Relative decline in density of Northern Hemisphere tree species in warm and arid regions of their climate niches.

Although climate change is expected to drive tree species toward colder and wetter regions of their distribution, broadscale empirical evidence is lacking. One possibility is that past and present human activities in forests obscure or alter the effects of climate. Here, using data from more than two million monitored trees from 73 widely distributed species, we quantify changes in tree species density within their climatic niches across Northern Hemisphere forests. We observe a reduction in mean density across species, coupled with a tendency toward increasing tree size. However, the direction and magnitude of changes in density exhibit considerable variability between species, influenced by stand development that results from previous stand-level disturbances. Remarkably, when accounting for stand development, our findings show a significant change in density toward cold and wet climatic conditions for 43% of the species, compared to only 14% of species significantly changing their density toward warm and arid conditions in both early- and late-development stands. The observed changes in climate-driven density showed no clear association with species traits related to drought tolerance, recruitment and dispersal capacity, or resource use, nor with the temperature or aridity affiliation of the species, leaving the underlying mechanism uncertain. Forest conservation policies and associated management strategies might want to consider anticipated long-term species range shifts alongside the integration of contemporary within-distribution density changes.

Accounting for trait variability and coordination in predictions of drought-induced range shifts in woody plants.

Functional traits offer a promising avenue to improve predictions of species range shifts under climate change, which will entail warmer and often drier conditions. Although the conceptual foundation linking traits with plant performance and range shifts appears solid, the predictive ability of individual traits remains generally low. In this review, we address this apparent paradox, emphasizing examples of woody plants and traits associated with drought responses at the species' rear edge. Low predictive ability reflects the fact not only that range dynamics tend to be complex and multifactorial, as well as uncertainty in the identification of relevant traits and limited data availability, but also that trait effects are scale- and context-dependent. The latter results from the complex interactions among traits (e.g. compensatory effects) and between them and the environment (e.g. exposure), which ultimately determine persistence and colonization capacity. To confront this complexity, a more balanced coverage of the main functional dimensions involved (stress tolerance, resource use, regeneration and dispersal) is needed, and modelling approaches must be developed that explicitly account for: trait coordination in a hierarchical context; trait variability in space and time and its relationship with exposure; and the effect of biotic interactions in an ecological community context.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Greater growth stability of trees in marginal habitats suggests a patchy pattern of population loss and retention in response to increased drought at the rear edge.

Species rear range edges are predicted to retract as climate warms, yet evidence of population persistence is accumulating. Accounting for this disparity is essential to enable prediction and planning for species' range retractions. At the Mediterranean edge of European beech-dominated temperate forest, we tested the hypothesis that individual performance should decline at the limit of the species' ecological tolerance in response to increased drought. We sampled 40 populations in a crossed factor design of geographical and ecological marginality and assessed tree growth resilience and decline in response to recent drought. Drought impacts occurred across the rear edge, but tree growth stability was unexpectedly high in geographically isolated marginal habitat and lower than anticipated in the species' continuous range and better-quality habitat. Our findings demonstrate that, at the rear edge, range shifts will be highly uneven and characterised by reduction in population density with local population retention rather than abrupt range retractions.

Refining predictions of population decline at species' rear edges.

According to broad-scale application of biogeographical theory, widespread retractions of species' rear edges should be seen in response to ongoing climate change. This prediction rests on the assumption that rear edge populations are "marginal" since they occur at the limit of the species' ecological tolerance and are expected to decline in performance as climate warming pushes them to extirpation. However, conflicts between observations and predictions are increasingly accumulating and little progress has been made in explaining this disparity. We argue that a revision of the concept of marginality is necessary, together with explicit testing of population decline, which is increasingly possible as data availability improves. Such action should be based on taking the population perspective across a species' rear edge, encompassing the ecological, geographical and genetic dimensions of marginality. Refining our understanding of rear edge populations is essential to advance our ability to monitor, predict and plan for the impacts of environmental change on species range dynamics.

Climate- and successional-related changes in functional composition of European forests are strongly driven by tree mortality.

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late-successional short-statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm-like strategies at low water availability in forests formerly dominated by broad-leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm-like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.

Anthropogenic land-use legacies underpin climate change-related risks to forest ecosystems.

Forest ecosystems with long-lasting human imprints can emerge worldwide as outcomes of land-use cessation. However, the interaction of these anthropogenic legacies with climate change impacts on forests is not well understood. Here, we set out how anthropogenic land-use legacies that persist in forest properties, following alterations in forest distribution, structure, and composition, can interact with climate change stressors. We propose a risk-based framework to identify anthropogenic legacies of land uses in forest ecosystems and quantify the impact of their interaction with climate-related stress on forest responses. Considering anthropogenic land-use legacies alongside environmental drivers of forest ecosystem dynamics will improve our predictive capacity of climate-related risks to forests and our ability to promote ecosystem resilience to climate change.

Structural and climatic determinants of demographic rates of Scots pine forests across the Iberian Peninsula.

The demographic rates of tree species typically show large spatial variation across their range. Understanding the environmental factors underlying this variation is a key topic in forest ecology, with far-reaching management implications. Scots pine (Pinus sylvestris L.) covers large areas of the Northern Hemisphere, the Iberian Peninsula being its southwestern distribution limit. In recent decades, an increase in severe droughts and a densification of forests as a result of changes in forest uses have occurred in this region. Our aim was to use climate and stand structure data to explain mortality and growth patterns of Scots pine forests across the Iberian Peninsula. We used data from 2392 plots dominated by Scots pine, sampled for the National Forest Inventory of Spain. Plots were sampled from 1986 to 1996 (IFN2) and were resampled from 1997 to 2007 (IFN3), allowing for the calculation of growth and mortality rates. We fitted linear models to assess the response of growth and mortality rates to the spatial variability of climate, climatic anomalies, and forest structure. Over the period of approximately 10 years between the IFN2 and IFN3, the amount of standing dead trees increased 11-fold. Higher mortality rates were related to dryness, and growth was reduced with increasing dryness and temperature, but results also suggested that effects of climatic stressors were not restricted to dry sites only. Forest structure was strongly related to demographic rates, suggesting that stand development and competition are the main factors associated with demography. In the case of mortality, forest structure interacted with climate, suggesting that competition for water resources induces tree mortality in dry sites. A slight negative relationship was found between mortality and growth, indicating that both rates are likely to be affected by the same stress factors. Additionally, regeneration tended to be lower in plots with higher mortality. Taken together, our results suggest a large-scale self-thinning related to the recent densification of Scots pine forests. This process appears to be enhanced by dry conditions and may lead to a mismatch in forest turnover. Forest management may be an essential adaptive tool under the drier conditions predicted by most climate models.