Sophia Ratcliffe

Tree diversity does not always improve resistance of forest ecosystems to drought

Significance In the context of climate change, expected drier and warmer environmental conditions will have drastic consequences on forest functions and services and may bring about important drought-induced die-off events. Biodiversity promotes forest ecosystem performance and resistance to insect pests and diseases, but whether or not diverse forests are also better adapted to deal with drought stress remains unknown. Within our study network of 160 forest stands across Europe, we found that mixed species forests are less exposed to drought stress in some regions only. Therefore, managing forest ecosystems for high tree species diversity does not necessarily assure improved resistance to the more severe and frequent drought events predicted for the future. Climate models predict an increase in the intensity and frequency of drought episodes in the Northern Hemisphere. Among terrestrial ecosystems, forests will be profoundly impacted by drier climatic conditions, with drastic consequences for the functions and services they supply. Simultaneously, biodiversity is known to support a wide range of forest ecosystem functions and services. However, whether biodiversity also improves the resistance of these ecosystems to drought remains unclear. We compared soil drought exposure levels in a total of 160 forest stands within five major forest types across Europe along a gradient of tree species diversity. We assessed soil drought exposure in each forest stand by calculating the stand-level increase in carbon isotope composition of late wood from a wet to a dry year (Δδ13CS). Δδ13CS exhibited a negative linear relationship with tree species diversity in two forest types, suggesting that species interactions in these forests diminished the drought exposure of the ecosystem. However, the other three forest types were unaffected by tree species diversity. We conclude that higher diversity enhances resistance to drought events only in drought-prone environments. Managing forest ecosystems for high tree species diversity does not necessarily assure improved adaptability to the more severe and frequent drought events predicted for the future.

Biotic homogenization can decrease landscape-scale forest multifunctionality

Significance Numerous studies have demonstrated the importance of biodiversity in maintaining multiple ecosystem functions and services (multifunctionality) at local spatial scales, but it is unknown whether similar relationships are found at larger spatial scales in real-world landscapes. Here, we show, for the first time to our knowledge, that biodiversity can also be important for multifunctionality at larger spatial scales in European forest landscapes. Both high local (α-) diversity and a high turnover in species composition between locations (high β-diversity) were found to be potentially important drivers of ecosystem multifunctionality. Our study provides evidence that it is important to conserve the landscape-scale biodiversity that is being eroded by biotic homogenization if ecosystem multifunctionality is to be maintained. Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

On the mechanisms of coexistence among annual-plant species, using neighbourhood techniques and simulation models

Many studies have investigated the density-dependent regulation ofannual-plant populations on coastal sand dunes, but few have explored theconsequences of competition for the coexistence of plants in these simplecommunities. We used neighbourhood techniques to parameterize competition anddispersal functions from field data collected for two species of dune annual(Aira praecox and Erodium cicutarium)over three successive years, and then combined these functions into spatiallyexplicit simulation models. The population size of Airavaried enormously between years, while Erodium remainedsteady. Competition with neighbours reduced the spike length ofAira plants only in one of the three years (when itspopulation density was highest), while competition with neighbouringErodium plants appeared to result in the local death ofAira plants. However, these density-dependent effects werefar too weak to generate the observed changes in the population size ofAira among years, or to maintain populations below theupper limits observed. The large-seeded Erodium wasaffected by intraspecific competition but was unaffected by small-seededAira plants. Therefore, the larger-seeded species wascompetitively superior to the smaller-seeded species, an affect that couldpromote coexistence (albeit weakly) by a competition-colonisation trade-off.Modal dispersion distances of Aira andErodium were 45 and 60 mm respectively,greater than the radius within which competitive interactions occurred (40mm). Theoretical studies suggest that under these conditions thespatial arrangement of plants should be nearly random. In factAira was spatially aggregated, especially when rare,suggesting that patchy mortality across the dunes was important in generatingspatial structure. The study suggests that density dependence only weaklyregulates dune annual communities, while year-to-year environmental variationexert major influences on population sizes and spatial structures.

Relationship between dew presence and Bassia dasyphylla plant growth

Dew has been recognized for its ecological significance and has also been identified as an additional source of water in arid zones. We used factorial control experiment, under dew presence in the field, to explore photosynthetic performance, water status and growth response of desert annual herbage. Bassia dasyphylla seedlings were grown in contrasting dew treatments (dew-absent and dew-present) and different watering regimes (normal and deficient). The effects of dew on the water status and photosynthetic performance of Bassia dasyphylla, grown in a desert area of the Hexi Corridor in Northwestern China, were evaluated. The results indicated the presence of dew significantly increased relative water content (RWC) of shoots and total biomass of plants in both water regimes, and enhanced the diurnal shoot water potential and stomatal conductance in the early morning, as well as photosynthetic rate, which reached its maximum only in the water-stressed regime. The presence of dew increased aboveground growth of plants and photosynthate accumulation in leaves, but decreased the root-to-shoot ratio in both water regimes. Dew may have an important role in improving plant water status and ameliorating the adverse effects of plants exposed to prolonged drought.

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.