Tree diversity is key for promoting the diversity and abundance of forest-associated taxa in Europe

Plant diversity is an important driver of diversity at other trophic levels, suggesting that cascading extinctions could reduce overall biodiversity. Most evidence for positive effects of plant diversity comes from grasslands. Despite the fact that forests are hotspots of biodiversity, the importance of tree diversity, in particular its relative importance compared to other management related factors, in affecting forest-associated taxa is not well known. To address this, we used data from 183 plots, located in different forest types, from Mediterranean to Boreal, and established along a climatic gradient across six European countries (FunDivEUROPE project). We tested the influence of tree diversity, tree functional composition (i.e. functional trait values), forest structure, climate and soil on the diversity and abundance/activity of nine taxa (bats, birds, spiders, microorganisms, earthworms, ungulates, foliar fungal pathogens, defoliating insects and understorey plants) and on their overall diversity and abundance/activity (multidiversity, multiabundance/activity). Tree diversity was a key driver of taxon-level and overall forest-associated biodiversity, along with tree functional composition, forest structure, climate and soil. Both tree species richness and functional diversity (variation in functional trait values) were important. The effects of tree diversity on the abundance/activity of forest-associated taxa were less consistent. Nonetheless, spiders, ungulates and foliar fungal pathogens were all more abundant/active in diverse forests. Tree functional composition and structure were also important drivers of abundance/activity: conifer stands had lower overall multidiversity (although the effect was driven by defoliating insects), while stands with potentially tall trees had lower overall multiabundance/activity. We found more synergies than tradeoffs between diversity and abundance/activity of different taxa, suggesting that forest management can promote high diversity across taxa. Our results clearly show the high value of mixed forest stands for multiple forest-associated taxa and indicate that multiple dimensions of tree diversity (taxonomic and functional) are important.     

Bat and bird diversity along independent gradients of latitude and tree composition in European forests

Species assemblages are shaped by local and continental-scale processes that are seldom investigated together, due to the lack of surveys along independent gradients of latitude and habitat types. Our study investigated changes in the effects of forest composition and structure on bat and bird diversity across Europe. We compared the taxonomic and functional diversity of bat and bird assemblages in 209 mature forest plots spread along gradients of forest composition and vertical structure, replicated in 6 regions spanning from the Mediterranean to the boreal biomes. Species richness and functional evenness of both bat and bird communities were affected by the interactions between latitude and forest composition and structure. Bat and bird species richness increased with broadleaved tree cover in temperate and especially in boreal regions but not in the Mediterranean where they increased with conifer abundance. Bat species richness was lower in forests with smaller trees and denser understorey only in northern regions. Bird species richness was not affected by forest structure. Bird functional evenness increased in younger and denser forests. Bat functional evenness was also influenced by interactions between latitude and understorey structure, increasing in temperate forests but decreasing in the Mediterranean. Covariation between bat and bird abundances also shifted across Europe, from negative in southern forests to positive in northern forests. Our results suggest that community assembly processes in bats and birds of European forests are predominantly driven by abundance and accessibility of feeding resources, i.e., insect prey, and their changes across both forest types and latitudes.     

Long-term changes in bryophyte diversity of central European managed forests depending on site environmental features

Cryptogamic diversity is a reliable indicator of the state of forest ecosystems. In this study we analysed the variations in both bryophyte species richness overall and number of hemerophobic bryophyte species in Central European managed forests over a 20-year time span, based on data collected in 132 plots scattered across Poland. We tested differences in species richness among five temporal replicates, as well as among site types grouped based on elevation a.s.l., dominant tree species and stand age. The analyses revealed no significant trend in species richness across years. Meanwhile, species richness significantly increased along with elevation a.s.l., especially in broadleaved forests. No significant difference in species number between spruce and pine dominated stands emerged for mature stands, while there was a strong difference for young stands, with spruce forest hosting a much higher number of species. Species richness exhibited a slight, but not significant, increase over time in broadleaved forests, no significant variations in pine dominated stands and significant fluctuations in spruce dominated stands, yet without a significant trend. Out of the tested drivers, dominant tree species exhibited the strongest impact on species community composition. Number of hemerophobic and strongly hemerophobic species did not undergo significant variations across years either. The lack of bryophyte diversity trends highlighted in this study suggests Central European managed forests are in an equilibrium sate, maintained by the opposing effects of climate changes, on one side and of more sustainable forest management and pollutant deposition decline, from the other.

     

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.     

Using niche-based modelling to assess the impact of climate change on tree functional diversity in Europe

Rapid anthropogenic climate change is already affecting species distributions and ecosystem functioning worldwide. We applied niche-based models to analyse the impact of climate change on tree species and functional diversity in Europe. Present-day climate was used to predict the distributions of 122 tree species from different functional types (FT). We then explored projections of future distributions under one climate scenario for 2080, considering two alternative dispersal assumptions: no dispersal and unlimited dispersal. The species-rich broadleaved deciduous group appeared to play a key role in the future of different European regions. Temperate areas were projected to lose both species richness and functional diversity due to the loss of broadleaved deciduous trees. These were projected to migrate to boreal forests, thereby increasing their species richness and functional diversity. Atlantic areas provided an intermediate case, with a predicted reduction in the numbers of species and occasional predicted gains in functional diversity. This resulted from a loss in species within the broadleaved deciduous FT, but overall maintenance of the group. Our results illustrate the fact that both species-specific predictions and functional patterns should be examined separately in order to assess the impacts of climate change on biodiversity and gain insights into future ecosystem functioning.     

Application of species richness estimators for the assessment of fungal diversity

Species richness and distribution patterns of wood-inhabiting fungi and mycetozoans (slime moulds) were investigated in the canopy of a Central European temperate mixed deciduous forest. Species richness was described with diversity indices and species-accumulation curves. Nonmetrical multidimensional scaling was used to assess fungal species composition on different tree species. Different species richness estimators were used to extrapolate species richness beyond our own data. The reliability of the abundance-based coverage estimator, Chao, Jackknife and other estimators of species richness was evaluated for mycological surveys. While the species-accumulation curve of mycetozoans came close to saturation, that of wood-inhabiting fungi was continuously rising. The Chao 2 richness estimator was considered most appropriate to predict the number of species at the investigation site if sampling were continued. Gray's predictor of species richness should be used if statements of the number of species in larger areas are required. Multivariate analysis revealed the importance of different tree species for the conservation and maintenance of fungal diversity within forests, because each tree species possessed a characteristic fungal community. The described mathematical approaches of estimating species richness possess great potential to address fungal diversity on a regional, national, and global scale.     

北京东灵山主要森林植被中苔藓植物的物种多样性

通过对北京东灵山地区32个样地植被及苔藓植物的调查,对主要森林植被中苔藓植物物种多样性进行分析。结果表明:东灵山主要森林植被中共有苔藓植物16科33属79种,优势科为青藓科(Brachytheciaceae)和绢藓科(Entodontaceae)等。密叶绢藓(Entodoncompressus)、小牛舌藓(Anomodon minor)等为优势种。地面生苔藓植物物种丰富度以落叶阔叶混交林中最大,白桦林中最小,苔藓植物群落相似性以辽东栎林与落叶阔叶林之间最高,以山杨林与油松林之间最低。树附生苔藓植物物种丰富度以落叶阔叶混交林中最大,油松林中最小;苔藓植物群落相似性以油松林和山杨林之间最高,华北落叶松林与白桦林之间最低。地面生和树附生苔藓植物多样性与林型、树种及其所形成的林内微环境相关。     

Forest floor plant diversity drives the use of mature spruce forests by European bison

The distribution of large ungulates in space is in large extent driven by the availability of forage, which in temperate forests depends on light availability, and associated plant diversity and cover. We hypothesized that the increased number of GPS fixes of European bison (Bison bonasus L.) in usually avoided spruce forests was an effect of higher plant species richness and cover of the forest floor, which developed owing to increased light availability enhanced by spruce mortality. We carried out 80 forest floor plant surveys combined with tree measurement on plots chosen according to the number of GPS locations of GPS‐collared European bison. The mean plant species richness per plot was higher on intensively visited plots (IV) than rarely visited (RV) plots (30 ± 5.75 (SD) versus. 26 ± 6.19 (SD)). The frequency of 34 plant species was higher on IV plots, and they were mainly herbaceous species (32 species), while a significant part of 13 species with higher frequency on RV plots was woody plants (5 species). The species richness of forbs was higher on IV plots, while other functional groups of plants did not differ. Tree stem density on the IV plots was lower than on the RV plots (17.94 ± 6.73 (SD) versus 22.9 ± 7.67 (SD)), and the mean value of Ellenberg's ecological indicator for light availability for all forest floor plant species was higher on IV plots. European bison visiting mature spruce forests was driven by higher forest floor plant cover and species richness, and high share and species richness of forbs. The two latter features may be translated into higher quality and diversity of forage. In spite of morphological characteristics suggesting that European bison is a species of mixed (mosaic) habitats, it seems to be well adapted to thrive in diverse forests.     

太白山北坡落叶阔叶林物种多样性特征

对太白山北坡落叶阔叶林从不同类型,层次的丰富度,均匀度和物种多样性指数及其与环境梯度的关系等方面进行了分析。结果表明,处于中山带的锐齿槲栎林及辽东栎林的多样性较高,高海拔与低海拔处的群落多样性较低。水热梯度对物种多样性有很大影响。植物生长型与物种多样性的关系表现为灌木层的多样性指数（Ｄ）和均匀度大于草本层,而草本层大于乔木层,草木层物种丰富度最大,灌木层次之,乔木层最小,不同群落间灌木层各多样性测     

The functional trait space of tree species is influenced by the species richness of the canopy and the type of forest

Analysing how species modify their trait expression along a diversity gradient brings insight about the role that intraspecific variability plays over species interactions, e.g. competition versus complementarity. Here, we evaluated the functional trait space of nine tree species dominant in three types of European forests (a continental‐Mediterranean, a mountainous mixed temperate and a boreal) growing in communities with different species richness in the canopy, including pure stands. We compiled whole‐plant and leaf traits in 1719 individuals, and used them to quantify species trait hypervolumes in communities with different tree species richness. We investigated changes along the species richness gradient to disentangle species responses to the neighbouring environment, in terms of hypervolume size (trait variance), shape (trait relative importance) and centroid translation (shifts of mean trait values) using null models. Our main results showed differences in trait variance and shifts of mean values along the tree diversity gradient, with shorter trees but with larger crowns in mixed stands. We found constrained functional spaces (trait convergence) in pure stands, suggesting an important intraspecific competition, and expanded functional spaces (trait divergence) in two‐species admixtures, suggesting competition release due to interspecific complementarity. Nevertheless, further responses to increasing species richness were different for each forest type, waning species complementarity in sites with limiting conditions for growth. Our results demonstrate that tree species phenotypes respond to the species richness in the canopy in European forests, boosting species complementarity at low level of canopy diversity and with a site‐specific pattern at greater level of species richness. These outcomes evidence the limitation of functional diversity measures based only on traits from pure stands or general trait database values.     

A novel comparative research platform designed to determine the functional significance of tree species diversity in European forests

One of the current advances in functional biodiversity research is the move away from short-lived test systems towards the exploration of diversity-ecosystem functioning relationships in structurally more complex ecosystems. In forests, assumptions about the functional significance of tree species diversity have only recently produced a new generation of research on ecosystem processes and services. Novel experimental designs have now replaced traditional forestry trials, but these comparatively young experimental plots suffer from specific difficulties that are mainly related to the tree size and longevity. Tree species diversity experiments therefore need to be complemented with comparative observational studies in existing forests. Here we present the design and implementation of a new network of forest plots along tree species diversity gradients in six major European forest types: the FunDivEUROPE Exploratory Platform. Based on a review of the deficiencies of existing observational approaches and of unresolved research questions and hypotheses, we discuss the fundamental criteria that shaped the design of our platform. Key features include the extent of the species diversity gradient with mixtures up to five species, strict avoidance of a dilution gradient, special attention to community evenness and minimal covariation with other environmental factors. The new European research platform permits the most comprehensive assessment of tree species diversity effects on forest ecosystem functioning to date since it offers a common set of research plots to groups of researchers from very different disciplines and uses the same methodological approach in contrasting forest types along an extensive environmental gradient.     

Markedly Divergent Tree Assemblage Responses to Tropical Forest Loss and Fragmentation across a Strong Seasonality Gradient

We examine the effects of forest fragmentation on the structure and composition of tree assemblages within three seasonal and aseasonal forest types of southern Brazil, including evergreen, Araucaria, and deciduous forests. We sampled three southernmost Atlantic Forest landscapes, including the largest continuous forest protected areas within each forest type. Tree assemblages in each forest type were sampled within 10 plots of 0.1 ha in both continuous forests and 10 adjacent forest fragments. All trees within each plot were assigned to trait categories describing their regeneration strategy, vertical stratification, seed-dispersal mode, seed size, and wood density. We detected differences among both forest types and landscape contexts in terms of overall tree species richness, and the density and species richness of different functional groups in terms of regeneration strategy, seed dispersal mode and woody density. Overall, evergreen forest fragments exhibited the largest deviations from continuous forest plots in assemblage structure. Evergreen, Araucaria and deciduous forests diverge in the functional composition of tree floras, particularly in relation to regeneration strategy and stress tolerance. By supporting a more diversified light-demanding and stress-tolerant flora with reduced richness and abundance of shade-tolerant, old-growth species, both deciduous and Araucaria forest tree assemblages are more intrinsically resilient to contemporary human-disturbances, including fragmentation-induced edge effects, in terms of species erosion and functional shifts. We suggest that these intrinsic differences in the direction and magnitude of responses to changes in landscape structure between forest types should guide a wide range of conservation strategies in restoring fragmented tropical forest landscapes worldwide.     

The variation of tree beta diversity across a global network of forest plots

Aims With the aim of understanding why some of the world's forests exhibit higher tree beta diversity values than others, we asked: (1) what is the contribution of environmentally related variation versus pure spatial and local stochastic variation to tree beta diversity assessed at the forest plot scale; (2) at what resolution are these beta‐diversity components more apparent; and (3) what determines the variation in tree beta diversity observed across regions/continents? Location World‐wide. Methods We compiled an unprecedented data set of 10 large‐scale stem‐mapping forest plots differing in latitude, tree species richness and topographic variability. We assessed the tree beta diversity found within each forest plot separately. The non‐directional variation in tree species composition among cells of the plot was our measure of beta diversity. We compared the beta diversity of each plot with the value expected under a null model. We also apportioned the beta diversity into four components: pure topographic, spatially structured topographic, pure spatial and unexplained. We used linear mixed models to interpret the variation of beta diversity values across the plots. Results Total tree beta diversity within a forest plot decreased with increasing cell size, and increased with tree species richness and the amount of topographic variability of the plot. The topography‐related component of beta diversity was correlated with the amount of topographic variability but was unrelated to its species richness. The unexplained variation was correlated with the beta diversity expected under the null model and with species richness. Main conclusions Because different components of beta diversity have different determinants, comparisons of tree beta diversity across regions should quantify not only overall variation in species composition but also its components. Global‐scale patterns in tree beta diversity are largely coupled with changes in gamma richness due to the relationship between the latter and the variation generated by local stochastic assembly processes.     

Relative importance of climate vs local factors in shaping the regional patterns of forest plant richness across northeast China

Northeast (NE) China covers three climatic zones and contains all the major forest types of NE Asia. We sampled 108 forest plots in six nature reserves across NE China to examine the influence of climate and local factors (canopy seasonality, successional stage, topography and forest structure) on geographic patterns of plant richness. We analyzed the relative effects of different factors at two spatial scales: the regional scale (across both latitude and altitude) and the local scale (along the altitudinal gradient within site). Our results showed that the relative importance of climate vs local factors differed remarkably depending on scale and functional group. While total and tree species richness were mainly limited by climate, herb and shrub richness was more related to local factors (especially at the local scale). In the climatic factors, heat sum was the major correlate of tree, shrub and total species richness, while herb richness was more associated with winter coldness. Precipitation was not a limiting factor for forest plant richness in NE China. Climate accounted for 34–76% of variation in richness at the regional scale, but explained only 0–44% at the local scale. Among the local factors, shrub species richness was sensitive to seasonal canopy openness, with higher richness in deciduous forests than in the evergreen needle-leaf forest. On the other hand, herb richness was sensitive to forest successional stage, with higher richness in middle- successional forests than in the early and late-sucessional forests. Local topography (aspect and position on slope) and forest structure (tree density) also showed remarkable influence on species richness. Our results suggest the importance of including local factors when examining large scale diversity gradient (especially for understory species), and the necessity of comparing diversity patterns among functional groups at different spatial scales.     

Patterns in woody species diversity, richness and partitioning of diversity in forest communities of tropical deciduous forest biome

The regressive succession model hypothesizes tropical savanna-woodlands to be a degraded stage of primary deciduous forests. Species diversity, richness and evenness of woody species in savanna-woodlands, secondary deciduous forests and mature deciduous forests of central India were compared to test if the regressive succession explained pattern in species richness, diversity, functional diversity and basal area. At the plot scale (0.1 ha) secondary deciduous forests and savanna-woodlands had similar species diversity, a pattern not consistent with the regressive model of deciduous forest succession, and mature deciduous forests had greater species diversity and richness (p<0.05). When examined at a larger scale or community scale by pooling all plots within a community type, the trend in diversity persisted even with greater effort allocated to sampling of secondary deciduous forests. Species richness at the community scale was greatest in secondary deciduous forest as expected from species area relationship. The communities shared 28 woody species but the species composition was significantly different between the communities. I suggest that conservation of tropical deciduous forests based on regressive succession model is problematic.     

Disentangling Biodiversity and Climatic Determinants of Wood Production

Background

Despite empirical support for an increase in ecosystem productivity with species diversity in synthetic systems, there is ample evidence that this relationship is dependent on environmental characteristics, especially in structurally more complex natural systems. Empirical support for this relationship in forests is urgently needed, as these ecosystems play an important role in carbon sequestration.

Methodology/Principal Findings

We tested whether tree wood production is positively related to tree species richness while controlling for climatic factors, by analyzing 55265 forest inventory plots in 11 forest types across five European countries. On average, wood production was 24% higher in mixed than in monospecific forests. Taken alone, wood production was enhanced with increasing tree species richness in almost all forest types. In some forests, wood production was also greater with increasing numbers of tree types. Structural Equation Modeling indicated that the increase in wood production with tree species richness was largely mediated by a positive association between stand basal area and tree species richness. Mean annual temperature and mean annual precipitation affected wood production and species richness directly. However, the direction and magnitude of the influence of climatic variables on wood production and species richness was not consistent, and vary dependent on forest type.

Conclusions

Our analysis is the first to find a local scale positive relationship between tree species richness and tree wood production occurring across a continent. Our results strongly support incorporating the role of biodiversity in management and policy plans for forest carbon sequestration.     

甘肃小陇山林区不同生境类型蝶类多样性研究

于2006～2008年对甘肃小陇山林区不同生境类型中蝴蝶多样性进行了调查研究,研究中依据植被的不同将该林区的蝴蝶生境划分为6种类型:人工林、灌木丛和次生林、居民农田、针阔混交林、落叶阔叶林、针叶林。共获得蝴蝶标本5365只,隶属于11科116属210种。计算了6种生境类型中蝶类物种丰富度、相似性系数和多样性指数。不同生境中,灌木丛和次生林蝶类物种的多样性指数、丰富度和个体数量较高,人工林物种多样性指数、丰富度和个体数量较低;人工林与针叶林之间的相似性系数(0.4194)最高,针阔混交林与落叶阔叶林的相似性系数(0.2951)次之,人工林与灌木丛和次生林之间的相似性系数(0.0769)最低,表明各生境之间蝶类相似性系数很低。     

Leaf shedding and weather in tropical dry-seasonal forest shape the phenology of fungi – Lessons from two years of monthly surveys in southwestern Panama

In the present study, conducted in a secondary dry-seasonal forest in the pacific lowlands of southwestern Panama over 2 years, fungal diversity is linked to plant phenology, litter, and climatic data. Agaricales fungi showed maximum species richness at the beginning of rainy seasons, probably due to the important litter accumulation during the dry season and the increase in humidity favoring fungal growth. Species richness declined during the wet season possibly due to torrential rains, moulds, and decreasing availability of nutrients. Occurrence of foliar pathogenic microfungi correlated negatively with flushing of new leaves at the beginning of the rainy season. Their incidence increased during the wet season and remained high during the dry season. Synchronization of leaf shedding in most tree species significantly reduced the yearly incidence of foliar pathogenic fungi causing an annual turn-over of fungal pathogens that probably contributes to maintain a high diversity of plant pathogenic species.     

Evenness of soil organic carbon chemical components changes with tree species richness,composition and functional diversity across forests in China

Higher tree species richness generally increases the storage of soil organic carbon (SOC). However, less attention is paid to the influence of varied tree species composition on SOC storage. Recently, the perspectives for the stronger persistence of SOC caused by the higher molecular diversity of organic compounds were proposed. Therefore, the influences of tree species richness and composition on the molecular diversity of SOC need to be explored. In this study, an index of the evenness of diverse SOC chemical components was proposed to represent the potential resistance of SOC to decomposition under disturbances. Six natural forest types were selected encompassing a diversity gradient, ranging from cold temperate to tropical forests. We examined the correlations of tree species richness, composition, and functional diversity, with the evenness of SOC chemical components at a molecular level by ¹³C nuclear magnetic resonance. Across the range, tree species richness correlated to the evenness of SOC chemical components through tree species composition. The negative correlation of evenness of SOC chemical components with tree species composition, and the positive correlation of evenness of SOC chemical components with tree functional diversity were found. These indicate the larger difference in tree species composition and the lower community functional diversity resulted in the higher heterogeneity of SOC chemical components among the communities. The positive correlation of the evenness of SOC chemical components with the important value of indicator tree species, further revealed the specific tree species contributing to the higher evenness of SOC chemical components in each forest type. Soil fungal and bacterial α-diversity had effect on the evenness of SOC chemical components. These findings suggest that the indicator tree species conservation might be preferrable to simply increasing tree species richness, for enhancing the potential resistance of SOC to decomposition.     

Fungal endophyte communities in two tropical forests of southern India: diversity and host affiliation

Fifteen tree species from a tropical dry thorn forest and fifteen tree species from a tropical dry deciduous forest in the Mudumalai Wildlife Sanctuary, Nilgiri Biosphere Reserve, southern India, were surveyed for their foliar endophyte communities during the dry and wet seasons. Surface sterilized leaf segments of uniform dimension were plated on nutrient agar and culturable endophytes growing from the segments were identified. Endophyte diversity was greater in the dry thorn forest than in the dry deciduous forest in the dry season. Although the isolation frequency of culturable endophytes increased for both forests during the wet season, the assemblages were represented not by any unique fungal species but by the commonly occurring ones. Furthermore, although individual leaves were densely colonized by endophytes, only a few species of endophytes colonized the whole leaves; and, only a few fungal species dominated the foliar endophytic communities and were common for both forests during both dry and wet seasons. Thus, even under wet conditions that favour dispersal and infection by fungi, the endophyte diversity increased only marginally, an indication that certain tropical forests are not hyperdiverse with reference to fungal endophytes. This should be considered when using culturable endophyte diversity as a surrogate for estimating global fungal diversity.