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Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.

Significance Although forest succession has been approached as a predictable process, successional trajectories vary widely, even among nearby stands with similar environmental conditions and disturbance histories. We quantified predictability and uncertainty during tropical forest succession using dynamical models describing the interactions among stem density, basal area, and species density over time. We showed that the trajectories of these forest attributes were poorly predicted by stand age and varied significantly within and among sites. Our models reproduced the general successional trends observed, but high levels of noise were needed to increase model predictability. These levels of uncertainty call into question the premise that successional processes are consistent over space and time, and challenge the way ecologists view tropical forest regeneration. Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributes—stem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.

Some disturbances can drive ecological systems to abrupt shifts between alternative stages (tipping points) when critical transitions occur. Drought‐induced tree death can be considered as a nonlinear shift in tree vigour and growth. However, at what point do trees become predisposed to drought‐related dieback and which factors determine this (tipping) point? We investigated these questions by characterizing the responses of three tree species, silver fir (Abies alba), Scots pine (Pinus sylvestris) and Aleppo pine (Pinus halepensis), to a severe drought event. We compared basal area increment (BAI) trends and responses to climate and drought in declining (very defoliated and dying) vs. non‐declining (slightly or not defoliated) trees by using generalized additive mixed models. Defoliation, BAI and sapwood production were related to functional proxies of tree vigour measured at the onset and end of the drought (non‐structural carbohydrate concentrations, needle N content and C isotopic discrimination, presence of wood‐inhabiting fungi). We evaluated whether early warning signals (increases in synchronicity among trees or in autocorrelation and standard deviation) could be extracted from the BAI series prior to tree death. Declining silver fir and Scots pine trees showed less growth than non‐declining trees one to three decades, respectively, before the drought event, whereas Aleppo pines showed growth decline irrespective of tree defoliation. At the end of the drought period, all species showed increased defoliation and a related reduction in the concentration of sapwood soluble sugars. Defoliation was constrained by the BAI of the previous 5 years and sapwood production. No specific wood‐inhabiting fungi were found in post‐drought declining trees apart from blue‐stain fungi, which extensively affected damaged Scots pines. Declining silver firs showed increases in BAI autocorrelation and variability prior to tree death. Synthesis. Early warning signals of drought‐triggered mortality seem to be species specific and reflect how different tree species cope with drought stress. Highly correlated declining growth patterns during drought can serve as a signal in silver fir, whereas changes in the content of sapwood soluble sugars are suitable vigour proxies for Scots and Aleppo pines. Longer growth and defoliation series, additional vigour parameters and multi‐species comparisons are required to understand and predict drought‐induced tree death.

Aim Climate change is expected to modify growth trends of forests around the world. However, this modification may vary in strength and intensity across a species' biogeographical range. Here, we study European populations of silver fir (Abies alba) across its southern distribution limits in Spain, Italy and Romania. We hypothesized that growth trends of silver fir will differ across its distribution range, with a marked decline in growth in drought-prone regions near the species' southernmost biogeographical limits. Location Europe (Spain, Italy, Romania). Methods We collected tree-ring data from at least 1300 silver fir trees located in 111 sites. The dataset was used to assess and model growth trends, quantified as changes in basal area increment, and to determine how growth responds to climate. Results We found contrasting patterns of basal area increments among countries and sites. Populations of silver fir located outside the Mediterranean area (e.g. northern Italy, Romania) have shown a clear increase in growth over the last two decades, whereas most populations in Spain and southern Italy have displayed a marked decline in growth since the 1980s. The growth of silver fir forests at the south-western distribution limit is severely constrained by low spring-summer water availability, whereas growth of silver fir forests in non-Mediterranean areas is limited by cold conditions in late winter to early spring. Main conclusions Climate warming is distinctly modifying growth patterns and responses to climate in silver fir across most of the species' European distribution area. In south-western Europe the reduction in growth of many populations is related to an observed increase in aridity, whereas in more temperate areas warming is enhancing growth. Our results confirm a decline in the growth of silver fir at its south-western distribution limits as a consequence of climate warming.

1. Mixed conifer-hardwood forests can be more productive than pure forests and they are increasingly considered as ecosystems that could provide adaptation strategies in the face of global change. However, the combined effects of tree-to-tree competition, rising atmospheric CO₂ concentrations and climate on such mixtures remain poorly characterized and understood. 2. To fill this research gap, we reconstructed 34-year series (1980-2013) of growth (basal area increment, BAI) and intrinsic water-use efficiency (iWUE) of Scots pine (Pinus sylvestris L.)-European beech (Fagus sylvatica L.) mixed stands at two climatically contrasting sites located in the southwestern Pyrenees. We also gathered data on tree-to-tree competition and climate variables in order to test the hypotheses that (1) radial growth will be greater when exposed to inter- than to intraspecific competition, that is, when species complementarity occurs and (2) enhanced iWUE could be linked to improved stem radial growth. 3. Growth of both species was reduced when intraspecific competition increased. Species complementarity was linked to improved growth of Scots pine at the continental site, while competition overrode any complementarity advantage at the drought-prone Mediterranean site. Beech growth did not show any significant response to pine admixture likely due to shade tolerance and the highly competitive nature of this species. Increasing interspecific competition drove recent iWUE changes, which increased in Scots pine but decreased in European beech. The iWUE enhancement did not involve any growth improvement in Scots pine. However, the positive BAI-iWUE relationship found for beech suggests an enhanced beech growth in drought-prone sites due to improved water use. 4. Synthesis. Complementarity may enhance growth in mixed forests. However, water scarcity can constrict light-related complementarity for shade intolerant species (Scots pine) in drought-prone sites. Basal area increment-intrinsic water-use efficiency relationships were negative for Scots pine and positive for European beech. These contrasting behaviours have got implications for coping with the expected increasing drought events in Scots pine-European beech mixtures located near the ecological limit of the two species. Complementarity effects between tree species should be considered to avoid overestimating the degree of future carbon uptake by mixed conifer-broadleaf forests.

Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60-80 year-old trees and 0.02-1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (+12 %), stand density (+20 %), basal area growth (+12 %), and stand volume growth (+8 %) were higher than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (+20 %) and height (+6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (-8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520-1175 mm year(-1)), mean annual temperature (6-10.5 A degrees C), and the drought index by de Martonne (28-61 mm A degrees C-1) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change.

Forest fragmentation creates forest edges, and the effect of those edges increases as the size of forest fragments decreases. Edge effects include changes to microclimatic conditions at the forest edge, which affect vegetation structure. No previous studies have directly tested the relationship between microclimate and vegetation structure (for instance, basal area, trees mean height, dead trees and damage trees) at the edge of forest fragments in the Atlantic Forest domain. We tested the following three hypotheses: (i) the microclimatic conditions differ between the edge and the interior of the forest, (ii) the forest structure differs between the edge and the interior of the forest and (iii) changes to microclimatic conditions at the forest edge negatively affect vegetation structure at the edges. Our results demonstrate that edge habitats are significantly more susceptible to strong winds, lower humidity and higher air temperatures than forest interiors. The microclimate may be considered the principal factor that explains the difference between the vegetation structure of the forest edge and the forest interior. Our results suggest that even large forest fragments in the Brazilian Atlantic Forest may be impacted by negative edge effects.

1.Tropical cyclones (TCs) are large-scale disturbances that regularly impact tropical forests. Although long-term impacts of TCs on forest structure have been proposed, a global test of the relationship between forest structure and TC frequency and intensity is lacking. We test on a pantropical scale whether TCs shape the structure of tropical and subtropical forests in the long-term. 2.We compiled forest structural features (stem density, basal area, mean canopy height and maximum tree size) for plants ≥ 10 cm in diameter at breast height from published forest inventory data (438 plots ≥ 0.1 ha, pooled into 250 1 x 1-degree grid cells) located in dry and humid forests. We computed maps of cyclone frequency and energy released by cyclones per unit area (power dissipation index, PDI) using a high-resolution historical database of TCs trajectories and intensities. We then tested the relationship between PDI and forest structural features using multivariate linear models, controlling for climate (mean annual temperature and water availability) and human disturbance (human foot print). 3.Forests subject to frequent cyclones (at least one TCs per decade) and high PDI exhibited higher stem density and basal area, and lower canopy heights. However, the relationships between PDI and basal area or canopy height were partially masked by lower water availability and higher human foot print in tropical dry forests. 4.Synthesis. Our results provide the first evidence that TCs have a long-term impact on the structure of tropical and subtropical forests in a globally consistent way. The strong relationship between PDI and stem density suggests that frequent and intense TCs reduce canopy cover through defoliation and tree mortality, encouraging higher regeneration and turnover of biomass. The projected increase in intensity and poleward extension of TCs due to anthropogenic climate change may therefore have important and lasting impacts on the structure and dynamics of forests in the future.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981?2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.

CONTEXT: An increase in the incidence of large wildfires worldwide has prompted concerns about the resilience of forest ecosystems, particularly in the western U.S., where recent changes are linked with climate warming and 20th-century land management practices. OBJECTIVES: To study forest resilience to recent wildfires, we examined relationships among fire legacies, landscape features, ecological conditions, and patterns of post-fire conifer regeneration. METHODS: We quantified regeneration across 182 sites in 21 recent large fires in dry mixed-conifer forests of the U.S. northern Rockies. We used logistic and negative binomial regression to predict the probability of establishment and abundance of conifers 5–13 years post-fire. RESULTS: Seedling densities varied widely across all sites (0–127,500 seedlings ha⁻¹) and were best explained by variability in distance to live seed sources (β = −0.014, p = 0.002) and pre-fire tree basal area (β = 0.072, p = 0.008). Beyond 95 m from the nearest live seed source, the probability of seedling establishment was low. Across all the fires we studied, 75 % of the burned area with high tree mortality was within this 95-m threshold, suggesting the presence of live seed trees to facilitate natural regeneration. CONCLUSIONS: Combined with the mix of species present within the burn mosaic, dry mixed-conifer forests will be resilient to large fires across our study region, provided that seedlings survive, fire do not become more frequent, high-severity patches do not get significantly larger, and post-fire climate conditions remain suitable for seedling establishment and survival.