(2011). Community assembly during secondary forest succession in a Chinese subtropical forest. Ecological Monographs, 81(1): 25-41. Community assembly during secondary forest succession in a Chinese subtropical forest Abstract Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30330 m plots (n=27) from five successional stages (<20, <40, <60, <80, and ≤80 yr), we investigated how the gradient in species composition reflects underlying processes of community assembly. In particular, we tested whether species richness of adult trees and shrubs decreased or increased and assessed to which degree this pattern was caused by negative density dependence or continuous immigration over time. Furthermore, we tested whether rare species were increasingly enriched and the species composition of adult trees and shrubs became more similar to species composition of seedlings during the course of succession. We counted the individuals of all adult species and shrubs >1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10310 m quadrant of each plot. In addition, we measured a number of environmental variables (elevation, slope, aspect, soil moisture, pH, C, N, and C/N ratio) and biotic structural variables (height and cover of layers). Adult species richness varied from 25 to 69 species per plot, and in total 148 woody species from 46 families were recorded. There was a clear successional gradient in species composition as revealed by nonmetric multidimensional scaling (NMDS), but only a poor differentiation of different successional stages with respect to particular species. Adult richness per 100 individuals (rarefaction method) increased with successional stage. None of the measured abiotic variables were significantly correlated with adult species richness. We found no evidence that rare species were responsible for the increasing adult species richness, as richness of rare species among both adults and recruits was independent of the successional stage. Furthermore, the similarity between established adults and recruits did not increase with successional stage. There was a constant number of recruit species and also of exclusive recruit species, i.e., those that had not been present as adult individuals, across all successional stages, suggesting a continuous random immigration over time. variables were significantly correlated with adult species richness. We found no evidence that 41 rare species were responsible for the increasing adult species richness, as richness of rare 42 species amongst both adults and recruits was independent of the successional stage.

Differences in herbivory among woody species can greatly affect the functioning of forest ecosystems, particularly in species-rich (sub)tropical regions. However, the relative importance of the different plant traits which determine herbivore damage remains unclear. Defence traits can have strong effects on herbivory, but rarely studied geographical range characteristics could complement these effects through evolutionary associations with herbivores. Herein, we use a large number of morphological, chemical, phylogenetic and biogeographical characteristics to analyse interspecific differences in herbivory on tree saplings in subtropical China. Unexpectedly, we found no significant effects of chemical defence traits. Rather, herbivory was related to the plants' leaf morphology, local abundance and climatic niche characteristics, which together explained 70% of the interspecific variation in herbivory in phylogenetic regression. Our study indicates that besides defence traits and apparency to herbivores, previously neglected measures of large-scale geographical host distribution are important factors influencing local herbivory patterns among plant species.

Experimental forest plantations to study biodiversity-ecosystem functioning (BEF) relationships have recently been established in different regions of the world, but subtropical biomes have not been covered so far. Here, we report about the initial survivorship of 26 tree species in the first such experiment in subtropical China. In the context of the joint Sino-German-Swiss Research Unit "BEFChina," 271 experimental forest plots were established using 24 naturally occurring tree species and two native commercial conifers. Based on the survival inventories carried out in November 2009 and June 2010, the overall survival rate was 87 % after the first 14 months. Generalized mixed-effects models showed that survival rates of seedlings were significantly affected by species richness, the species' leaf habit (deciduous or evergreen), species identity, planting date, and altitude. In the first survey, seedling establishment success decreased with increasing richness levels, a tendency that disappeared in the second survey after replanting. Though evergreen species performed less well than deciduous species with establishment rates of 84 versus 93 % in the second survey, their planting success exceeded the general expectation for subtropical broad-leaved evergreen species. These results have important implications for establishing mixed-species plantations for diversity conservation and improvement of ecosystem functioning in the Chinese subtropics and elsewhere. Additional costs associated with mixed-species plantations as compared to conventional plantations also demonstrate the potential of upscaling BEF experiments to large-scale afforestation projects. June 2010, the overall survival rate was 87% after the first 14 months. Generalized mixed-effects 22 models showed that survival rates of seedlings were significantly affected by species richness, the 23 species' leaf habit (deciduous or evergreen), species identity, planting date and altitude. In the first 24 survey, seedling establishment success decreased with increasing richness levels, a tendency that 25 disappeared in the second survey after replanting. Deciduous species had a significantly higher survival 26 than evergreen species (93% vs. 84% in the second survey). These results have implications for 27 establishing Though evergreen species performed less well than deciduous species with establishment 28 rates of 84% vs. 93% in the second survey, their planting success exceeded the general expectation for 29 subtropical broad-leave evergreen species. These results have important implications for establishing 30 mixed species plantations for diversity conservation and improvement of ecosystem functioning in the 31Chinese subtropics and elsewhere. Additional costs associated with mixed -species as compared to 32 conventional plantations also demonstrates the potential of upscaling BEF experiments to large-scale 33 afforestation projects. 34 35

Although trait analyses have become more important in community ecology, trait-environment correlations have rarely been studied along successional gradients. We asked which environmental variables had the strongest impact on intraspecific and interspecific trait variation in the community and which traits were most responsive to the environment. We established a series of plots in a secondary forest in the Chinese subtropics, stratified by successional stages that were defined by the time elapsed since the last logging activities. On a total of 27 plots all woody plants were recorded and a set of individuals of every species was analysed for leaf traits, resulting in a trait matrix of 26 leaf traits for 122 species. A Fourth Corner Analysis revealed that the mean values of many leaf traits were tightly related to the successional gradient. Most shifts in traits followed the leaf economics spectrum with decreasing specific leaf area and leaf nutrient contents with successional time. Beside succession, few additional environmental variables resulted in significant trait relationships, such as soil moisture and soil C and N content as well as topographical variables. Not all traits were related to the leaf economics spectrum, and thus, to the successional gradient, such as stomata size and density. By comparing different permutation models in the Fourth Corner Analysis, we found that the trait-environment link was based more on the association of species with the environment than of the communities with species traits. The strong species-environment association was brought about by a clear gradient in species composition along the succession series, while communities were not well differentiated in mean trait composition. In contrast, intraspecific trait variation did not show close environmental relationships. The study confirmed the role of environmental trait filtering in subtropical forests, with traits associated with the leaf economics spectrum being the most responsive ones.

No abstract

Biodiversity loss may alter ecosystem processes, such as herbivory, a key driver of ecological functions in species-rich (sub)tropical forests. However, the mechanisms underlying such biodiversity effects remain poorly explored, as mostly effects of species richness – a very basic biodiversity measure – have been studied. Here, we analyze to what extent the functional and phylogenetic diversity of woody plant communities affect herbivory along a diversity gradient in a subtropical forest.We assessed the relative effects of morphological and chemical leaf traits and of plant phylogenetic diversity on individual-level variation in herbivory of dominant woody plant species across 27 forest stands in south-east China.Individual-level variation in herbivory was best explained by multivariate, community-level diversity of leaf chemical traits, in combination with community-weighted means of single traits and species-specific phylodiversity measures. These findings deviate from those based solely on trait variation within individual species.Our results indicate a strong impact of generalist herbivores and highlight the need to assess food-web specialization to determine the direction of biodiversity effects. With increasing plant species loss, but particularly with the concomitant loss of functional and phylogenetic diversity in these forests, the impact of herbivores will probably decrease – with consequences for the herbivore-mediated regulation of ecosystem functions.

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

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity-Ecosystem Functioning experiment at Jiangxi (BEF-China). Information-theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi-layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi-predictor models for stomatal conductance (gs ) and xylem cavitation vulnerability. In addition, maximum gs was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum gs increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ50 ) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ50 values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.