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 spe...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.展开更多
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 Geom...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.展开更多
Aims Litterfall,as an important link between aboveground and belowground processes,plays a key role in forest ecosystems.Here,we test for effects of tree species richness on litter production and litter quality in sub...Aims Litterfall,as an important link between aboveground and belowground processes,plays a key role in forest ecosystems.Here,we test for effects of tree species richness on litter production and litter quality in subtropical forest.The study further encompasses a factorial gradient of secondary succession that resulted from human exploitation.Given that a large percentage of subtropical forests are in secondary successional stages,understanding the role of biodiversity on forest re-growth after disturbance appears critical.Methods From January 2009 to December 2014,we monitored forest litterfall in 27 Comparative Study Plots that spanned a gradient of tree species richness(3-20 species)and secondary successional ages(~20 to 120 years)in Gutianshan Natural Nature Reserve,Zhejiang Province,China.The experiment is part of the biodiversity-ecosystem functioning research platform‘BEF-China’.Tree litterfall was collected in monthly intervals using litter traps.Samples were separated into leaf and non-leaf components.Leaf litter was further sorted into dominant and other species.Community level monthly leaf litter C and N contents were analysed through a full year.General linear mixed-effects models were applied to test for effects of tree species richness and successional age on litter quantity and leaf litter C/N.Important Findings Litterfall increased with species richness among and within successional age and this effect was consistent across years.Successionally older stands had higher litterfall and this effect was related to increased tree species richness.However,species richness did not change the intra-and inter-annual temporal stability of litterfall.Increasing tree species richness increased leaf litter quality(decreased C/N),while successional age had no effect.Our study indicates that more diverse forest stands produce more leaf litter and that this litter has higher N concentrations,which could promote forest growth through accelerated nutrient re-cycling.展开更多
Aims Positive plant diversity-ecosystem function relations are ultimately driven by variation in functional traits among individuals that form a community.To date,research has largely focused on the role of species di...Aims Positive plant diversity-ecosystem function relations are ultimately driven by variation in functional traits among individuals that form a community.To date,research has largely focused on the role of species diversity for ecosystem functioning.However,substantial intraspecific trait variation is common and a significant part of this variation caused by genetic differences among individuals.Here,we studied the relative importance of species diversity and seed family(SF)diversity within species for growth and herbivory in experimental subtropical tree assemblages.Methods In 2010,we set up a field experiment in subtropical China,using four species from the local species pool.Trees were raised from seeds,with seeds from the same mother tree forming an SF.We established 23 plots containing one or four species(species diversity treatment)and one or four SFs per species(SF diversity treatment).Tree growth(stem diameter,plant height and crown expansion)and herbivory(percentage leaf loss due to leaf chewers)were monitored annually from 2011 to 2013.Important findings Tree species richness promoted growth but had no effect on herbivory.In contrast,SF diversity reduced growth and increased herbivory but only so in species mixtures.Most of the observed effects were time dependent,with the largest effect found in 2013.Our results suggest that biodiversity can affect plant performance directly via tree species-species interactions,or context dependent,via potential effects on inter-trophic interactions.Two important conclusions should be drawn from our findings.Firstly,in future studies regarding biodiversity and ecosystem functioning(BEF)relationships,intraspecific genetic diversity should be given similar weight as species diversity as it has often been neglected and its effects are not well understood.Secondly,we demonstrate opposite effects of biodiversity among and within species,stressing the importance to consider the effects of multiple levels of biodiversity simultaneously.展开更多
Aims Most biodiversity-ecosystem functioning research has been carried out in grassland ecosystems,and little is known about whether forest ecosystems,in particular outside the temperate zone,respond similarly.Here,we...Aims Most biodiversity-ecosystem functioning research has been carried out in grassland ecosystems,and little is known about whether forest ecosystems,in particular outside the temperate zone,respond similarly.Here,we tested whether productivity,assessed as leaf area index(LAI),increases with species richness in young experimental stands of subtropical trees,whether this response is similar for early-season leaf area(which is dominated by evergreens)and seasonal leaf area increase(which is dominated by deciduous species),and whether responses saturate at high species richness.Methods We used a planted tree biodiversity experiment in south-east China to test our hypotheses.LAI was determined three times by digital hemispheric photography in 144 plots that had been planted with 400 trees each,forming communities with 1,2,4,8 or 16 tree species.Important Findings LAI increased significantly with tree species richness in the fifth year of stand establishment.Similar,but weaker,statistically non-significant trends were observed 1 year before.We did not observe leaf area overyielding and the presence of particularly productive and unproductive species explained large amounts of variation in leaf area,suggesting that selection-type effects contributed substantially to the biodiversity effects we found in this early phase of stand establishment.Effects sizes were moderate to large and comparable in magnitude to the ones reported for grassland ecosystems.Subtropical(and tropical)forests harbor substantial parts of global net primary production and are critical for the Earth’s carbon and hydrological cycle,and our results suggest that tree diversity critically supports these ecosystem services.展开更多
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 Chi...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)≥5 cm 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 0.04-0.50 cm year^(−1)(mean=0.21),0.03-0.46 cm season^(−1)(mean=0.18)and 0.01-0.05 cm 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.展开更多
Aims Plants with extrafloral nectaries(EFNs)are common in tropical and subtropical habitats and,despite many other arthropods also forage for EFN,most studies solely focused on the defense mutualisms between EFN plant...Aims Plants with extrafloral nectaries(EFNs)are common in tropical and subtropical habitats and,despite many other arthropods also forage for EFN,most studies solely focused on the defense mutualisms between EFN plants and ants.This study aims at a quantitative assessment of the entire arthropod community that visits EFN trees to compare visitor communities between different tree species to disentangle the mechanisms that may drive EFN visitor community composition.We also test for tree diversity effects on EFN visitors,as it is unknown if local tree species richness relates to the abundance and species richness of arthropods foraging for EFN.Methods We sampled EFN-visiting arthropods in the experimental tree species richness gradient of the BEF-China Experiment,the currently largest forest diversity experiment in the world,and tested if tree species richness affects EFN visitors and if visitor community composition differs between EFN tree species.In a second step,we analyzed the EFN of Ailanthus altissima and Triadica cochinchinensis,the two EFN tree species with highest visitor abundance,for sugars and amino acids(AA)to test if tree species-specific differences in nectar chemistry translate to differing visitor communities.Lastly,we conducted a choice experiment using different artificial nectar solutions to test if nectar quality affects foraging decisions of ants,the most frequent EFN visitors in our study sites.Important Findings EFN trees in young successional forests in subtropical South-East China are visited by a diverse assemblage of arthropods including ants,beetles,flies,and spiders.Albeit ants accounted for about 75%of all individuals,non-ant visitors were by far more species rich.Visitor abundance and species richness declined with increasing tree species richness,suggesting a resource dilution effect,because plots with more tree species had proportionally less EFN tree individuals and thus lower nectar availability.Ailanthus altissima and T.cochinchinensis were visited by different arthropods and their nectar had species-specific AA composition and sugar concentration,indicating that differences in visitors may,at least partly,be explained by differences in nectar chemistry.These findings are supported by the choice experiment,in which artificial nectars containing sugar solutions supplemented with essential AAs attracted more ants than pure sugar solutions or sugar solutions supplemented with non-essential AAs.Our results improve the understanding of the complex ecology of EFN trees,a plant life form that might be crucial for understanding how tree diversity influences patterns of tree growth in young successional tropical and subtropical forests.展开更多
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 speci...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.展开更多
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 ...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.展开更多
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 ...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.展开更多
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 investig...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.展开更多
Sand-fflled splash cups were used to study the erosive power of rainfall and throughfall in the humid subtropics of Southeast China. The splash cup measurements yielded precise and reproducible results under both open...Sand-fflled splash cups were used to study the erosive power of rainfall and throughfall in the humid subtropics of Southeast China. The splash cup measurements yielded precise and reproducible results under both open field conditions and forest vegetation. The splash cups were exposed to specific forest stands of different ages and to selected species (Schima superba, Castanopsis eyrei, Daphniphyllum oldhamii, Lithocarpus glaber) in the Gutianshan (古田山) National Nature Reserve (GNNR). The results of the measurements under forest vegetation show that the erosive power of throughfall drops to be 2.59 times higher compared to the open field. This accentuates the importance of shrub, herb and litter layers in forest ecosystems to protect the soil against erosion. Coalescing drops from leaves and branches (drips) are responsible for this notable gain in erosive power. Moreover, differences in sandloss between the investigated tree species (deciduous, evergreen) revealed that the erosion potential and the spatial heterogeneity of throughfall are species-specific. This highlights the importance of selecting specific species for afforestation projects considering the prevention of soil erosion.展开更多
文摘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.
基金funded by the German Research Foundation(DFG FOR 891/1,2 and 3)funds from the National Natural Science Foundation of China(NSFC 30710103907,30930005,31170457 and 31210103910)+2 种基金the Swiss National Science Foundation(SNSF)financed by the Sino-German Centre for Research Promotion in Beijing(GZ 524,592,698,699 and 785)the University of Tübingen,Germany(PROMOS).
文摘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.
基金supported by the EU 7th FP Project BACCARA(grant number 226299 to J.-S.H.)the EU 7th FP Project IDPBRIDGE(grant number 608422 to B.S.)+1 种基金the National Basic Research Program of China(grant number 2014CB954004 to J.S.H.)the German Science Foundation(grant FOR-891 to P.A.N.and B.S.).
文摘Aims Litterfall,as an important link between aboveground and belowground processes,plays a key role in forest ecosystems.Here,we test for effects of tree species richness on litter production and litter quality in subtropical forest.The study further encompasses a factorial gradient of secondary succession that resulted from human exploitation.Given that a large percentage of subtropical forests are in secondary successional stages,understanding the role of biodiversity on forest re-growth after disturbance appears critical.Methods From January 2009 to December 2014,we monitored forest litterfall in 27 Comparative Study Plots that spanned a gradient of tree species richness(3-20 species)and secondary successional ages(~20 to 120 years)in Gutianshan Natural Nature Reserve,Zhejiang Province,China.The experiment is part of the biodiversity-ecosystem functioning research platform‘BEF-China’.Tree litterfall was collected in monthly intervals using litter traps.Samples were separated into leaf and non-leaf components.Leaf litter was further sorted into dominant and other species.Community level monthly leaf litter C and N contents were analysed through a full year.General linear mixed-effects models were applied to test for effects of tree species richness and successional age on litter quantity and leaf litter C/N.Important Findings Litterfall increased with species richness among and within successional age and this effect was consistent across years.Successionally older stands had higher litterfall and this effect was related to increased tree species richness.However,species richness did not change the intra-and inter-annual temporal stability of litterfall.Increasing tree species richness increased leaf litter quality(decreased C/N),while successional age had no effect.Our study indicates that more diverse forest stands produce more leaf litter and that this litter has higher N concentrations,which could promote forest growth through accelerated nutrient re-cycling.
基金German Science Foundation(DFG FOR 891/2,Du 404/3-2 to W.D.)is highly acknowledged.
文摘Aims Positive plant diversity-ecosystem function relations are ultimately driven by variation in functional traits among individuals that form a community.To date,research has largely focused on the role of species diversity for ecosystem functioning.However,substantial intraspecific trait variation is common and a significant part of this variation caused by genetic differences among individuals.Here,we studied the relative importance of species diversity and seed family(SF)diversity within species for growth and herbivory in experimental subtropical tree assemblages.Methods In 2010,we set up a field experiment in subtropical China,using four species from the local species pool.Trees were raised from seeds,with seeds from the same mother tree forming an SF.We established 23 plots containing one or four species(species diversity treatment)and one or four SFs per species(SF diversity treatment).Tree growth(stem diameter,plant height and crown expansion)and herbivory(percentage leaf loss due to leaf chewers)were monitored annually from 2011 to 2013.Important findings Tree species richness promoted growth but had no effect on herbivory.In contrast,SF diversity reduced growth and increased herbivory but only so in species mixtures.Most of the observed effects were time dependent,with the largest effect found in 2013.Our results suggest that biodiversity can affect plant performance directly via tree species-species interactions,or context dependent,via potential effects on inter-trophic interactions.Two important conclusions should be drawn from our findings.Firstly,in future studies regarding biodiversity and ecosystem functioning(BEF)relationships,intraspecific genetic diversity should be given similar weight as species diversity as it has often been neglected and its effects are not well understood.Secondly,we demonstrate opposite effects of biodiversity among and within species,stressing the importance to consider the effects of multiple levels of biodiversity simultaneously.
基金German Research Foundation grant(FOR 891)the University of Zürich.
文摘Aims Most biodiversity-ecosystem functioning research has been carried out in grassland ecosystems,and little is known about whether forest ecosystems,in particular outside the temperate zone,respond similarly.Here,we tested whether productivity,assessed as leaf area index(LAI),increases with species richness in young experimental stands of subtropical trees,whether this response is similar for early-season leaf area(which is dominated by evergreens)and seasonal leaf area increase(which is dominated by deciduous species),and whether responses saturate at high species richness.Methods We used a planted tree biodiversity experiment in south-east China to test our hypotheses.LAI was determined three times by digital hemispheric photography in 144 plots that had been planted with 400 trees each,forming communities with 1,2,4,8 or 16 tree species.Important Findings LAI increased significantly with tree species richness in the fifth year of stand establishment.Similar,but weaker,statistically non-significant trends were observed 1 year before.We did not observe leaf area overyielding and the presence of particularly productive and unproductive species explained large amounts of variation in leaf area,suggesting that selection-type effects contributed substantially to the biodiversity effects we found in this early phase of stand establishment.Effects sizes were moderate to large and comparable in magnitude to the ones reported for grassland ecosystems.Subtropical(and tropical)forests harbor substantial parts of global net primary production and are critical for the Earth’s carbon and hydrological cycle,and our results suggest that tree diversity critically supports these ecosystem services.
基金National Program on Key Basic Research Project(#2014CB954004)the National Natural Science Foundation of China(#31470486,31321061)+1 种基金the‘Strategic Priority Research Program’of Chinese Academy of Sciences(#XDA05050301)the Sino Swiss Science and Technology Cooperation Program(IZL CZ 123883).
文摘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)≥5 cm 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 0.04-0.50 cm year^(−1)(mean=0.21),0.03-0.46 cm season^(−1)(mean=0.18)and 0.01-0.05 cm 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.
基金German Research Foundation(DFG FOR 891/2,KL 1849/6-1)the Sino-German Center for Research Promotion(GZ 785)the German Academic Exchange Service(PROMOS scholarship to J.M.).
文摘Aims Plants with extrafloral nectaries(EFNs)are common in tropical and subtropical habitats and,despite many other arthropods also forage for EFN,most studies solely focused on the defense mutualisms between EFN plants and ants.This study aims at a quantitative assessment of the entire arthropod community that visits EFN trees to compare visitor communities between different tree species to disentangle the mechanisms that may drive EFN visitor community composition.We also test for tree diversity effects on EFN visitors,as it is unknown if local tree species richness relates to the abundance and species richness of arthropods foraging for EFN.Methods We sampled EFN-visiting arthropods in the experimental tree species richness gradient of the BEF-China Experiment,the currently largest forest diversity experiment in the world,and tested if tree species richness affects EFN visitors and if visitor community composition differs between EFN tree species.In a second step,we analyzed the EFN of Ailanthus altissima and Triadica cochinchinensis,the two EFN tree species with highest visitor abundance,for sugars and amino acids(AA)to test if tree species-specific differences in nectar chemistry translate to differing visitor communities.Lastly,we conducted a choice experiment using different artificial nectar solutions to test if nectar quality affects foraging decisions of ants,the most frequent EFN visitors in our study sites.Important Findings EFN trees in young successional forests in subtropical South-East China are visited by a diverse assemblage of arthropods including ants,beetles,flies,and spiders.Albeit ants accounted for about 75%of all individuals,non-ant visitors were by far more species rich.Visitor abundance and species richness declined with increasing tree species richness,suggesting a resource dilution effect,because plots with more tree species had proportionally less EFN tree individuals and thus lower nectar availability.Ailanthus altissima and T.cochinchinensis were visited by different arthropods and their nectar had species-specific AA composition and sugar concentration,indicating that differences in visitors may,at least partly,be explained by differences in nectar chemistry.These findings are supported by the choice experiment,in which artificial nectars containing sugar solutions supplemented with essential AAs attracted more ants than pure sugar solutions or sugar solutions supplemented with non-essential AAs.Our results improve the understanding of the complex ecology of EFN trees,a plant life form that might be crucial for understanding how tree diversity influences patterns of tree growth in young successional tropical and subtropical forests.
基金financed by the German Research Foundation(DFG FOR 891/1,2,3)in a grant to H.B.(Br1698/10-3)the Sino-German Centre for Research Promotion in Beijing for travel grants and the participation in a summer school on scientific writing(GZ 785)support through the cooperation group“Linkages between plant diversity,microbial diversity and ecosystem functioning in subtropical forest”(GZ 986).
文摘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.
基金This work was financially supported by the National Natural Science Foundation of China(31270496 and 31300353)the Strategic Priority Research Program of the Chinese Academy of Sciences(XDA 05050204).
文摘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.
基金the coordination team of the BEF-China project for their supportthe support of M.S.G.from the German Centre for Integrative Biodiversity Research(iDiv)Halle-Jena-Leipzig funded by the German Research Foundation(FZT 118)+1 种基金the Sino-German Centre for Research Promotion for the participation of M.S.G.in a Summer School in Jingdezhen(GZ 1146)funded by the German Research Foundation(DFG FOR 891/3)with a grant to A.E.(ER 573/1-3).
文摘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.
文摘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.
基金supported by the Deutsche Forschungsgemein-schaft (German Science Foundation) (No. DFG FOR 891/1)the National Natural Science Foundation of China (Nos. 30710103907, 30930005)
文摘Sand-fflled splash cups were used to study the erosive power of rainfall and throughfall in the humid subtropics of Southeast China. The splash cup measurements yielded precise and reproducible results under both open field conditions and forest vegetation. The splash cups were exposed to specific forest stands of different ages and to selected species (Schima superba, Castanopsis eyrei, Daphniphyllum oldhamii, Lithocarpus glaber) in the Gutianshan (古田山) National Nature Reserve (GNNR). The results of the measurements under forest vegetation show that the erosive power of throughfall drops to be 2.59 times higher compared to the open field. This accentuates the importance of shrub, herb and litter layers in forest ecosystems to protect the soil against erosion. Coalescing drops from leaves and branches (drips) are responsible for this notable gain in erosive power. Moreover, differences in sandloss between the investigated tree species (deciduous, evergreen) revealed that the erosion potential and the spatial heterogeneity of throughfall are species-specific. This highlights the importance of selecting specific species for afforestation projects considering the prevention of soil erosion.