• Energy Research

AbstractEcologists have limited understanding of how geographic variation in forest biomass arises from differences in growth and mortality at continental to global scales. Using forest inventories from across North America, we partitioned continental‐scale variation in biomass growth and mortality rates of 49 tree species groups into (1) species‐independent spatial effects and (2) inherent differences in demographic performance among species. Spatial factors that were separable from species composition explained 83% and 51% of the respective variation in growth and mortality. Moderate additional variation in mortality (26%) was attributable to differences in species composition. Age‐dependent biomass models showed that variation in forest biomass can be explained primarily by spatial gradients in growth that were unrelated to species composition. Species‐dependent patterns of mortality explained additional variation in biomass, with forests supporting less biomass when dominated by species that are highly susceptible to competition (e.g. Populus spp.) or to biotic disturbances (e.g. Abies balsamea).

AbstractClimate change effects on tree reproduction are poorly understood, even though the resilience of populations relies on sufficient regeneration to balance increasing rates of mortality. Forest‐forming tree species often mast, i.e. reproduce through synchronised year‐to‐year variation in seed production, which improves pollination and reduces seed predation. Recent observations in European beech show, however, that current climate change can dampen interannual variation and synchrony of seed production and that this masting breakdown drastically reduces the viability of seed crops. Importantly, it is unclear under which conditions masting breakdown occurs and how widespread breakdown is in this pan‐European species. Here, we analysed 50 long‐term datasets of population‐level seed production, sampled across the distribution of European beech, and identified increasing summer temperatures as the general driver of masting breakdown. Specifically, increases in site‐specific mean maximum temperatures during June and July were observed across most of the species range, while the interannual variability of population‐level seed production (CVp) decreased. The declines in CVp were greatest, where temperatures increased most rapidly. Additionally, the occurrence of crop failures and low seed years has decreased during the last four decades, signalling altered starvation effects of masting on seed predators. Notably, CVp did not vary among sites according to site mean summer temperature. Instead, masting breakdown occurs in response to warming local temperatures (i.e. increasing relative temperatures), such that the risk is not restricted to populations growing in warm average conditions. As lowered CVp can reduce viable seed production despite the overall increase in seed count, our results warn that a covert mechanism is underway that may hinder the regeneration potential of European beech under climate change, with great potential to alter forest functioning and community dynamics.

Abstract Forest stand densities are increasing in the boreal and temperate biomes, suggesting that tree‐tree competition is intensifying. Anticipating the consequences of this intensified competition is difficult because competition‐induced mortality may depend not only on the occurrence of extreme climatic events such as drought, but also on stand composition, since tree species differ in their ability to compete and tolerate competition. A better understanding of the effects of stand composition and drought on competition‐induced mortality would help to anticipate future changes in forest ecosystems. We studied the tree‐level probability of competition‐induced mortality using National Forest Inventory data from three European countries (Finland, France and Germany), covering a latitudinal gradient from the Mediterranean to the Arctic. We investigated how (i) the proportion of conspecifics, (ii) the shade tolerance (ST) of the focal tree and its competitors and (iii) drought events modify the effect of competition on tree mortality. We used a generalized linear mixed model on a dataset of 461,109 trees representing 39 species on 48,088 individual plots. Competition, measured as the basal area of larger trees, was a stronger driver of background mortality (BM) than tree size and climate. A higher proportion of conspecifics increased the competition effect on mortality, showing that conspecific individuals had a higher competitive effect compared to heterospecific individuals. The competition effect on mortality also increased as a function of the ST of neighbouring trees, suggesting an increased shading effect. A higher ST of a focal tree decreased the competition effect on mortality. Drought anomalies increased the competition effect, resulting in a higher mortality probability for the most suppressed trees. Synthesis. Competition was the main driver of background mortality. Increasing stand density increased competition‐induced tree mortality in both monospecific and mixed stands, but to different extents depending on the proportion of conspecifics and tree species shade tolerance (ST). Drought periods increase mortality, especially among the most suppressed trees, suggesting an interaction between competitive status and drought. Incorporating more detailed information on stand composition and tree species ST into tree mortality models will improve our understanding of forest dynamics in a changing climate.

Tree species composition is known to influence forest productivity, but its effect on forest resilience to disturbances such as storms remains largely unexplored. Furthermore, climate is likely to influence forest resilience directly but also to influence the effect of tree species composition on resilience. In Europe, storm-induced tree mortality is currently increasing across all climatic biomes. Understanding the drivers of forest resilience to storms and its consistency across climates appears to be crucial for predicting the consequences of climate change for European forests.In this study, we used a simulation approach with an integral projection model calibrated with National Forest Inventory (NFI) data at the European scale. We restricted our simulations to tree species assemblages observed in the NFI data, covering a species diversity gradient nested within a climate gradient. We quantified functional diversity and the mean position of each species assemblage at equilibrium on two functional axis: (i) conservative versus fast growing and (ii) low versus high recruitment. We disturbed each species assemblage from equilibrium using species-specific storm disturbance mortality probabilities and quantified the assemblages' resistance (inverse of immediate basal area loss), recovery (slope of post-disturbance increase in basal area) and resilience (inverse of the cumulative deviation of basal area from the undisturbed state).We found that on average, species-rich assemblages had higher recovery and resilience to storm disturbance, while functional diversity improved resistance and recovery. When analysing how this effect varied with climate, we found that diversity significantly increased resistance and resilience in the climatic margins only. Finally, we found that storm resilience was also driven by species mean position along both functional axes. In particular, the conservative-productive axis had an effect two to three times greater than diversity: forests dominated by conservative species were more resistant and resilient, but had lower recovery than species assemblages dominated by fast-growing species.Taken together, these results show that climate and tree species composition interact to control the ability of forests to resist and recover from a storm disturbance through both direct and indirect effects. As such, our findings should help to better anticipate climate change consequences for forest ecosystems. International audience

Recent climate warming has fueled interest into climate‐driven range shifts of tree species. A common approach to detect range shifts is to compare the divergent occurrences between juvenile and adult trees along environmental gradients using static data. Divergent occurrences between life stages can, however, also be caused by ontogenetic effects. These include shifts of the viable environmental conditions throughout development (‘ontogenetic niche shift') as well as demographic dependencies that constrain the possible occurrence of subsequent life stages. Whether ontogenetic effects are an important driver of divergent occurrences between juvenile and adult trees along large‐scale climatic gradients is largely unknown. It is, however, critical in evaluating whether impacts of environmental change can be inferred from static data on life stage occurrences. Here, we first show theoretically, using a two‐life stage simulation model, how both temporal range shift and ontogenetic effects can lead to similar divergent occurrences between adults and juveniles (juvenile divergence). We further demonstrate that juvenile divergence can unambiguously be attributed to ontogenetic effects, when juveniles diverge from adults in opposite direction to their temporal shift along the environmental gradient. Second, to empirically test whether ontogenetic effects are an important driver of divergent occurrences across Europe, we use repeated national forest inventories from Sweden, Germany and Spain to assess juvenile divergence and temporal shift for 40 tree species along large‐scale climatic gradients. About half of the species‐country combinations had significant juvenile divergences along heat sum and water availability gradients. Only a quarter of the tree species had significant detectable temporal shifts within the observation period. Furthermore, significant juvenile divergences were frequently associated with opposite temporal shifts, indicating that ontogenetic effects are a relevant cause of divergent occurrences between life stages. Our study furthers the understanding of ontogenetic effects and challenges the practice of inferring climate change impacts from static data.

AbstractIndirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.