Mingjian Yu

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%).

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

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.

Habitat fragmentation and biodiversity conservation: key findings and future challenges

Habitat loss and fragmentation has long been considered the primary cause for biodiversity loss and ecosystem degradation worldwide, and is a key research topic in landscape ecology (Wu 2013). Habitat fragmentation often refers to the reduction of continuous tracts of habitat to smaller, spatially distinct remnant patches, and habitat loss typically occurs concurrently with habitat fragmentation (Collinge 2009). Although some habitats are naturally patchy in terms of abiotic and biotic conditions (Wu and Loucks 1995), human actions have profoundly fragmented landscapes across the word (Haddad et al. 2015), altering the quality and connectivity of habitats. Therefore, understanding the causes and consequences of habitat fragmentation is critical to preserving biodiversity and ecosystem functioning. From May 4th to 10th, 2015, an International Workshop on Habitat Fragmentation and Biodiversity Conservation, held at the Thousand Island Lake, Zhejiang, China, discussed threats to biodiversity in fragmented landscapes and how fragmentation research can identify and help mitigate these threats. To meet these challenges, the Workshop had three goals. The first was to synthesize key findings in fragmentation science. Second was to identify important remaining research questions concerning the relationships between habitat fragmentation, biodiversity, and ecosystem functioning at local, regional, and global scales. Finally, we examined the unique roles of field-based fragmentation experiments in addressing these questions. The Workshop’s findings are relevant to the broader ecological community, and we present them here to stimulate research that will advance landscape ecology and conservation biology.

Determinants of plant species richness and patterns of nestedness in fragmented landscapes: Evidence from land-bridge islands

Land-bridge islands formed by dam construction are considered to be “experimental” systems for studying the effects of habitat loss and fragmentation, offering many distinct advantages over terrestrial fragments. The Thousand Island Lake in Southeast China is one such land-bridge system with more than 1000 islands. Based on a field survey of vascular plant richness on 154 land-bridge islands during 2007–2008, we examined the effects of island and landscape attributes on plant species richness and patterns of species nestedness. We also examined the different responses of plant functional groups (classified according to growth form and shade tolerance) to fragmentation. We found that island area explained the greatest amount of variation in plant species richness. Island area and shape index positively affected species diversity and the degree of nestedness exhibited by plant communities while the perimeter to area ratio of the islands had a negative effect. Shade-tolerant plants were the most sensitive species group to habitat fragmentation. Isolation negatively affected the degree of nestedness in herb and shade-intolerant plants including species with various dispersal abilities in the fragmented landscape. Based on these results, we concluded that the effects of habitat loss and fragmentation on overall species richness depended mostly on the degree of habitat loss, but patterns of nestedness were generated from different ecological mechanisms due to species-specific responses to different characteristics of habitat patches.

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.

The Effects of Landscape Variables on the Species-Area Relationship during Late-Stage Habitat Fragmentation

Few studies have focused explicitly on the later stages of the fragmentation process, or “late-stage fragmentation”, during which habitat area and patch number decrease simultaneously. This lack of attention is despite the fact that many of the anthropogenically fragmented habitats around the world are, or soon will be, in late-stage fragmentation. Understanding the ecological processes and patterns that occur in late-stage fragmentation is critical to protect the species richness in these fragments. We investigated plant species composition on 152 islands in the Thousand Island Lake, China. A random sampling method was used to create simulated fragmented landscapes with different total habitat areas and numbers of patches mimicking the process of late-stage fragmentation. The response of the landscape-scale species-area relationship (LSAR) to fragmentation per se was investigated, and the contribution of inter-specific differences in the responses to late-stage fragmentation was tested. We found that the loss of species at small areas was compensated for by the effects of fragmentation per se, i.e., there were weak area effects on species richness in landscapes due to many patches with irregular shapes and high variation in size. The study also illustrated the importance of inter-specific differences for responses to fragmentation in that the LSARs of rare and common species were differently influenced by the effects of fragmentation per se. In conclusion, our analyses at the landscape scale demonstrate the significant influences of fragmentation per se on area effects and the importance of inter-specific differences for responses to fragmentation in late-stage fragmentation. These findings add to our understanding of the effects of habitat fragmentation on species diversity.

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as βNRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.

Regeneration of different plant functional types in a Masson pine forest following pine wilt disease.

Pine wilt disease is a severe threat to the native pine forests in East Asia. Understanding the natural regeneration of the forests disturbed by pine wilt disease is thus critical for the conservation of biodiversity in this realm. We studied the dynamics of composition and structure within different plant functional types (PFTs) in Masson pine forests affected by pine wilt disease (PWD). Based on plant traits, all species were assigned to four PFTs: evergreen woody species (PFT1), deciduous woody species (PFT2), herbs (PFT3), and ferns (PFT4). We analyzed the changes in these PFTs during the initial disturbance period and during post-disturbance regeneration. The species richness, abundance and basal area, as well as life-stage structure of the PFTs changed differently after pine wilt disease. The direction of plant community regeneration depended on the differential response of the PFTs. PFT1, which has a higher tolerance to disturbances, became dominant during the post-disturbance regeneration, and a young evergreen-broad-leaved forest developed quickly after PWD. Results also indicated that the impacts of PWD were dampened by the feedbacks between PFTs and the microclimate, in which PFT4 played an important ecological role. In conclusion, we propose management at the functional type level instead of at the population level as a promising approach in ecological restoration and biodiversity conservation.

Disentangling the drivers of taxonomic and phylogenetic beta diversities in disturbed and undisturbed subtropical forests

Understanding the relative importance of dispersal limitation and environmental filtering processes in structuring the beta diversities of subtropical forests in human disturbed landscapes is still limited. Here we used taxonomic (TBD) and phylogenetic (PBD), including terminal PBD (PBDt) and basal PBD (PBDb), beta diversity indices to quantify the taxonomic and phylogenetic turnovers at different depths of evolutionary history in disturbed and undisturbed subtropical forests. Multiple linear regression model and distance-based redundancy analysis were used to disentangle the relative importance of environmental and spatial variables. Environmental variables were significantly correlated with TBD and PBDt metrics. Temperature and precipitation were major environmental drivers of beta diversity patterns, which explained 7–27% of the variance in TBD and PBDt, whereas the spatial variables independently explained less than 1% of the variation for all forests. The relative importance of environmental and spatial variables differed between disturbed and undisturbed forests (e.g., when Bray-Curtis was used as a beta diversity metric, environmental variable had a significant effect on beta diversity for disturbed forests but had no effect on undisturbed forests). We conclude that environmental filtering plays a more important role than geographical limitation and disturbance history in driving taxonomic and terminal phylogenetic beta diversity.