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 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.

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 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.

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 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 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 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 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 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.

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.

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.

Aims We conducted a simulated nitrogen (N) and sulfur (S) deposition experiment from 2006 to 2012 to answer the following questions: (i) does chronic N and S deposition decrease cation concentrations in the soil and foliage of understory plant species, and (ii) does chronic N and S deposition decrease plant diversity and alter species composition of the understory plant community in a boreal forest in western Canada where intensifying industrial activities are increasing N and S deposition?
Methods Our field site was a mixedwood boreal forest stand located ~100 km southeast of Fort McMurray, Alberta, Canada. The experiment involved a 2 × 2 factorial design, with two levels each of N (0 and 30 kg N ha^-1 yr^-1; applied as NH₄NO₃) and S addition (0 and 30 kg S ha^-1 yr^-1; applied as Na₂SO₄). Four blocks were established in July 2006, each with four plots of 20 × 20 m randomly assigned to the treatments. Soil and understory vegetation were sampled and cover (%) of individual species of herb (height ≤ 0.5 m) and shrub (height 0.5–1 m) layers was determined in August 2012.
Important findings Seven years after the treatments began, N addition increased dissolved organic carbon and N in the mineral soil (P < 0.05), whereas S addition decreased exchangeable cations (P < 0.05) in the forest floor. In the shrub layer, species evenness, and overall diversity were decreased by N addition (P < 0.05) due to increases in abundance of nitrophilous species and S addition (P < 0.01) due to decreased cation concentrations in soils. Total shrub cover decreased with S addition (P < 0.10). Nitrogen and S addition affected neither species richness nor evenness in the herb layer. However, permutational multivariate analysis of variance and non-metric multidimensional scaling analyses (based on plant cover) indicated that the effect of N and S addition on understory plant species composition in the both shrub and herb layers was species-specific. Addition of N decreased foliar phosphorus and potassium concentrations in some species, suggesting potential risk of N-meditated nutrient imbalance in those species. Our results indicate that long-term elevated levels of N and S deposition can negatively impact plant nutrition and decrease the diversity of the understory plant community in boreal forests in northern Alberta, Canada. However, considering that the current N and S deposition rates in northern Alberta are much lower than the rates used in this study, N and S deposition should not negatively affect plant diversity in the near future.

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 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.

Nitrogen (N) enrichment caused by human activities threatens biodiversity and alters plant community composition and structure. It has been found that heavy and infrequent N inputs may over-estimate species extinction, but it remains unclear whether plant community structure will equally respond to frequent reactive N enriched conditions.

We independently manipulated the rates and the frequencies of N addition in a temperate steppe, northern China, between 2008 and 2013.

We found that plant community structure changes, measured by ‘Euclidean distance’ involving species richness, composition and productivity, were significantly positively related to increasing N enrichment rates rather than frequencies. Changes in aboveground net primary productivity (ANPP), plant species richness and shifts in dominant species were observed. Community ANPP increased with N enrichment, whereas species richness reduced. The frequency of N enrichment increased species richness but had no impacts on community ANPP and the relative ANPP of the two dominant species, C₃perennial bunchgrass Stipa grandis and C₃ perennial rhizome grass Leymus chinensis. The ANPP and relative ANPP of the two dominant species were significantly negatively correlated with each other. Moreover, changes in the relative ANPP of S. grandis was negatively associated with the changes in community structure. After 5 years’ treatment, direct influence of the frequency of N enrichment on plant community structure was not observed, but the effects of the rate of N enrichment were apparent. Our results suggested that further study in various ecosystems and with long-term and well-controlled comparisons the frequency vs. the rate of N enrichment may still be needed.

Aims Dune building processes are affected by interactions between the growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and sea spray. Research investigating how species interactions influence dune community structure and functional trait responses in high abiotic stress environments is minimal. We investigated how species interactions influence the functional trait responses of three dominant dune grasses to common abiotic stressors.
Methods We performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata Fern., Uniola paniculata L. and Spartina patens Muhl.) in different interspecific combinations, using sand burial and sea spray as abiotic stressors. Sand burial was applied once at the beginning of the study. Sea spray was applied three times per week using a calibrated spray bottle. Morphological functional trait measurements (leaf elongation, maximum root length, aboveground biomass and belowground biomass) were collected at the end of the study. The experiment continued from May 2015 to August 2015.
Important findings Species interactions between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata, indicating that interactions with U. paniculata could alter dune community structure. Furthermore, A. breviligulata had a negative interaction with S. patens, which decreased S. patens functional trait responses to abiotic stress. When all species occurred together, the interactions among species brought about coexistence of all three species. Our data suggest that species interactions can change traditional functional trait responses of dominant species to abiotic stress.

Aims Lignin is generally considered as an important indicator of soil organic carbon (SOC) storage and dynamics. To evaluate the effects of plant communities and soil depth on soil lignin is critical to better understand forest carbon cycling.
Methods We compared lignin content and chemical signature in three soil depths of four major plant communities in a subtropical forest, which located in the north part of Wuling Mountains, China. Lignin was measured using CuO oxidation method.
Important findings Both lignin content and its biochemical signature in plant litter varied among communities. However, these differences were mostly no longer exist in the upper soil layers. Lignin chemistry in soils inherited some of the biochemical signature of lignin in litter, but in a diminished magnitude. These results suggest that different plant communities had similar decomposition process with varying rates, caused diminished differences in lignin content and its biochemical signature. Lignin content decreased with soil depth, but the biochemical signature of lignin was not significantly different among soil layers for all communities, which suggests that vertical movement of lignin within the soil profile is very likely a key process causing this similar biochemical signature. These results emphasized the important roles of lignin inputs and soil eluviation in shaping lignin characteristics and distribution in forest soils, which pinpoint the urgent need to consider hydrological processes in studying forest soil carbon cycling.

Aims As a unique geographical unit of the earth, the Tibetan Plateau is extensively covered by various Stipa communities. However, their vegetation features have not been reported systematically till now, especially in some scantily explored regions. In this study, we endeavor to reveal the community types, quantitative characteristics and climatic distribution patterns of Stipa steppes in these areas based on primary relevés obtained from fieldwork.
Methods We collected a total of 223 plots in 79 study sites in the Changthang Plateau and the Yarlung Zangbo Valley, ranging from 79°E to 91°E. The categories of Stipa formations were identified according to the classification scheme in Vegetation of China and then verified by Nonmetric Multidimensional Scaling. We performed detrended correspondence analysis and detrended canonical correspondence analysis to hunt for the alteration of Stipa communities along the precipitation gradient. Quantitative characteristics including species richness, coverage, biomass as well as importance values (IV) of dominant species were calculated and visualized, respectively.
Important findings Stipa steppes in scantily explored regions of the Tibetan Plateau are classified into 11 formations but major formations are rather limited in number. Formation (form.) Stipa purpurea is the most widespread Stipa assemblages not only in scantily explored regions but also across the whole Tibetan Plateau. The characteristics of Stipa communities, including coverage, species richness, productivity and IV of dominant species, demonstrate the features of typical alpine steppes on the Tibetan Plateau. Precipitation proves to be the prime climatic factor controlling the distribution patterns of Stipa assemblages. Form. Stipa subsessiliflora var. basiplumosa and form. Stipa glareosa normally distribute in arid habitats, but rainfall for the former is of greater variance. Form. Stipa roborowskyi and form. Stipa capillacea favor moderately moist environment. Form. Stipa purpurea and form. Stipa roborowskyi can tolerate a fairly broad range of precipitation.

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 In recent years, there has been an increased interest in examining changes in forest systems in response to drought, flooding, hurricanes and climate change. In the southern United States, forested wetlands are of special interest because of the extent of these forests. Coastal plain forested wetlands are among the most vulnerable to these climatic impacts. One of the problems in developing management practices for these coastal areas is the difficulty in adequately describing productivity relations and predicting how the structure and function of these communities might be affected by natural or anthropogenic disturbances. Community response to environmental change often occurs over a period of years, and the majority of reported studies are for 1–3 years in duration. This study documents long-term changes (10 years) in structure, composition and growth along a catena of high water table forested sites of an ancient beach ridge landscape in coastal South Carolina.
Methods Aboveground net primary production (ANPP) of trees was monitored from 2000 to 2009 on three sites within a longleaf pine-swamp blackgum forest system on the southern end of the Waccamaw Neck area of Georgetown County, SC. Permanent study plots (20 × 25 m) were established across a moisture gradient (Dry, Intermediate, and Wet). Water levels were continuously monitored, litterfall was measured monthly and growth of trees ≥10 cm diameter at breast height was monitored on an annual basis. Annual litterfall and tree production values were summed to provide estimates of ANPP.
Important findings The study site was under severe drought conditions July 2001 through late summer 2002 and again in 2007. Diameter growth was affected in all three sites, but with different patterns. It seems that diameter growth in the Wet site was more sensitive to drought conditions in 2001–02 and 2007 than either Dry or Intermediate sites. While droughts did not seem to have a significant impact on litterfall in the Wet site, litterfall in the Intermediate site was more sensitive to the drought than either Dry or Wet sites. ANPP was significantly lower in both Intermediate and Wet sites in 2001 at ≤602 g/m 2. Highest ANPP (>1?000 g/m 2) occurred in the Intermediate and Wet sites in 2003 following a return to more normal water levels at the end of the drought. Maximum tree production occurred on the Wet site in 2003 (657 g/m 2), which exceeded total ANPP of any site in 2001. In the Dry site, ANPP remained relatively consistent throughout the study when compared to Wet and Intermediate sites. While litterfall estimates are well defined with 3–5 years of data, data collection is continuing to assess impact of drought on stem growth across the gradient, which is still not clear with 10 years of data.

Carbon (C), nitrogen (N) and phosphorus (P) stoichiometry strongly affect functions and nutrient cycling within ecosystems. However, the related researches in shrubs were very limited. In this study, we aimed to investigate leaf stoichiometry and its driving factors in shrubs, and whether stoichiometry significantly differs among closely related species.

We analyzed leaf C, N and P concentrations and their ratios in 32 species of Ericaceae from 161 sites across southern China. We examined the relationships of leaf stoichiometry with environmental variables using linear regressions, and quantified the interactive and independent effects of climate, soil and species on foliar stoichiometry using general linear models (GLM).

The foliar C, N and P contents of Ericaceae were 484.66, 14.44 and 1.06 mg g?1, respectively. Leaf C, N and P concentrations and their ratios in Ericaceae were significantly related with latitude and altitude, except the N:P insignificantly correlated with latitude. Climate (mean annual temperature and precipitation) and soil properties (soil C, N and P and bulk density) were significantly influenced element stoichiometry. The GLM analysis showed that soil exerted a greater direct effect on leaf stoichiometry than climate did, and climate affected leaf traits mainly via indirect ways. Further, soil properties had stronger influences on leaf P than on leaf C and N. Among all independent factors examined, we found species accounted for the largest proportion of the variation in foliar stoichiometry. These results suggest that species can largely influence foliar stoichiometry, even at a lower taxonomic level.

Aims The impact of global warming on belowground processes, especially on fine root production, is poorly understood in comparison with its aboveground counterpart.
Methods Here, we compiled 227 measurements to assess the influence of temperature and precipitation on fine root biomass of Norway spruce (Picea abies [L.] Karst) forest ecosystems in the Eurasia boreal region.
Important findings We found that fine root biomass decreased significantly with latitudes. There was a biomass increase of 0.63 Mg ha^-1 and 0.32 Mg ha^-1 for fine roots <2 and <1 mm in diameter, respectively, with 1°C increase of mean annual temperature. There was an increase of 0.5 and 0.1 Mg ha^-1 per 100 mm year^-1 precipitation for the two size classes of fine roots. If the adaption of root production can match the pace of global warming and water is not a limiting factor for plant growth, fine root biomass would be expected to increase by 40–140% in response to the predicted increase in temperature (3–10°C) over the next century. Our analyses highlighted the strongly positive influences of temperature and precipitation on belowground function, suggesting that predicted future climate change could substantially enhance belowground biomass in the boreal region where the greatest warming is anticipated. This potential increase of belowground biomass, coupled with aboveground biomass, may provide a better understanding of climate–ecosystem feedbacks.

Aims Riparian forests in the Brazilian Cerrado, also known as gallery forests, are very heterogeneous in structure, species composition and ecological features due to strong and abrupt variations of soil, hydrological and topographic properties. However, what are the variables driving forest carbon stock and productivity, mortality and recruitment in disturbed gallery forests?
Methods We used 36 permanent plots data from a gallery forest in the Brazilian Cerrado. We investigated how tree community dynamics vary in a gallery forest on two contrasting disturbance levels—logged and non-logged—across a topographic gradient intrinsically related to differences in moisture conditions, edge effects, as well as soil fertility and texture.
Important findings Soil variables were reduced into principal components and we used structural equation modelling to disentangle covarying variables. We also included carbon stocks as a determinant variable of dynamics rates. Logged forest had 50% higher productivity than non-logged forest and streamside forest had aboveground carbon stocks 70% higher than the forest edge. Both logging and natural disturbance drove variation in the carbon stocks which contributed to shaping productivity and recruitment rates. Distance from the river also drove mortality and carbon stock rates. Areas with high-carbon stocks favoured higher competition and lessened productivity and recruitment rates. Although soil fertility and texture are considered crucial components shaping forest dynamics, there was no clear influence of those variables on the present forest, probably because the strong effects of soil moisture, forest edge and disturbance disrupted the correlation between soil and forest dynamics.

Aims Effects of climate change, especially changes in temperatures and precipitation patterns, are particularly pronounced in alpine regions. In response, plants may exhibit phenotypic plasticity in key functional traits allowing short-term adjustment to novel conditions. However, little is known about the degree of phenotypic plasticity of high elevation species relative to mid elevation congeners.
Methods We transplanted 14 herbaceous perennial species from high elevation into two common gardens (1050 and 2000 m.a.s.l.) in the Swiss Alps, and we examined plastic responses in key functional traits to changes in temperature and soil water availability. This design was replicated with 14 congeneric species from mid elevation to assess if the degree of phenotypic plasticity differs between mid and high elevation species. Survival was assessed across two growing seasons, while aboveground biomass and specific leaf area (SLA) were measured after the first growing season, and biomass allocation to belowground and reproductive structures after the second. Moreover, a phenotypic plasticity index was calculated for the functional traits to compare the degree of plasticity between mid and high elevation species.
Important findings Aboveground biomass was higher in mid elevation species relative to high elevation congeners in all treatments, yet decreased for both with elevation and drought. Similarly, SLA decreased with elevation and drought. Root mass fraction (RMF) was generally higher in high elevation species, and decreased with drought at the lower site. Drought increased the allocation to reproductive structures, especially when plants were grown at their elevation of origin. Interestingly, no difference was found in the degree of phenotypic plasticity averaged across mid and high elevation species for any of the studied functional traits. These results indicate that phenotypic plasticity in the focal traits did not depend on the elevation of origin of the species. Plasticity was not related to environmental heterogeneity, nor constrained by selective pressures at high elevation. However, both species groups showed a remarkable capacity for short-term acclimation to a prospective climate through rapid adjustments in key functional traits.

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 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.

Aims Plants with extrafloral nectaries (EFNs) are common in tropical and subtropical habitats and, despite many other arthropods also forage for EFN, most studies solely focused on the defense mutualisms between EFN plants and ants. This study aims at a quantitative assessment of the entire arthropod community that visits EFN trees to compare visitor communities between different tree species to disentangle the mechanisms that may drive EFN visitor community composition. We also test for tree diversity effects on EFN visitors, as it is unknown if local tree species richness relates to the abundance and species richness of arthropods foraging for EFN.
Methods We sampled EFN-visiting arthropods in the experimental tree species richness gradient of the BEF-China Experiment, the currently largest forest diversity experiment in the world, and tested if tree species richness affects EFN visitors and if visitor community composition differs between EFN tree species. In a second step, we analyzed the EFN of Ailanthus altissima and Triadica cochinchinensis, the two EFN tree species with highest visitor abundance, for sugars and amino acids (AA) to test if tree species-specific differences in nectar chemistry translate to differing visitor communities. Lastly, we conducted a choice experiment using different artificial nectar solutions to test if nectar quality affects foraging decisions of ants, the most frequent EFN visitors in our study sites.
Important findings EFN trees in young successional forests in subtropical South-East China are visited by a diverse assemblage of arthropods including ants, beetles, flies, and spiders. Albeit ants accounted for about 75% of all individuals, non-ant visitors were by far more species rich. Visitor abundance and species richness declined with increasing tree species richness, suggesting a resource dilution effect, because plots with more tree species had proportionally less EFN tree individuals and thus lower nectar availability. Ailanthus altissima and T. cochinchinensis were visited by different arthropods and their nectar had species-specific AA composition and sugar concentration, indicating that differences in visitors may, at least partly, be explained by differences in nectar chemistry. These findings are supported by the choice experiment, in which artificial nectars containing sugar solutions supplemented with essential AAs attracted more ants than pure sugar solutions or sugar solutions supplemented with non-essential AAs. Our results improve the understanding of the complex ecology of EFN trees, a plant life form that might be crucial for understanding how tree diversity influences patterns of tree growth in young successional tropical and subtropical forests.

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 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 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.

Cistus albidus reproductive traits have been studied on typical Mediterranean shrublands along a water availability gradient in Northeastern Iberian Peninsula. Germination of this species is known to be highly favoured by fire. Moreover, Mediterranean species are particularly dependent on water availability. Therefore, we establish the hypothesis that in addition to fire disturbance, seedling recruitment in this Mediterranean seeder will be improved in drought-induced episodes resulting in generalized canopy die-off.

Individuals of several populations of C. albidus were collected and the size, weight and number of fruits and seeds were measured. Germination tests were also carried out on five pre-germination treatments: seeds’ exposure to heat shock, imbibition, two cycles of imbibition/desiccation and the combination of heat shock and imbibition and imbibition/desiccation cycles. Moreover, the number of seedlings after a drought event was surveyed in the field and correlated with canopy die-off.

Our study shows the variability of the C. albidus reproductive traits, such as germination rate or fruit production, along the water availability gradient. This variability resulted in a decrease in fruit production but an increase in successful germination under drier conditions. Cistus albidus seeds increased germination with heat, demonstrating their ability to successfully establish after fire. However, recruitment was not exclusively fire dependent since seedling establishment was higher under C. albidus canopies that had collapsed after the extreme drought. Finally, adult density increased C. albidus die-off and mortality, as well as seedling establishment. These results suggest that this species exhibits a trade-off between different reproductive outcomes (i.e. seed production vs. viability), which in turn is determined by climate. This study also provides evidence of how intra-specific competition, climate, particularly drought events and fire disturbance, can determine the success of key early stages of the life history of a common, representative Mediterranean fire-prone seeder shrub.

Aims Facilitation by nurse plants is a common interaction in harsh environments and this positive plant–plant interaction may promote vegetation recovery in ecosystems affected by human activities. Determining the relevance of this process, however, requires assessing how nurse plants influence the establishment of other species, as well as the proportion of species in the regional species pool that would benefit from the presence of nurse plants in human-disturbed areas. Further, since vegetation recovery is a time-dependent process, the community-level consequences of facilitation are likely to vary among landscapes with different disturbance history. Thus, an integrative perspective of the relevance of nurse plants for vegetation recovery could be obtained by measuring their effects across different human-disturbed landscapes of the target region. This study focuses on these issues and uses a regional-scale approach to assess the community-level effects of a widespread nurse plant of American deserts, the creosotebush (Larrea tridentata).
Methods This study was conducted in the southernmost portion of Chihuahuan Desert because most floodplain valleys of this region have been affected by human activities during the past centuries. For this study, we selected 10 floodplain valleys differing in their age (i.e. the time elapsed after human activities were ceased). At each landscape, we measured the cover of creosotebushes and the proportion of plant species positively associated with them, as well as the density of seeds in the soil beneath creosotebush canopies. All these data were regressed against the age of the landscapes. Further, to assess whether positive association patterns were due to facilitation or other processes, we conducted field experiments and measured the ecophysiological performance of plant species established beneath and outside creosotebush canopies.
Important findings Most plant species from the target region were positively associated to creosotebushes, and our field experiments and ecophysiological measures indicated that these distribution patterns can be attributed to facilitative interactions. In most landscapes, the density of seeds was higher beneath creosotebushes than in the surrounding habitats, suggesting that these shrubs may also act as seed traps. The community-level effects of creosotebushes increased with landscape age and creosotebush cover, indicating that magnitude of these effects depends on the disturbance history of each site. These results highlight the relevance of performing large-scale assessments for identifying the consequences of facilitation on vegetation recovery across space and time. We then propose that this kind of large-scale approach should be taken into account in the development of conservation programs aimed at the recovery and preservation of plant biodiversity in harsh environments.

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 Soil respiration (Rs) is a major process controlling soil carbon loss in forest ecosystems. However, the underlying mechanisms leading to variation in Rs along forest successional gradients are not well understood. In this study, we investigated the effects of biotic and abiotic factors on Rs along a forest successional gradient in southeast China.
Methods We selected 16 plots stratified by forest age, ranging from 20 to 120 years. In each plot, six shallow collars and six deep collars were permanently inserted into the soil. Shallow and deep collars were used to measure Rs and heterotrophic respiration (Rh), respectively. Autotrophic soil respiration (Ra) was estimated as the difference between Rs and Rh. Litter layer respiration (R L) was calculated by subtracting soil respiration measured in collars without leaf litter layer (R NL) from Rs. Rs was measured every 2 months, and soil temperature (ST) and soil volumetric water content (SVWC) were recorded every hour for 19 months. We calculated daily Rs using an exponential model dependent on ST. Daily Rs was summed to obtain cumulative annual Rs estimates. Structural equation modelling (SEM) was applied to identify the drivers of Rs during forest succession.
Important findings Rs showed significant differences among three successive stages, and it was the highest in the young stage. Ra was higher in the young stage than in the medium stage. Cumulative annual Rs and Ra peaked in the young and old stages, respectively. Cumulative annual Rh and respiration measured from soil organic matter (R SOM) decreased, whereas R L increased with forest age. The SEM revealed that cumulative annual Rs was influenced by fine root biomass and SVWC. Our results indicated that the dominant force regulating Rs on a seasonal scale is ST; however, on a successional scale, belowground carbon emerges as the dominant influential factor.

To explore the pattern of the leaf functional traits of shrub species along a latitudinal gradient in eastern China and determine the driving factors of leaf trait variation at a large scale.

We investigated the leaf thickness (LT), leaf area (LA), specific leaf area (SLA) and leaf dry mass content (LDMC) of 185 shrub species from 13 sites across eastern China. The trends of these four-leaf traits were analyzed with respect to latitude, and the differences between different life forms (e.g., evergreen and deciduous) and habitats (e.g., understory and typical) were compared. We quantified the effects of the plant life forms and environmental factors on the leaf traits via mixed-model analyses.

The LT and LA decreased, whilst and the LDMC increased, as the latitude increased, and significant differences in these traits were observed between the different plant life forms. The LT and LA were smaller, whereas the SLA and LDMC were larger in deciduous shrubs than in evergreen shrubs. Among the different habitats, the LA and SLA were larger, while the LDMC was smaller in understory shrubs than in typical shrub species. These results indicate that typical shrub species are better adapted to drier environments, as indicated by a reduced LT and increased LDMC. Furthermore, general linear models showed that variations in the four-leaf traits with respect to latitude were mainly caused by a shift in plant life forms.

Aims Are there trends of increasing/decreasing dispersion of single, categorical traits related to early/late-successional species between stages of community development? If yes, are these trends dependent on species pool extension and habitat scale? Is there a consistent reduction in single trait convergence or divergence in any seral stage when scaling down from ecological to local species pool?
Methods Presence of all vascular species rooted within plots of 5 × 5 m was recorded in assemblages of exposed mining spoils (EMS) and heathlands (HTL), which form a chronosequence on two abandoned ore tailing heaps located close to each other in the south-eastern Carpathians (Romania). Fifteen nominal, trait attributes of plant species co-occurring in the two seral assemblages were collected from available databases and subsequently classified as either successionally 'pioneer' or 'mature'. The strength of single trait convergence or divergence was estimated by comparison with null plant assemblages at patch type (meta-community) level by reference to the ecological or local species pool, and at community level.
Important findings At patch type level, all pioneer and mature trait attributes (apart from short life span), with significant variation between the two seral stages, increased and, respectively, decreased in dispersion irrespective of species pool extension. However, these trends were more conspicuous when using the ecological species pool, very likely due to relaxation in abiotic filtering and dispersal limitation. At community level, no consistent trends were observed between EMS and HTL assemblages, probably because most trait attributes were sorted by microenvironmental filters displaying high variation, like topography or habitat patch geometry. In both seral stages, there was a general weakening of trait convergence or divergence at patch type level when scaling down from the ecological to the local species pool, which was due to niche space contraction. At community level, there was a trend of rise in dispersion of pioneer attributes along the observed chronosequence, presumably imputable to increasing competition for light and underground water, but an opposite trend of dispersion drop in mature attributes was not so evident. Based on these findings, we proposed two rules of thumb concerning the expected changes in dispersion of trait attributes at patch level along successions and between levels of species pool extension. In conclusion, trends in the successional dynamics of pioneer and mature trait dispersion are clearly detectable at meta-community level, especially by reference to the ecological species pool. Habitat scale and species pool extension are key factors to consider and report when estimating the magnitude of single trait dispersion.

Aim s: The long-term effects of changing fire regimes on the herbaceous component of savannas are poorly understood but essential for understanding savanna dynamics. We present results from one of the longest running (>44 years) fire experiments in savannas, the experimental burn plots (EBPs), which is located in the Kruger National Park (South Africa) and encompasses four major savanna vegetation types that span broad spatial gradients of rainfall (450–700 mm) and soil fertility.Methods: Herbaceous vegetation was sampled twice in the EBPs using a modified step-point method, once prior to initiation of the experiment (1954) and again after 44–47 years. Different combinations of three fire frequency (1-, 2- and 3-year return intervals) and five season (before the first spring rains, after the first spring rains, mid-summer, late summer and autumn) treatments, as well as a fire exclusion treatment, were applied at the plot level (~7 ha each), with each treatment (n = 12 total) replicated four times at each of the four sites (n = 192 plots total). The effects of long-term alterations to the fire regime on grass community structure and composition were analyzed separately for each site.Important Findings: Over the 44+ years duration of the experiment, fires were consistently more intense on sites with higher mean annual rainfall (>570 mm), whereas fires were not as intense or consistent for sites with lower and more variable rainfall (<510 mm) and potentially higher herbivory due to greater soil fertility. Because the plots were open to grazing, the impacts of herbivory along with more variable rainfall regimes likely minimized the effects of fire for the more arid sites. As a consequence, fire effects on grass community structure and composition were most marked for the higher rainfall sites and generally not significant for the more arid sites. For the high-rainfall sites, frequent dry season fires (1- to 3-year return intervals) resulted in high grass richness, evenness and diversity, whereas fire exclusion and growing season fires had the lowest of these measures and diverged the most in composition as the result of increased abundance of a few key grasses. Overall, the long-term cumulative impacts of altered fire regimes varied across broad climatic and fertility gradients, with fire effects on the grass community decreasing in importance and herbivory and climatic variability likely having a greater influence on community structure and composition with increasing aridity and soil fertility.

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 .