青藏高原高寒草地生物多样性与生态系统功能的关系

生物多样性和生态系统功能(BEF)之间的关系是目前陆地生态系统生态学研究的热点, 对于生态系统的高效利用与管理意义重大, 而且对于退化生态系统功能的恢复及生物多样性的保护有重要的指导作用。高寒草地是青藏高原生态系统的主体, 近年来, 在气候变化与人为干扰等因素的驱动下, 高寒草地生态系统功能严重衰退。为此, 本文在综述物种多样性和生态系统功能及其相互关系研究进展的基础上, 首先从地下生态学过程研究、全球变化对生态系统多功能性的影响等方面解析了目前关于草地生物多样性和生态系统功能研究中存在的问题。继而, 从不同草地类型、草地退化程度、放牧、模拟气候变化、刈割、施肥、封育和补播等干扰利用方式对高寒草地物种多样性与生态系统功能的影响进行了全面的评述。并指出了高寒草地BEF研究中存在的不足, 今后应基于物种功能多样性开展高寒草地BEF研究, 全面且综合地考虑非生物因子(养分资源、外界干扰、环境波动等)对生物多样性与生态系统功能之间关系的影响, 关注尺度效应和要素耦合在全球气候变化对高寒草地BEF研究中的作用。最后, 以高寒草地BEF研究进展和结论为支撑依据, 综合提出了高寒草地资源利用和生物多样性保护的措施与建议: 加强放牧管理, 保护生物多样性; 治理退化草地, 维持生物多样性功能; 加强创新保护理念, 增强生态系统功能。     

生物多样性与生态系统功能:最新的进展与动向

生物多样性与生态系统功能的关系及其内在机制是当前生态学领域的重大科学问题。 2 0 0 2年以来人们不再过多地纠缠于“抽样 -互补之争” ,对这一世纪课题的认识又有了新的进展。 (1)人们开始运用已有的知识揭示更大时间和空间尺度上的物种多样性 -生态系统功能关系。多样性作用机制可能存在着动态变化———“抽样向互补转型” :群落建立初期 ,抽样效应是主要的多样性作用机制 ;随时间推移 ,生态位互补成为主要机制。理论研究则预测 :局域尺度上生态系统功能与物种多样性呈现单峰曲线关系 ,在区域尺度上为单调上升关系 ;(2 )非生物因素与多样性 -生产力的交互关系吸引了许多实验研究。人们发现 :物种多样性 -生产力关系可能会受到资源供给率和环境扰动的修正 ,环境因素可能是多样性 -生产力关系的幕后操纵者 ;(3)人们开始重视营养级相互作用对于多样性 -生态系统功能关系的影响 ,生态位互补和抽样假说开始被扩展运用到消费者营养级上 ;(4 )人们开始认真思考物种共存机制在多样性 -生态系统功能关系的形成中所扮演的角色。理论模型研究表明 ,不同的物种共存机制会导致不同的多样性 -生产力关系     

Biodiversity acts as insurance of productivity of bacterial communities under abiotic perturbations

Anthropogenic disturbances are detrimental to the functioning and stability of natural ecosystems. Critical ecosystem processes driven by microbial communities are subjected to these disturbances. Here, we examine the stabilizing role of bacterial diversity on community biomass in the presence of abiotic perturbations such as addition of heavy metals, NaCl and warming. Bacterial communities with a diversity gradient of 1–12 species were subjected to the different treatments, and community biomass (OD₆₀₀) was measured after 24 h. We found that initial species richness and phylogenetic structure impact the biomass of communities. Under abiotic perturbations, the presence of tolerant species in community largely contributed in community biomass production. Bacterial diversity stabilized the biomass across the treatments, and differential response of bacterial species to different perturbations was the key reason behind these effects. The results suggest that biodiversity is crucial for maintaining the stability of ecosystem functioning and acts as ecological insurance under abiotic perturbations. Biodiversity in natural ecosystems may also uphold the ecosystem functioning under anthropogenic disturbance.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Biodiversity as a solution to mitigate climate change impacts on the functioning of forest ecosystems

Forest ecosystems are critical to mitigating greenhouse gas emissions through carbon sequestration. However, climate change has affected forest ecosystem functioning in both negative and positive ways, and has led to shifts in species/functional diversity and losses in plant species diversity which may impair the positive effects of diversity on ecosystem functioning. Biodiversity may mitigate climate change impacts on (I) biodiversity itself, as more‐diverse systems could be more resilient to climate change impacts, and (II) ecosystem functioning through the positive relationship between diversity and ecosystem functioning. By surveying the literature, we examined how climate change has affected forest ecosystem functioning and plant diversity. Based on the biodiversity effects on ecosystem functioning (B→EF), we specifically address the potential for biodiversity to mitigate climate change impacts on forest ecosystem functioning. For this purpose, we formulate a concept whereby biodiversity may reduce the negative impacts or enhance the positive impacts of climate change on ecosystem functioning. Further B→EF studies on climate change in natural forests are encouraged to elucidate how biodiversity might influence ecosystem functioning. This may be achieved through the detailed scrutiny of large spatial/long temporal scale data sets, such as long‐term forest inventories. Forest management strategies based on B→EF have strong potential for augmenting the effectiveness of the roles of forests in the mitigation of climate change impacts on ecosystem functioning.     

Nitrogen enrichment weakens ecosystem stability through decreased species asynchrony and population stability in a temperate grassland

Biodiversity generally promotes ecosystem stability. To assess whether the diversity–stability relationship observed under ambient nitrogen (N) conditions still holds under N enriched conditions, we designed a 6‐year field experiment to test whether the magnitude and frequency of N enrichment affects ecosystem stability and its relationship with species diversity in a temperate grassland. Results of this experiment showed that the frequency of N addition had no effect on either the temporal stability of ecosystem and population or the relationship between diversity and stability. Nitrogen addition decreased ecosystem stability significantly through decreases in species asynchrony and population stability. Species richness was positively associated with ecosystem stability, but no significant relationship between diversity and the residuals of ecosystem stability was detected after controlling for the effects of the magnitude of N addition, suggesting collinearity between the effects of N addition and species richness on ecosystem stability, with the former prevailing over the latter. Both population stability and the residuals of population stability after controlling for the effects of the magnitude of N addition were positively associated with ecosystem stability, indicating that the stabilizing effects of component populations were still present after N enrichment. Our study supports the theory predicting that the effects of environmental factors on ecosystem functioning are stronger than those of biodiversity. Understanding such mechanisms is important and urgent to protect biodiversity in mediating ecosystem functioning and services in the face of global changes.     

Species richness promotes ecosystem carbon storage: evidence from biodiversity-ecosystem functioning experiments

Plant diversity has a strong impact on a plethora of ecosystem functions and services, especially ecosystem carbon (C) storage. However, the potential context-dependency of biodiversity effects across ecosystem types, environmental conditions and carbon pools remains largely unknown. In this study, we performed a meta-analysis by collecting data from 95 biodiversity-ecosystem functioning (BEF) studies across 60 sites to explore the effects of plant diversity on different C pools, including aboveground and belowground plant biomass, soil microbial biomass C and soil C content across different ecosystem types. The results showed that ecosystem C storage was significantly enhanced by plant diversity, with stronger effects on aboveground biomass than on soil C content. Moreover, the response magnitudes of ecosystem C storage increased with the level of species richness and experimental duration across all ecosystems. The effects of plant diversity were more pronounced in grasslands than in forests. Furthermore, the effects of plant diversity on belowground plant biomass increased with aridity index in grasslands and forests, suggesting that climate change might modulate biodiversity effects, which are stronger under wetter conditions but weaker under more arid conditions. Taken together, these results provide novel insights into the important role of plant diversity in ecosystem C storage across critical C pools, ecosystem types and environmental contexts.     

Effects of plant diversity,community composition and environmental parameters on productivity in montane European grasslands

In the past years, a number of studies have used experimental plant communities to test if biodiversity influences ecosystem functioning such as productivity. It has been argued, however, that the results achieved in experimental studies may have little predictive value for species loss in natural ecosystems. Studies in natural ecosystems have been equivocal, mainly because in natural ecosystems differences in diversity are often confounded with differences in land use history or abiotic parameters. In this study, we investigated the effect of plant diversity on ecosystem functioning in semi-natural grasslands. In an area of 10×20 km, we selected 78 sites and tested the effects of various measures of diversity and plant community composition on productivity. We separated the effects of plant diversity on ecosystem functioning from potentially confounding effects of community composition, management or environmental parameters, using multivariate statistical analyses. In the investigated grasslands, simple measures of biodiversity were insignificant predictors of productivity. However, plant community composition explained productivity very well (R²=0.31) and was a better predictor than environmental variables (soil and site characteristics) or management regime. Thus, complex measures such as community composition and structure are important drivers for ecosystem functions in semi-natural grasslands. Furthermore, our data show that it is difficult to extrapolate results from experimental studies to semi-natural ecosystems, although there is a need to investigate natural ecosystems to fully understand the relationship of biodiversity and ecosystem functioning.     

Does disturbance regime change community assembly of angiosperm plant communities in the boreal forest?

Aims To examine if and how species and phylogenetic diversity change in relation to disturbance, we conducted a review of ecological literature by testing the consistency of the relationship between phylogenetic diversity and disturbance and compared taxonomic groups, type of disturbance and ecosystem/habitat context. We provide a case study of the phylogenetic diversity–disturbance relationship in angiosperm plant communities of a boreal forest region, compared with types of natural and anthropogenic disturbances and plant growth forms.Methods Using a large-scale sampling plot network along a complete (0–100%) anthropogenic disturbance gradient in the boreal biome, we compared the changes of angiosperm plant community structure and composition across plots. We estimated natural disturbance with historical records of major fires. We then calculated phylogenetic diversity indexes and determined species richness in order to compare linear and polynomial trends along disturbance gradients. We also compared the changes of community structure for different types of anthropogenic disturbances and examined how the relationships between species and phylogenetic diversity and disturbance regimes vary among three different life forms (i.e. forbs, graminoids and woody plants).Important findings Phylogenetic diversity was inconsistently related to disturbance in previous studies, regardless of taxon, disturbance type or ecosystem context. In the understudied boreal ecosystem, angiosperm plant communities varied greatly in species richness and phylogenetic diversity along anthropogenic disturbance gradients and among different disturbance types. In general, a quadratic curve described the relationship between species richness and anthropogenic disturbance, with the highest richness at intermediate anthropogenic disturbance levels. However, phylogenetic diversity was not related to disturbance in any consistent manner and species richness was not correlated with phylogenetic diversity. Phylogenetic relatedness was also inconsistent across plant growth forms and different anthropogenic disturbance types. Unlike the inconsistent patterns observed for anthropogenic disturbance, community assembly among localities varying in time since natural disturbance exhibited a distinct signature of phylogenetic relatedness, although those trends varied among plant growth forms.     

Ecosystem multifunctionality lowers as grasslands under restoration approach their target habitat type

Biodiversity is declining at a rapid pace and, with it, the ecosystem functions that support ecosystem services. To counter this, ecosystem restoration is necessary. While the relationship between biodiversity and ecosystem functioning has been studied in depth, the relationship between ecosystem restoration and ecosystem functioning is studied less. We performed an observational study in grasslands undergoing restoration management toward Nardus grassland. Eight ecosystem functions, representing flows of energy, matter or information between functional compartments, were measured across five successive restoration phases along the restoration gradient. The levels of functioning were then compared along the gradient for both the individual functions and a multifunctionality index. We hypothesized that plant richness increases when grasslands are more restored and this increase in biodiversity is paralleled by an increase in ecosystem functioning. In our study, the degraded grasslands, generally occurring on more nutrient-rich soils, were dominated by competitive fast-growing species, resulting in higher process rates and thus in higher, faster functioning. Likewise, more restored grasslands exhibited slower process rates and, thus, lower functioning. When studying ecosystem functioning, value judgments are easily made. Especially in a restoration context, high functioning does not necessarily equals well functioning, as this depends on the stakeholder perspective. We need to ask ourselves if a high functioning ecosystem is most desirable, especially in a restoration or conservation context. Policy frameworks will need to balance these goals.     

Biodiversity and ecosystem functioning at local and regional spatial scales

Local niche complementarity among species (the partitioning of species based upon niche differentiation) is predicted to affect local ecosystem functioning positively. However, recent theory predicts that greater local diversity may hinder local ecosystem functioning when diversity is enhanced through source–sink dynamics. We suggest community assembly as a way to incorporate both the local and regional processes that determine biodiversity and its consequent effects on ecosystem functioning. From this, we propose a hump-shaped relationship between diversity and ecosystem functioning at local scales, but a linear increase of functioning with diversity at regional scales due to regional complementarity.     

Biogeographic historical legacies in the net primary productivity of Northern Hemisphere forests

It has been suggested that biogeographic historical legacies in plant diversity may influence ecosystem functioning. This is expected because of known diversity effects on ecosystem functions, and impacts of historical events such as past climatic changes on plant diversity. However, empirical evidence for a link between biogeographic history and present‐day ecosystem functioning is still limited. Here, we explored the relationships between Late‐Quaternary climate instability, species‐pool size, local species and functional diversity, and the net primary productivity (NPP) of Northern Hemisphere forests using structural equation modelling. Our study confirms that past climate instability has negative effects on plant functional diversity and through that on NPP, after controlling for present‐day climate, soil conditions, stand biomass and age. We conclude that global models of terrestrial plant productivity need to consider the biogeographical context to improve predictions of plant productivity and feedbacks with the climate system.     

Regional zooplankton dispersal provides spatial insurance for ecosystem function

Changing environmental conditions are affecting diversity and ecosystem function globally. Theory suggests that dispersal from a regional species pool may buffer against changes in local community diversity and ecosystem function after a disturbance through the establishment of functionally redundant tolerant species. The spatial insurance provided by dispersal may decrease through time after environmental change as the local community monopolizes resources and reduces community invasibility. To test for evidence of the spatial insurance hypothesis and to determine the role dispersal timing plays in this response we conducted a field experiment using crustacean zooplankton communities in a subarctic region that is expected to be highly impacted by climate change – Churchill, Canada. Three experiments were conducted where nutrients, salt, and dispersal were manipulated. The three experiments differed in time‐since‐disturbance that the dispersers were added. We found that coarse measures of diversity (i.e. species richness, evenness, and Shannon–Weiner diversity) were generally resistant to large magnitude disturbances, and that dispersal had the most impact on diversity when dispersers were added shortly after disturbance. Ecosystem functioning (chl‐a) was degraded in disturbed communities, but dispersal recovered ecosystem function to undisturbed levels. This spatial insurance for ecosystem function was mediated through changes in community composition and the relative abundance of functional groups. Results suggest that regional diversity and habitat connectivity will be important in the future to maintain ecosystem function by introducing functionally redundant species to promote compensatory dynamics.     

Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands

Aims and Methods Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.Important findings Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites. Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.     

Environmental conditions influence the plant functional diversity effect on potential denitrification

Global biodiversity loss has prompted research on the relationship between species diversity and ecosystem functioning. Few studies have examined how plant diversity impacts belowground processes; even fewer have examined how varying resource levels can influence the effect of plant diversity on microbial activity. In a field experiment in a restored wetland, we examined the role of plant trait diversity (or functional diversity, (FD)) and its interactions with natural levels of variability of soil properties, on a microbial process, denitrification potential (DNP). We demonstrated that FD significantly affected microbial DNP through its interactions with soil conditions; increasing FD led to increased DNP but mainly at higher levels of soil resources. Our results suggest that the effect of species diversity on ecosystem functioning may depend on environmental factors such as resource availability. Future biodiversity experiments should examine how natural levels of environmental variability impact the importance of biodiversity to ecosystem functioning.     

Natural enemy diversity and pest control: patterns of pest emergence with agricultural intensification

Concern over declining biodiversity and the implications for continued provision of ecosystem services has led, recently, to intense research effort to describe relationships between biodiversity and ecosystem functioning. Here we extend this effort to the relationship between natural enemy species diversity and natural pest control. From simple modelled food‐webs and simulations of natural enemy species loss we derive specific predictions concerning the effect of herbivore life‐history traits, such as life‐cycle type and concealment, on the shape (reflecting diversity effects) and variance (reflecting species composition effects) of the relationship between natural enemy diversity and pest‐control. We show that these predictions are consistent with the emergence of different pest types following intensification of rice production in Asia. We suggest that basic biological insights can help define the structure of ecological processes and allow more accurate predictions of the effect of species loss on the delivery of ecosystem services.     

Testing the effects of plant species loss on multiple ecosystem functions based on extinction scenarios

Ecosystem functions are threatened by continuing global loss of biodiversity. We simultaneously investigated three ecosystem functions and forage nutrient values following potential species extinction scenarios (dominant species removal, rare species removal, end-member species removal and random species removal) in a Mongolian grassland. ANPP, forage nutrient values, litter decomposition, and soil respiration were measured one and/or two years after plant removal. DNA samples of microorganisms extracted from the soil were subjected to metagenomics analysis. Finally, we calculated the multifunctionality, and examined the relationship of multifunctionality with plant and microorganism diversity. Among ecosystem functions, ANPP and litter decomposition rate decreased under random and rare species extinction scenarios, respectively, and forage quality increased when only dominant species had been removed. Diversity and species composition of soil microorganism were not affected by plant species richness or removal scenario. Only genus-level diversity of bacteria and ANPP were significantly and positively correlated with microbial diversity. Taken together, decreasing species richness of plants and soil organisms rarely impaired multifunctionality. Ecosystem functions were relatively robust to realistic disturbances and species extinction in natural grasslands. However, as each function responded differently to the different sets of species removed, the consequences of a realistic non-random extinction scenario for multiple ecosystem functions should be critical to the management of biodiversity loss caused by different disturbances.     

Effects of land-use change on productivity depend on small-scale plant species diversity

Understanding the interplay between land-use change, species diversity and ecosystem function is critical for the prediction of global change impacts on ecosystem services. Biodiversity experiments with artificial species assemblages have shown that community-scale species richness may affect ecosystem productivity and spatial stability. However, the use of synthetic communities with controlled levels of species density for biodiversity experiments has been criticised and their relevance for natural communities has been questioned. Here, we use a land-use change experiment to investigate the biodiversity effects on production within managed, upland grasslands. We examine species diversity and productivity at both the small plant-neighbourhood scale (14×14 cm) and the field scale (15 m×25 m) for two land-use trajectories under field conditions: intensification through fertilisation, and extensification through the cessation of mowing. Both intensification and extensification were associated with a decrease in species number, but the magnitude of this decrease was greater at the small scale. Extensification was associated with a decrease in small-scale productivity whereas intensification had no significant effect on small-scale productivity. Effects of land-use treatments on biomass production were mediated by variation in small-scale species number; species number showed a significant positive relationship with small-scale productivity within each land-use treatment. Furthermore, species number was associated with a decrease in the variance of small-scale green biomass. In contrast, no species diversity effects were found on productivity at the field scale. Instead, field-scale species diversity decreased with increase in the total above-ground biomass (green biomass+litter). This study demonstrates that biodiversity effects can be observed under field conditions at the small scale and may play an important role for ecosystem functioning and stability even in low-diversity plant communities.     

关于蝗虫多样性与草原生态系统可持续发展的若干科学问题

蝗虫多样性既是草原生态系统演化的产物 ,反过来也影响着草原生态系统的结构与功能。蝗虫多样性状况与草原生态系统的持续发展有着密切的关联 ,这其中的科学问题包括 :(1 )蝗虫多样性及其空间变异机理 ;(2 )蝗虫多样性对草原生态系统过程的影响 ;(3 )蝗虫多样性空间格局与草原生态系统演化中的自然和人文因素的关联 ;(4 )蝗虫种群暴发的多样性阈值与草原生态系统的调控 ;(5 )蝗虫多样性和稳定性与草原生态系统的健康等。深入发掘生物多样性所包含的科学内涵 ,将一个物种的多样性与生态系统的结构和功能耦合起来研究 ,从重要物种蝗虫的生物多样性入手 ,深入探讨草原生态系统的持续发展问题 ,将为减少或减轻我国西部草原蝗灾发生提供科学的支持     

Soil fertility shifts the relative importance of saprotrophic and mycorrhizal fungi for maintaining ecosystem stability

Soil microbial communities are essential for regulating the dynamics of plant productivity. However, how soil microbes mediate temporal stability of plant productivity at large scales across various soil fertility conditions remains unclear. Here, we combined a regional survey of 51 sites in the temperate grasslands of northern China with a global grassland survey of 120 sites to assess the potential roles of soil microbial diversity in regulating ecosystem stability. The temporal stability of plant productivity was quantified as the ratio of the mean normalized difference vegetation index to its standard deviation. Soil fungal diversity, but not bacterial diversity, was positively associated with ecosystem stability, and particular fungal functional groups determined ecosystem stability under contrasting conditions of soil fertility. The richness of soil fungal saprobes was positively correlated with ecosystem stability under high-fertility conditions, while a positive relationship was observed with the richness of mycorrhizal fungi under low-fertility conditions. These relationships were maintained after accounting for plant diversity and environmental factors. Our findings highlight the essential role of fungal diversity in maintaining stable grassland productivity, and suggest that future studies incorporating fungal functional groups into biodiversity–stability relationships will advance our understanding of their linkages under different fertility conditions.