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.     

The role of resource dispersion in promoting the co‐occurrence of dominant and subordinate ant species

Dominant competitors govern resource use in many communities, leading to predictions of local exclusion and lower species diversity where dominant species are abundant. However, subordinate and dominant species frequently co‐occur. One mechanism that could facilitate resource sharing and co‐occurrence of dominant and subordinate competitors is fine‐scale resource dispersion. Here, we distributed 6 g of a food resource into 1, 2, 8, 32 or 64 units in small 0.40 m² areas centred on nests of the dominant ant Monomorium sydneyense. We tested three hypotheses. First, we hypothesized that the species richness and abundance of foraging ants would increase with increasing resource dispersion. Accordingly, species richness doubled and total ant abundance was two orders of magnitude higher in high resource dispersion treatments. Secondly, we hypothesized that increasing resource dispersion would reduce competitive interactions such as resource turnover events and lower the probability of food resources being occupied. Substantial support for this hypothesis was observed. Finally, we tested the hypothesis that the foraging time of each species would be proportional to the relative abundance of each species solely in high resource dispersion treatments. Expected and observed foraging times were statistically similar for only the dominant ant M. sydneyense. The subdominant Pheidole rugosula increased its foraging time much more than was expected, while two subordinate ants showed no relationship between observed and expected times. Thus, while increasing resource dispersion significantly increased overall species richness, this increase in co‐occurrence did not correlate with a significant increase in foraging time for the two subordinate species. Rather, changes in resource dispersion appeared to benefit only the subdominant species. Inter‐site variation appeared more important for other subordinate species indetermining co‐occurrence and foraging time. Multiple mechanisms facilitate co‐occurrence and resource sharing in this community, and probably in most other communities.     

Ecological traits influence the phylogenetic structure of bird species co‐occurrences worldwide

The extent to which species’ ecological and phylogenetic relatedness shape their co‐occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co‐occurrence patterns are linked – after accounting for regional effects – to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co‐occurrences of carnivorous, migratory and cold‐climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life‐history traits beyond the climatic niche. They further offer a novel species‐oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.     

Towards an understanding of the Holocene distribution of Fagus sylvatica L.

Aim Understanding the driving forces and mechanisms of changes in past plant distribution and abundance will help assess the biological consequences of future climate change scenarios. The aim of this paper is to investigate whether modelled patterns of climate parameters 6000 years ago can account for the European distribution of Fagus sylvatica at that time. Consideration is also given to the role of non‐climatic parameters as driving forces of the Holocene spread and population expansion of F. sylvatica. Location Europe. Methods European distributions were simulated using a physiologically‐based bioclimatic model (STASH) driven by three different atmospheric general circulation model (AGCM) outputs for 6000 years ago. Results The three simulations generally showed F. sylvatica to have potentially been as widespread 6000 years ago as it is today, which gives a profound mismatch with pollen‐based reconstructions of the F. sylvatica distribution at that time. The results indicate that drier conditions during the growing season 6000 years ago could have caused a restriction of the range in the south. Poorer growth conditions with consequently reduced competitive ability were modelled for large parts of France. Main conclusions Consideration of the entire European range of F. sylvatica showed that no single driving force could account for the observed distributional limits 6000 years ago, or the pattern of spread during the Holocene. Climatic factors, particularly drought during the growing season, are the likely major determinants of the potential range. Climatic factors are regionally moderated by competition, disturbance effects and the intrinsically slow rate of population increase of F. sylvatica. Dynamic vegetation modelling is needed to account for potentially important competitive interactions and their relationship with changing climate. We identify uncertainties in the climate and pollen data, as well as the bioclimatic model, which suggest that the current study does not identify whether or not climate determined the distribution of F. sylvatica 6000 years ago. Pollen data are better suited for comparison with relative abundance gradients rather than absolute distributional limits. These uncertainties from a study of the past, where we have information about plant distribution and abundance, argue for extreme caution in making forecasts for the future using equilibrium models.     

Local‐ to continental‐scale variation in the richness and composition of an aquatic food web

Aim We investigated patterns of species richness and composition of the aquatic food web found in the liquid‐filled leaves of the North American purple pitcher plant, Sarracenia purpurea (Sarraceniaceae), from local to continental scales. Location We sampled 20 pitcher‐plant communities at each of 39 sites spanning the geographic range of S. purpurea– from northern Florida to Newfoundland and westward to eastern British Columbia. Methods Environmental predictors of variation in species composition and species richness were measured at two different spatial scales: among pitchers within sites and among sites. Hierarchical Bayesian models were used to examine correlates and similarities of species richness and abundance within and among sites. Results Ninety‐two taxa of arthropods, protozoa and bacteria were identified in the 780 pitcher samples. The variation in the species composition of this multi‐trophic level community across the broad geographic range of the host plant was lower than the variation among pitchers within host‐plant populations. Variation among food webs in richness and composition was related to climate, pore‐water chemistry, pitcher‐plant morphology and leaf age. Variation in the abundance of the five most common invertebrates was also strongly related to pitcher morphology and site‐specific climatic and other environmental variables. Main conclusions The surprising result that these communities are more variable within their host‐plant populations than across North America suggests that the food web in S. purpurea leaves consists of two groups of species: (1) a core group of mostly obligate pitcher‐plant residents that have evolved strong requirements for the host plant and that co‐occur consistently across North America, and (2) a larger set of relatively uncommon, generalist taxa that co‐occur patchily.     

Examining the link between competition and negative co‐occurrence patterns

Negative species co‐occurrence patterns have long intrigued ecologists because of their potential link to competition. Although manipulative field experiments have consistently revealed evidence of competition in natural communities, there is little evidence that this competition produces negative co‐occurrence patterns. Evidence does suggest that abiotic variation, dispersal limitation and herbivory can contribute to patterns of negative co‐occurrence among species; it is possible these influences have obscured a link with competition. Here, we test for a connection between negative co‐occurrence and competition by examining a small‐scale, relatively homogeneous old‐field plant community where the influence of abiotic variation was likely to be minimal and we accounted for the impact of herbivory with an herbivore exclosure treatment. Using three years of data (two biennial periods), we tested whether negatively co‐occurring pairs of species, when occasionally found together, experienced asymmetric abundance decline more frequently than positively co‐occurring pairs, for which there is no such expectation. We found no evidence that negatively co‐occurring pairs consistently suffered asymmetric abundance decline more frequently than positively co‐occurring pairs, providing no evidence that competition is a primary driver of negative co‐occurrence patterns in this community. Our results were consistent across control and herbivore exclosure treatments, suggesting that herbivores are not driving patterns of negative species co‐occurrence in this community. Any influence of competition or herbivory on co‐occurrence patterns is small enough that it is obscured by other factors such as substrate heterogeneity, dispersal and differential species responses to climatic variation through time. We interpret our results as providing evidence that competition is not responsible for producing negative co‐occurrence patterns in our study community and suggest that this may be the case more broadly.     

Mutualism influences species distribution predictions for a bromeliad‐breeding anuran under climate change

Ecological niche models, or species distribution models, have been widely used to identify potentially suitable areas for species in future climate change scenarios. However, there are inherent errors to these models due to their inability to evaluate species occurrence influenced by non‐climatic factors. With the intuit to improve the modelling predictions for a bromeliad‐breeding treefrog (Phyllodytes melanomystax, Hylidae), we investigate how the climatic suitability of bromeliads influences the distribution model for the treefrog in the context of baseline and 2050 climate change scenarios. We used point occurrence data on the frog and the bromeliad (Vriesea procera, Bromeliaceae) to generate their predicted distributions based on baseline and 2050 climates. Using a consensus of five algorithms, we compared the accuracy of the models and the geographic predictions for the frog generated from two modelling procedures: (i) a climate‐only model for P. melanomystax and V. procera; and (ii) a climate‐biotic model for P. melanomystax, in which the climatic suitability of the bromeliad was jointly considered with the climatic variables. Both modelling approaches generated strong and similar predictive power for P. melanomystax, yet climate‐biotic modelling generated more concise predictions, particularly for the year 2050. Specifically, because the predicted area of the bromeliad overlaps with the predictions for the treefrog in the baseline climate, both modelling approaches produce reasonable similar predicted areas for the anuran. Alternatively, due to the predicted loss of northern climatically suitable areas for the bromeliad by 2050, only the climate‐biotic models provide evidence that northern populations of P. melanomystax will likely be negatively affected by 2050.     

Habitat distribution models for intertidal seaweeds: responses to climatic and non‐climatic drivers

Aim Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location North‐western Iberian Peninsula, southern Europe. Methods We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km² resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.     

Spatio‐temporal variation of biotic factors underpins contemporary range dynamics of congeners

Species’ ranges are complex often exhibiting multidirectional shifts over space and time. Despite the strong fingerprint of recent historical climate change on species’ distributions, biotic factors such as loss of vegetative habitat and the presence of potential competitors constitute important yet often overlooked drivers of range dynamics. Furthermore, short‐term changes in environmental conditions can influence the underlying processes of local extinction and local colonization that drive range shifts, yet are rarely considered at broad scales. We used dynamic state‐space occupancy models to test multiple hypotheses of the relative importance of major drivers of range shifts of Golden‐winged Warblers (Vermivora chrysoptera) and Blue‐winged Warblers (V. cyanoptera) between 1983 and 2012 across North America: warming temperatures; habitat changes; and occurrence of congeneric species, used here as proxy for biotic interactions. Dynamic occupancies for both species were most influenced by spatial relative to temporal variation in temperature and habitat. However, temporal variation in temperature anomalies and biotic interactions remained important. The two biotic factors considered, habitat change and biotic interactions, had the largest relative effect on estimated extinction rates followed by abiotic temperature anomalies. For the Golden‐winged Warbler, the predicted presence of the Blue‐winged Warbler, a hypothesized competitor, most influenced extinction probabilities, contributing to evidence supporting its role in site‐level species replacement. Given the overall importance of biotic factors on range‐wide dynamic occupancies, their consideration alongside abiotic factors should not be overlooked. Our results suggest that warming compounds the negative effect of habitat loss emphasizing species’ need for habitat to adapt to a changing climate. Notably, even closely related species exhibited individual responses to abiotic and biotic factors considered.     

Precipitation and winter temperature predict long‐term range‐scale abundance changes in Western North American birds

Predicting biodiversity responses to climate change remains a difficult challenge, especially in climatically complex regions where precipitation is a limiting factor. Though statistical climatic envelope models are frequently used to project future scenarios for species distributions under climate change, these models are rarely tested using empirical data. We used long‐term data on bird distributions and abundance covering five states in the western US and in the Canadian province of British Columbia to test the capacity of statistical models to predict temporal changes in bird populations over a 32‐year period. Using boosted regression trees, we built presence‐absence and abundance models that related the presence and abundance of 132 bird species to spatial variation in climatic conditions. Presence/absence models built using 1970–1974 data forecast the distributions of the majority of species in the later time period, 1998–2002 (mean AUC = 0.79 ± 0.01). Hindcast models performed equivalently (mean AUC = 0.82 ± 0.01). Correlations between observed and predicted abundances were also statistically significant for most species (forecast mean Spearman′s ρ = 0.34 ± 0.02, hindcast = 0.39 ± 0.02). The most stringent test is to test predicted changes in geographic patterns through time. Observed changes in abundance patterns were significantly positively correlated with those predicted for 59% of species (mean Spearman′s ρ = 0.28 ± 0.02, across all species). Three precipitation variables (for the wettest month, breeding season, and driest month) and minimum temperature of the coldest month were the most important predictors of bird distributions and abundances in this region, and hence of abundance changes through time. Our results suggest that models describing associations between climatic variables and abundance patterns can predict changes through time for some species, and that changes in precipitation and winter temperature appear to have already driven shifts in the geographic patterns of abundance of bird populations in western North America.     

Site‐adapted admixed tree species reduce drought susceptibility of mature European beech

Some forest‐related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long‐term performance, short‐term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co‐occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close‐by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.     

Changes in Patterns of Species Co‐occurrence across Two Tropical Cloud Forests Differing in Soil Nutrients and Air Temperature

Patterns of co‐occurrence of species are increasingly used to examine the contribution of biotic interactions to community assembly. We assessed patterns of co‐occurrence at four scales, in two types of tropical cloud forests in Hainan Island, China (tropical montane evergreen forests, TMEF and tropical dwarf forests, TDF) that varied significantly in soil nutrients and temperature. We tested if the patterns of co‐occurrence changed when we sorted species into classes by abundance and diameter at breast height (dbh). Co‐occurrence differed by forest type and with plot size, with significant species aggregation observed across larger plots in TDF and patterns of species segregation observed in smaller plots in TMEF. Analyses of differential abundance and dbh classes also showed that smaller plots in TMEF tend to have negative co‐occurrence patterns, but larger plots in TDF tend to show patterns of aggregation, suggesting competitive and facilitative interactions. This underscores the scale‐dependence of the processes contributing to community assembly. Furthermore, it is consistent with predictions of the stress gradient hypothesis that facilitation will be most important in biological systems subject to abiotic stress, while competition will be more important in less abiotically stressful habitats. Our results clearly demonstrate that these two types of tropical cloud forest exhibit different co‐occurrence patterns, and that these patterns are scale‐dependent, though independent of plant abundance and size class.     

Habitat Selection,the Included Niche,and Coexistence in Plant‐Specialist Frogs from Madagascar

Classic niche partitioning suggests that coexistence among asymmetric competitors is facilitated by differential resource use. Coexistence is also possible, however, when a species only has access to resources that are shared with a competitor, providing it is the superior competitor on that resource (the ‘included niche’). To test predictions of these two coexistence mechanisms, we studied habitat selection of two closely related sympatric plant‐specialist frogs from Madagascar (Guibemantis bicalcaratus and Guibemantis punctatus). Both species live and breed only in the water‐filled leaf axils of Pandanus plants, and previous experiments with their tadpoles demonstrated asymmetric competition. In a 3‐yr field study, we: (1) monitored the biotic and abiotic conditions of 348 plants; (2) surveyed these plants for frogs; (3) undertook a mark‐recapture study; and (4) conducted an experiment where we manipulated the amount of detritus in plants. We identified several differences in the conditions of the habitats selected by both species (e.g., plant height, canopy cover over the plant, abundance of heterospecifics). Co‐occurrence was nevertheless common, and G. punctatus persisted almost exclusively in plants it shared with G. bicalcaratus. As predicted by theory, G. punctatus was the superior competitor, at least in the tadpole stage. The inferior competitor (G. bicalcaratus) had access to exclusive resources not available to G. punctatus, by virtue of a faster developmental rate that permitted reproduction in lower quality plants. While there was some evidence of weak niche partitioning, we conclude that coexistence in these plant‐specialist frogs is primarily via an included niche mechanism. Abstract in French is available at http://www.blackwell‐synergy.com/loi/btp .     

Growing‐season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity

Stem xylem‐specific hydraulic conductivity (K_S) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in K_S has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global‐scale patterns of K_S and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species‐at‐site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how K_S varied with climatic variables, plant functional types, and biomes. Growing‐season temperature and growing‐season precipitation drove global variation in K_S independently. Both the mean and the variation in K_S were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for K_S in predicting shifts in community composition in the face of climate change.     

The evolution of climate tolerance in conifer‐feeding aphids in relation to their host's climatic niche

Climate adaptation has major consequences in the evolution and ecology of all living organisms. Though phytophagous insects are an important component of Earth's biodiversity, there are few studies investigating the evolution of their climatic preferences. This lack of research is probably because their evolutionary ecology is thought to be primarily driven by their interactions with their host plants. Here, we use a robust phylogenetic framework and species‐level distribution data for the conifer‐feeding aphid genus Cinara to investigate the role of climatic adaptation in the diversity and distribution patterns of these host‐specialized insects. Insect climate niches were reconstructed at a macroevolutionary scale, highlighting that climate niche tolerance is evolutionarily labile, with closely related species exhibiting strong climatic disparities. This result may suggest repeated climate niche differentiation during the evolutionary diversification of Cinara. Alternatively, it may merely reflect the use of host plants that occur in disparate climatic zones, and thus, in reality the aphid species' fundamental climate niches may actually be similar but broad. Comparisons of the aphids' current climate niches with those of their hosts show that most Cinara species occupy the full range of the climatic tolerance exhibited by their set of host plants, corroborating the hypothesis that the observed disparity in Cinara species' climate niches can simply mirror that of their hosts. However, 29% of the studied species only occupy a subset of their hosts' climatic zone, suggesting that some aphid species do indeed have their own climatic limitations. Our results suggest that in host‐specialized phytophagous insects, host associations cannot always adequately describe insect niches and abiotic factors must be taken into account.     

Division within the North American boreal forest: Ecological niche divergence between the Bicknell's Thrush (Catharus bicknelli) and Gray‐cheeked Thrush (C. minimus)

Sister species that diverged in allopatry in similar environments are expected to exhibit niche conservatism. Using ecological niche modeling and a multivariate analysis of climate and habitat data, I test the hypothesis that the Bicknell's Thrush (Catharus bicknelli) and Gray‐cheeked Thrush (C. mimimus), sister species that breed in the North American boreal forest, show niche conservatism. Three tree species that are important components of breeding territories of both thrush species were combined with climatic variables to create niche models consisting of abiotic and biotic components. Abiotic‐only, abiotic+biotic, and biotic‐only models were evaluated using the area under the curve (AUC) criterion. Abiotic+biotic models had higher AUC scores and did not over‐project thrush distributions compared to abiotic‐only or biotic‐only models. From the abiotic+biotic models, I tested for niche conservatism or divergence by accounting for the differences in the availability of niche components by calculating (1) niche overlap from ecological niche models and (2) mean niche differences of environmental values at occurrence points. Niche background similarity tests revealed significant niche divergence in 10 of 12 comparisons, and multivariate tests revealed niche divergence along 2 of 3 niche axes. The Bicknell's Thrush breeds in warmer and wetter regions with a high abundance of balsam fir (Abies balsamea), whereas Gray‐cheeked Thrush often co‐occurs with black spruce (Picea mariana). Niche divergence, rather than conservatism, was the predominant pattern for these species, suggesting that ecological divergence has played a role in the speciation of the Bicknell's Thrush and Gray‐cheeked Thrush. Furthermore, because niche models were improved by the incorporation of biotic variables, this study validates the inclusion of relevant biotic factors in ecological niche modeling to increase model accuracy.     

Responses of foliage‐living spider assemblage composition and traits to a climatic gradient in Themeda grasslands

For speciose, but poorly known groups, such as terrestrial arthropods, functional traits present a potential avenue to assist in predicting responses to environmental change. Species turnover is common along environmental gradients, but it is unclear how this is reflected in species traits. Community‐level change in arthropod traits, other than body size, has rarely been explored across spatial scales comparable to those examined here. We hypothesized that the composition and morphological traits of spider assemblages would differ across a gradient of climate and habitat structure. We examined foliage‐living spider assemblages associated with Themeda triandra grasslands along a 900 km climatic gradient in south‐eastern Australia. We used sweep‐netting to collect T. triandra‐associated spiders and counted juveniles and identified adults. We also measured morphological traits of adult spiders and noted their hunting mode. Associations with measures of habitat structure were less consistent than relationships with climate. Both juvenile and adult spiders were more abundant in warmer sites, although species richness was not affected by temperature. We found distinct turnover in species composition along the climatic gradient, with hunting spiders, particularly crab spiders (Thomisidae), making up a greater proportion of assemblages in warmer climates. A range of traits of spiders correlated with the climatic gradient. For example, larger spider species and species that were active hunters were more common in warmer climates. Changes in morphological traits across species, rather than within species drove the morphology‐climate relationship. Strong climate‐trait correlations suggest that it may be possible to predict changes in functional traits of assemblages in response to anthropogenic disturbances such as climate change.     

Historical and event‐based bioclimatic suitability predicts regional forest vulnerability to compound effects of severe drought and bark beetle infestation

Vulnerability to climate change, and particularly to climate extreme events, is expected to vary across species ranges. Thus, we need tools to standardize the variability in regional climatic legacy and extreme climate across populations and species. Extreme climate events (e.g., droughts) can erode populations close to the limits of species' climatic tolerance. Populations in climatic‐core locations may also become vulnerable because they have developed a greater demand for resources (i.e., water) that cannot be enough satisfied during the periods of scarcity. These mechanisms can become exacerbated in tree populations when combined with antagonistic biotic interactions, such as insect infestation. We used climatic suitability indices derived from Species Distribution Models (SDMs) to standardize the climatic conditions experienced across Pinus edulis populations in southwestern North America, during a historical period (1972–2000) and during an extreme event (2001–2007), when the compound effect of hot drought and bark beetle infestation caused widespread die‐off and mortality. Pinus edulis climatic suitability diminished dramatically during the die‐off period, with remarkable variation between years. P. edulis die‐off occurred mainly not just in sites that experienced lower climatic suitability during the drought but also where climatic suitability was higher during the historical period. The combined effect of historically high climatic suitability and a marked decrease in the climatic suitability during the drought best explained the range‐wide mortality. Lagged effects of climatic suitability loss in previous years and co‐occurrence of Juniperus monosperma also explained P. edulis die‐off in particular years. Overall, the study shows that past climatic legacy, likely determining acclimation, together with competitive interactions plays a major role in responses to extreme drought. It also provides a new approach to standardize the magnitude of climatic variability across populations using SDMs, improving our capacity to predict population's or species' vulnerability to climatic change.     

Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests

Tree populations at the low‐altitudinal or ‐latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree‐ring widths and δ¹³C and δ¹⁸O chronologies to compare the growth rates and long‐term ecophysiological responses to climate in the temperate‐Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO₂. Tree‐ring δ¹⁸O for both species were mostly correlated with δ¹⁸O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree‐ring δ¹⁸O but had a positive effect on Q. ilex tree‐ring δ¹⁸O. Furthermore, tree‐ring δ¹⁸O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO₂ fertilization, in contrast to the water‐limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.