Altitudinal biodiversity patterns of seed plants along Gongga Mountain in the southeastern Qinghai–Tibetan Plateau

The mechanisms underlying elevation patterns in species and phylogenetic diversity remain a central issue in ecology and are vital for effective biodiversity conservation in the mountains. Gongga Mountain, located in the southeastern Qinghai–Tibetan Plateau, represents one of the longest elevational gradients (ca. 6,500 m, from ca. 1,000 to 7,556 m) in the world for studying species diversity patterns. However, the elevational gradient and conservation of plant species diversity and phylogenetic diversity in this mountain remain poorly studied. Here, we compiled the elevational distributions of 2,667 native seed plant species occurring in Gongga Mountain, and estimated the species diversity, phylogenetic diversity, species density, and phylogenetic relatedness across ten elevation belts and five vegetation zones. The results indicated that species diversity and phylogenetic diversity of all seed plants showed a hump‐shaped pattern, peaking at 1,800–2,200 m. Species diversity was significantly correlated with phylogenetic diversity and species density. The floras in temperate coniferous broad‐leaved mixed forests, subalpine coniferous forests, and alpine shrublands and meadows were significantly phylogenetically clustered, whereas the floras in evergreen broad‐leaved forests had phylogenetically random structure. Both climate and human pressure had strong correlation with species diversity, phylogenetic diversity, and phylogenetic structure of seed plants. Our results suggest that the evergreen broad‐leaved forests and coniferous broad‐leaved mixed forests at low to mid elevations deserve more conservation efforts. This study improves our understanding on the elevational gradients of species and phylogenetic diversity and their determinants and provides support for improvement of seed plant conservation in Gongga Mountain.     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.     

Functional and genetic diversity changes through time in a cloud forest ant assemblage

Climate change in the Neotropics is causing upslope range shifts. We used arrays of ant species collected in a cloud forest at 1,500 m in Área de Conservación Guanacaste (ACG), northwestern Costa Rica, collected in two time periods (1998–2000 and 2008–2011) to measure changes in species richness and diversity over a decade. Using metrics of community structure, we found that the species assemblage in the collections from the 1990s was significantly phylogenetically clustered and functionally less diverse as compared to collections from the early 2000s. At both time points, the assemblages were significantly phylogenetically clustered and while the difference in functional diversity between the time points was not significant, the ant assemblage has become lighter in color (on average) over time. When individual species are considered, the overall pattern of replacement is consistent with the cloud forest ant assemblage being colonized by arrivals from lower elevation forests. The invertebrate communities on cloud forested mountain tops are especially vulnerable to a changing climate as there are two factors working together; no higher terrain to which they can move and the invasion of more and more taxa from lower downslope. This vulnerability is already measurable.     

Spider diversity across an elevation gradient in Área de Conservación Guanacaste (ACG), Costa Rica

Throughout the Neotropics, temperature and precipitation change with elevation and these changes affect the assemblage of species at any particular elevation. We documented the diversity of litter-inhabiting spiders, (Arachnida: Araneae) along a Costa Rican elevational gradient as it relates to covarying abiotic factors such as temperature and precipitation. The spiders we collected were principally unidentifiable juveniles, and so we used Barcode Index Numbers (BINs) derived from DNA barcodes as proxies for species-level interim names. We contrasted these taxon-based estimates with phylogenetic measures of alpha- and beta-diversity derived from both the mitochondrial DNA barcode region and a multi-gene phylogeny of spiders and found that neither the abundance nor the species richness of spiders was significantly correlated with elevation, temperature, or precipitation. However, we did find that spider assemblages in the upper elevation cloud forests were phylogenetically clustered, (and this pattern was unrelated to whether the phylogenetic patterns were derived mitochondrially or from a multi-gene analysis). One standard explanation for such a pattern is that harsh abiotic conditions in higher elevation forests have selected for particular spider lineages; however, this remains to be tested fully. The diversity of leaf-litter spider species we uncovered was high and further sampling of spider abundance and diversity across the ACG is likely to yield many new species.     

High specialization and limited structural change in plant‐herbivore networks along a successional chronosequence in tropical montane forest

Secondary succession is well‐understood, to the point of being predictable for plant communities, but the successional changes in plant‐herbivore interactions remains poorly explored. This is particularly true for tropical forests despite the increasing importance of early successional stages in tropical landscapes. Deriving expectations from successional theory, we examine properties of plant‐herbivore interaction networks while accounting for host phylogenetic structure along a succession chronosequence in montane rainforest in Papua New Guinea. We present one of the most comprehensive successional investigations of interaction networks, equating to > 40 person years of field sampling, and one of the few focused on montane tropical forests. We use a series of nine 0.2 ha forest plots across young secondary, mature secondary and primary montane forest, sampled almost completely for woody plants and larval leaf chewers (Lepidoptera) using forest felling. These networks comprised of 12 357 plant‐herbivore interactions and were analysed using quantitative network metrics, a phylogenetically controlled host‐use index and a qualitative network beta diversity measure. Network structural changes were low and specialisation metrics surprisingly similar throughout succession, despite high network beta diversity. Herbivore abundance was greatest in the earliest stages, and hosts here had more species‐rich herbivore assemblages, presumably reflecting higher palatability due to lower defensive investment. All herbivore communities were highly specialised, using a phylogenetically narrow set of hosts, while host phylogenetic diversity itself decreased throughout the chronosequence. Relatively high phylogenetic diversity, and thus high diversity of plant defenses, in early succession forest may result in herbivores feeding on fewer hosts than expected. Successional theory, derived primarily from temperate systems, is limited in predicting tropical host‐herbivore interactions. All succession stages harbour diverse and unique interaction networks, which together with largely similar network structures and consistent host use patterns, suggests general rules of assembly may apply to these systems.     

Fate of epiphytes on phorophytes with different architectural characteristics along the perturbation gradient of Sabal mexicana forests in Veracruz,Mexico

Question: Vascular epiphytes and hemiepiphytes (E/HE) in neotropical forests account for a large fraction of plant richness, but little is known of how the interplay between phorophyte architectural characteristics and habitat perturbation affect communities of E/HE. Location: Sabal mexicana forests in a coastal area of Veracruz, Mexico. Methods: We compared communities of E/HE on phorophytes with different architectural characteristics – the palm S. mexicana and non‐palm phorophytes – in three environments: conserved sites, perturbed sites and small regenerated forest fragments. We combined traditional (abundance, species richness, similarity and complementarity indices) and more recent (phylogenetic diversity) metrics to describe the communities of E/HE. Results: Overall, we recorded 924 E/HE individuals (nine families, 16 genera and 21 species). The abundance and species richness of E/HE was higher on palms than on non‐palm phorophytes. Abundance‐based complementarities between phorophytes and sites were high. We detected clear changes in community structure of E/HE with habitat perturbation, but there were no effects on the phylogenetic diversity of the E/HE community. Palm phorophytes hosted a more phylogenetically diverse community of E/HE than did non‐palm phorophytes. Conclusions: Palm phorophytes are key elements supporting the conservation of resilient communities of E/HE in S. mexicana forest. Habitat fragmentation has a strong effect on the structure of the E/HE community in S. mexicana forests. Ferns are the group of epiphytes most severely affected by habitat perturbation, but we detected no significant effect on the phylogenetic diversity of the community.     

Soil phosphorus heterogeneity promotes tree species diversity and phylogenetic clustering in a tropical seasonal rainforest

The niche theory predicts that environmental heterogeneity and species diversity are positively correlated in tropical forests, whereas the neutral theory suggests that stochastic processes are more important in determining species diversity. This study sought to investigate the effects of soil nutrient (nitrogen and phosphorus) heterogeneity on tree species diversity in the Xishuangbanna tropical seasonal rainforest in southwestern China. Thirty‐nine plots of 400 m² (20 × 20 m) were randomly located in the Xishuangbanna tropical seasonal rainforest. Within each plot, soil nutrient (nitrogen and phosphorus) availability and heterogeneity, tree species diversity, and community phylogenetic structure were measured. Soil phosphorus heterogeneity and tree species diversity in each plot were positively correlated, while phosphorus availability and tree species diversity were not. The trees in plots with low soil phosphorus heterogeneity were phylogenetically overdispersed, while the phylogenetic structure of trees within the plots became clustered as heterogeneity increased. Neither nitrogen availability nor its heterogeneity was correlated to tree species diversity or the phylogenetic structure of trees within the plots. The interspecific competition in the forest plots with low soil phosphorus heterogeneity could lead to an overdispersed community. However, as heterogeneity increase, more closely related species may be able to coexist together and lead to a clustered community. Our results indicate that soil phosphorus heterogeneity significantly affects tree diversity in the Xishuangbanna tropical seasonal rainforest, suggesting that deterministic processes are dominant in this tropical forest assembly.     

The role of habitat and daily activity patterns in explaining the diversity of mountain Neotropical dung beetle assemblages

The interaction between land use and climate change is expected to strongly affect species distributions along high elevation landscapes. We aimed to test the effect of climatic variables on community metrics among five types of land use in a high elevation landscape. We described dung beetle spatial and temporal taxonomic and functional diversity patterns, and partitioned β‐diversity into turnover and nestedness components. The interaction between land use and daily period of activity mostly drives abundance, functional richness and functional diversity, but not dung beetle species richness. Unlike Neotropical lowlands, species richness and abundance in open environments are similar to those existing in forests. Temperature is an important predictor of abundance and functional divergence. There is a higher spatial component of the taxonomic β‐diversity, which is highly driven by species turnover. The temporal component of the taxonomic β‐diversity was strongly driven by nestedness, where night assemblages are sub‐sets, although not entirely, of diurnal assemblages. For functional diversity, the temporal β‐diversity was much higher than the spatial β‐diversity, but both were similarly represented by functional group turnover and nestedness. The composition of nocturnal and diurnal assemblages is clearly different, even more than the differences observed between habitats. However, taxonomic turnover is the dominant force between sampling sites while nestedness dominates the daily pattern. This means that forest habitats are unlikely to act as shelters for grassland species under a scenario of rising temperature.     

Consistent patterns of high alpha and low beta diversity in tropical parasitic and free‐living protists

Tropical animals and plants are known to have high alpha diversity within forests, but low beta diversity between forests. By contrast, it is unknown whether microbes inhabiting the same ecosystems exhibit similar biogeographic patterns. To evaluate the biogeographies of tropical protists, we used metabarcoding data of species sampled in the soils of three lowland Neotropical rainforests. Taxa–area and distance–decay relationships for three of the dominant protist taxa and their subtaxa were estimated at both the OTU and phylogenetic levels, with presence–absence and abundance‐based measures. These estimates were compared to null models. High local alpha and low regional beta diversity patterns were consistently found for both the parasitic Apicomplexa and the largely free‐living Cercozoa and Ciliophora. Similar to animals and plants, the protists showed spatial structures between forests at the OTU and phylogenetic levels, and only at the phylogenetic level within forests. These results suggest that the biogeographies of macro‐ and micro‐organismal eukaryotes in lowland Neotropical rainforests are partially structured by the same general processes. However, and unlike the animals and plants, the protist OTUs did not exhibit spatial structures within forests, which hinders our ability to estimate the local and regional diversity of protists in tropical forests.     

Bioindicators of edge effects within Atlantic Forest remnants: Conservation implications in a threatened biodiversity hotspot

Aim

Deforestation of the Atlantic Forest of eastern Paraguay has been recent but extensive, resulting in a fragmented landscape highly influenced by forest edges. We examined edge effects on multiple dimensions of small mammalian diversity.

Location

Forest fragments of eastern Paraguayan Atlantic Forest.

Methods

We trapped small mammal species at different distances from the forest edge (DTE) in reserves and estimated multiple dimensions of diversity per site. Similarity analysis identified species clusters that best described the patterns of diversity across reserves. Multivariate ordination and linear mixed models were used to determine the influence of DTE on various dimensions of small mammal diversity.

Results

There was an increase in richness and abundance along a DTE gradient, and remnants with higher edge:area ratios showed higher richness and abundance, independent of remnant size. Species at edges were generalists, open-habitat species or exotic species (spillover effect). We found higher phylogenetic diversity and functional richness and divergence towards forest edges. Spillover of non-forest and invasive species best explained richness, generalist forest species best explained total abundance, abundance of Hylaeamys megacephalus best explained diversity and evenness metrics and the presence of Marmosa paraguayana best explained various phylogenetic diversity models. None of the models that included megafauna or social factors were shown to be important in explaining patterns as a function of DTE.

Main Conclusions

We found strong support for a spillover effect and mixed support for complementary resource use and enhanced habitat resources associated with ecotones. Generalists characterized edge assemblages but not all generalists were equivalent. Edges showed more phylogenetically and functionally distinct assemblages than the interior of remnants. There was a conservation of functional diversity; however, open-habitat species, habitat generalists and exotic species boosted diversity near forest edges. Mechanisms governing diversity along forest edges are complex; disentangling those mechanisms necessitates the use of multiple dimensions of diversity.     

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.     

Contrasting changes in the abundance and diversity of North American bird assemblages from 1971 to 2010

Although it is generally recognized that global biodiversity is declining, few studies have examined long‐term changes in multiple biodiversity dimensions simultaneously. In this study, we quantified and compared temporal changes in the abundance, taxonomic diversity, functional diversity, and phylogenetic diversity of bird assemblages, using roadside monitoring data of the North American Breeding Bird Survey from 1971 to 2010. We calculated 12 abundance and diversity metrics based on 5‐year average abundances of 519 species for each of 768 monitoring routes. We did this for all bird species together as well as for four subgroups based on breeding habitat affinity (grassland, woodland, wetland, and shrubland breeders). The majority of the biodiversity metrics increased or remained constant over the study period, whereas the overall abundance of birds showed a pronounced decrease, primarily driven by declines of the most abundant species. These results highlight how stable or even increasing metrics of taxonomic, functional, or phylogenetic diversity may occur in parallel with substantial losses of individuals. We further found that patterns of change differed among the species subgroups, with both abundance and diversity increasing for woodland birds and decreasing for grassland breeders. The contrasting changes between abundance and diversity and among the breeding habitat groups underscore the relevance of a multifaceted approach to measuring biodiversity change. Our findings further stress the importance of monitoring the overall abundance of individuals in addition to metrics of taxonomic, functional, or phylogenetic diversity, thus confirming the importance of population abundance as an essential biodiversity variable.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

Clade composition of a plant community indicates its phylogenetic diversity

Phylogenetic diversity quantification is based on indices computed from phylogenetic distances among species, which are derived from phylogenetic trees. This approach requires phylogenetic expertise and available molecular data, or a fully sampled synthesis‐based phylogeny. Here, we propose and evaluate a simpler alternative approach based on taxonomic coding. We developed metrics, the clade indices, based on information about clade proportions in communities and species richness of a community or a clade, which do not require phylogenies. Using vegetation records from herbaceous plots from Central Europe and simulated vegetation plots based on a megaphylogeny of vascular plants, we examined fit accuracy of our proposed indices for all dimensions of phylogenetic diversity (richness, divergence, and regularity). For real vegetation data, the clade indices fitted phylogeny‐based metrics very accurately (explanatory power was usually higher than 80% for phylogenetic richness, almost always higher than 90% for phylogenetic divergence, and often higher than 70% for phylogenetic regularity). For phylogenetic regularity, fit accuracy was habitat and species richness dependent. For phylogenetic richness and divergence, the clade indices performed consistently. In simulated datasets, fit accuracy of all clade indices increased with increasing species richness, suggesting better precision in species‐rich habitats and at larger spatial scales. Fit accuracy for phylogenetic divergence and regularity was unreliable at large phylogenetic scales, suggesting inadvisability of our method in habitats including many distantly related lineages. The clade indices are promising alternative measures for all projects with a phylogenetic framework, which can trade‐off a little precision for a significant speed‐up and simplification, such as macroecological analyses or where phylogenetic data is incomplete.     

Taxonomic,functional, and phylogenetic dimensions of rodent biodiversity along an extensive tropical elevational gradient

Relationships among taxonomic, functional, and phylogenetic dimensions of biodiversity provide insight about the relative contributions of ecological and evolutionary processes in structuring local assemblages. We used data for rodent species distributions from an extensive tropical elevational gradient to 1) describe elevational gradients for each of three dimensions of biodiversity, 2) evaluate the sufficiency of species richness as a surrogate for other dimensions, and 3) quantify the relative support for mechanisms that increase or decrease phylogenetic or functional dispersion. Taxonomic biodiversity was quantified by species richness, as well as by richness, evenness, diversity, dominance, and rarity at generic and familial levels. Morphological and categorical traits were used to estimate functional biodiversity, and an ultrametric mammalian supertree was used as the basis for estimating phylogenetic biodiversity. Elevational gradients of each dimension of biodiversity were strong, with significant linear and non‐linear components based on orthogonal polynomial regression. Empirical linear and non‐linear regression components were consistently different than those expected based on species richness for generic, familial, and phylogenetic biodiversity, but not for functional biodiversity. Nevertheless, the congruence of dimensions of biodiversity based on correlation analyses indicated that any one dimension is a useful surrogate for the other dimensions for rodents at Manu. Given variation in species richness, assemblages from lowland rainforests comprised more biodiversity than expected, whereas assemblages from cloud and elfin forests represented less biodiversity than expected. Warm temperatures, vertical complexity of the vegetation, and high productivity likely facilitate niche differentiation in rainforests, whereas cricetid rodents are competitively superior to other clades in the less structurally complex, less productive, and colder, high elevation habitats.     

Small montane cloud forest fragments are important for conserving tree diversity in the Ecuadorian Andes

Montane tropical cloud forests, with their complex topography, biodiversity, high numbers of endemic species, and rapid rates of clearing, are a top global conservation priority. However, species distributions at local and landscape scales in cloud forests are still poorly understood, in part because few regions have been surveyed. Empirical work has focused on species distributions along elevation gradients, but spatial variation among forests at the same elevation is less commonly investigated. In this study, the first to compare tree communities across multiple Andean cloud forests at similar elevations, we surveyed trees in five ridge‐top forest reserves at the upper end of the ‘mid‐elevation diversity bulge’ (1900–2250 masl) in the Intag Valley, a heavily deforested region in the Ecuadorian Andes. We found that tree communities were distinct in reserves located as close as 10 to 35 km apart, and that spatially closer forests were not more similar to one another. Although larger (1500 to 6880 ha), more intact forests contained significantly more tree species (108–120 species/0.1 ha) than smaller (30 to 780 ha) ones (56–87 species/0.1 ha), each reserve had unique combinations of more common species, and contained high proportions of species not found in the others. Results thus suggest that protecting multiple cloud forest patches within this narrow elevational band is essential to conserve landscape‐level tree diversity, and that even small forest reserves contribute significantly to biodiversity conservation. These findings can be applied to create management plans to conserve and restore cloud forests in the Andes and tropical montane cloud forests elsewhere.     

海南岛3个林区热带云雾林植物多样性变化

以分布在海南岛西部(霸王岭国家级自然保护区, 21个样方)、西南部(尖峰岭国家级自然保护区, 12个样方)和中部(黎母山省级自然保护区, 15个样方)的热带云雾林为研究对象, 研究α及β物种多样性、功能多样性、谱系多样性的变化, 为植物多样性的保护提供科学依据。结果表明: 尖峰岭群落树木个体多度、物种丰富度最大, 黎母山群落树木个体多度、物种丰富度最小; 黎母山群落间物种组成差异最大, 霸王岭群落间物种组成差异最小, 海南岛霸王岭(西部)、尖峰岭(西南部)和黎母山(中部) 3个林区热带云雾林物种多样性差异可能与空气温度和相对湿度有关。尖峰岭群落内功能丰富度、Rao’s二次熵最低, 功能均匀度最高, 群落间平均成对性状距离最小, 反映群落构建主要受环境筛影响; 霸王岭群落Rao’s二次熵最高, 功能均匀度最低, 群落间平均成对性状距离最大, 反映群落构建主要受限制相似性影响; 黎母山群落内功能丰富度最高, 群落间平均最近性状距离最大, 表明限制相似性在黎母山热带云雾林群落构建中的作用更重要。霸王岭群落内谱系多样性、物种间平均最近相邻谱系距离均较大, 反映物种间谱系关系趋于发散; 黎母山群落内谱系多样性、物种间平均成对谱系距离及群落间平均成对谱系距离均最小, 反映物种间谱系关系趋于聚集; 而尖峰岭群落内物种间平均成对谱系距离、群落间平均成对谱系距离最大, 但物种间平均最近相邻谱系距离最小, 反映物种间谱系关系既具有发散又具有聚集的共存格局。因此, 海南岛热带云雾林群落植物多样性变化格局与环境及物种间相互作用有关。     

Structural and taxonomic diversity predict above-ground biomass better than functional measures of maximum height in mixed-species forests

Aims: Mixed-species forests are known to be highly productive systems because of their high species diversity, including taxonomic diversity (species richness) and structural diversity. Recent empirical evidence also points to plant maximum height, as a functional trait that potentially drives forest above-ground biomass (AGB). However, the interrelations between these biotic variables are complex, and it is not always predictable if structural diversity attributes or functional metrics of plant maximum height would act as the most important determinant of stand biomass. Here we evaluated the relative importance of structural diversity attributes and functional metrics of plant maximum height (Hmax) in predicting and mediating AGB response to variation in species richness in mixed-species forests, while also accounting for fine-scale environmental variation. Location: Northern Benin. Methods: We used forest inventory data from mixed-species stands of native and exotic species. We quantified structural diversity as coefficient of variation of tree diameter at breast height (CVdbh) and of height (CVHt). For plant Hmax, we computed three metrics: functional range (FRHmax), functional divergence (FDHmax) and community-weighted mean (CWMHmax). We used topographical variables such as elevation and slope to account for possible environmental effects. Simple and multiple mixed-effects models, and structural equation models were performed to assess the direct and indirect links of AGB with species richness through structural diversity attributes and functional metrics of plant Hmax. Results: Species richness and CVdbh were positively related to AGB, while functional metrics of plant Hmax were not. Structural equation models revealed that species richness influenced AGB indirectly via CVdbh, which alone strongly promoted AGB. Elevation only had a positive direct effect on AGB. While increasing species richness enhanced CVdbh and functional measures of plant Hmax, there was no support for the latter mediating the effects of species richness on AGB. Conclusion: Structural diversity has a significant advantage in predicting and mediating the positive effect of species richness on AGB more so than functional measures of plant Hmax. We argue that structural diversity acts as a mechanism for the species richness–AGB relationship, and that maintaining high structural diversity would enhance biomass in mixed-species forests.     

Spatiotemporal patterns and dynamics of species richness and abundance of woody plant functional groups in a tropical forest landscape of Hainan Island, South China

Tropical forests are among the most species-diverse ecosystems on Earth. Their structures and ecological functions are complex to understand. Functional group is defined as a group of species that play similar roles in an ecosystem. The functional group approach has been regarded as an effective way of linking the compositions of complex ecosystems with their ecological functions. To understand the variation of functional groups in species-rich ecosystems after disturbance, the present study investigated the spatial pattern and temporal dynamics of woody plants in a typically fragmented natural forest landscape of Hainan Island in South China. The study area was classified into eight landscape types based on vegetation type, disturbance manner and the time of recovery. The woody plant species were aggregated into seven functional groups based on the growth form, successional status and plant size. The results gained from the present study showed that all functional groups, except for the emergent and canopy tree species, were present in all eight landscape types. Each landscape type had different numbers of dominant functional groups. There are similar species richness and stem abundance structure among functional groups between mid-successional clear cut lowland rainforest and old growth tropical coniferous forest. This similarity exists in selective logged lowland rainforest and old-growth lowland rainforest, as well as among landscape types of montane rainforest. The functional groups with the same successional status had similar patterns of species richness and stem abundance ratios among different landscape types. The variation patterns of functional groups along the successional stages in terms of species richness and stem abundance among the tropical lowland rainforest landscape types were more similar to each other than those in the tropical montane reinforest landscape types. This study provides further support for the competition-colonization tradeoff and successional niche theory as opposed to models of neutrality and ecological equivalence.     

The role of phytoplankton diversity metrics in shallow lake and river quality assessment

Ecological water quality problems are frequently connected to increment of phytoplankton productivity and overdominance of some phytoplankton species. Metrics that show monotonously increasing or decreasing tendencies along stressor gradients is recommended for ecological state assessment. Diversity metrics are influenced by various physical disturbances and show high within-year variability; thus, there is no agreement on the usefulness of these metrics as state indicators.To test the usefulness of phytoplankton diversity in ecological state assessment we investigated the productivity–diversity relationships for lakes and rivers in the Carpathian Basin (Hungary). We demonstrated that the shape of productivity–diversity relationship depends on the investigated water body type. Regarding lakes, hump-shaped relationship was found for all computed metrics. Parallel with the increase in phytoplankton productivity values, diversity metrics showed monotonously increasing tendencies in rhithral and decreasing tendencies in large potamal rivers. We found no systematic relationship in the case of small lowland rivers.Changes of diversity metrics calculated for species and functional groups showed similar tendencies within the types, only the slopes of regression lines differ each other.The use of diversity metrics as ecological state indicators should be restricted to water body types where diversity decreases or increases monotonously with phytoplankton biomass. Regarding the lakes the use of diversity metrics is not recommended for ecological state assessment. In rhithral and large potamal river assessment, application of diversity metrics should be strongly considered. We demonstrated that diversity metrics can be useful components of multimetric indices proposed to use by the Water Framework Directive.