Moose and vole browsing patterns in experimentally assembled pure and mixed forest stands

Plants growing in diverse communities are believed to exhibit associational resistance to herbivores, but this hypothesis has seldom been tested experimentally for vertebrate herbivores in forest ecosystems. We examined browsing patterns of the two principal mammalian herbivores of Finnish boreal forests, moose and voles, in young stands where tree species diversity and composition were experimentally manipulated. The stands were composed either of monocultures or different 2–5 species mixtures of Norway spruce, Scots pine, Siberian larch, silver birch, and black alder. Voles and moose showed contrasting responses to stand diversity and species composition. In accordance with the predictions of the associational resistance hypothesis, vole damage was higher in tree monocultures than in mixed stands, although stand diversity effects were statistically significant only at one of the three study areas. Voles also damaged more trees in coniferous than in deciduous stands. In contrast, moose browsing tended to increase with the number of tree species in a stand and with the presence of the preferred tree species, birch, in a mixture. The observed differences in vole and moose responses to stand diversity and species composition are likely to be due to different feeding specialisation, foraging patterns, and movement ability of these herbivores. Voles switched to trees only when the supply of a more preferred food (grasses and forbs) was depleted and restricted their feeding choice only to the most palatable tree species regardless of the number of tree species present per stand. In contrast, tree branches and foliage represented an important part of moose diet throughout the year; moose may be able to tolerate secondary plant metabolites of different tree species better than voles and may thus benefit from diet broadening when more tree species are available. Furthermore, the home range size and foraging ability of these two very differently sized herbivores may partly explain the observed differences in utilisation of different tree species. Finally, both moose and voles showed high spatial and temporal variation in their feeding; in particular, vole damage was more influenced by tree species diversity in areas and years with high vole densities. Thus, diversification of forest stands may have very different effects on mammalian browsing depending on the herbivores present, their densities, and the tree species used in reforestation.     

Global species–energy relationship in forest plots: role of abundance,temperature and species climatic tolerances

Aim To evaluate the strength of evidence for hypotheses explaining the relationship between climate and species richness in forest plots. We focused on the effect of energy availability which has been hypothesized to influence species richness: (1) via the effect of productivity on the total number of individuals (the more individuals hypothesis, MIH); (2) through the effect of temperature on metabolic rate (metabolic theory of biodiversity, MTB); or (3) by imposing climatic limits on species distributions. Location Global. Methods We utilized a unique ‘Gentry‐style’ 370 forest plots data set comprising tree counts and individual stem measurements, covering tropical and temperate forests across all six forested continents. We analysed variation in plot species richness and species richness controlled for the number of individuals by using rarefaction. Ordinary least squares (OLS) regression and spatial regressions were used to explore the relative performance of different sets of environmental variables. Results Species richness patterns do not differ whether we use raw number of species or number of species controlled for number of individuals, indicating that number of individuals is not the proximate driver of species richness. Productivity‐related variables (actual evapotranspiration, net primary productivity, normalized difference vegetation index) perform relatively poorly as correlates of tree species richness. The best predictors of species richness consistently include the minimum temperature and precipitation values together with the annual means of these variables. Main conclusion Across the world's forests there is no evidence to support the MIH, and a very limited evidence for a prominent role of productivity as a driver of species richness patterns. The role of temperature is much more important, although this effect is more complex than originally assumed by the MTB. Variation in forest plot diversity appears to be mostly affected by variation in the minimum climatic values. This is consistent with the ‘climatic tolerance hypothesis’ that climatic extremes have acted as a strong constraint on species distribution and diversity.     

Tree species richness decreases while species evenness increases with disturbance frequency in a natural boreal forest landscape

Understanding species diversity and disturbance relationships is important for biodiversity conservation in disturbance‐driven boreal forests. Species richness and evenness may respond differently with stand development following fire. Furthermore, few studies have simultaneously accounted for the influences of climate and local site conditions on species diversity. Using forest inventory data, we examined the relationships between species richness, Shannon''s index, evenness, and time since last stand‐replacing fire (TSF) in a large landscape of disturbance‐driven boreal forest. TSF has negative effect on species richness and Shannon''s index, and a positive effect on species evenness. Path analysis revealed that the environmental variables affect richness and Shannon''s index only through their effects on TSF while affecting evenness directly as well as through their effects on TSF. Synthesis and applications. Our results demonstrate that species richness and Shannon''s index decrease while species evenness increases with TSF in a boreal forest landscape. Furthermore, we show that disturbance frequency, local site conditions, and climate simultaneously influence tree species diversity through complex direct and indirect effects in the studied boreal forest.     

Tree diversity promotes generalist herbivore community patterns in a young subtropical forest experiment

Stand diversification is considered a promising management approach to increasing the multifunctionality and ecological stability of forests. However, how tree diversity affects higher trophic levels and their role in regulating forest functioning is not well explored particularly for (sub)tropical regions. We analyzed the effects of tree species richness, community composition, and functional diversity on the abundance, species richness, and beta diversity of important functional groups of herbivores and predators in a large-scale forest biodiversity experiment in south-east China. Tree species richness promoted the abundance, but not the species richness, of the dominant, generalist herbivores (especially, adult leaf chewers), probably through diet mixing effects. In contrast, tree richness did not affect the abundance of more specialized herbivores (larval leaf chewers, sap suckers) or predators (web and hunting spiders), and only increased the species richness of larval chewers. Leaf chemical diversity was unrelated to the arthropod data, and leaf morphological diversity only positively affected oligophagous herbivore and hunting spider abundance. However, richness and abundance of all arthropods showed relationships with community-weighted leaf trait means (CWM). The effects of trait diversity and CWMs probably reflect specific nutritional or habitat requirements. This is supported by the strong effects of tree species composition and CWMs on herbivore and spider beta diversity. Although specialized herbivores are generally assumed to determine herbivore effects in species-rich forests, our study suggests that generalist herbivores can be crucial for trophic interactions. Our results indicate that promoting pest control through stand diversification might require a stronger focus on identifying the best-performing tree species mixtures.     

The diversity of beetle assemblages in different habitat types in Sabah, Malaysia

The diversity of beetle assemblages in different habitat types (primary forest, logged forest, acacia plantation and oil palm plantation) in Sabah, Malaysia was investigated using three different methods based on habitat levels (Winkler sampling, flight-interception-trapping and mist-blowing). The overall diversity was extremely high, with 1711 species recorded from only 8028 individuals and 81 families (115 family and subfamily groups). Different degrees of environmental changes had varying effects on the beetle species richness and abundance, with oil palm plantation assemblage being most severely affected, followed by acacia plantation and then logged forest. A few species became numerically dominant in the oil palm plantation. In terms of beetle species composition, the acacia fauna showed much similarity with the logged forest fauna, and the oil palm fauna was very different from the rest. The effects of environmental variables (number of plant species, sapling and tree densities, amount of leaf litter, ground cover, canopy cover, soil pH and compaction) on the beetle assemblage were also investigated. Leaf litter correlated with species richness, abundance and composition of subterranean beetles. Plant species richness, tree and sapling densities correlated with species richness, abundance and composition of understorey beetles while ground cover correlated only with the species richness and abundance of these beetles. Canopy cover correlated only with arboreal beetles. In trophic structure, predators represented more than 40% of the species and individuals. Environmental changes affected the trophic structure with proportionally more herbivores (abundance) but fewer predators (species richness and abundance) in the oil palm plantation. Biodiversity, conservation and practical aspects of pest management were also highlighted in this study.     

Predictors of moss and liverwort species diversity of microsites in conifer‐dominated boreal forest

Patterns of moss and liverwort species diversity — species richness and species turnover (β‐diversity) — in three conifer‐dominated boreal forest stands of northern Alberta, Canada are described. We examined the relationship between bryophyte species diversity and micro‐environment at two sample grains, the microsite — substrate types for moss colonization: logs, stumps, tree bases, undisturbed patches of forest floor (dominated by feather moss species), and disturbed patches of forest floor — and the mesosite (25 m × 25 m plots). Microsite type and properties (e.g. decay class, hardwood vs softwood, pH) were the principal predictors of bryophyte species diversity and not micro‐environment variation among mesosites. Microsite type was the strongest predictor of microsite species richness and β‐diversity was higher among microsites and types and within microsites than among mesosites or stands. Microsite properties were significant predictors of species richness for all microsite types. Log and stump decay classes, influenced also by hardwood vs softwood predicted species richness of woody microsite types and soil pH and moisture predicted species richness of forest floor microsites. β‐diversity was highest for tree bases and disturbed patches of forest floor and lowest for logs. Mesosite β‐diversity was lower than that among microsites, and mesosite species richness was not well explained by measured environmental parameters. Results suggest that in conifer‐dominated boreal stands, species richness of microsites is only negligibly influenced by within‐stand variation at the mesosite grain and that substrate characteristics are the most important predictors of bryophyte species diversity in this ecosystem.     

The effect of biodiversity on tree productivity: from temperate to boreal forests

Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning. Experimental work has led to the proposal of mechanisms such as niche complementarity. However, few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions. (1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps. (1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables, as well as density and biodiversity indices, were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large‐scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity–ecosystem functioning (B‐EF) research.     

Contrasting effects of tree diversity on young tree growth and resistance to insect herbivores across three biodiversity experiments

Tree diversity is an important driver of forest ecosystem functioning, hypothesised to enhance tree growth and resistance to herbivores. To test this, we assessed the relative importance of tree species richness and functional diversity on tree height growth and insect herbivore damage across three tree diversity experiments in Finland, France and Germany, established within the last fifteen years. These experiments encompass species richness gradients from monocultures up to five species mixtures, with compositions drawn from a pool of eleven tree species. Tree height growth and total insect herbivory were evaluated at both the tree species and forest plot scales. Trees in mixtures tended to grow taller, but on average received more insect herbivory relative to monocultures. Gradients of tree species richness or functional diversity had only weak impact on the magnitude of these effects. Community weighted means of specific leaf area alone captured diversity effects on tree height growth, with stronger positive effects of diversity in mixtures with high community SLA. Tree species‐specific responses were highly variable. No species significantly benefited both in terms of increased growth and reduced herbivory when grown in mixtures. More species showed positive height growth responses in mixed assemblages, but only the two exotic conifers experienced associational resistance to herbivores. This large‐scale study shows that tree height growth in young forest plantations tends to be higher in species mixtures than in monocultures, but incremental increases in functional diversity have, at best, weak marginal growth benefits. Moreover, there appear to be contrasting effects at forest plot versus individual species scales. Thus, while some species show lower herbivore damage in mixtures, this is not a consistent trend and contrasts the higher overall damage in mixtures observed at the forest plot scale. To improve both tree growth and resistance to herbivores in tree species mixtures seems therefore challenging.     

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.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

The role of regional and local scale predictors for plant species richness in Mediterranean forests

Abstract

Both local and regional predictors play a role in determining plant community structure and composition. Climate, soil features as well as different local history and management affect forest understorey and tree species composition, but to date their specific role is relatively unknown. Few studies have addressed the importance of these predictors, especially in the Mediterranean area, where environmental conditions and human impacts have generated heterogeneous forest communities. In this study, the relationships between environmental variables and species richness of different groups of vascular plants (vascular species, woody species and open habitat species) and bryophytes were investigated in Tuscan forests. A total of 37 environmental variables were used by generalised linear model fitting in order to find parsimonious sub-sets of environmental factors (predictors) that are able to explain species diversity patterns at the local scale. Moreover, the role of regional and local variable groups on species richness of the considered plant groups was estimated by using the variance partitioning approach. We found that local variables, such as forest management and structure, explained more variance than regional variables for total species richness, open habitat species richness and bryophyte species richness. On the other hand, regional variables (such as elevation) played a central role for woody species richness.     

Evidence of history in explaining diversity patterns in tropical riverine fish

Aim Documentation of the ongoing effect of rain forest refuges at the last glacial maximum (LGM) on patterns of tropical freshwater fish diversity. Location Tropical South and Central America, and West Africa. Methods LGM rain forest regions and species richness by drainage were compiled from published data. GIS mapping was applied to compile drainage area and contemporary primary productivity. We used multiple regression analyses, applied separately for Tropical South America, Central America and West Africa, to assess differences in species richness between drainages that were connected and disconnected to rain forest refuge zones during the LGM. Spatial autocorrelation of the residuals was tested using Moran's I statistic. We added an intercontinental comparison to our analyses to see if a historical signal would persist even when a regional historical effect (climate at the LGM) had already been accounted for. Results Both area and history (contact with LGM rain forest refuge) explained the greatest proportions of variance in the geographical pattern of riverine species richness. In the three examined regions, we found highest richness in drainages that were connected to the rain forest refuges. No significant residual spatial autocorrelation was detected after considering area, primary productivity and LGM rain forest refuges. These results show that past climatic events still affect West African and Latin American regional and continental freshwater fish richness. At the continental scale, we found South American rivers more species‐rich than expected on the basis of their area, productivity and connectedness to rain forest refuge. Conversely, Central American rivers were less species‐diverse than expected by the grouped model. African rivers were intermediate. Therefore, a historical signal persists even when a regional historical effect (climate at the LGM) had already been accounted for. Main conclusions It has been hypothesized that past climatic events have limited impact on species richness because species have tracked environmental changes through range shifts. However, when considering organisms with physically constrained dispersal (such as freshwater fish), past events leave a perceptible imprint on present species diversity. Furthermore, we considered regions that have comparable contemporary climatic and environmental characteristics, explaining the absence of a productivity effect. From the LGM to the present day (a time scale of 18,000 years), extinction processes should have played a predominant role in shaping the current diversity pattern. By contrast, the continental effects could reflect historical contingencies explained by differences in speciation and extinction rates between continents at higher time scales (millions of years).     

Patterns of diversity in the strata of boreal montane forest in British Columbia

Abstract. Patterns of diversity were analyzed in a boreal coniferous forest and its strata (tree, shrub, herb and bryophyte layers): number of species per community — α-diversity, total species richness — γ-diversity, mean similarity — β-diversity, and mosaic diversity, a measure of complexity. These four measures of diversity consistently decreased from lower to upper vegetation layers. To study the effect of juveniles of larger life forms on diversity of lower layers, they were removed from the data and the measures of diversity reanalyzed. Number of species per community and mosaic diversity decreased substantially, but β-diversity did not change. So, the effect of juveniles on γ-diversity is due to the greater number of species per community. Multiple regression models revealed that the relationships between α-diversity and the environmental variables were the same for the whole forest and for the herb layer. Elevation and soil pH were the major variables explaining α-diversity in the whole community. Climate was the only environmental gradient related to species richness in all individual strata. Tree and herb richness values were negatively related to soil drainage and acidity, respectively. Species richness of the plant community was affected by environmental variability mostly through the herb layer. Various explanations of the observed diversity patterns included: environmental constraints, resource competition, generation time, and colonization processes.     

Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

The environmental basis of North American species richness patterns among Epicauta (Coleoptera: Meloidae)

Understanding regional variability in species richness is necessary for conservation efforts to succeed in the face of large-scale environmental deterioration. Several analyses of North American vertebrates have shown that climatic energy provides the best explanation of contemporary species richness patterns. The paucity of analyses of insect diversity patterns, however, remains a serious obstacle to a general hypothesis of spatial variation in diversity. We collected species distribution data on a North American beetle genus, Epicauta (Coleoptera: Meloidae) and tested several major diversity hypotheses. These beetles are generally grasshopper egg predators as larvae, and angiosperm herbivores as adults. Epicauta richness is highest in the hot, dry American southwest, and decreases north and east, consistent with the species richness-energy hypothesis. Potential evapotranspiration, which is also the best predictor of richness patterns among North American vertebrates, explains 80.2% of the variability in Epicauta species richness. Net primary productivity and variables measuring climatic heat energy only (such as PET) are not generally comparable, though they are sometimes treated as if they were equivalent. We conclude that the species richness-energy hypothesis currently provides a better overall explanation for Epicauta species richness patterns in North America than other major diversity hypotheses. The observed relationship between climatic energy and regional species richness may provide significant insight into the response of ecological communities to climate change.     

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.     

Phylogenetic structure illuminates the mechanistic role of environmental heterogeneity in community organization

1. Diversity begets diversity. Numerous published positive correlations between environmental heterogeneity and species diversity indicate ubiquity of this phenomenon. Nonetheless, most assessments of this relationship are phenomenological and provide little insight into the mechanism whereby such positive association results. 2. Two unresolved issues could better illuminate the mechanistic basis to diversity begets diversity. First, as environmental heterogeneity increases, both productivity and the species richness that contributes to that productivity often increase in a correlated fashion thus obscuring the primary driver. Second, it is unclear how species are added to communities as diversity increases and whether additions are trait based. 3. We examined these issues based on 31 rodent communities in the central Mojave Desert. At each site, we estimated rodent species richness and characterized environmental heterogeneity from the perspectives of standing primary productivity and number of seed resources. We further examined the phylogenetic structure of communities by estimating the mean phylogenetic distance (MPD) among species and by comparing empirical phylogenetic distances to those based on random assembly from a regional species pool. 4. The relationship between rodent species diversity and environmental heterogeneity was positive and significant. Moreover, diversity of resources accounted for more unique variation than did total productivity, suggesting that variety and not total amount of resource was the driver of increased rodent diversity. Relationships between environmental heterogeneity and phylogenetic distance were negative and significant; species were significantly phylogenetically over-dispersed in communities of low environmental heterogeneity and became more clumped as environmental heterogeneity increased. 5. Results suggest that species diversity increases with environmental heterogeneity because a wider variety of resources allow greater species packing within communities.     

The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

Diversity and species distribution of ectomycorrhizal fungi along productivity gradients of a southern boreal forest

Coniferous forests with diverse ectomycorrhizal fungus (EMF) communities are associated with nutrient-poor, acidic soils but there is some debate whether EMF can be equally adapted to more productive, nitrogen-rich sites. We compared EMF species distribution and diversity along a replicated productivity gradient in a southern boreal forest of British Columbia (Canada). Roots from subalpine fir (Abies lasiocarpa) saplings of the understory were sampled and EMF species were identified by morphotypes supplemented with ITS rDNA analysis. There were significant changes in the distribution and abundance of 74 EMF species along the productivity gradient, with as little as 24% community similarity among contrasting sites. Species richness per plot increased asymptotically with foliar nitrogen concentrations of subalpine fir, demonstrating that many EMF species were well suited to soils with high rates of nitrogen mineralization. EMF species abundance in relation to site productivity included parabolic, negative linear, and positive exponential curves. Both multi-site and more narrowly distributed EMF were documented, and a diverse mix of mantle exploration types was present across the entire productivity gradient. The results demonstrate strong associations of EMF fungal species with edaphic characteristics, especially nitrogen availability, and a specialization in EMF communities that may contribute to the successful exploitation of such contrasting extremes in soil fertility by a single tree host.     

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.