.     

Climate change vulnerability of forest biodiversity: climate and competition tracking of demographic rates

Forest responses to climate change will depend on demographic impacts in the context of competition. Current models used to predict species responses, termed climate envelope models (CEMs), are controversial, because (i) calibration and prediction are based on correlations in space (CIS) between species abundance and climate, rather than responses to climate change over time (COT), and (ii) they omit competition. To determine the relative importance of COT, CIS, and competition for light, we applied a longitudinal analysis of 27 000 individual trees over 6–18 years subjected to experimental and natural variation in risk factors. Sensitivities and climate and resource tracking identify which species are vulnerable to these risk factors and in what ways. Results show that responses to COT differ from those predicted based on CIS. The most important impact is the effect of spring temperature on fecundity, rather than any input variable on growth or survival. Of secondary importance is growing season moisture. Species in the genera Pinus, Ulmus, Magnolia, and Fagus are particularly vulnerable to climate variation. However, the effect of competition on growth and mortality risk exceeds the effects of climate variation in space or time for most species. Because sensitivities to COT and competition are larger than CIS, current models miss the most important effects. By directly comparing sensitivity to climate in time and space, together with competition, the approach identifies which species are sensitive to climate change and why, including the heretofore overlooked impact on fecundity.     

Species,Climate and Landscape Physiography Drive Variable Growth Trends in Subalpine Forests

Forests around the world are undergoing rapid changes due to changing climate and increasing physiological stress, but forest response to climate at the ecosystem scale can be highly variable due to the mixed responses of different trees across heterogeneous landscapes. To determine the response of ecosystems in the Rocky Mountains to climate stress, we investigated the response of subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii), two widely distributed subalpine forest species of Rocky Mountains, to climate warming across a region characterized by gradients of elevation, aspect and soil type. We investigated the growth trend of individual trees through time, determined the climate variables most important for driving growth and quantified the interactions between climate and topography that influence long-term growth trends and potential ecological changes across the study region. Growth trends of these two species are similar through the first part of the century, but diverge during the last several decades. Since 1975, subalpine fir growth decreased through time, while Engelmann spruce growth increased. We find that aspect and warm summer temperatures are the most important factors determining growth in subalpine fir, and subalpine fir growth declines are greatest on east- and south-facing aspects. In contrast, Engelmann spruce growth is uniformly unresponsive to climate. In addition to highlighting the importance of species-level differences in growth response to climate, our results also identify interactions between climate and local physiography as controls on long-term growth trends and suggest that the local landscape physiography can mediate climate-related stress in forested ecosystems. This work advances our understanding of how forest stress is mitigated by landscape factors at the ecosystem scale, and how interactions of species, landscape and climate will control future ecosystem composition and forest growth dynamics.     

Demographic synthesis for global tree species conservation

Conserving the tree species of the world requires syntheses on which tree species are most vulnerable to pressing threats, such as climate change, invasive pests and pathogens, or selective logging. Here, we review the population and forest dynamics models that, when parameterized with data from population studies, forest inventories, or tree rings, have been used for identifying life-history strategies of species and threat-related changes in population demography and dynamics. The available evidence suggests that slow-growing and/or long-lived species are the most vulnerable. However, a lack of comparative, multi-species studies still challenges more precise predictions of the vulnerability of tree species to threats. Improving data coverage for mortality and recruitment, and accounting for interactions among threats, would greatly advance vulnerability assessments for conservation prioritizations of trees worldwide.     

Species interactions weakly modify climate‐induced tree co‐occurrence patterns

Aims

Species distributions are hypothesized to be underlain by a complex association of processes that span multiple spatial scales including biotic interactions, dispersal limitation, fine‐scale resource gradients and climate. Species disequilibrium with climate may reflect the effects of non‐climatic processes on species distributions, yet distribution models have rarely directly considered non‐climatic processes. Here, we use a Joint Species Distribution Model (JSDM) to investigate the influence of non‐climatic factors on species co‐occurrence patterns and to directly quantify the relative influences of climate and alternative processes that may generate correlated responses in species distributions, such as species interactions, on tree co‐occurrence patterns.

Location

US Rocky Mountains.

Methods

We apply a Bayesian JSDM to simultaneously model the co‐occurrence patterns of ten dominant tree species across the Rocky Mountains, and evaluate climatic and residual correlations from the fitted model to determine the relative contribution of each component to observed co‐occurrence patterns. We also evaluate predictions generated from the fitted model relative to a single‐species modelling approach.

Results

For most species, correlation due to climate covariates exceeded residual correlation, indicating an overriding influence of broad‐scale climate on co‐occurrence patterns. Accounting for covariance among species did not significantly improve predictions relative to a single‐species approach, providing limited evidence for a strong independent influence of species interactions on distribution patterns.

Conclusions

Overall, our findings indicate that climate is an important driver of regional biodiversity patterns and that interactions between dominant tree species contribute little to explain species co‐occurrence patterns among Rocky Mountain trees.     

Climatic Stress during Stand Development Alters the Sign and Magnitude of Age-Related Growth Responses in a Subtropical Mountain Pine

The modification of typical age-related growth by environmental changes is poorly understood, In part because there is a lack of consensus at individual tree level regarding age-dependent growth responses to climate warming as stands develop. To increase our current understanding about how multiple drivers of environmental change can modify growth responses as trees age we used tree ring data of a mountain subtropical pine species along an altitudinal gradient covering more than 2,200 m of altitude. We applied mixed-linear models to determine how absolute and relative age-dependent growth varies depending on stand development; and to quantify the relative importance of tree age and climate on individual tree growth responses. Tree age was the most important factor for tree growth in models parameterised using data from all forest developmental stages. Contrastingly, the relationship found between tree age and growth became non-significant in models parameterised using data corresponding to mature stages. These results suggest that although absolute tree growth can continuously increase along tree size when trees reach maturity age had no effect on growth. Tree growth was strongly reduced under increased annual temperature, leading to more constant age-related growth responses. Furthermore, young trees were the most sensitive to reductions in relative growth rates, but absolute growth was strongly reduced under increased temperature in old trees. Our results help to reconcile previous contrasting findings of age-related growth responses at the individual tree level, suggesting that the sign and magnitude of age-related growth responses vary with stand development. The different responses found to climate for absolute and relative growth rates suggest that young trees are particularly vulnerable under warming climate, but reduced absolute growth in old trees could alter the species’ potential as a carbon sink in the future.     

The impact of climate change measured at relevant spatial scales: new hope for tropical lizards

Much attention has been given to recent predictions that widespread extinctions of tropical ectotherms, and tropical forest lizards in particular, will result from anthropogenic climate change. Most of these predictions, however, are based on environmental temperature data measured at a maximum resolution of 1 km², whereas individuals of most species experience thermal variation on a much finer scale. To address this disconnect, we combined thermal performance curves for five populations of Anolis lizard from the Bay Islands of Honduras with high‐resolution temperature distributions generated from physical models. Previous research has suggested that open‐habitat species are likely to invade forest habitat and drive forest species to extinction. We test this hypothesis, and compare the vulnerabilities of closely related, but allopatric, forest species. Our data suggest that the open‐habitat populations we studied will not invade forest habitat and may actually benefit from predicted warming for many decades. Conversely, one of the forest species we studied should experience reduced activity time as a result of warming, while two others are unlikely to experience a significant decline in performance. Our results suggest that global‐scale predictions generated using low‐resolution temperature data may overestimate the vulnerability of many tropical ectotherms to climate change.     

A climate change‐induced threat to the ecological resilience of a subtropical monsoon evergreen broad‐leaved forest in Southern China

Recent studies have suggested that tropical forests may not be resilient against climate change in the long term, primarily owing to predicted reductions in rainfall and forest productivity, increased tree mortality, and declining forest biomass carbon sinks. These changes will be caused by drought‐induced water stress and ecosystem disturbances. Several recent studies have reported that climate change has increased tree mortality in temperate and boreal forests, or both mortality and recruitment rates in tropical forests. However, no study has yet examined these changes in the subtropical forests that account for the majority of China's forested land. In this study, we describe how the monsoon evergreen broad‐leaved forest has responded to global warming and drought stress using 32 years of data from forest observation plots. Due to an imbalance in mortality and recruitment, and changes in diameter growth rates between larger and smaller trees and among different functional groups, the average DBH of trees and forest biomass have decreased. Sap flow measurements also showed that larger trees were more stressed than smaller trees by the warming and drying environment. As a result, the monsoon evergreen broad‐leaved forest community is undergoing a transition from a forest dominated by a cohort of fewer and larger individuals to a forest dominated by a cohort of more and smaller individuals, with a different species composition, suggesting that subtropical forests are threatened by their lack of resilience against long‐term climate change.