Paloma Ruiz-Benito

Plant functional traits have globally consistent effects on competition

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits—wood density, specific leaf area and maximum height—consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

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.

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

Abstract 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.

Biodiversity and ecosystem functioning relations in European forests depend on environmental context

The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests.

Stand Structure and Recent Climate Change Constrain Stand Basal Area Change in European Forests: A Comparison Across Boreal, Temperate, and Mediterranean Biomes

European forests have a prominent role in the global carbon cycle and an increase in carbon storage has been consistently reported during the twentieth century. Any further increase in forest carbon storage, however, could be hampered by increases in aridity and extreme climatic events. Here, we use forest inventory data to identify the relative importance of stand structure (stand basal area and mean d.b.h.), mean climate (water availability), and recent climate change (temperature and precipitation anomalies) on forest basal area change during the late twentieth century in three major European biomes. Using linear mixed-effects models we observed that stand structure, mean climate, and recent climatic change strongly interact to modulate basal area change. Although we observed a net increment in stand basal area during the late twentieth century, we found the highest basal area increments in forests with medium stand basal areas and small to medium-sized trees. Stand basal area increases correlated positively with water availability and were enhanced in warmer areas. Recent climatic warming caused an increase in stand basal area, but this increase was offset by water availability. Based on recent trends in basal area change, we conclude that the potential rate of aboveground carbon accumulation in European forests strongly depends on both stand structure and concomitant climate warming, adding weight to suggestions that European carbon stocks may saturate in the near future.

Functional diversity underlies demographic responses to environmental variation in European forests

Aim Biodiversity loss and climate-driven ecosystem modification are leading to substantial changes in forest structure and function. However, the effects of diversity on demographic responses to the environment are poorly understood. We tested the diversity hypothesis (measured through functional diversity) and the mass ratio hypothesis (measured through functional identity) in relation to tree growth, tree mortality and sapling abundance. We sought to determine whether functional diversity underlies demographic responses to environmental variation in European forests. Location Europe (Spain, Germany, Wallonia, Finland and Sweden). Methods We used data from five European national forest inventories from boreal to Mediterranean biomes (c. 700,000 trees in 54,000 plots and 143 tree species) and the main forest types across Europe (i.e. from needle-leaved evergreen forests to broad-leaved deciduous forests). For each forest type, we applied maximum likelihood techniques to quantify the relative importance of stand structure, climate and diversity (i.e. functional diversity and functional identity) as determinants of growth, mortality and sapling abundance. We also tested whether demographic responses to environmental conditions (including stand density, evapotranspiration and temperature anomalies) varied with functional diversity. Results Our results suggest that functional diversity has a positive effect on sapling abundance and growth rates in forests across Europe, while no effect was observed on tree mortality. Functional identity has a strong effect on mortality and sapling abundance, with greater mortality rates in forests dominated by needle-leaved individuals and a greater abundance of saplings in forests dominated by broad-leaved individuals. Furthermore, we observed that functional diversity modified the effects of stand density on demographic responses in Mediterranean forests and the influence of evapotranspiration and temperature anomalies in forests widely distributed across Europe. Main conclusion Our results suggest that functional diversity may play a key role in forest dynamics through complementarity mechanisms, as well as by modulating demographic responses to environmental variation.

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.

Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe

Neglecting tree size and stand structure dynamics might bias the interpretation of the diversity-productivity relationship in forests. Here we show evidence that complementarity is contingent on tree size across large-scale climatic gradients in Europe. We compiled growth data of the 14 most dominant tree species in 32,628 permanent plots covering boreal, temperate and Mediterranean forest biomes. Niche complementarity is expected to result in significant growth increments of trees surrounded by a larger proportion of functionally dissimilar neighbours. Functional dissimilarity at the tree level was assessed using four functional types: i.e. broad-leaved deciduous, broad-leaved evergreen, needle-leaved deciduous and needle-leaved evergreen. Using Linear Mixed Models we show that, complementarity effects depend on tree size along an energy availability gradient across Europe. Specifically: (i) complementarity effects at low and intermediate positions of the gradient (coldest-temperate areas) were stronger for small than for large trees; (ii) in contrast, at the upper end of the gradient (warmer regions), complementarity is more widespread in larger than smaller trees, which in turn showed negative growth responses to increased functional dissimilarity. Our findings suggest that the outcome of species mixing on stand productivity might critically depend on individual size distribution structure along gradients of environmental variation.

Functional diversity underlies demographic responses to environmental variation in European forests (Forthcoming/Available Online)

Aim Biodiversity loss and climate-driven ecosystem modification are leading to substantial changes in forest structure and function. However, the effects of diversity on demographic responses to the environment are poorly understood. We tested the diversity hypothesis (measured through functional diversity) and the mass ratio hypothesis (measured through functional identity) in relation to tree growth, tree mortality and sapling abundance. We sought to determine whether functional diversity underlies demographic responses to environmental variation in European forests. Location Europe (Spain, Germany, Wallonia, Finland and Sweden). Methods We used data from five European national forest inventories from boreal to Mediterranean biomes (c. 700,000 trees in 54,000 plots and 143 tree species) and the main forest types across Europe (i.e. from needle-leaved evergreen forests to broad-leaved deciduous forests). For each forest type, we applied maximum likelihood techniques to quantify the relative importance of stand structure, climate and diversity (i.e. functional diversity and functional identity) as determinants of growth, mortality and sapling abundance. We also tested whether demographic responses to environmental conditions (including stand density, evapotranspiration and temperature anomalies) varied with functional diversity. Results Our results suggest that functional diversity has a positive effect on sapling abundance and growth rates in forests across Europe, while no effect was observed on tree mortality. Functional identity has a strong effect on mortality and sapling abundance, with greater mortality rates in forests dominated by needle-leaved individuals and a greater abundance of saplings in forests dominated by broad-leaved individuals. Furthermore, we observed that functional diversity modified the effects of stand density on demographic responses in Mediterranean forests and the influence of evapotranspiration and temperature anomalies in forests widely distributed across Europe. Main conclusion Our results suggest that functional diversity may play a key role in forest dynamics through complementarity mechanisms, as well as by modulating demographic responses to environmental variation.

Forest productivity in southwestern Europe is controlled by coupled North Atlantic and Atlantic Multidecadal Oscillations

The North Atlantic Oscillation (NAO) depicts annual and decadal oscillatory modes of variability responsible for dry spells over the European continent. The NAO therefore holds a great potential to evaluate the role, as carbon sinks, of water-limited forests under climate change. However, uncertainties related to inconsistent responses of long-term forest productivity to NAO have so far hampered firm conclusions on its impacts. We hypothesize that, in part, such inconsistencies might have their origin in periodical sea surface temperature anomalies in the Atlantic Ocean (i.e., Atlantic Multidecadal Oscillation, AMO). Here we show strong empirical evidence in support of this hypothesis using 120 years of periodical inventory data from Iberian pine forests. Our results point to AMO+ NAO+ and AMO−NAO− phases as being critical for forest productivity, likely due to decreased winter water balance and abnormally low winter temperatures, respectively. Our findings could be essential for the evaluation of ecosystem functioning vulnerabilities associated with increased climatic anomalies under unprecedented warming conditions in the Mediterranean.The North Atlantic Oscillation (NAO) drives biological responses in terrestrial ecosystems through oscillatory modes of climatic variability. Here, the authors show how landscape scale productivity responses to NAO are contingent upon the Atlantic Multidecadal Oscillation in southwestern Europe.