Lower Within-Community Variance of Negative Density Dependence Increases Forest Diversity

Local abundance of adult trees impedes growth of conspecific seedlings through host-specific enemies, a mechanism first proposed by Janzen and Connell to explain plant diversity in forests. While several studies suggest the importance of this mechanism, there is still little information of how the variance of negative density dependence (NDD) affects diversity of forest communities. With computer simulations, we analyzed the impact of strength and variance of NDD within tree communities on species diversity. We show that stronger NDD leads to higher species diversity. Furthermore, lower range of strengths of NDD within a community increases species richness and decreases variance of species abundances. Our results show that, beyond the average strength of NDD, the variance of NDD is also crucially important to explain species diversity. This can explain the dissimilarity of biodiversity between tropical and temperate forest: highly diverse forests could have lower NDD variance. This report suggests that natural enemies and the variety of the magnitude of their effects can contribute to the maintenance of biodiversity.     

The genetic dimension of pest pressure in the tropical rainforest

Wet tropical forests are among the most diverse ecosystems on Earth and can host several hundreds of tree species per hectare. To maintain such diversity, the community must contain large numbers of relatively rare species rather than be dominated by a few very common trees, as is often the case in temperate forests. Explaining the mechanisms preventing dominance by common species has been a major task of tropical forest ecology. One of the most promising mechanisms is negative density dependence (NDD) of tree abundance driven by pests, including fungal diseases (‘pest pressure’). NDD entails that the chance of survival of a sapling increases with the distance from a mature tree of the same species, thus preventing species from becoming locally dominant. Curiously, the strength of NDD is negatively correlated with abundance, meaning that tree species that are more common generally show weaker NDD (Comita et al. 2010 ). Interactions between plants and soil pathogens have been shown to play an important role in NDD (Klironomos 2002 ), and rare species are apparently more strongly affected (Mangan et al. 2010 ). However, the genetic mechanisms underlying this phenomenon have remained obscure. In this issue of Molecular Ecology, Marden et al. ( 2017 ) suggest that reduced diversity of the genes involved in pathogen recognition (Resistance genes or R genes) could explain why NDD is stronger in locally rare species.     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.     

Density‐dependent pre‐dispersal seed predation and fruit set in a tropical tree

Negative density‐dependent demographic processes operating at post‐dispersal seed, seedling, and juvenile stages are the dominant explanation for the coexistence of high numbers of tree species in tropical forests. At adult stages, the effect of pollinators and pre‐dispersal fruit predators are often dependent on the density or abundance of flowers and fruit in the canopy, but each have opposite effects on individual realized reproduction. We studied the effect of density on total and mature fruit set and pre‐dispersal predation rates within individual tree canopies in a common canopy tree species, Jacaranda copaia in a 50‐ha forest census plot in central Panama. We sampled all reproductive sized trees in the plot (n = 188) across three years and estimated fruit set and predation rates. Population‐wide pre‐dispersal seed predation averaged between 6–37% across years. Using linear mixed effects models, we found that increased density and fecundity of conspecific neighbours increased focal tree fruit set, but also the rate of pre‐dispersal predation. An interaction between individual and neighbourhood fruit production predicted lower predation rates at high individual and neighbourhood fecundities, which suggests predator satiation at high fruit abundance levels. However, the rate at which fruit set increased with conspecific neighbour fruit production was greater than the rate at which fruit were lost to predation, resulting in an overall positive effect of neighbour density on mature fruit production in focal trees. Our results run counter to the expectation of a uniformly negative effect of density across all life stages in tropical trees and suggest further exploration of the role of spatial clumping, pollen dispersal limitation, and predation at pre‐dispersal adult stages in maintenance of species diversity in plant communities.     

Tree diversity drives abundance and spatiotemporal β‐diversity of true bugs (Heteroptera)

1. Spatiotemporal patterns of canopy true bug diversity in forests of different tree species diversity have not yet been disentangled, although plant diversity has been shown to strongly impact the diversity and distribution of many insect communities. 2. Here we compare species richness of canopy true bugs across a tree diversity gradient ranging from simple beech to mixed forest stands. We analyse changes in community composition by additive partitioning of species diversity, for communities on various tree species, as well as for communities dwelling on beech alone. 3. Total species richness (γ‐diversity) and α‐diversity, and abundance of true bugs increased across the tree diversity gradient, while diversity changes were mediated by increased true bug abundance in the highly diverse forest stands. The same pattern was found for γ‐diversity in most functional guilds (e.g. forest specialists, herbivores, predators). Temporal and even more, spatial turnover (β‐diversity) among trees was closely related to tree diversity and accounted for ～90% of total γ‐diversity. 4. Results for beech alone were similar, but species turnover could not be related to the tree diversity gradient, and monthly turnover was higher compared to turnover among trees. 5. Our findings support the hypothesis that with increasing tree diversity and thereby increasing habitat heterogeneity, enhanced resource availability supports a greater number of individuals and species of true bugs. Tree species identity and the dissimilarity of true bug communities from tree to tree determine community patterns. 6. In conclusion, understanding diversity and distribution of insect communities in deciduous forests needs a perspective on patterns of spatiotemporal turnover. Heterogeneity among sites, tree species, as well as tree individuals contributed greatly to overall bug diversity.     

Niche assembly of epiphytic bryophyte communities in the Guianas: a regional approach

Aim Epiphytic bryophyte communities of tropical forests show a gradient in species composition from the base to the top of the host trees, indicating a strong role of niche assembly. This pattern, however, has never been tested at a regional scale. The aim of this study was to test whether niche assembly, rather than dispersal limitation, predominantly drives species composition of bryophyte communities across large spatial scales. Location Three lowland forests in the Guianas: one near Saul, French Guiana; and two near Mabura Hill, Guyana. Methods Communities of epiphytic bryophytes were sampled from six different height zones of several trees in three lowland forests. We analysed the composition of these communities using detrended correspondence analysis in order to find the best explanatory variable for the variation in community composition. A multi‐response permutation procedure was used to test the significance of grouping communities by height zone. We conducted an indicator species analysis to classify species as specialists or generalists and then tested, through weighted averaging, if specialists would indeed maintain their preferred height zone across the Guianas. Results Community composition was explained mainly by height zone. The similarity among communities inhabiting the same height zone of trees, across a distance of up to 640 km, was higher than the similarity among communities established along the vertical gradient of a single standing tree (30–50 m). More than half (57%) of the species had a preferred height zone, and the preference was consistent: species occupied roughly the same height zone on host trees in the different localities. The three local communities investigated were found to belong to the same regional species pool. Main conclusions Throughout the Guianas, epiphytic bryophyte communities are drawn from the same regional species pool, and their composition is shaped by micro‐environmental conditions. The predominance of niche assembly over dispersal assembly rules is consistently found at both local and regional scales.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Characteristics of snags and forest structure in southern mistbelt forests of the Amatole region,South Africa

Standing dead trees (or snags) are an important component of forest ecosystems, especially for tree cavity‐nesting vertebrate species, but their prevalence in South African forests remains under studied. Consequently, we investigated forest structure, and the presence and abundance of snags in six southern mistbelt forests in the Eastern Cape, South Africa. These forests have had varying levels of timber extraction over the past 150 years or more. We found snags were relatively rare in all six forests (<4.3% of trees sampled). Mean diameter at breast height (dbh) of snags ranged from 52 to 82 cm across the forests, with smaller snags in Kologha Forest and larger snags in Tyume Forest. A bimodal distribution of snag successional stages was found, with frequencies peaking at early and late stages, and few in the intermediate stages. Tree species diversity in the forests was relatively low (twelve–nineteen species across forests; only 28 species in total). There was no significant difference in dbh of trees between forests, with most occurring in the 20–29‐cm dbh size class. Future studies are required to identify trees that most likely support suitable cavities for tree cavity‐nesting bird species, and to determine cavity‐nester assemblage requirements in southern African forests.     

两种珍稀植物群落物种多度分布的核方法研究

首次提出物种多度分布的非参数核密度估计方法,介绍了此方法的构造和主要性质。珍稀濒危植物观光木群落和长苞铁杉群落的乔木层、灌木层、所有木本植物物种多度分布实例拟合结果表明,核方法能很好地描述群落物种多度分布。非参数核估计方法是群落物种多度分布模拟的一种有效方法,它丰富了物种多度分布拟合方法,为珍稀濒危植物的管理与保护提供了理论参考。     

Do foliar fungal communities of Norway spruce shift along a tree species diversity gradient in mature European forests?

Foliar fungal species are diverse and colonize all plants, though whether forest tree species composition influences the distribution of these fungal communities remains unclear. Fungal communities include quiescent taxa and the functionally important and metabolically active taxa that respond to changes in the environment. To determine fungal community shifts along a tree species diversity gradient, needles of Norway spruce were sampled from trees from four mature European forests. We hypothesized that the fungal communities and specific fungal taxa would correlate with tree species diversity. Furthermore, the active fungal community, and not the total community, would shift along the tree diversity gradient. High-throughput sequencing showed significant differences in the fungal communities in the different forests, and in one forest, tree diversity effects were observed, though this was not a general phenomenon. Our study also suggests that studying the metabolically active community may not provide additional information about community composition or diversity.     

Limited contributions of plant pathogens to density‐dependent seedling mortality of mast fruiting Bornean trees

Fungal pathogens are implicated in driving tropical plant diversity by facilitating strong, negative density‐dependent mortality of conspecific seedlings (C‐NDD). Assessment of the role of fungal pathogens in mediating coexistence derives from relatively few tree species and predominantly the Neotropics, limiting our understanding of their role in maintaining hyper‐diversity in many tropical forests. A key question is whether fungal pathogen‐mediated C‐NDD seedling mortality is ubiquitous across diverse plant communities. Using a manipulative shadehouse experiment, we tested the role of fungal pathogens in mediating C‐NDD seedling mortality of eight mast fruiting Bornean trees, typical of the species‐rich forests of South East Asia. We demonstrate species‐specific responses of seedlings to fungicide and density treatments, generating weak negative density‐dependent mortality. Overall seedling mortality was low and likely insufficient to promote overall community diversity. Although conducted in the same way as previous studies, we find little evidence that fungal pathogens play a substantial role in determining patterns of seedling mortality in a SE Asian mast fruiting forest, questioning our understanding of how Janzen‐Connell mechanisms structure the plant communities of this globally important forest type.     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

The contribution of rare species to community phylogenetic diversity across a global network of forest plots

Niche differentiation has been proposed as an explanation for rarity in species assemblages. To test this hypothesis requires quantifying the ecological similarity of species. This similarity can potentially be estimated by using phylogenetic relatedness. In this study, we predicted that if niche differentiation does explain the co-occurrence of rare and common species, then rare species should contribute greatly to the overall community phylogenetic diversity (PD), abundance will have phylogenetic signal, and common and rare species will be phylogenetically dissimilar. We tested these predictions by developing a novel method that integrates species rank abundance distributions with phylogenetic trees and trend analyses, to examine the relative contribution of individual species to the overall community PD. We then supplement this approach with analyses of phylogenetic signal in abundances and measures of phylogenetic similarity within and between rare and common species groups. We applied this analytical approach to 15 long-term temperate and tropical forest dynamics plots from around the world. We show that the niche differentiation hypothesis is supported in six of the nine gap-dominated forests but is rejected in the six disturbance-dominated and three gap-dominated forests. We also show that the three metrics utilized in this study each provide unique but corroborating information regarding the phylogenetic distribution of rarity in communities.     

Fungal disease incidence along tree diversity gradients depends on latitude in European forests

European forests host a diversity of tree species that are increasingly threatened by fungal pathogens, which may have cascading consequences for forest ecosystems and their functioning. Previous experimental studies suggest that foliar and root pathogen abundance and disease severity decrease with increasing tree species diversity, but evidences from natural forests are rare. Here, we tested whether foliar fungal disease incidence was negatively affected by tree species diversity in different forest types across Europe. We measured the foliar fungal disease incidence on 16 different tree species in 209 plots in six European countries, representing a forest‐type gradient from the Mediterranean to boreal forests. Forest plots of single species (monoculture plots) and those with different combinations of two to five tree species (mixed species plots) were compared. Specifically, we analyzed the influence of tree species richness, functional type (conifer vs. broadleaved) and phylogenetic diversity on overall fungal disease incidence. The effect of tree species richness on disease incidence varied with latitude and functional type. Disease incidence tended to increase with tree diversity, in particular in northern latitudes. Disease incidence decreased with tree species richness in conifers, but not in broadleaved trees. However, for specific damage symptoms, no tree species richness effects were observed. Although the patterns were weak, susceptibility of forests to disease appears to depend on the forest site and tree type.     

The effects of individual tree species on species diversity in a tropical dry forest change throughout ontogeny

Understanding how diversity is maintained in species‐rich communities, such as tropical forests, remains a challenge in ecology. Recent work suggests that the controversy between competing theories could be better resolved by considering the spatial scale at which different processes rule community assembly. Here we use individual species–area relationships (ISAR) to evaluate the spatial organization of tree diversity around individuals of different species in a completely‐mapped tropical dry forest in south Ecuador. We test two hypotheses. First, stressful environmental conditions promote facilitative interactions that will generate spatial signals of accumulation of diversity around individual trees – contrary to what has been reported in humid tropical forests. Second, spatial signals will shift through ontogeny. As, as larger, older trees generate new microsite conditions that affect the recruitment of younger, smaller trees. We compute ISAR functions for adult trees, for young trees and a new crossed‐ISAR function measuring the accumulation of diversity of young trees around the old trees. We compare observed ISARs to the expectations of inhomogeneous Poisson (i.e. null) models controlling for the effects of environmental variation and habitat association on tree distribution. Although the prevalent response among adult trees was not different from null expectations, which means that the organization of diversity in this size class could be explained by environmental heterogeneity alone, most species accumulated more diversity than expected over short spatial scales in the small stem size class. Only two species accumulated significant diversity in the crossed‐ISARs. Our study indicates the role of facilitation in the organization of plant diversity in this dry forest, but that facilitation is limited to some key species acting on early life stages and accumulating diversity around them. Our results demonstrate the benefit of considering different life‐stages and crossed analyses to disentangle the processes affecting community assembly in tropical dry forests.     

Heterogeneity in soil water and light environments and dispersal limitation: what facilitates tree species coexistence in a temperate forest?

In the present study, we analysed the habitat association of tree species in an old‐growth temperate forest across all life stages to test theories on the coexistence of tree species in forest communities. An inventory for trees was implemented at a 6‐ha plot in Ogawa Forest Reserve for adults, juveniles, saplings and seedlings. Volumetric soil water content (SMC) and light levels were measured in 10‐m grids. Relationships between the actual number of stems and environmental variables were determined for 35 major tree species, and the spatial correlations within and among species were analysed. The light level had no statistically significant effect on distribution of saplings and seedlings of any species. In contrast, most species had specific optimal values along the SMC gradient. The optimal values were almost identical in earlier life stages, but were more variable in later life stages among species. However, no effective niche partitioning among the species was apparent even at the adult stage. Furthermore, results of spatial analyses suggest that dispersal limitation was not sufficient to mitigate competition between species. This might result from well‐scattered seed distribution via wind and bird dispersal, as well as conspecific density‐dependent mortality of seeds and seedlings. Thus, both niche partitioning and dispersal limitation appeared less important for facilitating coexistence of species within this forest than expected in tropical forests. The tree species assembly in this temperate forest might be controlled through a neutral process at the spatial scale tested in this study.     

The influence of anthropogenic disturbances on the structure of arboreal arthropod communities

In the framework of our research, aimed at understanding the processes structuring tropical arthropod communities, we investigated the changes occurring in tree crown communities of forests of different disturbance levels. These were a mixed dipterocarp primary lowland rain forest in Kinabalu National Park (in Sabah, Malaysia) and, some kilometres away, three forests of regeneration periods 5, 15, and 40 years following a clear-cut. These disturbed forest sites were adjacent to one another and merged into mature forest. From each forest at least ten individuals of one tree species were sampled using the fogging method. In the primary forest relative proportions of some arthropod taxa differed on the ordinal and familial level significantly within trees. The dominance of Formicidae was characteristic as was the almost complete lack of less mobile arthropods such as Lepidoptera larvae. In the five-year- old forest, differences in relative proportions among most taxa had almost disappeared. Formicidae abundances had declined drastically which coincided with an increase of Lepidoptera larvae. With progressing forest succession, arthropod communities increasingly converged on the pattern of primary forest, and total ant abundance as well as diversity increased significantly. Ant communities in the most disturbed forest were of low structural complexity, and to a large degree predictable in species arrangement, but became more and more unpredictable as the complexity of the forest increased. Several species of Coleoptera and non-formicine Hymenoptera occurred in high numbers in the youngest forest, contrasting with the mature forest where all species were typically rare. These changes may indicate a change in the structuring mechanisms from predominantly deterministic processes in disturbed forests to stochastic processes in mature forest.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.