Species packing and the latitudinal gradient in beta-diversity.

The decline in species richness at higher latitudes is among the most fundamental patterns in ecology. Whether changes in species composition across space (beta-diversity) contribute to this gradient of overall species richness (gamma-diversity) remains hotly debated. Previous studies that failed to resolve the issue suffered from a well-known tendency for small samples in areas with high gamma-diversity to have inflated measures of beta-diversity. Here, we provide a novel analytical test, using beta-diversity metrics that correct the gamma-diversity and sampling biases, to compare beta-diversity and species packing across a latitudinal gradient in tree species richness of 21 large forest plots along a large environmental gradient in East Asia. We demonstrate that after accounting for topography and correcting the gamma-diversity bias, tropical forests still have higher beta-diversity than temperate analogues. This suggests that beta-diversity contributes to the latitudinal species richness gradient as a component of gamma-diversity. Moreover, both niche specialization and niche marginality (a measure of niche spacing along an environmental gradient) also increase towards the equator, after controlling for the effect of topographical heterogeneity. This supports the joint importance of tighter species packing and larger niche space in tropical forests while also demonstrating the importance of local processes in controlling beta-diversity.

Leaf functional traits of Parrotia subaequalis from different environments in eastern China.

Functional traits are important in understanding how plants respond and adapt to their immediate environment. Parrotia subaequalis is a highly endangered arbor species found throughout eastern China, primarily inhabiting hillsides and valleys, yet, little is known about the variation in leaf traits across these environments. In the present study, we tested this by comparing leaf surface area, leaf weight, leaf length, leaf symmetry and leaf mass per unit area, as well as the relationship between leaf traits and environmental factors and the scaling relationship between leaf surface area versus leaf dry mass. We observed significant differences in leaf surface area, weight, and length among the population sites, and these variables were strongly affected by environmental factors, especially high mean annual temperatures in hillside habitats and high mean annual precipitation in valley habitats. The scaling exponents remained numerically variant among the 10 populations, with different slopes greater than 1.0, and the scaling exponents increased significantly with hillside habitats. These metrics correlated with soil thickness associated with different habitat types. The areal ratio (AR) values in all populations deviated from 1, indicating that the two lamina sides were asymmetrical. The standardized symmetry index (SI) values displayed significant variation, especially in leaves from hillside habitats with a high degree of asymmetry. Collectively, our findings demonstrated that leaf functional traits exhibit considerable variability in response to different environmental contexts and provide valuable reference data that could be useful for conserving this endangered species.

Intraspecific leaf trait variation mediates edge effects on litter decomposition rate in fragmented forests.

There is strong trait dependence in species-level responses to environmental change and their cascading effects on ecosystem functioning. However, there is little understanding of whether intraspecific trait variation (ITV) can also be an important mechanism mediating environmental effects on ecosystem functioning. This is surprising, given that global change processes such as habitat fragmentation and the creation of forest edges drive strong trait shifts within species. On 20 islands in the Thousand Island Lake, China, we quantified intraspecific leaf trait shifts of a widely distributed shrub species, Vaccinium carlesii, in response to habitat fragmentation. Using a reciprocal transplant decomposition experiment between forest edge and interior on 11 islands with varying areas, we disentangled the relative effects of intraspecific leaf trait variation versus altered environmental conditions on leaf decomposition rates in forest fragments. We found strong intraspecific variation in leaf traits in response to edge effects, with a shift toward recalcitrant leaves with low specific leaf area and high leaf dry matter content from forest interior to the edge. Using structural equation modeling, we showed that such intraspecific leaf trait response to habitat fragmentation had translated into significant plant afterlife effects on leaf decomposition, leading to decreased leaf decomposition rates from the forest interior to the edge. Importantly, the effects of intraspecific leaf trait variation were additive to and stronger than the effects from local environmental changes due to edge effects and habitat loss. Our experiment provides the first quantitative study showing that intraspecific leaf trait response to edge effects is an important driver of the decrease in leaf decomposition rate in fragmented forests. By extending the trait-based response-effect framework toward the individual level, intraspecific variation in leaf economics traits can provide the missing functional link between environmental change and ecological processes. These findings suggest an important area for future research on incorporating ITV to understand and predict changes in ecosystem functioning in the context of global change.

Higher trophic levels and species with poorer dispersal traits are more susceptible to habitat loss on island fragments.

Ongoing habitat loss and fragmentation caused by human activities represent one of the greatest causes of biodiversity loss. However, the effects of habitat loss and fragmentation are not felt equally among species. Here, we examined how habitat loss influenced the diversity and abundance of species from different trophic levels, with different traits, by taking advantage of an inadvertent experiment that created habitat islands from a once continuous forest via the creation of the Thousand Island Lake, a large reservoir in China. On 28 of these islands with more than a 9000-fold difference in their area (0.12-1154 ha), we sampled plants, herbivorous insects, and predatory insects using effort-controlled sampling and analyses. This allowed us to discern whether any observed differences in species diversity were due to passive sampling alone or to demographic effects that disproportionately influenced some species relative to others. We found that while most metrics of sampling effort-controlled diversity increased with island area, the strength of the effect was exacerbated for species in higher trophic levels. When we more explicitly examined differences in species composition among islands, we found that the pairwise difference in species composition among islands was dominated by species turnover but that nestedness increased with differences in island area, indicating that some species are more likely to be absent from smaller islands. Furthermore, by examining trends of several dispersal-related traits of species, we found that species with lower dispersal propensity tended to be those that were lost from smaller islands, which was observed for herbivorous and predatory insects. Our results emphasize the importance of incorporating within-patch demographic effects, as well as the taxa and traits of species when understanding the influence of habitat loss on biodiversity.

Quantifying effects of habitat heterogeneity and other clustering processes on spatial distributions of tree species.

Spatially explicit consideration of species distribution can significantly add to our understanding of species coexistence. In this paper, we evaluated the relative importance of habitat heterogeneity and other clustering processes (e.g., dispersal limitation, collectively called the non-habitat clustering process) in explaining the spatial distribution patterns of 341 tree species in three stem-mapped 25-50 ha plots of tropical, subtropical, and temperate forests. Their relative importance was estimated by a method that can take one mechanism into account when estimating the effects of the other mechanism and vice versa. Our results demonstrated that habitat heterogeneity was less important in explaining the observed species patterns than other clustering processes in plots with flat topography but was more important in one of the three plots that had a complex topography. Meanwhile, both types of clustering mechanisms (habitat or non-habitat) were pervasive among species at the 50-ha scale across the studied plots. Our analyses also revealed considerable variation among species in the relative importance of the two types of mechanism within each plot and showed that this species-level variation can be partially explained by differences in dispersal mode and growth form of species in a highly heterogeneous environment. Our findings provide new perspectives on the formation of species clustering. One important finding is that a significant species-habitat association does not necessarily mean that the habitat heterogeneity has a decisive influence on species distribution. The second insight is that the large species-level variation in the relative importance of the two types of clustering mechanisms should not be ignored. Non-habitat clustering processes can play an important role on species distribution.

Climate and fragment area jointly affect the annual dynamics of seedlings in different functional groups in the Thousand Island Lake.

Habitat fragmentation and climate change are the two main threats to global biodiversity. Understanding their combined impact on plant community regeneration is vital for predicting future forest structure and conserving biodiversity. This study monitored the seed production, seedling recruitment and mortality of woody plants in the Thousand Island Lake, a highly fragmented anthropogenic archipelago, for 5 years. We analyzed the seed-seedling transition, seedling recruitment and mortality of different functional groups in the fragmented forests and conducted correlation analyses involving climatic factors, island area, and plant community abundance. Our results showed that: 1) shade-tolerant and evergreen species had higher seed-seedling transition, seedling recruitment and survival rate than shade-intolerant and deciduous species in time and space, and these advantages increased with the island area. 2) Seedlings in different functional groups responded differently to island area, temperature and precipitation. 3) Increasing active accumulated temperature (the sum of the mean daily temperature above 0 °C) significantly increased seedling recruitment and survival, and warming climate favored the regeneration of evergreen species. 4) The seedling mortality rate of all plant functional groups increased with the increase of island area, but the increasing strength weakened significantly with the increase of the annual maximum temperature. These results suggested that the dynamics of woody plant seedlings varied among functional groups, and can be regulated separately and jointly by fragmentation and climate.

Linkages between Plant Community Composition and Soil Microbial Diversity in Masson Pine Forests.

Plant species identity influences soil microbial communities directly by host specificity and root exudates, and indirectly by changing soil properties. As a native pioneer species common in early successional communities, Masson pine (Pinus massoniana) forests are widely distributed in subtropical China, and play a key role in improving ecosystem productivity. However, how pine forest composition, especially the dominance of plant functional groups, affects soil microbial diversity remains unclear. Here, we investigated linkages among woody plant composition, soil physicochemical properties, and microbial diversity in forests along a dominance gradient of Masson pine. Soil bacterial and fungal communities were mainly explained by woody plant community composition rather than by woody species alpha diversity, with the dominance of tree (without including shrub) species and ectomycorrhizal woody plant species accounting for more of the variation among microbial communities than pine dominance alone. Structural equation modeling revealed that bacterial diversity was associated with woody plant compositional variation via altered soil physicochemical properties, whereas fungal diversity was directly driven by woody plant composition. Bacterial functional groups involved in carbohydrate and amino acid metabolism were negatively correlated with the availability of soil nitrogen and phosphorus, whereas saprotrophic and pathogenic fungal groups showed negative correlations with the dominance of tree species. These findings indicate strong linkages between woody plant composition than soil microbial diversity; meanwhile, the high proportion of unexplained variability indicates great necessity of further definitive demonstration for better understanding of forest-microbe interactions and associated ecosystem processes.

Plant life history strategies vary in subtropical forests with different disturbance histories: an assessment of biodiversity, biomass, and functional traits.

Disturbance alters environmental conditions in forests. Plants growing in forests with different disturbance histories in diverse environments may adopt varying life history strategies, but few studies focus on this effect. This study comprehensively investigated plant biodiversity, biomass, and functional traits in subtropical forests with two different disturbance histories in east China to explore differences in life history strategies. Biodiversity was slightly higher in disturbed compared to conserved forests. Significantly higher biomass was measured in conserved relative to disturbed evergreen broadleaved forests (P < 0.05). In conserved forests, leaf tissue density (LTD) was significantly higher and leaf thickness (LT), leaf dry matter content (LDMC), twig tissue density (TTD), twig dry matter content (TDMC), bark tissue density (BTD) and dry matter content (BDMC), and stem tissue density (STD) and dry matter content (SDMC) were significantly lower than in disturbed forests (P < 0.05). In terms of associated plant biodiversity, biomass, and functional traits, conserved forests adopted a resource acquisition strategy, reducing biodiversity and developing multiple functional traits such as high leaf area and specific leaf area and low LT, LDMC, TTD, TDMC, BTD, BDMC, STD, and SDMC to support a high biomass accumulation rate. Disturbed forests adopted a resource conservation strategy, enhancing biodiversity and developing converse trait combinations to lower the rate of biomass accumulation. A comprehensive investigation of plant biodiversity, biomass, and functional traits and subsequent assessment of plant life history strategies in conserved and disturbed forests will aid investigations of regional biodiversity and carbon reserves, contribute data to the TRY and Chinese plant trait databases, and improve ecological management and restoration efforts in east China.

Carex borealifujianica (Cyperaceae), a new species of the core Carex clade from Fujian, southeastern China.

A new species, Carex borealifujianica Y.F. Lu & X.F. Jin (Cyperaceae, sect. Occlusae of core Carex clade) is described and illustrated from northern Fujian, China. In addition to morphological comparisons with its relatives, comparative micromorphology of utricles and achenes of seven species in Carex sect. Occlusae was examined. Micromorphology of utricles and achenes revealed the similarity of Carex borealifujianica and C. ligulata. Morphologically, this new species is similar to Carex ligulata in having lateral spikes remote and densely flowered, as well as utricles densely hispidulous, but differs in having 2 or 3 narrowly clavate staminate spikes, leaves 2.5-5 mm wide with sheaths sparsely pilose, and achenes emarginate at the apex. The phylogenetic analysis from two nuclear DNA regions (ETS and ITS) and two chloroplast DNA regions (matK and trnL-F) of 68 taxa resolved C. borealifujianica as a distinct species.

Differences in Density Dependence among Tree Mycorrhizal Types Affect Tree Species Diversity and Relative Growth Rates.

Conspecific negative density dependence (CNDD) may vary by tree mycorrhizal type. However, whether arbuscular mycorrhizal (AM)-associated tree species suffer from stronger CNDD than ectomycorrhizal (EcM) and ericoid mycorrhizal (ErM)-associated tree species at different tree life stages, and whether EcM tree species can promote AM and ErM saplings and adults growth, remain to be studied. Based on the subtropical evergreen broad-leaved forest data in eastern China, the generalized linear mixed-effects model was used to analyze the effects of the conspecific density and heterospecific density grouped by symbiont mycorrhizal type on different tree life stages of different tree mycorrhizal types. The results showed that compared to other tree mycorrhizal types at the same growth stage, EcM saplings and AM adults experienced stronger CNDD. Heterospecific EcM density had a stronger positive effect on AM and ErM individuals. Species diversity and average relative growth rate (RGR) first increased and then decreased with increasing basal area (BA) ratios of EcM to AM tree species. These results suggested that the stronger CNDD of EcM saplings and AM adults favored local species diversity over other tree mycorrhizal types. The EcM tree species better facilitated the growth of AM and ErM tree species in the neighborhood, increasing the forest carbon sink rate. Interestingly, species diversity and average RGR decreased when EcM or AM tree species predominated. Therefore, our study highlights that manipulating the BA ratio of EcM to AM tree species will play a nonnegligible role in maintaining biodiversity and increasing forest carbon sink rates.

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It is of great significance to understand the pattern of soil respiration rate in fragmented forests for further revealing terrestrial ecosystem carbon cycling. With different habitats (island vs. mainland, island edge vs. island interior) of the artificial land-bridge island system in Thousand Island Lake (TIL) region as the objects, we analyzed the seasonal dynamics of soil respiration rate and its relationships with soil physicochemical factors. The results showed that: 1) Soil respiration rates varied significantly across different seasons, with an order of summer (3.74 µmol·m-2·s-1) > autumn (2.30 µmol·m-2·s-1) > spring (1.82 µmol·m-2·s-1) > winter (1.40 µmol·m-2·s-1). 2) Forest fragmentation had significant effects on soil respiration rate, with soil respiration rate of island (2.37 µmol·m-2·s-1) being significantly higher than that of mainland (2.08 µmol·m-2·s-1) and the soil respiration rate of island edge (2.46 µmol·m-2·s-1) being significantly higher than that of island interior (2.03 µmol·m-2·s-1). 3) Soil temperature significantly promoted soil respiration rate, explaining 56.1% of the total variation. 4) There was a significant positive correlation between soil respiration rate and soil total carbon, ammo-nium nitrogen content, and vegetation coverage. The soil total carbon and ammonium nitrogen content of island edge were significantly higher than those of island interior. In all, forest fragmentation promoted soil respiration rate, with soil physicochemical factors as the drivers for its variation.

Phylogenetic Diversity of Wetland Plants across China.

Accelerating and severe wetland loss has made wetland restoration increasingly important. Current wetland restorations do not take into consideration the ecological adaptability of wetland plants at large scales, which likely affects their long-term restoration success. We explored the ecological adaptability, including plant life forms and phylogenetic diversity, of plants across 28 wetlands in China. We found that perennial herbs were more common than annual herbs, with the proportion of perennial herbs accounting for 40-50%, 45-65%, 45-70%, 50-60%, and 60-80% of species in coastal wetlands, human-made wetlands, lake wetlands, river wetlands, and marsh wetlands, respectively. A ranking of phylogenetic diversity indices (PDIs) showed an order of marsh < river < coastal < lake < human-made, meaning that human-made wetlands had the highest phylogenetic diversity and marsh wetlands had the lowest phylogenetic diversity. The nearest taxon index (NTI) was positive in 23 out of 28 wetlands, indicating that species were phylogenetically clustered in wetland habitats. Dominant species tended to be distantly related to non-dominant species, as were alien invasive species and native species. Our study indicated that annual herbs and perennial herbs were found in different proportions in different types of wetlands and that species were phylogenetically clustered in wetland habitats. To improve wetland restoration, we suggest screening for native annual herbs and perennial herbs in proportions that occur naturally and the consideration of the phylogenetic similarity to dominant native species.

Alien, Naturalized and Invasive Plants in China.

Invasive species are a subset of naturalized species, and naturalized species are a subset of alien species. Determining the relationship among these three assemblages would be helpful in predicting and preventing biological invasion. Here, we reviewed the families, lifeforms, origins, introduction pathways and phylogenetic diversity of alien, naturalized and invasive vascular plants in China. The results show that species in the Asteraceae, Fabaceae and Poaceae families had a high dominance among alien, naturalized and invasive species. Moreover, almost all alien species in the Amaranthaceae, Solanaceae, Convolvulaceae and Euphorbiaceae families became naturalized species, and about 26.7% of the naturalized species became invasive species. Perennial herbs comprised a higher proportion of alien species than did annual herbs, though annual herbs were more suited to becoming invasive than perennial herbs. A considerable proportion (57.8%) of invasive species were introduced from America. More than half (56.5%) of alien species were introduced for their ornamental value, and half of these have become naturalized in China. Moreover, about half (55.2%) of all invasive species were introduced for their economic value (including ornamental, foraging and medicinal purposes). Invasive species were phylogenetically clustered and phylogenetically distant from alien and naturalized species, which indicates that phylogenetic differences could be helpful in becoming invasive. There is no doubt that human activity plays a significant role in biological invasion. This study suggests that when introducing alien species to a region, decision-makers should certainly consider the species' phylogeny, beyond just its fundamental characteristics.

Disentangling biotic and abiotic drivers of intraspecific trait variation in woody plant seedlings at forest edges.

In fragmented forests, edge effects can drive intraspecific variation in seedling performance that influences forest regeneration and plant composition. However, few studies have attempted to disentangle the relative biotic and abiotic drivers of intraspecific variation in seedling performance. In this study, we carried out a seedling transplant experiment with a factorial experimental design on three land-bridge islands in the Thousand Island Lake, China, using four common native woody plant species. At different distances from the forest edge (2, 8, 32, 128 m), we transplanted four seedlings of each species into each of three cages: full-cage, for herbivore exclusion; half-cage, that allowed herbivore access but controlled for caging artifacts; and no-cage control. In the 576 cages, we recorded branch architecture, leaf traits, and seedling survival for each seedling before and after the experimental treatment. Overall, after one full growing season, edge-induced abiotic drivers and varied herbivory pressure led to intraspecific variation in seedling performance, including trade-offs in seedling architecture and resource-use strategies. However, responses varied across species with different life-history strategies and depended on the driver in question, such that the abiotic and biotic effects were additive across species, rather than interactive. Edge-induced abiotic variation modified seedling architecture of a shade-tolerant species, leading to more vertical rather than lateral growth at edges. Meanwhile, increased herbivory pressure resulted in a shift toward lower dry matter investment in leaves of a light-demanding species. Our results suggest that edge effects can drive rapid directional shifts in the performance and intraspecific traits of some woody plants from early ontogenetic stages, but most species in this study showed negligible phenotypic responses to edge effects. Moreover, species-specific responses suggest the importance of interspecific differences modulating the degree of trait plasticity, implying the need to incorporate individual-level responses when understanding the impact of forest fragmentation on plant communities.

Spatiotemporal patterns and ecological consequences of a fragmented landscape created by damming.

BACKGROUND: Damming disrupts rivers and destroys neighboring terrestrial ecosystems through inundation, resulting in profound and long-lasting impacts on biodiversity and ecosystem processes far beyond the river system itself. Archipelagos formed by damming are often considered ideal systems for studying habitat fragmentation. METHODS: Here we quantified the island attributes and landscape dynamics of the Thousand Island Lake (TIL) in China, which is one of the several long-term biodiversity/fragmentation research sites around the world. We also synthesized the major findings of relevant studies conducted in the region to further ecological understanding of damming and landscape fragmentation. RESULTS: Our results show that the vegetations on islands and the neighboring mainland were both recovering between 1985 and 2005 due to reforestation and natural succession, but the regeneration was partly interrupted after 2005 because of increasing human influences. While major changes in landscape composition occurred primarily in the lakefront areas and near-lakeshore islands, landscape patterns became structurally more complex and fragmented on both islands and mainland. About 80 studies from the TIL region show that the genetic, taxonomic, functional, and phylogenetic diversity on these islands were mainly influenced by island area at the patch scale, but fragmentation per se also affected species composition and related ecological processes at patch and landscape scales. In general, islands had lower species diversity but a steeper species-area relationship than the surrounding mainland. Fragmentation and edge effects substantially hindered ecological succession towards more densely vegetated forests on the islands. Environmental heterogeneity and filtering had a major impact on island biotic communities. We hypothesize that there are multiple mechanisms operating at different spatial scales that link landscape fragmentation and ecological dynamics in the TIL region, which beg for future studies. By focusing on an extensive spatiotemporal analysis of the island-mainland system and a synthesis of existing studies in the region, this study provides an important foundation and several promising directions for future studies.

Partitioning beta diversity in a subtropical broad-leaved forest of China.

The classical environmental control model assumes that species distribution is determined by the spatial variation of underlying habitat conditions. This niche-based model has recently been challenged by the neutral theory of biodiversity which assumes that ecological drift is a key process regulating species coexistence. Understanding the mechanisms that maintain biodiversity in communities critically depends on our ability to decompose the variation of diversity into the contributions of different processes affecting it. Here we investigated the effects of pure habitat, pure spatial, and spatially structured habitat processes on the distributions of species richness and species composition in a recently established 24-ha stem-mapping plot in the subtropical evergreen broad-leaved forest of Gutianshan National Nature Reserve in East China. We used the new spatial analysis method of principal coordinates of neighbor matrices (PCNM) to disentangle the contributions of these processes. The results showed that (1) habitat and space jointly explained approximately 53% of the variation in richness and approximately 65% of the variation in species composition, depending on the scale (sampling unit size); (2) tree diversity (richness and composition) in the Gutianshan forest was dominantly controlled by spatially structured habitat (24%) and habitat-independent spatial component (29%); the spatially independent habitat contributed a negligible effect (6%); (3) distributions of richness and species composition were strongly affected by altitude and terrain convexity, while the effects of slope and aspect were weak; (4) the spatial distribution of diversity in the forest was dominated by broad-scaled spatial variation; (5) environmental control on the one hand and unexplained spatial variation on the other (unmeasured environmental variables and neutral processes) corresponded to spatial structures with different scales in the Gutianshan forest plot; and (6) five habitat types were recognized; a few species were statistically significant indicators of three of these habitats, whereas two habitats had no significant indicator species. The results suggest that the diversity of the forest is equally governed by environmental control (30%) and neutral processes (29%). In the fine-scale analysis (10 x 10 m cells), neutral processes dominated (43%) over environmental control (20%).

Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.

Species-area relationships (SARs) characterize the spatial distribution of species diversity in community ecology, but the biological mechanisms underlying the SARs have not been fully explored. Here, we examined the roles of dispersal limitation and habitat heterogeneity in shaping SARs in two large-scale forest plots. One is a 24-ha subtropical forest in Gutianshan National Nature Reserve, China. The other is a 50-ha tropical rain forest in Barro Colorado Island, Panama. Spatial point pattern models were applied to investigate the contributions of dispersal and habitat heterogeneity and their interactions to the formation of the SARs in the two sites. The results showed that, although dispersal and habitat heterogeneity each could significantly contribute to the SARs, each alone was insufficient to explain the SARs. Their joint effects sufficiently explained the real SARs, suggesting that heterogeneous habitat and dispersal limitation are two predominant mechanisms for maintaining the spatial distributions of the species in these two forests. These results add to our understanding of the ecological processes underlying the spatial variation of SARs in natural forest communities.

Decoupling species richness variation and spatial turnover in beta diversity across a fragmented landscape.

BACKGROUND: How habitat fragmentation affects the relationship between local richness and the variation in community composition across space is important to both ecology and conservation biology, but this effect remains poorly understood. METHODS: Here, we present an empirical study to address this topic in a fragmented landscape, the Thousand Island Lake (TIL), an artificial land-bridge island system with more than 1,000 islands, which provides an "experimental" fragmented landscape with a homogeneous matrix and similar successional history. We measured species composition and plant functional type (PFT) on 29 islands, and tested the effects of island area and isolation on the relationship between α- and ß-diversity. General Linear Models were applied to test the impact of habitat fragmentation. In addition, variation partitioning was used to decouple α-diversity dependent and α-diversity independent spatial turnover in ß-diversity of the plant community and across different PFTs. RESULTS: We found habitat fragmentation influences ß-diversity of plants primarily by modifying local α-diversity, not spatial turnover in the TIL system. We also found area-dependent environmental filtering and differential plant responses across functional types were the most likely underlying driving mechanisms. DISCUSSION: These results highlight the importance of hierarchical linkages between components of biodiversity across scales in fragmented landscapes, and have practical conservation implications.

Diversity and density patterns of large old trees in China.

Large old trees are keystone ecological structures that provide vital ecosystem services to humans. However, there are few large-scale empirical studies on patterns of diversity and density of large old trees in human-dominated landscapes. We present the results of the first nationwide study in China to investigate the patterns of diversity and density of large old trees in human-dominated landscapes. We collated data on 682,730 large trees ≥100 years old from 198 Chinese regions to quantify tree species diversity, tree density and maximum tree age patterns. We modelled the effects of natural environmental variables (e.g. climate and topography) and anthropogenic variables (e.g. human population density and city age) on these measures. We found a low density of large old trees across study regions (0.36 trees/km2), and large variation in species richness among regions (ranging from 1 to 232 species). More than 95% of trees were <500 years old. The best fit models showed that: (1) Species diversity (species richness adjusted by region size) was positively associated with mean annual rainfall and city age; (2) Density of clustered trees, which are mostly remnants of ancient woods, was negatively influenced by human population density and rural population (% of total population). In contrast, the density of scattered trees, which are mostly managed by local people, was positively correlated with mean annual rainfall and human population density. To better protect large old trees in cities and other highly-populated areas, conservation policy should protect ancient wood remnants, mitigate the effects environmental change (e.g. habitat fragmentation), minimize the negative effects of human activities (e.g. logging), and mobilize citizens to participate in conservation activities (e.g. watering trees during droughts).

Forest fragmentation in China and its effect on biodiversity.

Land-use change is fragmenting natural ecosystems, with major consequences for biodiversity. This paper reviews fragmentation trends - historical and current - in China, the fourth largest country on Earth, and explores its consequences. Remote sensing makes it possible to track land-use change at a global scale and monitor fragmentation of dwindling natural landscapes. Yet few studies have linked fragmentation mapped remotely with impacts on biodiversity within human-modified landscapes. Recent reforestation programs have caused substantial increases in forest cover but have not stopped fragmentation, because the new forests are mostly monocultures that further fragment China's remnant old-growth lowland forests that harbour the highest levels of biodiversity. Fragmentation - and associated biodiversity declines - is unevenly distributed in China's forests, being most problematic where agricultural expansion is occurring in the southwest and northeast, serious in the densely populated eastern regions where urbanisation and transport infrastructure are modifying landscapes, but less of a problem in other regions. Analyses of temporal trends show that the drivers of forest fragmentation are shifting from mainly agricultural expansion to urbanisation and infrastructure development. Most of China's old-growth forests persist in small, isolated fragments from which many native species have disappeared, on land unsuitable for human utilisation. Fragmentation throughout China is likely to have major consequences on biodiversity conservation, but few studies have considered these large-scale processes at the national level. Our review fills this research gap and puts forward a systematic perspective relevant to China and beyond.