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 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.

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 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 Grasslands are the world's most extensive terrestrial ecosystem, and are a major feed source for livestock. Meeting increasing demand for meat and other dairy products in a sustainable manner is a big challenge. At a field scale, Global Positioning System and ground-based sensor technologies provide promising tools for grassland and herd management with high precision. With the growth in availability of spaceborne remote sensing data, it is therefore important to revisit the relevant methods and applications that can exploit this imagery. In this article, we have reviewed the (i) current status of grassland monitoring/observation methods and applications based on satellite remote sensing data, (ii) the technological and methodological developments to retrieve different grassland biophysical parameters and management characteristics (i.e. degradation, grazing intensity) and (iii) identified the key remaining challenges and some new upcoming trends for future development.
Important findings The retrieval of grassland biophysical parameters have evolved in recent years from classical regression analysis to more complex, efficient and robust modeling approaches, driven by satellite data, and are likely to continue to be the most robust method for deriving grassland information, however these require more high quality calibration and validation data. We found that the hypertemporal satellite data are widely used for time series generation, and particularly to overcome cloud contamination issues, but the current low spatial resolution of these instruments precludes their use for field-scale application in many countries. This trend may change with the current rise in launch of satellite constellations, such as RapidEye, Sentinel-2 and even the microsatellites such as those operated by Skybox Imaging. Microwave imagery has not been widely used for grassland applications, and a better understanding of the backscatter behaviour from different phenological stages is needed for more reliable products in cloudy regions. The development of hyperspectral satellite instrumentation and analytical methods will help for more detailed discrimination of habitat types, and the development of tools for greater end-user operation.

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 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 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 In forests, the herbaceous understory vegetation plays an important role for ecosystem functioning as it represents a significant component of total aboveground productivity. In addition, the herb layer contributes to overall forest species richness and controls tree species regeneration. Vice versa, trees in the overstory control understory herb and shrub growth through competition for resources. Using an experimental forest plantation with manipulated tree richness, we asked to which degree tree species richness and identity affect herb layer composition, richness and productivity and how these relationships across strata change with abiotic environmental conditions and competition intensity.
Methods In the context of the Biodiversity-Ecosystem Functioning project in subtropical China (BEF-China), we made use of the integrated BEFmod experiment arranged along a tree species richness gradient at two sites, with additional subplot treatments of phosphorus addition, herb layer weeding and no weeding. We recorded the understory vegetation and determined herb layer biomass production on a total of 201 subplots.
Important findings We found only minor effects of tree layer richness on herb layer species composition and no significant effect on herb layer richness or productivity yet. However, there were strong tree layer identity effects on all response variables, which were partly explained by differences in leaf area index and by a high share of woody species both in total herb layer species richness and biomass. There were strong treatment effects, which were largest in the 'no weeding' treatment but we did not find any treatment × tree layer richness interaction in herb layer responses. Thus, these effects are mainly explained by increased competition intensity within the herb layer in the absence of weeding. Despite the young age of the experiment, the interactions between tree species identity, tree richness and the herb layer did already emerge and can be expected to become stronger with ongoing runtime of the experiment.

Aims Studies have showed that arbuscular mycorrhizal fungi (AMF) can greatly promote the growth of host plants, but how AMF affect flowering phenology of host plants is not well known. Here, we conducted a pot experiment to test whether life cycle and flowering phenology traits of host plant Medicago truncatula Gaertn can be altered by AMF under low and high soil phosphorus (P) levels.
Methods The experiment was conducted in a greenhouse at Zhejiang University in China (120°19′E, 30°26′N) and had a completely randomized design with two factors: AMF treatments and soil P levels. Six AMF species (Acaulospora scrobiculata, As; Gigaspora margarita, Gma; Funneliformis geosporum, Fg; Rhizophagus intraradices, Ri; Funneliformis mosseae, Fmo and Glomus tortuosum, Gt.) were used, and two soil P levels (24.0 and 5.7 mg kg^-1 Olsen-soluble P) were designed. The six AMF species were separately inoculated or in a mixture (Mix), and a non-AMF control (NAMF) was included. When plants began to flower, the number of flowers in each pot was recorded daily. During fruit ripening, the number of mature fruits was also recorded daily. After ~4 months, the biomass, biomass P content and AMF colonization of host plant were measured. Correlation between root colonization and first flowering time, or P content and first flowering time was analyzed.
Important findings Under the low P level, first flowering time negatively correlated with root colonization and biomass P. Only host plants with AMF species As, Fg, Ri, or Mix were able to complete their life cycle within 112 days after sowing. And treatment with AMF species Fg, Gt, or As resulted in two periods of rapid flower production while other fungi treatments resulted in only one within 112 days after sowing. The cumulative number of flowers produced and biomass P content were highest with species Fg. Host biomass allocation significantly differed depending on the species of AMF. Under both soil P levels, the host plant tended to allocate more biomass to fruits in the Mix treatment than in the other treatments. These results indicated that the effects of AMF on host flowering phenology and biomass allocation differed depending on AMF species and soil P levels.

Aims Lianas are expected to influence composition, structure and functioning of forest systems due to unequal distribution across the potential set of host plants. However, our understanding of mechanisms associated with preferences for specific hosts is still limited, and so is our ability to discern between endogenous and exogenous forces driving forest dynamics in the long run. In this paper, we evaluated whether the dominant liana Hedera helix can indirectly contribute to the eventual dominance of the small multi-stemmed tree Corylus avellana in a remnant temperate forest in central Iberian Peninsula from comparatively reduced liana infestation on C. avellana relative to co-occurring woody species.
Methods Through principal component analysis and co-occurrence analysis, we studied the distribution and spatial association between woody species and the liana H. helix. We analyzed the relationship between the number of species in a plot and the number of species infested by the liana to test the hypothesis that H. helix is a generalist liana. Through generalized linear mixed models, we tested the dynamic-multi-stemmed growth form of C. avellana as a plausible life strategy to withstand, in the long run, the liana infestation. In particular, we tested (i) the relationship between stem size and the probability of H. helix infestation including all the tree species within plots and (ii) the relationship between stem size and mortality as evidence of the stem turn over in the tree C. avellana .
Important findings Our results indicate that H. helix and C. avellana significantly co-occur in mature stands of this remnant temperate forest where pioneer woody species are absent. Hedera helix severely infests all the woody species whenever stem size exceed ≈25 cm perimeter and there is physical contact at the base of the stem. This implies that all the trees in the community are potential hosts for H. helix. Mixed models indicate that both, infestation by H. helix and stem mortality, are positively related to C. avellana stem perimeter. Reduced long-term infestation of the liana by means of a multi-stemmed growth form with high stem turnover in C. avellana might be an advantage with respect to unipodial tree species. Thus, the liana-tree coexistence pattern may be interpreted as an indirect positive interaction that, contrary to previous findings, results here in species dominance instead of species coexistence.

Aims Litter decomposition is a fundamental process within ecosystem functioning, and it is largely dependent on the biodiversity of ecosystems. We explored the effects of species diversity and genetic diversity of litter on the litter decomposition rate.
Methods We used laboratory microcosms to determine whether species diversity and genetic diversity and their interaction affect leaf litter decomposition. We set up 8 treatments containing 1, 2, 4 diversity levels of four broad-leaf species (Alniphyllum fortunei, Idesia polycarpa, Cinnamomum camphora and Daphniphyllum oldhamii) both in species and genetic sense. Totally 246 microcosms containing same amount of soil and litter of prescribed diversity treatment were stored in the dark at 25°C for 12 weeks.
Important findings The effect of litter species diversity on litter decomposition was largely dependent on species composition of the litter mixture in terms of species identity. Overall, the decomposition rate increased linearly with the richness of seed family when the species identity was disregarded. However, no interactive effect of species diversity and genetic diversity on mass loss was detected. The litter decomposition rate was found to be unrelated to the initial carbon (C), whereas it was negatively correlated with the initial total nitrogen (N) and N:P ratio. However, the regression curves of the litter decomposition rate against the total P and C:N ratio displayed quadratic parabolas opening upward and downward, respectively. This study demonstrated how species and/or genetic diversity and the stoichiometry of litter per se affect litter decomposition. Further studies should be performed in the long term to ascertain how such effects operate and how they change during the decomposition process, particularly in response to varying composition and diversity of standing plants in the environments.

Aims Plants directly and indirectly interact with many abiotic and biotic soil components. Research so far mostly focused on direct, individual abiotic or biotic effects on plant growth, but only few studies tested the indirect effects of abiotic soil factors on plant growth. Therefore, we investigated how abiotic soil conditions affect plant performance, via changes induced by soil biota.
Methods In a full-factorial experiment, we grew the widespread grass Dactylis glomerata either with or without soil biota and investigated the impact of soil temperature, fertility and moisture on the soil biota effects on plant growth. We measured biomass production, root traits and colonization by arbuscular mycorrhizal fungi as well as microbial respiration.
Important findings We found significant interaction effects between abiotic soil conditions and soil biota on plant growth for fertility, but especially for soil temperature, as an increase of 10°C significantly changed the soil biota effects on plant growth from positive to neutral. However, if tested individually, an increase in soil temperature and fertility per se positively affected plant biomass production, whereas soil biota per se did not affect overall plant growth, but both influenced root architecture. By affecting soil microbial activity and root architecture, soil temperature might influence both mutualistic and pathogenic interactions between plants and soil biota. Such soil temperature effects should be considered in soil feedback studies to ensure greater transferability of results from artificial and experimental conditions to natural environmental conditions.

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.
Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.
Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.

Aims We aim to understand how small-scale genotypic richness and genotypic interactions influence the biomass and potential invasiveness of the invasive grass, Phalaris arundinacea under two different disturbance treatments: intact plots and disturbed plots, where all the native vegetation has been removed. Specifically, we address the following questions (i) Does genotypic richness increase biomass production? (ii) Do genotypic interactions promote or reduce biomass production? (iii) Does the effect of genotypic richness and genotypic interactions differ in different disturbance treatments? Finally (iv) Is phenotypic variation greater as genotypic richness increases?
Methods We conducted a 2-year common garden experiment in which we manipulated genotype richness using eight genotypes planted under both intact and disturbed conditions in a wetland in Burlington, Vermont (44°27′23″N, 73°11′29″W). The experiment consisted of a randomized complete block design of three blocks, each containing 20 plots (0.5 m 2) per disturbed treatment. We calculated total plot biomass and partitioned the net biodiversity effect into three components: dominance effect, trait-dependent complementarity and trait-independent complementarity. We calculated the phenotypic variance for each different genotype richness treatment under the two disturbance treatments.
Important findings Our results indicate that local genotypic richness does not increase total biomass production of the invasive grass P. arundinacea in either intact or disturbed treatments. However, genotypic interactions underlying the responses showed very different patterns in response to increasing genotypic richness. In the intact treatment, genotypic interactions resulted in the observed biomass being greater than the predicted biomass from monoculture plots (e.g., overyielding) and this was driven by facilitation. However, facilitation was reduced as genotypic richness increased. In the disturbed treatment, genotypic interactions resulted in underyielding with observed biomass being slightly less than expected from the performance of genotypes in monocultures; however, underyielding was reduced as genotypic richness increased. Thus, in both treatments, higher genotypic richness resulted in plot biomass nearing the additive biomass from individual monocultures. In general, higher genotypic richness buffered populations against interactions that would have reduced biomass and potentially spread. Phenotypic variance also had contrasting patterns in intact and disturbed treatments. In the intact treatment, phenotypic variance was low across all genotypic richness levels, while in the disturbed treatment, phenotypic variance estimates increased as genotypic richness increased. Thus, under the disturbed treatment, plots with higher genotypic richness had a greater potential response to selection. Therefore, limiting the introduction of new genotypes, even if existing genotypes of the invasive species are already present, should be considered a desirable management strategy to limit the invasive behavior of alien species.

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 Pigment composition is an important functional trait that can be affected by environmental factors. The objective of this study was to investigate the effect of soil salinity on pigment composition in Suaeda salsa by comparing chlorophyll and betacyanin content in the Liaohe estuary wetland, a typical coastal wetland in northeast China.
Methods We investigated the plant biomass, percentage of red leaves and pigment content (chlorophyll a, chlorophyll b and betacyanins) in S. salsa in intertidal and supratidal zones of the upper, middle and lower reaches of the Liaohe estuary wetlands. The Na + content of both the soil and plant was also measured. Full analysis of variance and multivariate analysis were used to compare differences in pigment content and Na + content between the supratidal and intertidal zones.
Important findings Pigment composition was significantly affected by soil salinity. With increasing soil salinity, the percentage of red leaves was higher in the intertidal zone than in the supratidal zone. In all three reaches, plants had lower chlorophyll a content and higher betacyanin content in the intertidal zone than in the supratidal zone. Compared to chlorophyll a, chlorophyll b was less sensitive to soil salinity. There were no differences in chlorophyll b content between the intertidal and supratidal zones in the upper and lower reaches. Furthermore, pigment composition was associated with both the plant tissue and soil Na + content. Compared to the supratidal zone, the intertidal zone had a higher Na + content in plants. There was a negative relationship between plant chlorophyll content and soil Na + content, but a positive relationship between betacyanin content and soil Na + content. Overall, the results indicated that there might be a trade-off between leaf chlorophyll and betacyanin content in S. salsa to maintain its growth and survival in high salinity environments.

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 Community assembly persists as a key topic in ecology due to the complex variation in the relative importance of assembly forces and mechanisms across spatio-temporal scales and ecosystems. Here we address a forest–savanna vegetation mosaic in the Brazilian Atlantic forest to examine the role played by soil attributes as determinants of community assembly and organization at a landscape spatial scale.
Methods We examined soil and plant assemblage attributes across 23 plots of forest and savanna in a 1600 km 2 landscape exposed to the same climatic conditions in the Atlantic forest region of northeast Brazil. Assemblage attributes included species richness, taxonomic and functional composition (community weighted mean, CWM) and functional diversity (quadratic diversity; Rao's quadratic entropy index) relative to plant leaf area, specific leaf area, leaf dry matter content, thickness and succulence.
Important findings Our results suggest that forest and savanna patches exposed to the same climatic conditions clearly differ in terms of soil attributes, plant assemblage structure, taxonomic and functional composition. By selecting particular plant strategies relative to resource economy, soil potentially affects community structure, with forest assemblages bearing more acquisitive resource-use strategies, while conservative plant strategies are more frequent in savannas. Accordingly, savanna–forest mosaics in the Atlantic forest region represent spatially organized plant assemblages in terms of taxonomic and functional features, with a signal of trait convergence in both vegetation types. Soil-mediated filtering thus emerges as a potential deterministic assembly force affecting the spatial organization of savanna–forest boundaries and mosaics.

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.

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 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.

null

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.

Aims Aquatic plants play an important role in freshwater ecosystems. Previous works have largely focused on the functional significance of plant above ground parts, with much less attention on the root structures of aquatic plants. In this study, we divided 21 aquatic plants (including five introduced plants) into multiple plant groups (different life forms, monocot/eudicot and introduced/native) with the goal of addressing two questions: (i) what root structures do aquatic plants exhibit, and (ii) are there differences among these plant groups?
Methods Twenty-one aquatic plants belonging to four life forms (free-floating, emergent, floating-leaved and submerged) were collected at the near flowering stage from a typical macrophyte-dominated lake in the Yangtze River Basin, China. The following root topological parameters were quantified: altitude (a), path length (p e), magnitude (M), mean topological length (b), topological index (TI) and normed indices q a and q b .
Important findings The root topological indices TI, q a and q b for the 21 aquatic plants were 0.724 ± 0.013, 0.290 ± 0.031 and 0.152 ± 0.024 (means ± S.E.), respectively, revealing a general pattern of dichotomous branching, except for the aquatic root of Myriophyllum aquaticum (Vell.) Verdc., which displays herringbone branching. All three topological indices were significantly lower for monocots (TI = 0.700 ± 0.130, q a = 0.191 ± 0.149 and q b = 0.086 ± 0.236) than eudicots (TI = 0.752 ± 0.206, q a = 0.405 ± 0.569 and q b = 0.229 ± 0.393), indicating that the roots of monocots are typically more dichotomous-like than those of eudicots. Among the four life forms, the three topological indices for emergent plants (TI = 0.832 ± 0.006, q a = 0.616 ± 0.018 and q b = 0.381 ± 0.014) were significantly higher than those of the other three life forms. Overall, there was no difference between the topological indices of introduced and native aquatic plants, but the introduced species M. aquaticum and Alternanthera philoxeroides (Mart.) Griseb. had both aquatic and edaphic roots as well as unusual functions, which may help explain their strong viability.

Aims It is known that taxonomic diversity can be predicted by the spatial configuration of the habitat, in particular by its area and degree of isolation. However, taxonomic diversity is a poor predictor of ecosystem functioning. While functional diversity is strongly linked to the functionality and stability of ecosystems, little is known about how changes in the spatial configuration of the habitat affect functional diversity. In this study, we evaluated whether the spatial configuration of forest patches predicts the functional diversity of plants in a fragmented forest.
Methods Five functional leaf traits (leaf dry matter content, leaf punch force, specific leaf area, leaf size and leaf thickness) were measured for 23 dominant plant species in 20 forest patches in a naturally fragmented forest on the Yucatan Peninsula. Abundance-weighted multivariate and individual trait metrics of functional diversity were calculated and correlated with size, degree of isolation and the shape of forest patches.
Important findings Patch shape was negatively correlated with multivariate and individual trait (leaf dry matter content and leaf size) metrics of functional diversity. Patch isolation measures were also negatively correlated with individual trait (leaf dry matter content, leaf punch force and leaf size) metrics of functional diversity. In other words, greater patch shape irregularity and isolation degree impoverish plant functional variability. This is the first report of the negative effects of patch shape irregularity and isolation on the functional diversity of plant communities in a forest that has been fragmented for a long time.

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.

Aims The mature meadows (MMs) and the swamp meadows (SMs) are the two most important ecosystems in the eastern Tibetan Plateau, China. Besides their substantial differences in terms of soil water conditions and thereby the soil oxygen and nutrients, however, little is known about the differences in community composition, structure, traits and productivity between these two meadows. We particularly ask whether light availability mediated by physical structure heterogeneity is a key determinant of the difference in community composition and productivity between these two meadows.
Methods We examined the community structure, composition, aboveground net primary productivity (ANPP), light availability in understory and the community-weighted means (CWMs) for leaf morphological and physiological traits in 12 random plots (5 m × 5 m) for each of the studied habitats.
Important findings The results showed that plant community in the MM had higher variation in both vertical and horizontal structure and thus had more light availability in the understory. The MM had higher species richness and greater ANPP than the SM. The CWMs of leaf morphological and physiological traits for species in the MM featured a fast-growing strategy (i.e. higher height, leaf area and net photosynthesis rate and lower nitrogen:phosphorus ratio), in contrast to those in the SM. We also found that there were significant correlations between the CWM of traits and the ANPP, indicating that some key traits in these habitats have linked to community productivity. Our study also suggests that the heterogeneity in the community structure, which affects light availability in the understory, may play an important role in determining the community composition and productivity. In conclusion, our study revealed significant differences in community structure, composition and traits between the MM and the SM, and the light availability that related closely to community structure is the key factor to determine the composition and productivity of the community of these two habitats.

Aim The ability to quantitatively measure the continuum of macroscale patterns of species invasion is a first step toward deeper understanding of their causal factors. We took advantage of two centuries worth of herbarium data, to evaluate a set of metrics to measure macroscale patterns, allowing cross-species comparisons of invasive expansion across large geographic areas.
Methods We used herbarium specimens to reconstruct county-level invasion histories for two non-native plants (Alliaria petiolata and Lonicera japonica), with distinct spatiotemporal distribution patterns over the past two centuries. Using county centroids from species' initial occurrences, we quantified point pattern metrics from multiple disciplines (e.g. urban crime analysis, landscape ecology etc.) that are historically used at smaller spatial scales, to evaluate their ability to detect macroscale spatial diffusion and amount of directional expansion. Metrics were further assessed for their ease of use, data requirements, independence from other metrics and intuitiveness of interpretation.
Important findings We identified four suitable metrics for distinguishing differences in spatial patterns: (i) standard distance, (ii) number of patches, (iii) Euclidean nearest neighbor summary class statistic coefficient of variation and (iv) mean center that when applied to county-level presence data allowed us to determine the directions by which distributions expanded and if distributions increased via outward expansion, infilling and/or jump dispersal events. These metrics when compared during the same invasion phase are capable of quantifying macroscale variability among species in their distributional and dispersal patterns. Being able to quantify differences among species in these patterns is important in understanding the drivers of species dispersal patterns. These metrics therefore represent a simple yet thorough toolset for achieving this goal.

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 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 The Cape Peninsula is a small area (471 km 2) situated on the south-westernmost tip of the Core Cape Subregion (CCR) of South Africa. Within the Cape Peninsula, Fabaceae are the third most species-rich plant family (162 species) and they have the second highest number of endemic species after the Ericaceae. However, legumes are not the dominant taxa in the vegetation. They tend to show patchy distributions within the landscape and different species assemblages usually occupy particular niches at any given locality. The present study undertook to establish if edaphic factors influence legume species distribution in the Cape Peninsula and to determine the key indicator species for the different assemblages.
Methods Soils from 27 legume sites, spanning all major geological substrates of the Cape Peninsula, were analysed for 31 chemical and physical properties. Legume species present at each site were recorded and a presence/absence matrix was generated. Cluster analysis and discriminant function analysis (DFA) were run to group the sites based on overall similarity in edaphic characteristics and to identify the soil parameters contributing towards discriminating the groups. Canonical correspondence analysis (CCA) was used to test for a correlation between legume species compositions and edaphic factors. The strength of the association between legume species and site groupings based on edaphic properties was assessed using indicator species analysis.
Important findings Based on similarity in overall soil characteristics, the sites formed three clusters: one comprising sites of sandstone geology, one with dune sand sites and the third cluster comprising sites of both shale and granite geologies (hereafter referred to as soil types). The DFA confirmed the distinctness of these clusters and the CCA showed a significant correlation between legume species composition and edaphic factors. The key edaphic parameters were clay content, iron (Fe), potassium (K), sulphur (S) and zinc (Zn). These findings reveal that the Cape Peninsula is edaphically heterogeneous and edaphically distinct habitats contain discrete legume species assemblages that can be distinguished by unique indicator species. Furthermore, multiple soil parameters, rather than a single parameter, are involved. Therefore, edaphic factors play a significant role in driving the distribution of legume species in the Cape Peninsula and discrete legume species assemblages occupy distinct habitats.

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 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.

null

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 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.

Aims Mycorrhizal fungi can re-distribute nutrients among plants through formation of underground common mycorrhizal networks and therefore may alter interspecific plant competition. However, the effect of ectomycorrhizal (EM) fungi on interspecific plant competition in subtropical forests is poorly understood. In this study, we investigated the effects of EM fungal identity and diversity on the outcome of interspecific competition of plant species in relation to different successional stages in a Chinese subtropical forest.
Materials and Methods This study selected four woody plant species, i.e. a pioneer tree Pinus massoniana, a late-pioneer tree Quercus serrata, a mid- successional tree Cyclobalanopsis glauca and a late-successional tree Lithocarpus glaber in a Chinese subtropical forest. The outcomes of interspecific competition were investigated in the seedlings of three plant pairs, i.e. between Cy. glauca and Pin. massoniana, between Q. serrata and Pin. massoniana, and between Li. glaber and Q. serrata in a pot experiment. In the Cy. glauca – Pin. massoniana combination, plants in monoculture and two-species mixture were uninoculated or inoculated with EM fungi Paxillus involutus, Pisolithus tinctorius, Cenococcum geophilum, Laccaria bicolor and a mixture of these four fungal species. In the Q. serrata – Pin. massoniana and Li. glaber – Q. serrata combinations, plants in monocultures and two-species mixtures were uninoculated or inoculated with EM fungi Pis. tinctorius, Ce. geophilum, La. bicolor and a mixture of these three fungal species. EM root colonization rate and seedling biomass of each plant species were measured, and the outcomes of interspecific competition were estimated using competitive balance index after 6-month cultivation.
Important findings All EM fungal inoculation significantly promoted a competitive ability of the mid-successional tree Cy. glauca over the pioneer tree Pin. massoniana compared with the uninoculated control treatment, and the extent to which EM fungi affected the outcome of interspecific competition was dependent on EM fungal identity in the Cy. glauca and Pin. massoniana combination. EM fungal inoculation had no significant effect on the outcomes of interspecific competition between the late-pioneer tree Q. serrata and Pin. massoniana combination and between the late-successional tree Li. glaber and Q. serrata combination, compared with the uninoculated control treatment. However, amongst the EM fungal inoculation treatments the competitive ability of Q. serrata over Pin. massoniana was significantly higher in EM fungi Ce. geophilum and La. bicolor treatments than in Pis. tinctorius treatment. EM fungal diversity did not show a complementary effect on the outcomes of interspecific competition in all three plant pairs. This study demonstrated that the effect of EM fungi on the outcome of interspecific competition was dependent on the plant pairs tested in the subtropical forest ecosystem.

Aims Phenotypic optimality models neglect genetics. However, especially when heterozygous genotypes are fittest, evolving allele, genotype and phenotype frequencies may not correspond to predicted optima. This was not previously addressed for organisms with complex life histories.
Methods Therefore, we modelled the evolution of a fitness-relevant trait of clonal plants, stolon internode length. We explored the likely case of an asymmetric unimodal fitness profile with three model types. In constant selection models (CSMs), which are gametic, but not spatially explicit, evolving allele frequencies in the one-locus and five-loci cases did not correspond to optimum stolon internode length predicted by the spatially explicit, but not gametic, phenotypic model. This deviation was due to the asymmetry of the fitness profile. Gametic, spatially explicit individual-based (SEIB) modeling allowed us relaxing the CSM assumptions of constant selection with exclusively sexual reproduction.
Important findings For entirely vegetative or sexual reproduction, predictions of the gametic SEIB model were close to the ones of spatially explicit non-gametic phenotypic models, but for mixed modes of reproduction they approximated those of gametic, not spatially explicit CSMs. Thus, in contrast to gametic SEIB models, phenotypic models and, especially for few loci, also CSMs can be very misleading. We conclude that the evolution of traits governed by few quantitative trait loci appears hardly predictable by simple models, that genetic algorithms aiming at technical optimization may actually miss the optimum and that selection may lead to loci with smaller effects in derived compared with ancestral lines.

Aims Floral traits are frequently used in traditional plant systematics because of their assumed constancy. One potential reason for the apparent constancy of flower size is that effective pollen transfer between flowers depends on the accuracy of the physical fit between the flower and pollinator. Therefore, flowers are likely to be under stronger stabilizing selection for uniform size than vegetative plant parts. Moreover, as predicted by the pollinator-mediated stabilizing selection (PMSS) hypothesis, an accurate fit between flowers and their pollinators is likely to be more important for specialized pollination systems as found in many species with bilaterally symmetric (zygomorphic) flowers than for species with radially symmetric (actinomorphic) flowers.
Methods In a comparative study of 15 zygomorphic and 13 actinomorphic species in Switzerland, we tested whether variation in flower size, among and within individuals, is smaller than variation in leaf size and whether variation in flower size is smaller in zygomorphic compared to actinomorphic species.
Important findings Indeed, variation in leaf length was significantly larger than variation in flower length and width. Within-individual variation in flower and leaf sizes did not differ significantly between zygomorphic and actinomorphic species. In line with the predictions of the PMSS, among-individual variation in flower length and flower width was significantly smaller for zygomorphic species than for actinomorphic species, while the two groups did not differ in leaf length variation. This suggests that plants with zygomorphic flowers have undergone stronger selection for uniform flowers than plants with actinomorphic flowers. This supports that the relative uniformity of flowers compared to vegetative structures within species, as already observed in traditional plant systematics, is, at least in part, a consequence of the requirement for effective pollination.