Aims Mangrove species are classified as true mangroves and mangrove associates. However, as for some fringe species found mainly on the landward transitional zones of mangroves, no consensus among scientists could be reached in favor of this classification and much debate arises. We hypothesized that true mangroves differ from mangrove associates physiologically and ecologically in their ability to survive in mangrove environment.
Methods To test this hypothesis, leaf structural traits and osmotic properties were used to describe variation in 33 mangrove species (17 true mangroves, 6 mangrove associates and 10 controversial species).
Important findings Specific leaf area (SLA) of true mangroves as well as leaf nitrogen concentration on a leaf mass (Nmass) were lower than that of mangrove associates; leaf succulence was, in general, twice as high in true mangroves compared to mangrove associates; true mangroves accumulated 8–9 times more Na and Cl than mangrove associates and the former had K/Na ratios <0.5, but the latter had K/Na ratios>0.5. These results indicated that true mangroves differed reliably from mangrove associates in leaf traits and osmotic properties. True mangroves are true halophytes and mangrove associates are glycophytes with certain salt tolerance. Combining distribution pattern information, the 10 controversial species were reclassified.

Aims The aim of this article is 3-fold. First, we present an updated version of a published megaphylogeny of vascular plants that can be used in studies of plant ecology and biogeography. Second, we develop a tool that can be used by botanists and plant ecologists to generate phylogenetic hypotheses in three scenarios. Third, we use a set of regional assemblages of angiosperm trees in North America as a model system to evaluate the effect of differences in phylogenies generated using the three scenarios on the quantification of phylogenetic properties and the relationship between measures of phylogenetic properties and environment.
Methods The taxonomy and nomenclature of plant species in the megaphylogeny were standardized according to The Plant List (version 1.1). A tool for generating phylogenies was created using the R language. The robustness of derived phylogenies was evaluated using correlation and regression analyses.
Important findings An updated megaphylogeny of vascular plants (PhytoPhylo) and a tool for reconstructing phylogenies of seed plants (S.PhyloMaker) were generated. Our study shows that phylogenies generated by S.PhyloMaker using the PhytoPhylo megaphylogeny as a backbone are nearly as good as phylogeny resolved at the species level when using derived phylogenies to quantify phylogenetic properties (e.g. phylogenetic diversity and phylogenetic relatedness) of biological assemblages, and that S.PhyloMaker-generated phylogenies are robust for studies of community ecology and biogeography, particularly those seeking for patterns of phylogenetic properties along environmental gradients.

Aims The aim of this guide is to provide practical help for ecologists who analyze data from biodiversity–ecosystem functioning experiments. Our approach differs from others in the use of least squares-based linear models (LMs) together with restricted maximum likelihood-based mixed models (MMs) for the analysis of hierarchical data. An original data set containing diameter and height of young trees grown in monocultures, 2- or 4-species mixtures under ambient light or shade is used as an example.
Methods Starting with a simple LM, basic features of model fitting and the subsequent analysis of variance (ANOVA) for significance tests are summarized. From this, more complex models are developed. We use the statistical software R for model fitting and to demonstrate similarities and complementarities between LMs and MMs. The formation of contrasts and the use of error (LMs) or random-effects (MMs) terms to account for hierarchical data structure in ANOVAs are explained.
Important findings Data from biodiversity experiments can be analyzed at the level of entire plant communities (plots) and plant individuals. The basic explanatory term is species composition, which can be divided into contrasts in many ways depending on specific biological hypotheses. Typically, these contrasts code for aspects of species richness or the presence of particular species. For significance tests in ANOVAs, contrast terms generally are compared with remaining variation of the explanatory terms from which they have been 'carved out'. Once a final model has been selected, parameters (e.g. means or slopes for fixed-effects terms and variance components for error or random-effects terms) can be estimated to indicate the direction and size of effects.

Aims Soil plays an important role in the formation and heterogeneity of habitats and thus can cause changes in vegetation structure and plant diversity. The differentiation between Cerrado/savanna and forest is well known, but the relationship between soil and habitats from savannic or forest formations still needs to be better understood, particularly in tropical ecotonal areas. We studied the association between attributes of plant communities, namely structure and diversity, and physicochemical characteristics of soils in the Caatinga domain at the transition to Cerrado in Brazil.
Methods Chemical and physical analyses of soils were performed in samples of 38 plots from savannic formations and 30 plots from forest formations. Vegetation was characterized floristically and structurally in all plots, five habitats being assessed in each plant formation. Soil features and vegetation parameters were highly distinct among the different habitats.
Important findings In general, forest habitats were more nutrient rich than savannic formation. Furthermore, soil variables showed effects both on vegetation structure and on its species diversity, more pronouncedly in the savannic formations. Habitats were structurally distinct, and diversity differed between savannic and forest communities; however, a higher differentiation occurred when the savannic formation habitats were compared among them. Although plant diversity did not differ among forest formation habitats, soil attributes showed a close relationship with edaphic factors and can contribute for similar vegetation. The soil–vegetation relationship in highly diverse ecotonal landscapes is important from the conservation biology point of view and aid in the execution of proactive plans for the maintenance of biodiversity. Thus, we noticed that diversity and soil behaves distinctly between savannic and forest communities.

Morphological variation of leaves is a key indicator of plant response to climatic change. Leaf size and shape are associated with carbon, water and energy exchange of plants with their environment. However, whether and how leaf size and shape responded to climate change over the past decades remains poorly studied. Moreover, many studies have only explored inter- but not intraspecific variation in leaf size and shape across space and time.

We collected >6000 herbarium specimens spanning 98 years (1910–2008) in China for seven representative dicot species and measured their leaf length and width. We explored geographical patterns and temporal trends in leaf size (i.e. leaf length, leaf width and length × width product) and shape (i.e. length/width ratio), and investigated the effects of changes in precipitation and temperature over time and space on the variation in leaf size and shape.

After accounting for the effects of sampling time, leaf size decreased with latitude for all species combined, but the relationship varied among species. Leaf size and shape were positively correlated with temperature and precipitation across space. After accounting for the effects of sampling locations, leaf size of all species combined increased with time. Leaf size changes over time were mostly positively correlated with precipitation, whereas leaf shape changes were mostly correlated with temperature. Overall, our results indicate significant spatial and temporal intraspecific variation in leaf size and shape in response to climate. Our study also demonstrates that herbarium specimens collected over a considerable period of time provide a good resource to study the impacts of climate change on plant morphological traits.

Aims Soil sample preservation is a challenging aspect in molecular studies on soil microbial communities. The demands for specialized sample storage equipment, chemicals and standardized protocols for nucleic acid extraction often require sample processing in a home laboratory that can be continents apart from sampling sites. Standard sampling procedures, especially when dealing with RNA, comprise immediate snap freezing of soils in liquid nitrogen and storage at ?80°C until further processing. For these instances, organizing a reliable cooling chain to transport hundreds of soil samples between continents is very costly, if possible at all. In this study we tested the effect of soil sample preservation by freeze-drying with subsequent short-term storage at 4°C or ambient temperatures compared to ?80°C freezing by comparative barcoding analyses of soil microbial communities.
Methods Two grassland soil samples were collected in Central Germany in the Biodiversity Exploratory Hainich-Dün. Samples were freeze-dried or stored at ?80°C as controls. Freeze-dried samples were stored at 4°C or ambient temperature. Investigated storage times for both storage temperatures were 1 and 7 days. Total DNA and RNA were extracted and bacterial and arbuscular mycorrhizal (AM) fungal communities were analyzed by amplicon 454 pyrosequencing of the 16S (V4-V5 variable region) and 18S (NS31-AM1 fragment) of ribosomal RNA (rRNA) marker genes, respectively.
Important findings Bacterial communities were sufficiently well preserved at the rDNA and rRNA level although storage effects showed as slightly decreased alpha diversity indices for the prolonged storage of freeze-dried samples for 7 days. AM fungal communities could be studied without significant changes at the rDNA and rRNA level. Our results suggest that proper sampling design followed by immediate freeze-drying of soil samples enables short-term transportation of soil samples across continents.

Aims Interspecific and intraspecific variation in flower color in natural populations provides an opportunity for us to understand the evolution and maintenance of diversity of floral traits. Compared to corolla color, little is known about the color polymorphism of sexual organs in flowering plants. To explore evolutionary transitions of androecium color and polymorphism within species, interspecific and intraspecific variation in androecium (anther and pollen) color in the genus Epimedium (Berberidaceae) was investigated.
Methods To explore the geographical patterns of anther/pollen color variation in Epimedium species, data of 45 species were collected and their phylogeny was constructed based on available DNA sequences. To investigate whether intraspecific variation in androecium color relates to habitat preference, three environmental factors were measured in the field population of Epimedium pubescens in northeastern Sichuan, China, which plants had green or yellow androecia. Vegetative and reproductive traits of this species were compared between the two color morphs.
Important findings Androecium (anther and pollen) color polymorphism in field populations of Epimedium pubescens is reported here where nine populations are monomorphic with a green androecium but three populations are dimorphic with individuals having either a green or a yellow androecium. Inflorescence stalk height, stalk diameter, leaf number, flower number and spur length (as well as spur and nectar volume) were not significantly different between two morphs. Compared to the yellow morph, the green morph had relatively larger leaves and anthers, but smaller sepals. The green morph produced more pollen and larger seeds, but the same number of ovules. Seed set was not significantly different between green and yellow morph. Investigations of environmental factors in the color dimorphic populations of E. pubescens indicated that the green morph was more likely to occur in habitats with relatively lower light intensity. The distribution survey of 45 Epimedium species showed that species with a green androecium tended to appear at lower elevations. Comparative phylogenetic analysis showed that transitions from yellow to a green androecium or to androecial color dimorphism occurred at least seven times. This genus, characterized by anther color diversity and containing some species with anther color polymorphism, provides a model system in which to study the evolution and maintenance of colorful sexual organs in flowering plants.

Grassland degradation represents a major challenge in the maintenance of grassland productivity. This process has dramatic impacts on energy flows and soil nutrient dynamics, thus directly or indirectly influencing soil microbes. Here, we aim to (i) examine changes in soil microbial composition, diversity and functionality in response to different levels of grassland degradation (i.e. non-degraded, moderately and severely degraded) in a temperate grassland in Inner Mongolia, and (ii) elucidate biotic and abiotic factors that are responsible for these changes.

The composition structure of soil microbial community was determined by high-throughput sequencing. The functionality of bacterial communities was examined using the tool of FAPROTAX, and functional guilds of fungal communities were quantified using the FUNGuild pipeline.

Grassland degradation significantly decreased soil bacterial diversity but it did not affect fungal diversity. Belowground biomass, soil organic carbon and total nitrogen were positively related to changes in diversity of bacterial community. Grassland degradation significantly increased the relative abundance of Chloroflexi (from 2.48% to 8.40%) and decreased Firmicutes (from 3.62% to 1.08%) of bacterial community. Degradation also significantly increased the relative abundance of Glomeromycota (from 0.17% to 1.53%) and decreased Basidiomycota (from 19.30% to 4.83%) of fungal community. The relative abundance of pathogenic fungi (Didymella and Fusarium) was decreased significantly by degradation. In addition, degradation had a significant impact on putative functionality of soil bacteria related to soil carbon and nitrogen cycling. Our results suggest that soil bacterial community is more sensitive than fungal community in response to degradation in the temperate grassland.

Aims Invasive alien plants can greatly affect native communities and ecosystem processes but only a small fraction of alien plant species become invasive. Barriers to establishment and invasion include reproductive limitations. Clematis vitalba L. has been a popular horticultural species for the past century and is widely distributed and can be highly invasive. In Ireland, it is considered naturalized and potentially invasive. Despite this, little is known about its reproductive biology.
Methods We carried out manipulative field experiments in Ireland and compared fruit and seed set from a number of pollination treatments, namely cross-pollination, geitonogamy, autogamy and natural pollination. We also recorded floral visitation to C. vitalba through a series of timed observations.
Important findings We found that C. vitalba is capable of uniparental reproduction via geitonogamy and autonomous selfing, albeit at a reduced rate compared with outcrossing treatments. Clematis vitalba was visited by at least 10 native pollinator taxa, with hoverflies dominating visitation. Neither fruit set nor seed set in our study population was pollen limited. Given the lack of reproductive constraint, C. vitalba may easily spread in suitable habitats. This is of concern in Ireland, given its prevalence in some of the country's most floristically diverse regions.

Aims The vegetation on Brazilian Páramos consists of assemblages that are driven mainly by the influence of strong environmental filtering. It is very important to understand the effect of environmental variation on taxonomic diversity and on functional diversity. Considering the lack of information about the functional diversity in Brazilian Páramos, we analyzed for the first time the effects of altitude and edaphic attributes on functional traits, as well as on taxonomic and functional diversity. We also wanted to answer the questions: Which ecological strategies are favorable in high-altitude grassland? Does soil attributes determine distributions of traits in high-altitude grassland? Considering the studied altitudinal gradient is altitude an important variable in the community assembly?
Methods The study was conducted on three mountains: Mammoth (1850 m), Elephant (1790 m) and Totem (1690 m) in Serra do Brigadeiro State Park, Minas Gerais State, Brazil. Those mountains represent the 'Serra das Cabe?as', a smaller ridge that is surrounded by the Atlantic Forest, one of the 25 hotspot of biodiversity. The samples were taken using 100 plots of 1 m 2 per mountain that were randomly distributed. All plants except mosses were sampled. The taxonomic diversity was evaluated using richness, Shannon diversity, effective number of species and Pielou evenness. For the functional diversity, we considered the functional richness, functional evenness and functional divergence. Generalized linear models (GLM/Poisson and quasi-Poisson) were used to evaluate the effect of abiotic variables (altitude, soil depth and soil chemical attributes) on biotic variables (number of species and individuals, life form, dispersal and fruit type) and ordinary least squares regression to evaluate the effect of abiotic variables on the functional and taxonomic diversity.
Important findings The soil variables presented a considerable edaphic gradient associated with altitude. The soil in Serra das Cabe?as plays an important role for the plant diversity: richness and diversity index were positively related with fertility. With regard to the life form, nanophanerophytes tended to increase with altitude and soil depth, while therophytes tended to decrease with altitude. The dispersal type was also associated with the abiotic variables: autochory decreases with altitude, while zoochory increases. Functional richness increases with fertility and the functional evenness with altitude. The studied gradient showed that altitude is working as a filter for functional traits and indices and is, together with soil attributes, an important determinant for the distribution of plants on Brazilian Páramos .

Aims Mediterranean coastal dunes are habitats of great conservation interest, with a distinctive and rich flora. In the last century, Acacia spp., native from Australia, have been introduced in Portugal, with the objective of stabilizing sand dunes, and since have become dominant in numerous sand dune habitats. This invasion process led to the reduction of native plant species richness, changed soil characteristics and modified habitat's microclimatic characteristics. The aim of this research was to typify and compare, in Mediterranean sand dune ecosystems, the ecophysiological responses to drought of Helichrysum italicum and Corema album, two native species, and Acacia longifolia, an exotic invasive species. We addressed the following specific objectives: (i) to compare water relations and water use efficiencies, (ii) to evaluate water stress, (iii) to assess water use strategies and water sources used by plants and (iv) to evaluate the morphological adaptations at leaf and phyllode level.
Methods In order to obtain an integrative view of ecophysiological patterns, water relations and performance measuring methods have been applied: predawn (ψ PD) and midday (ψ MD) water potential, chlorophyll a fluorescence, oxygen isotopic composition of xylem, rain and groundwater (δ 18 O) and leaf carbon isotopic discrimination (Δ 13 C). The leaf characteristics of the three species, as well as the histochemistry of non-glandular trichome cell walls, were also studied to identify morpho-traits related to drought resistance.
Important findings The results support our initial hypothesis: although A. longifolia clearly possesses a degree of resistance to water stress, such ability is provided by a different water strategy, when compared to native species. Natives relied on morphological adaptations to restrict water loss, whereas the invasive species adjusted the water uptake as a way to balance their limited ability of restricting water loss. We corroborate that woody native species (i) have a conservative water-saving strategy and minor seasonal variations relative to invasive species, (ii) use enriched water sources during drought periods, indicating different water sources and root systems comparing with invasive species and (iii) present drought leaf morpho-functional adaptations related with limiting water loss. Comparing the physiological performance of invasive and native species can offer causal explanations for the relative success of alien plant invasions on sand dunes ecosystems.

Aims Carbon (C) and nitrogen (N) stoichiometry contributes to understanding elemental compositions and coupled biogeochemical cycles in ecosystems. However, we know little about the temporal patterns of C:N stoichiometry during forest development. The goal of this study is to explore the temporal patterns of intraspecific and ecosystem components' variations in C:N stoichiometry and the scaling relationships between C and N at different successional stages.
Methods Along forest development in a natural temperate forest, northeastern China, four age gradients were categorized into ca. 10-, 30-, 70- and 200-year old, respectively, and three 20 m × 20 m plots were set up for each age class. Leaves, branches, fine roots and fresh litter of seven dominant species as well as mineral soil at depth of 0–10 cm were sampled. A Universal CHN Elemental Analyzer was used to determine the C and N concentrations in all samples.
Important findings Intraspecific leaf C, N and C:N ratios remained stable along forest development regardless of tree species; while C, N concentrations and C:N ratios changed significantly either in branches or in fine roots, and they varied with tree species except Populus davidiana (P < 0.05). For ecosystem components, we discovered that leaf C:N ratios remained stable when stand age was below ca. 70 years and dominant tree species were light-demanding pioneers such as Betula platyphylla and Populus davidiana, while increased significantly at the age of ca. 200 years with Pinus koraiensis as the dominant species. C:N ratios in branches and fresh litter did not changed significantly along forest development stages. C concentrations scaled isometrically with respect to N concentrations in mineral soil but not in other ecosystem components. Our results indicate that, leaf has a higher intraspecific C:N stoichiometric stability compared to branch and fine root, whereas for ecosystem components, shifts in species composition mainly affect C:N ratios in leaves rather than other components. This study also demonstrated that C and N remain coupled in mineral soils but not in plant organs or fresh litter during forest development.

Aims To quantify the seasonal differences in effects of leaf habit, species identity, initial diameter, neighborhood interaction and stand environment on tree absolute diameter growth rates in a subtropical forest in China.
Methods We used man-made dendrometer bands to record radial increments of all trees with diameter at breast height (DBH) ≥5cm and height ≥3 m within 25 comparative study plots (30×30 m for each) of the 'Biodiversity–Ecosystem Functioning Experiment China' (BEF-China) in the Gutianshan National Nature Reserve, Zhejiang Province, China. We measured stem circumferences twice a year from 2011 to 2014 to calculate absolute diameter growth rate of a warm and wet season (WWS, April to September) and a dry and cold season (DCS, October to the next March) for each individual tree: annual growth (GR year), growth during the WWS (GR WWS) and growth during the DCS (GR DCS). We firstly tested the differences in growth rates between different seasons using paired t -tests with Bonferroni correction. Then we applied linear mixed models to explore the effects of leaf habit, species identity, initial diameter, neighborhood interaction (indicated by richness, density and total basal area of all neighboring trees within a radius of 5 m around target trees), stand age and topography (elevation, slope and aspect) on tree growth rates of the two different seasons in three deciduous and 14 evergreen species.
Important findings GR year, GR WWS and GR DCS varied between 0.04–0.50cm year^-1 (mean = 0.21), 0.03–0.46cm season^-1 (mean = 0.18) and 0.01–0.05cm season^-1 (mean = 0.03) across the 17 species, respectively. GR WWS was significantly higher than GR DCS for all species. Growth rates of faster growing species tended to have larger absolute differences between the WWS and DCS. Tree growth rates of both seasons and of the year (GR year, GR WWS and GR DCS) varied significantly among leaf habit and species, and increased allometrically with initial diameter, decreased with stand age, but were not significantly related to topography and neighborhood richness or density. GR WWS decreased with neighborhood total basal area, while GR DCS did not. In conclusion, species might the temporally complementary, contributing to plot growth at different times of the year.

Biomass allocation to different organs is a fundamental plant ecophysiological process to better respond to changing environments; yet, it remains poorly understood how patterns of biomass allocation respond to nitrogen (N) additions across terrestrial ecosystems worldwide.

We conducted a meta-analysis using 5474 pairwise observations from 333 articles to assess how N addition affected plant biomass and biomass allocation among different organs. We also tested the ‘ratio-based optimal partitioning’ vs. the ‘isometric allocation’ hypotheses to explain potential N addition effects on biomass allocation.

We found that (i) N addition significantly increased whole plant biomass and the biomass of different organs, but decreased root:shoot ratio (RS) and root mass fraction (RMF) while no effects of N addition on leaf mass fraction and stem mass fraction at the global scale; (ii) the effects of N addition on ratio-based biomass allocation were mediated by individual or interactive effects of moderator variables such as experimental conditions, plant functional types, latitudes and rates of N addition and (iii) N addition did not affect allometric relationships among different organs, suggesting that decreases in RS and RMF may result from isometric allocation patterns following increases in whole plant biomass. Despite alteration of ratio-based biomass allocation between root and shoot by N addition, the unaffected allometric scaling relationships among different organs (including root vs. shoot) suggest that plant biomass allocation patterns are more appropriately explained by the isometric allocation hypothesis rather than the optimal partitioning hypothesis. Our findings contribute to better understand N-induced effects on allometric relationships of terrestrial plants, and suggest that these ecophysiological responses should be incorporated into models that aim to predict how terrestrial ecosystems may respond to enhanced N deposition under future global change scenarios.

Plant height is a key functional trait related to aboveground biomass, leaf photosynthesis and plant fitness. However, large-scale geographical patterns in community-average plant height (CAPH) of woody species and drivers of these patterns across different life forms remain hotly debated. Moreover, whether CAPH could be used as a predictor of ecosystem primary productivity is unknown.

We compiled mature height and distributions of 11 422 woody species in eastern Eurasia, and estimated geographic patterns in CAPH for different taxonomic groups and life forms. Then we evaluated the effects of environmental (including current climate and historical climate change since the Last Glacial Maximum (LGM)) and evolutionary factors on CAPH. Lastly, we compared the predictive power of CAPH on primary productivity with that of LiDAR-derived canopy-height data from a global survey.

Geographic patterns of CAPH and their drivers differed among taxonomic groups and life forms. The strongest predictor for CAPH of all woody species combined, angiosperms, all dicots and deciduous dicots was actual evapotranspiration, while temperature was the strongest predictor for CAPH of monocots and tree, shrub and evergreen dicots, and water availability for gymnosperms. Historical climate change since the LGM had only weak effects on CAPH. No phylogenetic signal was detected in family-wise average height, which was also unrelated to the tested environmental factors. Finally, we found a strong correlation between CAPH and ecosystem primary productivity. Primary productivity showed a weaker relationship with CAPH of the tallest species within a grid cell and no relationship with LiDAR-derived canopy height reported in the global survey. Our findings suggest that current climate rather than historical climate change and evolutionary history determine the geographical patterns in CAPH. However, the relative effects of climatic factors representing environmental energy and water availability on spatial variations of CAPH vary among plant life forms. Moreover, our results also suggest that CAPH can be used as a good predictor of ecosystem primary productivity.

Aims Although shrubs are an important component of forests, their role has not yet been considered in forest biodiversity experiments. In the biodiversity–ecosystem functioning (BEF) experiment with subtropical tree species in south-east China (BEF-China), we factorially combined tree with shrub species-diversity treatments. Here, we tested the hypotheses that shrub survival differs between the 10 planted shrub species, with lower survival rates of late- than early-successional species and is affected by environmental conditions, such as topography and top soil characteristics, as well as by biotic factors, represented by tree, shrub and herb layer characteristics.
Methods We analyzed the survival of 42 000 shrub individuals in 105 plots varying in tree and shrub species richness of the BEF-China project four years after planting. Shrub survival was analyzed with generalized linear mixed effects models at the level of individuals and with variance partitioning at the plot level. Random intercept and random slope models of different explanatory variables were compared with respect to the Bayesian Information Criterion (BIC).
Important findings Survival rates differed largely between the 10 shrub species, ranging from 26% to 91% for Ardisia crenata and Distylium buxifolium, respectively. Irrespective of species identity, single abiotic factors explained up to 5% of species survival, with a negative effect of altitude and slope inclination and a positive effect of the topsoil carbon to nitrogen ratio, which pointed to drought as the major cause of shrub mortality. In contrast, neither tree nor shrub richness affected shrub survival at this early stage of the experiment. Among the biotic predictors, only herb layer species richness and cover of the dominant fern species (Dicranopteris pedata) affected shrub survival. Overall, our models that included all variables could explain about 65% in shrub survival, with environmental variables being most influential, followed by shrub species identity, while tree species diversity (species richness and identity) and herb layer characteristics contributed much less. Thus, in this early stage of the experiment the biotic interactions among shrubs and between shrubs and trees have not yet overruled the impact of abiotic environmental factors.

Aims Growth rates of plants are driven by factors that influence the amount of resources captured and the efficiency of resource use. In trees, the amount of light captured and the efficiency of light use strongly depends on crown characteristics and leaf traits. Although theory predicts that both crown and leaf traits affect tree growth, few studies have yet to integrate these two types of traits to explain species-specific growth rates. Using 37 broad-leaved tree species of subtropical forests in SE China, we investigated how interspecific differences in wood volume growth rates were affected by crown and leaf traits. We tested the hypotheses that (i) larger crown dimensions promote growth rates, (ii) species-specific growth rates are positively related to leaf stomatal conductance, leaf water potential and leaf chemical components, and negatively related to leaf C/N and leaf toughness and (iii) the two sets of traits better explain growth rates in combination than either alone.
Methods Our study was conducted in a large-scale forest Biodiversity and Ecosystem Functioning experiment in China (BEF-China), located in a mountainous region in Jiangxi Province. We related 17 functional traits (two crown dimension and three crown structure traits; six physiological and six morphological leaf traits) to the mean annual growth rate of wood volume of young trees of the studied species. Interrelationships between crown and leaf traits were analyzed using principal component analysis. Simple linear regression analysis was used to test the effect of each trait separately. We used multiple regression analysis to establish the relationship of growth rate to each set of traits (crown traits, physiological and morphological leaf traits) and to the combination of all types of traits. The coefficients of determination (R 2 adj) of the best multiple regression models were compared to determine the relative explanatory power of crown and leaf traits and a combination of both.
Important findings The species-specific growth rates were not related to any of the single crown traits, but were related positively to leaf stomatal conductance and leaf water potential individually, and negatively to leaf toughness, with approximately 13% variance explained by each of the traits. Combinations of different crown traits did not significantly explain the species-specific growth rates, whereas combinations of either physiological or morphological leaf traits explained 24% and 31%, respectively. A combination of both crown and leaf traits explained 42% of variance in species-specific growth rates. We concluded that sets of traits related to carbon assimilation at the leaf-level and to overall amount of leaves exposed at the crown-level jointly explained species-specific growth rates better than either set of traits alone.

Aims Plants use a variety of hydraulic strategies to adapt to seasonal drought that differ by species and environmental conditions. The early-diverging Magnoliaceae family includes two closely related genera with contrasting leaf habits, Yulania (deciduous) and Michelia (evergreen), which naturally inhabit temperate and tropical regions, respectively. Here, we evaluate the hydraulic strategy of species from both genera that have been ex situ conserved in a subtropical region to determine how they respond to the novel cool–dry season climatic pattern.
Methods We measured ecophysiological traits in five Michelia and five Yulania species conserved in the South China Botanical Garden in both wet and dry season conditions and monitored the whole-year sap flow for four of these species.
Important findings We found that Magnoliaceae species that have been ex situ conserved in a subtropical climate did not suffer from excessive water stress due to the mild drought conditions of the dry season and the ecophysiological adjustments the species made to avoid this stress, which differed by leaf habit. Specifically, deciduous species completely shed their leaves during the dry season, while evergreen species decreased their turgor loss points, dry mass based photosynthetic rates, stomatal conductance and specific leaf areas (SLAs) compared to wet season measurements. In comparing the two distinct leaf habits during the wet season, the leathery-leaved evergreen species had higher leaf hydraulic conductance and leaf to sapwood area ratios than the papery-leaved deciduous species, while the deciduous species had greater hydraulic conductivity calculated on both a stem and leaf area basis, dry mass based photosynthetic rates, leaf nutrients, SLAs and stomatal sizes than the evergreen species. Interestingly, species from both genera maintained similar sap flow in the wet season. Both photosynthetically active radiation and vapour pressure deficit affected the diurnal patterns of sap flow in the wet season, while only vapour pressure deficit played a dominant role in the dry season. This study reveals contrasting hydraulic strategies in Yulania and Michelia species under subtropical seasonal conditions, and suggests that these ecophysiological adjustments might be affected more by leaf habit than seasonality, thus reflecting the divergent evolution of the two closely related genera. Furthermore, we show that Magnoliaceae species that are ex situ conserved in a subtropical climate are hydraulically sound, a finding that will inform future conservation efforts of this ancient family under the threat of climatic change.

Aims Seed dormancy and the soil seed bank are crucial to plant regeneration strategy, especially in semiarid ecosystems with unpredictable precipitation. The aim of this study was to investigate how seed dormancy is controlled by environmental factors and how it is correlated with the soil seed bank and regeneration of the perennial legume Oxytropis racemosa, a dominant perennial herb in Mu Us Sandland of semiarid China.
Methods Germination and imbibition experiments on fresh intact and scarified seeds of O. racemosa were used to identify physical dormancy (PY) in seeds of this species. Soil seed bank dynamics, timing of seedling emergence and the fate of buried seeds in the natural habitat were investigated.
Important findings PY was broken by mechanical scarification or wet heat/ice water cycles but not solely by dry heat or wet heat treatment. The soil seed bank exhibited seasonal changes in the number of seeds, which was highest in September and lowest in July. Seeds buried at different sand depths gradually lost dormancy; 20–42% of the seeds remained dormant after 20 months of burial. Dormancy break occurs gradually throughout the year. Our results indicate that O. racemosa exhibits hardcoatedness heterogeneity that spreads germination of a seed cohort between seasons and years in the semiarid environment, where the amount of precipitation during the growing season is highly variable.

Aims The aim of our research was to understand small-scale effects of topography and soil fertility on tree growth in a forest biodiversity and ecosystem functioning (BEF) experiment in subtropical SE China.
Methods Geomorphometric terrain analyses were carried out at a spatial resolution of 5×5 m. Soil samples of different depth increments and data on tree height were collected from a total of 566 plots (667 m 2 each). The soils were analyzed for carbon (soil organic carbon [SOC]), nitrogen, acidity, cation exchange capacity (CEC), exchangeable cations and base saturation as soil fertility attributes. All plots were classified into geomorphological units. Analyses of variance and linear regressions were applied to all terrain, soil fertility and tree growth attributes.
Important findings In general, young and shallow soils and relatively small differences in stable soil properties suggest that soil erosion has truncated the soils to a large extent over the whole area of the experiment. This explains the concurrently increasing CEC and SOC stocks downslope, in hollows and in valleys. However, colluvial, carbon-rich sediments are missing widely due to the convexity of the footslopes caused by uplift and removal of eroded sediments by adjacent waterways. The results showed that soil fertility is mainly influenced by topography. Monte–Carlo flow accumulation (MCCA), curvature, slope and aspect significantly affected soil fertility. Furthermore, soil fertility was affected by the different geomorphological positions on the experimental sites with ridge and spur positions showing lower exchangeable base cation contents, especially potassium (K), due to leaching. This geomorphological effect of soil fertility is most pronounced in the topsoil and decreases when considering the subsoil down to 50cm depth. Few soil fertility attributes affect tree height after 1–2 years of growth, among which C stocks proved to be most important while pH KCl and CEC only played minor roles. Nevertheless, soil acidity and a high proportion of Al on the exchange complex affected tree height even after only 1–2 years growth. Hence, our study showed that forest nutrition is coupled to a recycling of litter nutrients, and does not only depend on subsequent supply of nutrients from the mineral soil. Besides soil fertility, topography affected tree height. We found that especially MCCA as indicator of water availability affected tree growth at small-scale, as well as aspect. Overall, our synthesis on the interrelation between fertility, topography and tree growth in a subtropical forest ecosystem in SE China showed that topographic heterogeneity lead to ecological gradients across geomorphological positions. In this respect, small-scale soil–plant interactions in a young forest can serve as a driver for the future development of vegetation and biodiversity control on soil fertility. In addition, it shows that terrain attributes should be accounted for in ecological research.

Aims We aim to construct a comprehensive global database of litter decomposition rate (k value) estimated by surface floor litterbags, and investigate the direct and indirect effects of impact factors such as geographic factors (latitude and altitude), climatic factors (mean annual tempePlrature, MAT; mean annual precipitation, MAP) and litter quality factors (the contents of N, P, K, Ca, Mg and C:N ratio, lignin:N ratio) on litter decomposition.
Methods We compiled a large data set of litter decomposition rates (k values) from 110 research sites and conducted simple, multiple regression and path analyses to explore the relationship between the k values and impact factors at the global scale.
Important findings The k values tended to decrease with latitude (LAT) and lignin content (LIGN) of litter but increased with temperature, precipitation and nutrient concentrations at the large spatial scale. Single factor such as climate, litter quality and geographic variable could not explain litter decomposition rates well. However, the combination of total nutrient (TN) elements and C:N accounted for 70.2% of the variation in the litter decomposition rates. The combination of LAT, MAT, C:N and TN accounted for 87.54% of the variation in the litter decomposition rates. These results indicate that litter quality is the most important direct regulator of litter decomposition at the global scale. This data synthesis revealed significant relationships between litter decomposition rates and the combination of climatic factor (MAT) and litter quality (C:N, TN). The global-scale empirical relationships developed here are useful for a better understanding and modeling of the effects of litter quality and climatic factors on litter decomposition rates.

Aims In ecology and conservation biology, the number of species counted in a biodiversity study is a key metric but is usually a biased underestimate of total species richness because many rare species are not detected. Moreover, comparing species richness among sites or samples is a statistical challenge because the observed number of species is sensitive to the number of individuals counted or the area sampled. For individual-based data, we treat a single, empirical sample of species abundances from an investigator-defined species assemblage or community as a reference point for two estimation objectives under two sampling models: estimating the expected number of species (and its unconditional variance) in a random sample of (i) a smaller number of individuals (multinomial model) or a smaller area sampled (Poisson model) and (ii) a larger number of individuals or a larger area sampled. For sample-based incidence (presence–absence) data, under a Bernoulli product model, we treat a single set of species incidence frequencies as the reference point to estimate richness for smaller and larger numbers of sampling units.
Methods The first objective is a problem in interpolation that we address with classical rarefaction (multinomial model) and Coleman rarefaction (Poisson model) for individual-based data and with sample-based rarefaction (Bernoulli product model) for incidence frequencies. The second is a problem in extrapolation that we address with sampling-theoretic predictors for the number of species in a larger sample (multinomial model), a larger area (Poisson model) or a larger number of sampling units (Bernoulli product model), based on an estimate of asymptotic species richness. Although published methods exist for many of these objectives, we bring them together here with some new estimators under a unified statistical and notational framework. This novel integration of mathematically distinct approaches allowed us to link interpolated (rarefaction) curves and extrapolated curves to plot a unified species accumulation curve for empirical examples. We provide new, unconditional variance estimators for classical, individual-based rarefaction and for Coleman rarefaction, long missing from the toolkit of biodiversity measurement. We illustrate these methods with datasets for tropical beetles, tropical trees and tropical ants.
Important findings Surprisingly, for all datasets we examined, the interpolation (rarefaction) curve and the extrapolation curve meet smoothly at the reference sample, yielding a single curve. Moreover, curves representing 95% confidence intervals for interpolated and extrapolated richness estimates also meet smoothly, allowing rigorous statistical comparison of samples not only for rarefaction but also for extrapolated richness values. The confidence intervals widen as the extrapolation moves further beyond the reference sample, but the method gives reasonable results for extrapolations up to about double or triple the original abundance or area of the reference sample. We found that the multinomial and Poisson models produced indistinguishable results, in units of estimated species, for all estimators and datasets. For sample-based abundance data, which allows the comparison of all three models, the Bernoulli product model generally yields lower richness estimates for rarefied data than either the multinomial or the Poisson models because of the ubiquity of non-random spatial distributions in nature.

Aims The functional advantages of arsenic (As) hyperaccumulation by plants are poorly understood. One proposed benefit, termed elemental allelopathy, occurs when hyperaccumulated As is cycled from the plant back into the top layer of soil, allowing As hyperaccumulators to gain an advantage over intolerant species by increasing soil As concentrations ([ As]) underneath their canopy. To date, there are no studies that detail the presence of increased soil [ As] associated with As hyperaccumulators. In this study, we documented variation in the soil [ As] associated with the Chinese brake fern, Pteris vittata L. and also compared the effects of environmentally relevant soil and solution [ As] on competitor plant growth.
Methods Four populations of P. vittata were identified in central Florida, USA. P. vittata tissue samples and soil samples were collected at the base of and at 3 m away from ferns in each population (n = 36). Five sample locations were randomly selected from each site, and soils from the base and 3 m away from each fern were collected to examine the effects of naturally occurring soil [ As] on the germination and growth of a potential competitor plant (Oxalis stricta). Solutions with increasing [ As] were also used to examine the threshold for negative effects of [ As] on O. stricta growth. [ As] were measured using inductively coupled plasma mass spectrometry (ICP-MS).
Important findings Overall, soil [ As] from the base of ferns was nearly twice that of soil 3 m away indicating that ferns hyperaccumulate As. However, ferns and their associated soil, contained different [ As] depending on their collection site, indicating that these populations accumulate and use [ As] differently. O. stricta growth decreased and germination was delayed as solution and soil [ As] increased. However, the relative distance from the fern that the soil was collected from did not affect growth, which would be expected with elemental allelopathy. Our results show that P. vittata is associated with higher soil [ As] and these concentrations are sufficient to inhibit growth of competitors. However, the absence of a strong inhibitory relationship associated with proximity to the fern across all locations suggests that the possible functional advantages of elemental allelopathy may depend on site specific characteristics.

Long-term heavy grazing reduces plant diversity and ecosystem function by intensifying nitrogen (N) and water limitation. In contrast, the absence of biomass removal can cause species loss by elevating light competition and weakening community stability, which is exacerbated by N and water enrichment. Hence, how to maintain species diversity and community stability is still a huge challenge for sustainable management of worldwide grasslands.

We conducted a 4-year manipulated experiment in six long-term grazing blocks to explore combination of resource additions and biomass removal (increased water, N and light availability) on species richness and community stability in semiarid grasslands of Inner Mongolia, China.

In all blocks treated with the combination of resource additions and biomass removal, primary productivity increased and species richness and community stability were maintained over 4 years of experiment. At both species and plant functional group (PFG) levels, the aboveground biomass of treated plants remained temporally stable in treatments with the combination of N and/or water addition and biomass removal. The maintenance of species richness was primarily caused by the biomass removal, which could increase the amount of light exposure for grasses under resource enrichment. Both species asynchrony and stability of PFGs contributed to the high temporal stability observed in these communities. Our results indicate that management practices of combined resource enrichment with biomass removal, such as grazing or mowing, could not only enhance primary productivity but also maintain plant species diversity, species asynchrony and community stability. Furthermore, as overgrazing-induced degradation and resource enrichment-induced biodiversity loss continue to be major problems worldwide, our findings have important implications for adaptive management in semiarid grasslands and beyond.

Aims Positive plant diversity–ecosystem function relations are ultimately driven by variation in functional traits among individuals that form a community. To date, research has largely focused on the role of species diversity for ecosystem functioning. However, substantial intraspecific trait variation is common and a significant part of this variation caused by genetic differences among individuals. Here, we studied the relative importance of species diversity and seed family (SF) diversity within species for growth and herbivory in experimental subtropical tree assemblages.
Methods In 2010, we set up a field experiment in subtropical China, using four species from the local species pool. Trees were raised from seeds, with seeds from the same mother tree forming an SF. We established 23 plots containing one or four species (species diversity treatment) and one or four SFs per species (SF diversity treatment). Tree growth (stem diameter, plant height and crown expansion) and herbivory (percentage leaf loss due to leaf chewers) were monitored annually from 2011 to 2013.
Important findings Tree species richness promoted growth but had no effect on herbivory. In contrast, SF diversity reduced growth and increased herbivory but only so in species mixtures. Most of the observed effects were time dependent, with the largest effect found in 2013. Our results suggest that biodiversity can affect plant performance directly via tree species–species interactions, or context dependent, via potential effects on inter-trophic interactions. Two important conclusions should be drawn from our findings. Firstly, in future studies regarding biodiversity and ecosystem functioning (BEF) relationships, intraspecific genetic diversity should be given similar weight as species diversity as it has often been neglected and its effects are not well understood. Secondly, we demonstrate opposite effects of biodiversity among and within species, stressing the importance to consider the effects of multiple levels of biodiversity simultaneously.

Aims Fine roots play an important role in the biogeochemical cycles of terrestrial ecosystems and are vital for understanding forest ecosystem functioning and services. Higher plant species diversity has been largely reported to increase aboveground community biomass, but how biodiversity affects fine-root production and the related mechanisms in forests remain unclear. In this study, we aim to answer two questions: (i) does fine-root production increase with tree species richness? (ii) Can this effect be explained by niche complementarity among species?
Methods We analyzed data from a large forest biodiversity experiment (BEF-China) with 5-year-old trees. Fine-root growth was measured as standing biomass and annual fine-root regrowth was estimated using ingrowth cores. Moreover, relative yield was calculated to test whether over- or under-yielding occurred when mixtures were compared with the average monoculture of the species included in the mixtures. We calculated functional diversity for fine-root (≤2mm in diameter) traits by Rao's quadratic entropy index for each species mixture. The effects of manipulated tree species richness and identity on fine-root traits were analyzed with linear mixed-effects models. Mixed models were also used to test the relationships between tree species richness and fine-root standing biomass, annual regrowth and vertical heterogeneity.
Important findings Fine roots of more than one species were found in half of the soil cores in mixtures indicating that belowground interactions in these young forest stands occurred much earlier than canopy closure. We found significant differences among species in fine-root traits such as diameter and specific root length (SRL), which suggested different resource-use strategies and niche partitioning among species. Mean fine-root diameter of species ranged from 0.31 to 0.74mm, mean SRL ranged from 12.43 m·g^-1 to 70.22 m·g^-1 and mean vertical distribution index β ranged from 0.68 to 0.93. There was a significant positive relationship between species richness and the evenness of the vertical distribution of fine-root standing biomass. Moreover, marginally significant positive relationships existed between species richness and standing biomass as well as annual regrowth of fine roots. Relative yields and Rao's quadratic entropy index were both not significantly affected by species richness. However, the relative yield of fine-root standing biomass was marginally correlated with Rao's quadratic entropy index, implying that belowground niche complementarity between species does contribute to diversity effects. In conclusion, our study showed positive effects of species richness on the filling of soil volume by fine roots in the studied experimental forest communities. This has positive effects on fine-root standing biomass and may also lead to increased aboveground biomass.

Aims Beta diversity is the variation in species composition among sites in a geographic region. Beta diversity is a key concept for understanding the functioning of ecosystems, for the conservation of biodiversity and for ecosystem management. The present report describes how to analyse beta diversity from community composition and associated environmental and spatial data tables.
Methods Beta diversity can be studied by computing diversity indices for each site and testing hypotheses about the factors that may explain the variation among sites. Alternatively, one can carry out a direct analysis of the community composition data table over the study sites, as a function of sets of environmental and spatial variables. These analyses are carried out by the statistical method of partitioning the variation of the diversity indices or the community composition data table with respect to environmental and spatial variables. Variation partitioning is briefly described herein.
Important findings Variation partitioning is a method of choice for the interpretation of beta diversity using tables of environmental and spatial variables. Beta diversity is an interesting 'currency' for ecologists to compare either different sampling areas or different ecological communities co-occurring in an area. Partitioning must be based upon unbiased estimates of the variation of the community composition data table that is explained by the various tables of explanatory variables. The adjusted coefficient of determination provides such an unbiased estimate in both multiple regression and canonical redundancy analysis. After partitioning, one can test the significance of the fractions of interest and plot maps of the fitted values corresponding to these fractions.

Aims Most biodiversity–ecosystem functioning research has been carried out in grassland ecosystems, and little is known about whether forest ecosystems, in particular outside the temperate zone, respond similarly. Here, we tested whether productivity, assessed as leaf area index (LAI), increases with species richness in young experimental stands of subtropical trees, whether this response is similar for early-season leaf area (which is dominated by evergreens) and seasonal leaf area increase (which is dominated by deciduous species), and whether responses saturate at high species richness.
Methods We used a planted tree biodiversity experiment in south-east China to test our hypotheses. LAI was determined three times by digital hemispheric photography in 144 plots that had been planted with 400 trees each, forming communities with 1, 2, 4, 8 or 16 tree species.
Important findings LAI increased significantly with tree species richness in the fifth year of stand establishment. Similar, but weaker, statistically non-significant trends were observed 1 year before. We did not observe leaf area overyielding and the presence of particularly productive and unproductive species explained large amounts of variation in leaf area, suggesting that selection-type effects contributed substantially to the biodiversity effects we found in this early phase of stand establishment. Effects sizes were moderate to large and comparable in magnitude to the ones reported for grassland ecosystems. Subtropical (and tropical) forests harbor substantial parts of global net primary production and are critical for the Earth's carbon and hydrological cycle, and our results suggest that tree diversity critically supports these ecosystem services.

Generalized linear mixed models (GLMMs) have been widely used in contemporary ecology studies. However, determination of the relative importance of collinear predictors (i.e. fixed effects) to response variables is one of the challenges in GLMMs. Here, we developed a novel R package, glmm.hp, to decompose marginal R² explained by fixed effects in GLMMs. The algorithm of glmm.hp is based on the recently proposed approach ‘average shared variance’ i.e. used for multivariate analysis. We explained the principle and demonstrated the use of this package by simulated dataset. The output of glmm.hp shows individual marginal R²s that can be used to evaluate the relative importance of predictors, which sums up to the overall marginal R². Overall, we believe the glmm.hp package will be helpful in the interpretation of GLMM outcomes.

Non-structural carbohydrates (NSCs) are plant storage compounds used for metabolism, transport, osmoregulation and regrowth following the loss of plant tissue. Even in conditions suitable for optimal growth, plants continue to store NSCs. This storage may be due to passive accumulation from sink-inhibited growth or active reserves that come at the expense of growth. The former pathway implies that NSCs may be a by-product of sink limitation, while the latter suggests a functional role of NSCs for use during poor conditions.

Using 13C pulse labelling, we traced the source of soluble sugars in stem and root organs during drought and everwet conditions for seedlings of two tropical tree species that differ in drought tolerance to estimate the relative allocation of NSCs stored prior to drought versus NSCs assimilated during drought. We monitored growth, stomatal conductance, stem water potential and NSC storage to assess a broad carbon response to drought.

We found that the drought-sensitive species had reduced growth, conserved NSC concentrations in leaf, stem and root organs and had a larger proportion of soluble sugars in stem and root organs that originated from pre-drought storage relative to seedlings in control conditions. In contrast, the drought-tolerant species maintained growth and stem and root NSC concentrations but had reduced leaf NSCs concentrations with a larger proportion of stem and root soluble sugars originated from freshly assimilated photosynthates relative to control seedlings. These results suggest the drought-sensitive species passively accumulated NSCs during water deficit due to growth inhibition, while the drought-tolerant species actively responded to water deficit by allocating NSCs to stem and root organs. These strategies seem correlated with baseline maximum growth rates, which supports previous research suggesting a trade-off between growth and drought tolerance while providing new evidence for the importance of plasticity in NSC allocation during drought.

Aims Mapping vegetation through remotely sensed images involves various considerations, processes and techniques. Increasing availability of remotely sensed images due to the rapid advancement of remote sensing technology expands the horizon of our choices of imagery sources. Various sources of imagery are known for their differences in spectral, spatial, radioactive and temporal characteristics and thus are suitable for different purposes of vegetation mapping. Generally, it needs to develop a vegetation classification at first for classifying and mapping vegetation cover from remote sensed images either at a community level or species level. Then, correlations of the vegetation types (communities or species) within this classification system with discernible spectral characteristics of remote sensed imagery have to be identified. These spectral classes of the imagery are finally translated into the vegetation types in the image interpretation process, which is also called image processing. This paper presents an overview of how to use remote sensing imagery to classify and map vegetation cover.
Methods Specifically, this paper focuses on the comparisons of popular remote sensing sensors, commonly adopted image processing methods and prevailing classification accuracy assessments.
Important findings The basic concepts, available imagery sources and classification techniques of remote sensing imagery related to vegetation mapping were introduced, analyzed and compared. The advantages and limitations of using remote sensing imagery for vegetation cover mapping were provided to iterate the importance of thorough understanding of the related concepts and careful design of the technical procedures, which can be utilized to study vegetation cover from remote sensed images.

Aims Litterfall, as an important link between aboveground and belowground processes, plays a key role in forest ecosystems. Here, we test for effects of tree species richness on litter production and litter quality in subtropical forest. The study further encompasses a factorial gradient of secondary succession that resulted from human exploitation. Given that a large percentage of subtropical forests are in secondary successional stages, understanding the role of biodiversity on forest re-growth after disturbance appears critical.
Methods From January 2009 to December 2014, we monitored forest litterfall in 27 Comparative Study Plots that spanned a gradient of tree species richness (3–20 species) and secondary successional ages (~20 to 120 years) in Gutianshan Natural Nature Reserve, Zhejiang Province, China. The experiment is part of the biodiversity–ecosystem functioning research platform 'BEF-China'. Tree litterfall was collected in monthly intervals using litter traps. Samples were separated into leaf and non-leaf components. Leaf litter was further sorted into dominant and other species. Community level monthly leaf litter C and N contents were analysed through a full year. General linear mixed-effects models were applied to test for effects of tree species richness and successional age on litter quantity and leaf litter C/N.
Important findings Litterfall increased with species richness among and within successional age and this effect was consistent across years. Successionally older stands had higher litterfall and this effect was related to increased tree species richness. However, species richness did not change the intra- and inter-annual temporal stability of litterfall. Increasing tree species richness increased leaf litter quality (decreased C/N), while successional age had no effect. Our study indicates that more diverse forest stands produce more leaf litter and that this litter has higher N concentrations, which could promote forest growth through accelerated nutrient re-cycling.

Aims European frogbit (Hydrocharis morsus-ranae L.) is an aquatic plant originating from Europe that has emerged as an invasive species, spreading in the USA and Canada since it was first brought to North America in 1932. It can now be found in many water bodies, from small ponds and long rivers to large lakes such as Lake Ontario and Lake Erie. The continuous spread of this species indicates its success as an invasive species despite legislative attempts to limit its distribution. Catling et al. (Catling PM, Miltrow G, Haber E, et al. (2003) The biology of Canadian weeds. 124. Hydrocharis morsus-ranae L. Can J Plant Sci 83:1001–16) wrote a thorough review about this invasive species in North America. Our review aims for a compilation of the most recent available data and recent studies on H. morsus-ranae L. and focuses primarily on its environmental uses, ecological impacts and management. The purpose of this review is to offer an organized and updated report on European frogbit that can be used towards future studies with the goal of eradicating this invasive species and providing insights on management of other invasive plants.
Important findings Our findings reveal that European forgbit's ecological effects on other species and the invaded environment were shown to be less harmful than previously feared. European frogbit had negative impacts on native plants and reduced dissolved oxygen concentration. However, water chemistry, phytoplankton and zooplankton communities were actually not affected by European frogbit. For fungi, bacteria and macroinvertebrates, studies have showed complex and sometimes conflicting results. We also specifically discussed the new method to control this species using shading and the more recent studies on biological control. Shading with a shade cloth has been shown to effectively remove European frogbit and had minor environmental effects. However, using biological control to combat the spread of the invasive frogbit seems not as successful as we wished.

Recent studies have revealed heritable phenotypic plasticity through vegetative generations. In this sense, changes in gene regulation induced by the environment, such as DNA methylation (i.e. epigenetic changes), can result in reversible plastic responses being transferred to the offspring generations. This trans-generational plasticity is expected to be especially relevant in clonal plants, since reduction of sexual reproduction can decrease the potential for adaptation through genetic variation. Many of the most aggressive plant invaders are clonal, and clonality has been suggested as key to explain plant invasiveness. Here we aim to determine whether trans-generational effects occur in the clonal invader Alternanthera philoxeroides, and whether such effects differ between populations from native and non-native ranges.

In a common garden experiment, parent plants of A. philoxeroides from populations collected in Brazil (native range) and Iberian Peninsula (non-native range) were grown in high and low soil nutrient conditions, and offspring plants were transplanted to control conditions with high nutrients. To test the potential role of DNA methylation on trans-generational plasticity, half of the parent plants were treated with the demethylating agent, 5-azacytidine.

Trans-generational effects were observed both in populations from the native and the non-native ranges. Interestingly, trans-generational effects occurred on growth variables (number of ramets, stem mass, root mass and total mass) in the population from the native range, but on biomass partitioning in the population from the non-native range. Trans-generational effects of the population from the native range may be explained by a ‘silver-spoon’ effect, whereas those of the population from the non-native range could be explained by epigenetic transmission due to DNA methylation. Our study highlights the importance of trans-generational effects on the growth of a clonal plant, which could help to understand the mechanisms underlying expansion success of many clonal plants.

Aims Distyly is one of the most widespread floral polymorphisms promoting cross-fertilization. Evolutionary transition from obligate cross-fertilized distyly to predominantly self-fertilized homostyly is frequently documented in various groups. However, empirical studies concerning the ecological factors connected with this transition are still lacking. Primula chungensis, suggested to be evolving from distyly to homostyly, provides an ideal model for the study of the ecological factors concerned with this transition. We study P. chungensis to understand if autonomous self-fertilization would provide reproductive assurance for the self-fertilized homo-styled morph in the field.
Methods The incompatibility features of P. chungensis were tested with hand-pollination experiments. We compared the capacity of autonomous self-fertilization between the distylous and homo-styled morph of P. chungensis in the field by excluding the pollinators with bags. In addition, the degrees of herkogamy of some P. chungensis plants were between the short-styled and homo-styled morphs. These plants were studied to understand whether they were able to obtain greater reproductive assurance when the herkogamy in the flowers was reduced.
Important findings All three morphs of P. chungensis were highly self- and intra-morph compatible. The degree of herkogamy positively correlated with the capacity for autonomous self-fertilization. A negative correlation between the degree of herkogamy and the magnitude of pollen limitation was found, but no significant correlation was found between the degree of herkogamy and the contribution of cross-fertilization to overall fertilization. This study suggests that reducing the degree of herkogamy can significantly increase the reproductive assurance for a self-compatible plant. Our results provided evidence that the homo-styled morph of P. chungensis had the highest capacity for autonomous self-fertilization and the highest seed production in the field, because autonomous self-fertilization provided reproductive assurance for the homo-styled morph. This may cause selection towards the transition from distyly to homostyly.

Aims Several studies have shown that plant height changes along environmental gradients. However, altitudinal patterns of plant height across species are still unclear, especially in regions sensitive to climate change. As canopy height decreases dramatically near the tree line in alpine areas, we hypothesize that plant height across all species also decreases with increasing altitude, and distinct thresholds exist along this gradient.
Methods Using a large dataset of maximum plant height and elevation range (400 to 6000 m a.s.l.) of 4295 angiosperms from the regional flora of the Tibetan Plateau, we regressed plant height for every 100 m belt against elevation to explore the relationships. To identify the approximate boundaries where dramatic changes in plant height occurs for herbaceous plants, shrubs, trees, woody plants and all angiosperms, we used piecewise linear regression. Phylogenetically independent contrast was used to test the potential evolutionary influences on altitudinal patterns at the family level.
Important findings Results showed that for herbaceous plants, shrubs, trees, woody plants and all angiosperms, plant height decreases significantly as altitude increases. In addition, we found that altitude, a proxy for many environmental factors, had obvious thresholds (breakpoints) dictating patterns of plant height. The results of phylogenetically independent contrast also emphasized the importance of evolutionary history in determining the altitudinal patterns of plant height for some growth forms. Our results highlight the relative intense filtering effect of environmental factors in shaping patterns of functional traits and how this could vary for different ranges of environmental variables.

Grasslands are dominant vegetation of China, support outstanding biodiversity and sequester bulk amount of atmospheric CO₂. These grasslands are highly degraded and fragmented due to remarkable anthropogenic and grazing loads. Chinese Government has made great attempt to restore by grazing exclusion. The relations of carbon fluxes with species composition and diversity in the communities sensitive to grazing by large herbivores are needed to be analysed under the global climate change scenario. The objective of present study was to comprehend the effects of grazing and fencing on the ecosystem structure and function of the typical steppe grassland.

To meet the objectives, overgrazed and fenced (since year 2001) systems were selected in typical steppe grassland at the Duolun Restoration Ecology Research Station, Inner Mogolia, China. Within each system, three dominant communities with three replicates were selected. In each replicate community, three 1 × 1 m plots, were randomly located. Each plot was divided into four 50 × 50 cm quadrats. A total of 216, 50 × 50 cm quadrats were sampled. From each quadrat, number of individuals and above-ground herbaceous biomass for each species, soil respiration (SR), ecosystem respiration (ER), net (NEE) as well as gross (GEE) ecosystem CO₂ exchanges were recorded in June 2015. Data were well analysed using statistical software. Canonical correspondence analysis showed differential responses of communities to the structure and function of the typical steppe grassland.

Across the communities, fencing reduced the soil temperature by 12% and at the same time increased the soil moisture by 44.30%, thus, increased the species richness by 28%, evenness by 21%, above-ground biomass by 19% and plant carbon by 20%. Interestingly, fencing increased NEE by 128%, GEE by 77%, SR by 65% and ER by 39%. Under fencing, species composition partially governed the CO₂ exchange processes.

Fencing reduces soil temperature and thereby improves species diversity and more efficient CO₂ sequestration and long-term and in-depth study is desirable for a better understanding of the relationship between species diversity and ecosystem carbon uptake.

Aims The positive relationship between plant biodiversity and community productivity is well established. However, our knowledge about the mechanisms underlying these positive biodiversity effects is still limited. One of the main hypotheses is that complementarity in resource uptake is responsible for the positive biodiversity effects: plant species differ in resource uptake strategy, which results in a more complete exploitation of the available resources in space and time when plant species are growing together. Recent studies suggest that functional diversity of the community, i.e. the diversity in functional characteristics ('traits') among species, rather than species richness per se, is important for positive biodiversity effects. However, experimental evidence for specific trait combinations underlying resource complementarity is scarce. As the root system is responsible for the uptake of nutrients and water, we hypothesize that diversity in root traits may underlie complementary resource use and contribute to the biodiversity effects.
Methods In a common garden experiment, 16 grassland species were grown in monoculture, 4-species mixtures differing in root trait diversity and 16-species mixtures. The 4-species mixtures were designed to cover a gradient in average rooting depth. Above-ground biomass was cut after one growing season and used as a proxy for plant productivity to calculate biodiversity effects.
Important findings Overall, plant mixtures showed a significant increase in biomass and complementarity effects, but this varied greatly between communities. However, diversity in root traits (measured in a separate greenhouse experiment and based on literature) could not explain this variation in complementarity effects. Instead, complementarity effects were strongly affected by the presence and competitive interactions of two particular species. The large variation in complementarity effects and significant effect of two species emphasizes the importance of community composition for positive biodiversity effects. Future research should focus on identifying the traits associated with the key role of particular species for complementarity effects. This may increase our understanding of the links between functional trait composition and biodiversity effects as well as the relative importance of resource complementarity and other underlying mechanisms for the positive biodiversity effects.

Aims Although the net biodiversity effect (NE) can be statistically partitioned into complementarity and selection effects (CE and SE), there are different underlying mechanisms that can cause a certain partitioning. Our objective was to assess the role of resource partitioning and species interactions as two important mechanisms that can bring about CEs by interspecific and intraspecific trait variation.
Methods We measured tree height of 2493 living individuals in 57 plots and specific root length (SRL) on first-order roots of 368 of these individuals across different species richness levels (1, 2, 4, 8 species) in a large-scale forest biodiversity and ecosystem functioning experiment in subtropical China (BEF-China) established in 2009. We describe the effects of resource partitioning between species by a fixed component of interspecific functional diversity (RaoQ) and further effects of species interactions by variable components of interspecific and intraspecific functional diversity (community weighted trait similarity and trait dissimilarity, CWS and CWD). Finally, we examined the relationships between biodiversity effects on stand-level tree height and functional diversity (RaoQ, CWS and CWD) in SRL using linear regression and assessed the relative importance of these three components of functional diversity in explaining the diversity effects.
Important findings Our results show that species richness significantly affected SRL in five and tree height in ten out of 16 species. A positive NE was generally brought about by a positive CE on stand-level tree height and related to high values of RaoQ and CWS in SRL. A positive CE was related to high values of all three components of root functional diversity (RaoQ, CWS and CWD). Our study suggests that both resource partitioning and species interactions are the underlying mechanisms of biodiversity effects on stand-level tree growth in subtropical forest.

Aims Why invasive plants are more competitive in their introduced range than native range is still an unanswered question in plant invasion ecology. Here, we used the model invasive plant Solidago canadensis to test a hypothesis that enhanced production of allelopathic compounds results in greater competitive ability of invasive plants in the invaded range rather than in the native range. We also examined the degree to which the allelopathy contributes increased competitive ability of S. canadensis in the invaded range.
Methods We compared allelochemical production by S. canadensis growing in its native area (the USA) and invaded area (China) and also by populations that were collected from the two countries and grown together in a 'common garden' greenhouse experiment. We also tested the allelopathic effects of S. canadensis collected from either the USA or China on the germination of Kummerowia striata (a native plant in China). Finally, we conducted a common garden, greenhouse experiment in which K. striata was grown in monoculture or with S. canadensis from the USA or China to test the effects of allelopathy on plant–plant competition with suitable controls such as adding activated carbon to the soil to absorb the allelochemicals and thereby eliminating any corresponding allopathic effects.
Important findings Allelochemical contents (total phenolics, total flavones and total saponins) and allelopathic effects were greater in S. canadensis sampled from China than those from the USA as demonstrated in a field survey and a common garden experiment. Inhibition of K. striata germination using S. canadensis extracts or previously grown in soil was greater using samples from China than from the USA. The competitive ability of S. canadensis against K. striata was also greater for plants originating from China than those from the USA. Allelopathy could explain about 46% of the difference. These findings demonstrated that S. canadensis has evolved to be more allelopathic and competitive in the introduced range and that allelopathy significantly contributes to increased competitiveness for this invasive species.