Aims and Methods Mostly due to land use changes, European heathlands have become increasingly rare. In addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses and a loss in species diversity. Despite many investigations, information about the precise environmental parameters that determine the development and maintenance of heathland vegetation is still insufficient. In order to determine the environmental factors that control heath succession and grass encroachment, and to develop appropriate management schemes, we studied the influence of several soil and microclimate parameters on species composition and vegetation characteristics in five successional stages in a coastal heathland on the island of Hiddensee, north-east Germany, where the encroachment of Carex arenaria has become a major problem.
Important findings We recorded the highest plant species richness in grey dune and birch forest plots, while the encroachment of C. arenaria let to a significant decline in plant species richness. The most important environmental factors influencing species richness and distribution of single species were microclimate, soil moisture, soil pH and the C/N ratio. While many studies reported the importance of differences in nutrient availability, we found no significant correlations between soil nutrient availability and vegetation pattern. Environmental conditions in dense C. arenaria stands, especially soil properties (e.g. soil pH), showed great differences in comparison to the other successional stages. However, no correlations between the encroachment of C. arenaria and single environmental factors were found. Our results show that not only soil nutrients are important abiotic factors in heaths but that also microclimate and soil moisture play an important role and that many factors are involved in heath succession and in the promotion of grass encroachment. Management plans for the conservation and restoration of heathlands should therefore focus on the specific site conditions and should take several abiotic and biotic factors into account.

Aims The assembly of plant communities is a complex process which combines impacts from the species pool, dispersal and propagule pressure, niche requirements of colonizing species and the niche structure of the community. Recent theory development has incorporated all these aspects, e.g. in 'stochastic niche theory'. We investigated recruitment into a species-rich grassland community, using an experimental approach where we manipulated the trait composition of the community and examined the success of colonizing species entering with various propagule pressure. Specifically, we examined two predictions: (i) colonization success increases with increasing difference between traits of the colonizing species and the trait profile of the community and (ii) colonization success increases with increasing propagule pressure.
Methods The examined communities were species-rich semi-natural grasslands located in southern Sweden. After a careful documentation of the composition of the plant communities at the experimental sites, we manipulated the trait profile of species-rich grassland plots based on the plant functional trait specific leaf area (SLA), which is correlated with several key life history functions. In addition to SLA, seed mass was also used to describe the trait profile of grassland plots. Seeds of 12 plant species from the regional species pool, varying in SLA and seed mass, were sown into plots using four different levels of propagule pressure. Recruitment was examined after 1 year. We also planted juvenile 'plug plants' of the same species which allowed us to examine survivorship and growth beyond the seedling stage.
Important findings Overall we found very limited evidence for relationships between the traits of the colonizing species and the trait profile of the community and for recruitment after sowing these relationships were contrary to the prediction. Survival of plug plants after two seasons of growth was high irrespective of the trait profile of the community, but growth of plug plants was affected by the trait profile of the surrounding community. For four of the species there was a positive effect of increased propagule pressure on colonization. The results suggest that species assembly in species-rich grasslands is not strongly dependent on the niche structure of the community. However, the finding that colonization of only a third of the species responded positively to increased propagule pressure indicates that there might be niche-related effects that were not captured by our treatments. Overall, our results indicate that the factors determining colonization in this community are species specific. Some species are able to colonize irrespective of niche relationships, provided that the propagule pressure is sufficiently high to overcome stochastic mortality after seed arrival. For other species, however, we cannot exclude that niche assembly occurred, but we failed to identify the relevant niche factor.

Aims The dispersal of pollen and seeds is spatially restricted and may vary among plant populations because of varying biotic interactions, population histories or abiotic conditions. Because gene dispersal is spatially restricted, it will eventually result in the development of spatial genetic structure (SGS), which in turn can allow insights into gene dispersal processes. Here, we assessed the effect of habitat characteristics like population density and community structure on small-scale SGS and estimate historical gene dispersal at different spatial scales.
Methods In a set of 12 populations of the subtropical understory shrub Ardisia crenata, we assessed genetic variation at 7 microsatellite loci within and among populations. We investigated small-scale genetic structure with spatial genetic autocorrelation statistics and heterogeneity tests and estimated gene dispersal distances based on population differentiation and on within-population SGS. SGS was related to habitat characteristics by multiple regression.
Important findings The populations showed high genetic diversity (H e = 0.64) within populations and rather strong genetic differentiation (F ′ ST = 0.208) among populations, following an isolation-by-distance pattern, which suggests that populations are in gene flow–drift equilibrium. Significant SGS was present within populations (mean Sp = 0.027). Population density and species diversity had a joint effect on SGS with low population density and high species diversity leading to stronger small-scale SGS. Estimates of historical gene dispersal from between-population differentiation and from within-population SGS resulted in similar values between 4.8 and 22.9 m. The results indicate that local-ranged pollen dispersal and inefficient long-distance seed dispersal, both affected by population density and species diversity, contributed to the genetic population structure of the species. We suggest that SGS in shrubs is more similar to that of herbs than to trees and that in communities with high species diversity gene flow is more restricted than at low species diversity. This may represent a process that retards the development of a positive species diversity–genetic diversity relationship.

Aims With the continuing increase in the impact of human activities on ecosystems, ecologists are increasingly interested in understanding the effects of high temperature on litter decomposition since litter decomposition and the accompanying release of nutrients and carbon dioxide are key processes in ecosystem nutrient cycling and carbon flux. This study was conducted to evaluate the temperature sensitivity of forest litter decomposition and soil enzymes during litter decomposition in subtropical forest in China.
Methods Two dominant litter types were chosen from Zijin Mountain in China: Quercus acutissima leaves from a broadleaf forest (BF) and Pinus massoniana needles from a coniferous forest (CF). The litter samples were incubated in soil microcosms at ambient control temperature (20°C) and 10°C warmer. During a 5-month incubation, chemical composition of litter samples, litter mass losses, and related soil enzyme activities were determined.
Important findings Three main results were found: (i) high temperature accelerated decomposition rates of both litter types, and the temperature sensitivities of litter decomposition for BF leaves and that for CF needles are equivalent basically, (ii) high temperature enhanced soil enzyme activities in the two forest types, and the temperature sensitivities of polyphenol oxidase were significantly higher than those of the other soil enzymes and (iii) the temperature sensitivities of nitrate reductase were significantly higher in the CF soil than in the BF soil, while there was no significant difference in the temperature sensitivities of the other soil enzymes between BF and CF. As a long-term consequence, the high-temperature-induced acceleration of litter decomposition rates in these subtropical forests may cause carbon stored belowground to be transferred in the atmosphere, which may alter the balance between carbon uptake and release, and then alter the global carbon cycle in the coming decades.

Aims In eroded lands of the French Southern Alps, burial of early established seedlings under marly sediment weakens the effect of vegetation on soil stabilization and sediment trapping. Therefore, this protective role is largely dependent on species' resistance to burial, and the understanding of species' tolerance to this environmental disturbance is highly valuable for basic knowledge on plant succession and for ecological restoration purposes.
Methods The response of five woody species with contrasting ecological requirements and natural habitats—three tree species, Pinus nigra, Robinia pseudoacacia and Acer campestre, and two shrubs, Ononis fruticosa and Hippophae rhamnoides —to experimental burial under marly sediment was studied. Seedlings were exposed to three burial levels: no burial (control), partial burial (50% of seedling height) and complete burial (100% of seedling height). Burial tolerance was evaluated based on seedling survival, height and biomass. Biomass allocation to shoots and roots and soluble sugar and starch contents in roots and stems were measured to identify plant traits that determine species response to burial.
Important findings All species survived partial burial but only A. campestre seedlings emerged from complete burial. Tree species were more tolerant to burial and buried plants showed no significant differences with control. The two shrubs were found less tolerant and buried plants showed slower growth than controls. The results showed that species response was not related to initial soluble and starch content in roots and stems, but instead to biomass allocation pattern flexibility.

Aims Nighttime sap flow of trees may indicate transpiration and/or recharge of stem water storage at night. This paper deals with the water use of Acacia mangium at night in the hilly lands of subtropical South China. Our primary goal was to reveal and understand the nature of nighttime sap flow and its functional significance.
Methods Granier's thermal dissipation method was used to determine the nighttime sap flux of A. mangium. Gas exchange system was used to estimate nighttime leaf transpiration and stomatal conductance of studied trees.
Important findings Nighttime sap flow was substantial and showed seasonal variation similar to the patterns of daytime sap flow in A. mangium. Mean nighttime sap flow was higher in the less precipitation year of 2004 (1122.4 mm) than in the more precipitation year of 2005 (1342.5 mm) since more daytime transpiration and low soil water availability in the relatively dry 2004 can be the cause of more nighttime sap flow. Although vapor pressure deficit and air temperature were significantly correlated with nighttime sap flow, they could only explain a small fraction of the variance in nighttime sap flow. The total accumulated water loss (E L) by transpiration of canopy leaves was only ~2.6–8.5% of the total nighttime sap flow (E t) during the nights of July 17–18 and 18–19, 2006. Therefore, it is likely that the nighttime sap flow was mainly used for refilling water in the trunk. The stem diameter at breast height, basal area and sapwood area explained much more variance of nighttime water recharge than environmental factors and other tree form features, such as tree height, stem length below the branch, and canopy size. The contribution of nighttime water recharge to the total transpiration ranged from 14.7 to 30.3% depending on different DBH class and was considerably higher in the dry season compared to the wet season.

Aims Many observations concerning biological and ecological differentiation between narrow endemic and widespread congeneric plant species suggest that narrow endemic species are constrained to colonize marginal habitats because of a low tolerance to competition. Despite this topic being an important issue both for understanding evolutionary processes leading to endemism and for conservation purposes, few studies have been performed to compare competitive abilities between endemic and widespread species. Here, we present two independent experiments performed under controlled conditions using two different pairs of endemic and widespread congeneric species: Centaurea corymbosa / Centaurea maculosa and Arenaria provincialis / Arenaria serpyllifolia, both endemic species occurring in rocky calcareous habitats .
Methods Mature seeds of C. corymbosa and C. maculosa were sown in pots containing ramets of the common grass, Brachypodium retusum. Pots were sorted in three treatments according to grass cover (low, intermediate and high). A control treatment (without competition) was also used. Germination, seedling survival and rosette growth were followed. For the comparisons between A. provincialis and A. serpyllifolia, seeds from natural populations were first sown without a competitor. One week after germination, healthy seedlings were transplanted in pots without Brachypodium seedling (control) or containing two Brachypodium seedlings (low competition) or four seedlings (high competition). We checked the number of capsules per individual, and we harvested the biomass after capsule maturation.
Important findings Despite differences in the protocol design, results are congruent, and in both cases, endemic species are highly affected by the presence of a competitor, as are the widespread species, although we did not detect any differences between species for response to competition. The results are discussed in relation to processes leading to endemism, suggesting that the specialist model is more likely for both the study species. The present study also contributes to guidelines for the conservation of rare species in relation to landscape modification in the Mediterranean area.

Aims Our aim was to study how diversity and dominance of plant species and plant functional types (PFTs) change and covary across three dune fixation stages in the Chinese steppe zone.
Methods In the Chinese steppe zone, we measured coverage, mean height and density of each plant species in three types of dunes (mobile, semi-fixed and fixed dunes) in four sites (Mu Us, Otindag, Hulunbeir and Horqin). Plant species were grouped into 24 PFTs according to their lifespan, photosynthetic pathway, reproductive mode and life form. Dominance of each plant species and PFT were determined, and species diversity and PFT diversity were quantified using Shannon–Wiener index.
Important findings PFT diversity was positively related to plant species diversity in each dune stage, but PFT diversity increased more with increasing plant species diversity in the mobile and semi-fixed dunes than in the fixed dunes. Dune fixation stage explained 87.2% of the variation in plant species diversity and 84.8% of the variation in PFT diversity. Dominant species and PFTs differed among the three dune fixation stages; the more fixed the dunes were, the more perennial, shrubby, clonal and C₃ species co-dominated. Specifically, in mobile dunes annual C₄ non-clonal herbs were the most dominant, and in semi-fixed and fixed dunes perennial C₃ clonal shrubs were most dominant.

Aims Better understanding of microbial compositional and physiological acclimation mechanisms is critical for predicting terrestrial ecosystem responses to global change. The aim is to assess variations in soil microbial communities under future scenarios of changing precipitation and N deposition in a semiarid grassland of northern China.
Methods In order to explicitly estimate microbial responses, a field experiment with water and N addition was established in April 2005 and continuously conducted for 4 years. Specifically, soil microbial community composition and microbial C utilization potential were determined by phospholipid fatty acid (PLFA) and community-level physiological profiles, respectively.
Important findings Water addition had no effects on the PLFA concentrations of gram-positive (GP) and negative bacteria (GN), total bacteria and fungi. However, N addition caused significant reductions in the PLFA concentrations of GP, GN, total bacteria and fungi and thus decreased total PLFA of microbial communities. Moreover, there were interactive effects of water and N addition on GN/GP and the ratio of fungal to bacterial PLFA (F/B). In addition, synergistic effects were found between water and nitrogen in affecting microbial C utilization potentials, which implies that microbial C utilization potentials tend to be enhanced when both N and water availability are sufficient. Overall, the microbial responses to water and N addition support our hypothesis that water and N addition may be combined together to affect microbial communities in the semiarid grassland.

Aims Rhamnus lycioides L. subsp. Oleoides (Rhamnaceae) is a perennial shrub native to the Mediterranean Basin distributed along an altitudinal gradient, from sea level up to 1?000 m a.s.l. The specific goals of our study were (i) to compare plant morphology between two contrasting populations, (ii) to determine the reproductive system of R. lycioides, quantifying the relative importance of insects and wind as pollen vectors, (iii) to test if pollen limitation differs between populations, (iv) to study the main factors influencing fruit set and (v) to compare plant reproductive performance (mass allocation to flowers, fruits and seeds) between the two habitats.
Methods In the present study, we examined plant morphology and the reproductive performance of R. lycioides L. in contrasting environments in two populations located at the extremes of its altitudinal range in the island of Mallorca (Balearic Islands, Western Mediterranean Basin) along a 3-year period. Plant morphology, the relative importance of insects and wind as pollen vectors, the pollen limitation to seed production and the plant reproductive performance (mass allocation to flowers, fruits and seeds) were determined.
Important findings Rhamnus lycioides individuals showed a higher plant surface/plant height ratio at the mountain than at the coast. This species appeared to be ambophilous despite its inconspicuous flowers, although the relative importance of wind as a pollination vector was higher at the mountain than at the coastal site. Fruit set was much higher at the mountain, where pollen appeared to be a limiting factor. By contrast, fruit set was not limited by pollen availability at the coastal population, where resource (water and nutrients) limitation seemed to be more determining. Flower size was greater at the coast, in contrast to fresh fruit weight that was higher at the mountain. Despite the relatively few differences between sites in precipitation patterns along the study period, water availability appeared to be the key factor explaining not only fruit set but also the reproductive performance of this species in the study populations.

Aims Preserving and restoring Tamarix ramosissima is urgently required in the Tarim Basin, Northwest China. Using species distribution models to predict the biogeographical distribution of species is regularly used in conservation and other management activities. However, the uncertainty in the data and models inevitably reduces their prediction power. The major purpose of this study is to assess the impacts of predictor variables and species distribution models on simulating T. ramosissima distribution, to explore the relationships between predictor variables and species distribution models and to model the potential distribution of T. ramosissima in this basin.
Methods Three models—the generalized linear model (GLM), classification and regression tree (CART) and Random Forests—were selected and were processed on the BIOMOD platform. The presence/absence data of T. ramosissima in the Tarim Basin, which were calculated from vegetation maps, were used as response variables. Climate, soil and digital elevation model (DEM) data variables were divided into four datasets and then used as predictors. The four datasets were (i) climate variables, (ii) soil, climate and DEM variables, (iii) principal component analysis (PCA)-based climate variables and (iv) PCA-based soil, climate and DEM variables.
Important findings The results indicate that predictive variables for species distribution models should be chosen carefully, because too many predictors can reduce the prediction power. The effectiveness of using PCA to reduce the correlation among predictors and enhance the modelling power depends on the chosen predictor variables and models. Our results implied that it is better to reduce the correlating predictors before model processing. The Random Forests model was more precise than the GLM and CART models. The best model for T. ramosissima was the Random Forests model with climate predictors alone. Soil variables considered in this study could not significantly improve the model's prediction accuracy for T. ramosissima. The potential distribution area of T. ramosissima in the Tarim Basin is ~3.57 × 10 4 km 2, which has the potential to mitigate global warming and produce bioenergy through restoring T. ramosissima in the Tarim Basin.

Aims The spatial segregation hypothesis and the low-frequency hypothesis are two important proposed mechanisms that delay or prevent competitive exclusion in ecosystems. Because tree species interact with their neighbors, the importance of these potential processes can be investigated by analyzing the spatial structures of tree species.
Methods The distribution of the adults of 27 common tree species in a fully mapped 5-ha subtropical forest plot in Baishanzu, eastern China, was analyzed to investigate the community-level intra- and interspecific spatial association patterns. We first tested for the overall spatial pattern in the 5- to 40-m neighborhoods and classified first-order bivariate associations with a diametric scheme based on Ripley's K and nearest-neighbor statistic (G -function). Then heterogeneous Poisson null models were used to distinguish second-order interactions from overall spatial associations (including first-order effects). Finally, we analyzed correlations between the existence of species interactions and some attributes of the species involved.
Important findings Partial overlap and segregation increased with scale, whereas mixing decreased. Nearly 70% of the species pairs occurred less than expected at random, and only 3.4% of the species pairs were well mixed; 11.0% of all species pairs showed significant small-scale interactions, which was a greater frequency than expected by chance if species are abundant or prefer the same habitat, but less frequent than expected if species are highly aggregated. This suggests that both spatial segregation and low frequency of species facilitate species coexistence by reducing the opportunity that trees of two species encounter each other. The study also revealed that positive interactions were more prevalent than negative interactions in the forest, which indicates that positive interactions may have important effects on forest species assemblies.