Effects of tree sapling diversity and nutrient addition on herb-layer invasibility in communities of subtropical species

Exotic species are assumed to alter ecosystem functioning. However, little is known of the relationships within vertically structured plant communities such as forests, where tree saplings interact with herbaceous species, especially in the early phases of succession. This relationship was tested in a common garden experiment which assessed the impacts on tree saplings and herbaceous species following nutrient addition and the introduction of exotic herb species. The experiment was established in South- East China using four broad-leaved tree species (Elaeocarpus decipiens, Schima superba, Castanea henryi and Quercus serrata) to study the relationships between tree sapling diversity, herb-layer productivity and invasibility. Tree saplings were planted in monoculture and in mixtures of two and four species. A full factorial design was applied, within which species composition was crossed with nutrient and exotic seed-addition treatments. The seed-addition treatment included mixtures of seeds from eight exotic herb species, and herb community attributes were assessed after a four month growing season. Results indicate that certain tree species negatively affect native as well as exotic herbs;however, the high productivity of native herbs had a stronger negative impact on exotic species than tree saplings. Nutrient addition increased the productivity of exotic herbs but had no effect on native herbs. Remarkably, exotic species introduction had a negative feedback effect on the growth of tree saplings, which highlights the potential of exotic herbs to diminish tree recruitment. Although tree saplings reduced invasive effects on the herb-layer during the earliest phase of forest succession, nutrient addition had a more profound and opposite effect on these invaders.

Crown and leaf traits as predictors of subtropical tree sapling growth rates

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.

Interspecific and intraspecific variation in specific root length drives aboveground biodiversity effects in young experimental forest stands

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

On the combined effect of soil fertility and topography on tree growth in subtropical forest ecosystems—a study from SE China

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 m2 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 50 cm 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.

Positive effects of tree species richness on fine-root production in a subtropical forest in SE-China

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(≤2 mm 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 0.74 mm,mean SRL ranged from 12.43 m·g^(−1)to 70.22 m·g^(−1)and mean vertical distribution indexβranged 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.

Soil respiration is driven by fine root biomass along a forest chronosequence in subtropical China

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.

Limited tree richness effects on herb layer composition,richness and productivity in experimental forest stands

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.

Impact of tree diversity and environmental conditions on the survival of shrub species in a forest biodiversity experiment in subtropical China

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 42000 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.

Shifts in community leaf functional traits are related to litter decomposition along a secondary forest succession series in subtropical China

Aims We investigated shifts in community-weighted mean traits(CWm)of 14 leaf functional traits along a secondary successional series in an evergreen broadleaf forest in subtropical southeast China.most of the investigated traits have been reported to affect litter decomposition in previous studies.We asked whether changes in CWms along secondary succession followed similar patterns for all investigated traits and whether the shifts in CWm indicated a change in resource use strategy along the successional gradient.using community decomposition rates(k-rates)estimated from annual lit-ter production and standing litter biomass,we asked whether the dynamics of litter decomposition were related to changes in leaf functional traits along the successional series.Methods twenty-seven plots were examined for shifts in leaf CWm traits as well as in k-rates along a series of secondary forest succession cov-ered in the framework of the bEF-China project.We investigated whether the changes in CWms followed similar patterns for all traits with ongoing succession.three alternative linear models were used to reveal the general patterns of shifts in CWm trait values.moreover,multiple regression analysis was applied to investigate whether there were causal relationships between the changes in leaf functional traits and the dynamics of litter decomposition along secondary succession.We furthermore assessed which traits had the highest impact on community litter decomposition.Important Findingsshifts in CWm values generally followed logarithmic patterns for all investigated traits,whereas community k-rates remained stable along the successional gradient.In summary,the shifts in CWm values indicate a change in community resource use strategy from high nutrient acquisition to nutrient retention with ongoing succession.stands with higher CWm values of traits related to nutrient acquisition had also higher CWm values of traits related to chemical resistance,whereas stands with higher CWm values of traits related to nutrient retention exhibited higher CWm values in leaf physical defense.moreover,high values in CWm values related to nutritional quality(such as high leaf phosphorus concentrations)were found to promote com-munity k-rates,whereas high values in physical or chemical defense traits(such as high contents in polyphenols or high leaf toughness)decreased litter decomposition rates.In consequence,litter decom-position,which was simultaneously affected by these characteristics,did not change significantly along succession.our findings show that leaf decomposition within the investigated communities is dependent on the interplay of several traits and is a result from interactions of traits that affect decomposition in opposing directions.

Early positive effects of tree species richness on soil organic carbon accumulation in a large-scale forest biodiversity experiment

Aims tree species richness has been reported to have positive effects on aboveground biomass and productivity,but little is known about its effects on soil organic carbon(SOC)accumulation.Methods to close this gap,we made use of a large biodiversity-ecosystem functioning experiment in subtropical china(BEF-china)and tested whether tree species richness enhanced SOC accumulation.In 2010 and 2015,vertically layered soil samples were taken to a depth of 30 cm from 57 plots ranging in tree species richness from one to eight species.Least squares-based linear models and analysis of variance were used to investigate tree diversity effects.Structural equation modeling was used to explore hypothesized indirect relationships between tree species richness,leaf-litter biomass,leaf-litter carbon content,fine-root biomass and SOC accumulation.Important Findings Overall,SOC content decreased by 5.7 and 1.1 g C kg^(-1) in the top 0-5 and 5-10 cm soil depth,respectively,but increased by 1.0 and 1.5 g C kg^(-1) in the deeper 10-20 and 20-30 cm soil depth,respect-ively.converting SOC content to SOC stocks using measures of soil bulk density showed that tree species richness did enhance SOC accumulation in the different soil depths.these effects could only to some extent be explained by leaf-litter biomass and not by fine-root biomass.Our findings suggest that carbon storage in new forests in china could be increased by planting more diverse stands,with the potential to contribute to mitigation of climate warming.

Biodiversity-ecosystem functioning research in Chinese subtropical forests

Worldwide,forests provide habitat for a large diversity of plants,animals and microbes.At the same time,forest ecosystems are essential providers of multiple ecosystem services important for human well-being.However,the relationship between biodiversity and ecosystem functioning has only been little researched in forests and therefore its role for the functioning of forest ecosystems and their services is not yet understood.If results from comparable studies in other ecosystems are considered(e.g.Balvanera et al.2006),it is conceivable that diverse forests would,for example,grow faster,produce more biomass,store more carbon and better maintain soil fertility and plant-insect interactions and functional stability than less diverse forests or monoculture tree plantations.The hypothesis that biodiversity increases and stabilizes interactions and functions is the common theme of the papers in the present issue of the Journal of Plant Ecology.

林下生物量影响因素:幼龄林树种特性比丰富度更重要

生物多样性与生态系统功能的正相关关系己被广泛报道,其主要来源于对草原生态系统的研究。然而,该结论并不一定适用于更复杂的环境,例如具有不同垂直层次的森林。举例而言,己有研究表明上层乔木树种丰富度与林下生产力降低有关。树种丰富度是否会通过增加(由于生境异质性)或降低(通过增强竞争)资源的可利用性进而影响林下生产力,以及林下生产力是否受树种特性的影响更大,这些影响机制都可能会随着时间的推移而改变。此外,研究还表明,丰富度-生产力关系随着环境背景的变化而改变。本研究利用可以操控树种丰富度的实验林场研究了这些不同垂直层位里的时间和环境动态。在中国亚热带森林生物多样性与生态系统功能(BEF-China)研究计划的框架下,我们在3年时间里沿树种丰富度梯度反复采集林下生物量样本,研究了不同环境处理中树种丰富度、树种特性和时间对林下生物量的影响。尽管我们发现乔木层特性对林下生物量有显著和一致的影响,但是树种丰富度对后者却不具有这种影响。另外,在森林结构层之间,可能并不存在单一的、具有普遍性的上层乔木树种丰富度与林下生产力的相关关系,并且与上层乔木相关的环境因素(如透光率)对林下生产力的贡献程度会随着时间而变化。总体而言,我们的结果表明,在研究森林结构层之间的关系时应将时间动态变化考虑在内。

Changes in carbon storages of Fagus forest ecosystems along an elevational gradient on Mt. Fanjingshan in Southwest China

森林生态系统碳库通常包括植被、土壤、凋落物与木质残体等几个组分。本研究旨在探讨中国一种重要的森林生态系统—水青冈林(Fagus L.)中这些碳库的沿海拔梯度格局变化及其驱动因子。在中国贵州省梵净山,沿海拔梯度(1095–1930 m)调查了9个水青冈林各碳库的碳储量。采用方差分解探讨了林龄、气候及其他因子对碳储量的影响,同时对梵净山与贵州和全球其他地区水青冈林的碳储量进行了比较。梵净山水青冈林生态系统碳储量在190.5–504.3 Mg C ha^(–1)之间,其主要组分包括植被碳库(33.7–73.9%)和土壤碳库(23.9–65.5%),而木质残体(0.05–3.1%)和凋落物(0.2–0.7%)对该生态系统碳储量的贡献不超过4%。随海拔升高,生态系统碳储量呈增加趋势,其中植被与木质残体碳库增加,而凋落物与土壤碳库无明显的变化趋势。对梵净山水青冈林,气候与林龄是其各组分碳储量海拔格局形成的主要原因;而对于全球水青冈林,林龄是其碳储量变化的主导因子。相比全球其他地区的水青冈林,贵州水青冈林具有较高的植被碳储量积累速率,这可能与贵州较高的降水量有关。本研究结果有助于理解中国水青冈林碳收支及其在区域碳循环中的可能作用,同时强调了林龄与气候对碳积累的重要性。

Does higher ploidy level increase the risk of invasion?A case study with two geo-cytotypes of Solidago gigantea Aiton(Asteraceae)

Aims Understanding the role of genetics in biological invasions has become an important aspect for modern plant ecology.Many studies suggest that increased ploidy level benefits the success of an invasive species,but the basis for this phenomenon is not fully understood.in its native,North American range,Solidago gigantea has three geo-cytotypes comprising di-,tetra-and hexa-ploid populations,while in Europe,where it is highly invasive,S.gigantea stands are composed primarily of tetraploid individu-als.Our study investigates whether North American hexaploids can induce a greater risk of invasion,due to their higher perfor-mance in a non-native range,as compared to the existing tetra-ploids of that range.Methods We performed greenhouse and common garden experiments along with microsatellite analyses to test whether differences in chromo-some number and origin of the species mean superior fitness in the introduced range.Important Findings Genetic diversity was significantly higher in the native hexaploid popula-tions(A_(R)=6.04;H_(e)=0.7794),rather than the non-native tetraploid pop-ulations(A_(R)=4.83;H_(e)=0.6869).Furthermore,differentiation between geo-cytotypes was moderate(ρST=0.1838),which was also confirmed by their clear segregation in principal component analysis and structure analyses,proving their different genetic structure.in contrast to genetic diversity,the non-native tetraploid geo-cytotype performed better in the common garden experiment,implying that higher genetic diversity does not always mean better success.Our results suggest that native hexa-ploids do not present a greater risk,as assessed by their performance in the introduced range,when compared to the non-native tetraploids,as was suggested by previous studies.Nevertheless,their introduction is still undesirable due to their different genetic structure,which,through hybridization,could give a new drive to the invasion of S.gigantea.