物种多样性与生态系统功能的关系研究进展

物种的空前丧失促使人们越来越多地开始研究物种多样性与生态系统功能的关系,并探讨其潜在的作用机制.本文根据最新研究进展,归纳了微宇宙实验、"生态箱"实验、Cedar Creek草地多样性实验和欧洲草地实验等代表性实验中关于物种多样性与生产力、稳定性、抗入侵性等生态系统功能的焦点问题,介绍了去除实验在多样性与生态系统功能研究中的应用.在此基础上,提出未来研究所面临的挑战,并对研究趋势进行了展望.主要挑战和趋势有:将小尺度上开展的实验研究扩展到较大的时空尺度上;综合考虑生物因素和非生物因素对多样性变化、生态系统功能的交互作用;营养级之间的相互作用、物种共存机制对物种多样性与生态系统功能关系的影响.     

亚热带森林生物多样性与生态系统功能实验研究基地(BEF-China)研究进展

生物多样性与生态系统功能的关系(BEF)及其内在机制是当前生物多样性研究领域的热点问题。长期以来,以草地生态系统为主的BEF研究积累了大量研究成果,而基于森林生态系统的相关研究则相对较少。亚热带森林生物多样性与生态系统功能实验研究基地(BEF-China)是目前包含树种最多、涉及多样性水平最高的大型森林控制实验样地。该文总结了基于BEF-China平台的研究进展,特别是生物多样性对生态系统生产力、养分循环以及多营养级相互作用关系等方面的影响,并提出了未来BEF-China的研究应注重高通量测序和遥感等新兴技术的应用,在生物多样性的多维度、生态系统的多种组分与多种功能以及BEF研究的多种尺度等交叉方向上持续开展深入研究。针对BEF-China研究成果的梳理有助于理解驱动亚热带森林生物多样性与生态系统功能关系的内在机理,为生物多样性保护和生态修复提供科学依据。     

植物功能多样性与功能群研究进展

综述了植物功能多样性与功能群研究的最新进展。介绍了植物功能群的定义及植物功能群的划分方法。在功能多样性与生态系统资源动态关系方面．抽样效应和生态位互补效应用来解释植物多样性在生态系统资源动态中的作用。功能多样性与生态系统的稳定性间的关系可以用生态冗余或生态保险概念来解释,这两个概念是一个问题的两个侧面,是多样性与生态系统功能争论的焦点。     

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。     

生态系统多功能性的指标选择与驱动因子: 研究现状与展望

全球变化和人类活动正以空前的速度在世界范围内改变着生物多样性, 这导致了全球生物多样性的锐减以及生产力的下降、病虫害的增加和抗入侵能力的减弱等生态问题。近30年来, 生态学家开始对于生物多样性的持续丧失是否以及如何影响生态系统功能的问题越来越感兴趣, 生物多样性与生态系统功能(biodiversity and ecosystem functioning, BEF)关系的研究应运而生, 并成为生态学研究的热点之一。但长期以来, 研究者更多地关注单一生态系统功能, 而忽略了生态系统能够同时提供多种生态系统功能的能力, 即生态系统多功能性(ecosystem multifunctionality, EMF)。本文综述了EMF研究中功能指标的选择、生物多样性的不同维度、微生物多样性对EMF的影响以及其他非生物因子对EMF的驱动等进展。因只考虑单一功能可能会低估生物多样性对整体生态系统功能的影响, 故生物多样性与生态系统多功能性(BEMF)关系的研究成为BEF关系研究的重点。近年来, BEMF关系的研究发展较快, 在不同生态系统(包括水生、草地、森林、旱地、农业等)、不同研究尺度(从区域到全球尺度)、BEMF关系的驱动机制(从单一驱动机制到多种驱动机制共同作用)、研究方法(包括新概念以及新的量化方法的提出和应用)等方面均取得了新的进展。但仍有不足之处, 如对于EMF研究中功能指标的选取没有统一的标准、对地下微生物多样性的关注度不够、涉及多营养级水平下的BEMF关系研究较少、驱动EMF的机制仍存在争论等。未来应加强对于功能指标选取的标准研究, 综合分析地上、地下生物多样性以及非生物因子对EMF的整体影响, 加强生态系统多服务性(ecosystem multiserviceability, EMS)方法的研究和应用。     

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     

Variation in the methods leads to variation in the interpretation of biodiversity–ecosystem multifunctionality relationships

生物多样性常常和生态系统多功能性(生态系统同时提供多个生态系统功能的能力)正相关。然而,生物多样性与生态系统多功能性的关系是否依赖于生态系统功能的数目有诸多争议。其中,生物多样性对生态系统多功能性的影响或许不随生态系统功能数目的变化而变化,或者随生态系统功能数目的增多而增强。我们期望通过研究不同生态系统多功能性指数的统计原理来解决这些争议。 我们使用了模型模拟和一系列来自不同空间尺度(从局域到全球)和不同生物群系(温带和高寒草地、森林和干旱地)的经验数据。我们回顾了量化生态系统多功能性的三种方法,包括平均值法、加和法和阈值法。我们发现随着生态系统功能数目的增加,生物多样性与生态系统多功能性的关系要么不变,要么增强。这些结果可由平均和加和的多功能性指数的统计原理来解释。具体来讲,当利用生态系统功能的平均值计算多功能性指数时,由于多样性对多功能性的效应等于多样性对单个生态系统功能效应的平均值,所以不会随生态系统功能数目的变化而变化。同样的道理,当利用单个生态系统的加和值计算多功能性指数时,多样性的效应会随着生态系统功能数目的增加而增强。我们提出了一个改进的多功能性指数,将平均或加和多功能性指数转化为标准化的多功能性指数, 以便于对不同研究的结果进行比较。此外,我们提出了基于变量数值范围的标准化方法来解决阈值法的数学假象问题(多样性效应随生态系统功能数目的增加而增强)。我们的研究结果表明,量化多功能性指数的方法不同,结果也不同。因此,有必要加深对不同方法数理基础的理解。而标准化的多功能性指数为比较不同研究中的生物多样性与生态系统多功能性的关系提供了有效的方法。     

青藏高原高寒草地地下生物多样性:进展、问题与展望

栖息于土壤中的微生物和微型动物种类繁多、数量巨大,在对地上生物多样性的调控和在生态系统功能与服务的维系中,具有举足轻重的作用。虽然对土壤微生物以及土壤动物已经开展了广泛的调查,但是整体上对于地下生物多样性的分布格局、驱动机制及其对全球变化的响应与适应过程,仍缺乏深刻的认识。青藏高原是全球变化的敏感区域,其中高寒草地是高原最主要的植被类型,占高原面积的60%左右,在高寒生态系统生物多样性维持中具有重要意义。近年来,已有大量研究关注于高寒草地地下生物多样性,但是缺乏系统的总结与论述。基于此,本文从细菌、真菌、古菌、线虫、节肢动物五大土壤生物类群出发,阐述了青藏高原高寒草地的地下物种丰富度、分布格局及其影响因素,重点探讨了它们对气候变化和人类活动的响应,并就未来高寒草地地下生物多样性亟需关注的关键问题进行了展望,包括:(1)地下各个生物类群的分布格局、各类群之间的联系及驱动机制;(2)地上与地下生物多样性耦联的机制;(3)地下生物多样性对生态系统功能和健康的影响;(4)地下生物多样性的调控实验研究。     

生产力、可靠度与物种多样性:微宇宙实验研究

近年来,生物多样性与生态系统功能的关系成为生态领域内一个重大科学问题。有一些实验研究表明,物种多样性的降低会使生态系统的生产力、稳定性等功能受损,然而对这些实验结果的解释却产生了激烈的争论,因为有两种机制-“生态位互补”和“抽样效应”都可能会产生这种结果。本项研究通过微宇宙实验探讨了物种多样性与生态系统生产力及其可靠度的关系。在10种单细胞藻类中随机抽取物种,构建具有不同物种丰富度的水生群落,并使同一物种丰富度水平的群落之间没有物种交叉,然后检测物种丰富度对群落生产力及其可靠度的作用,群落生产力以藻类干重表示,自实验开始后第4周起,每周测定1次,共测5次。结果显示：物种丰富度对群落生产力有正效应,并且这种正效应随时间推移而增强;许多混合群落的生产力超过了该群落内所有物种的单产,即发生了超产现象,在实验初期某些特定物种对一些混合群落生产力有主要贡献,而在实验后期却没有任何多物种群落的生产力受个别物种存在与否的影响,群落生产力的可靠度与物种丰富度之间不存在显著相关。从以上结果可以得知：物种多样性对群落生产力有着逐渐增强的正效应;物种多样性对生产力的正效应是生态位互补和抽样效应共同作用的结果,但随时间推移,抽取效应逐渐减弱,本顶研究支持了关于生态位互补与抽样效应在多样性正效应中共同起作用的认识,并说明了这两种机制的相对重要性随时间推移而发生改变。     

植物遗传多样性与生态系统功能关系的研究进展

全球变化和人类活动导致物种生境的萎缩,造成很多植物种群数量缩减,遗传多样性快速丧失。对于物种多样性低的生态系统,优势种的遗传多样性可能比物种多样性对生态系统功能产生更大的影响。因此,了解遗传多样性和生态系统功能的关系(GD-EF)及其机制对生物多样性保护、应对环境变化和生态修复具有指导意义。该文综述了植物遗传多样性对生态系统结构(高营养级生物群落结构)和生态系统功能(初级生产力、养分循环和稳定性)的影响及机制、功能多样性对GD-EF的影响、遗传多样性效应和物种多样性效应的比较,以及GD-EF在生态修复等实际应用的研究进展。最后指出当前研究的不足之处,以期为后续研究提供参考:1)还需深入研究GD-EF机制;2)未评估遗传多样性对生态系统多功能性的影响;3)不同遗传多样性测度对生态系统功能的影响不明确;4)缺少长期的和多空间尺度结合的GD-EF实验;5)遗传多样性效应相对于其他因子的作用不清楚。     

Biodiversity effects on ecosystem functioning: emerging issues and their experimental test in aquatic environments

Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity‐ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity‐ecosystem functioning relationships. These include (1) investigating the effects of non‐random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non‐random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness‐centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.     

生物多样性-生产力关系研究进展——基于文献计量分析

生态系统的结构和功能是生态学研究的核心内容。早期基于野外调查的生态学研究强调生产力表征的环境梯度对生态系统结构的影响,而基于控制试验的生态学研究则强调生态系统结构变化对生态系统功能的影响。围绕这两类研究所支持理论间的争论是当前生态学的前沿、热点和难点,其中最具代表性的科学问题是生物多样性与以生产力为代表的生态系统功能间是否存在一般性关系。为深入了解生物多样性-生产力关系研究脉络,分析其对生态学研究范式与理论发展的影响以及对未来研究方向的启示,以Web of Science核心合集数据库中的相关文献为数据源,结合文献计量分析和文献综述,系统总结了多样性-生产力关系研究进展。结果表明：(1)生物多样性-生产力关系研究推动了生态学研究范式由以样带调查为主的观察性研究向以控制试验为主的实验性研究的转变,促进了全球联网控制试验研究的发展。(2)研究聚焦的生态系统类型由最初的北美普列利草原逐渐向其它草地、灌丛、森林等多样的生态系统过渡,研究结论及其生态学理论的普适性逐渐增强。(3)该研究推动了对生物多样性不同维度(如功能多样性和系统发育多样性)在生态系统中作用的认识,促进了学界对除生产功能外的生态...     

Genotypic richness and dissimilarity opposingly affect ecosystem functioning

Biodiversity is an essential determinant of ecosystem functioning. Numerous studies described positive effects of diversity on the functioning of communities arising from complementary resource use and facilitation. However, high biodiversity may also increase competitive interactions, fostering antagonism and negatively affecting community performance. Using experimental bacterial communities we differentiated diversity effects based on genotypic richness and dissimilarity. We show that these diversity characteristics have opposite effects on ecosystem functioning. Genotypic dissimilarity governed complementary resource use, improving ecosystem functioning in complex resource environments. Contrastingly, genotypic richness drove allelopathic interactions, mostly reducing ecosystem functioning. The net biodiversity effect on community performance resulted from the interplay between the genetic structure of the community and resource complexity. These results demonstrate that increasing richness, without concomitantly increasing dissimilarity, can decrease ecosystem functioning in simple environments due to antagonistic interactions, an effect insufficiently considered so far in mechanistic models of the biodiversity-ecosystem functioning relationship.     

Biodiversity and ecosystem functioning: recent theoretical advances

The relationship between biodiversity and ecosystem functioning has emerged as a major scientific issue today. As experiments progress, there is a growing need for adequate theories and models to provide robust interpretations and generalisations of experimental results, and to formulate new hypotheses. This paper provides an overview of recent theoretical advances that have been made on the two major questions in this area: (1) How does biodiversity affect the magnitude of ecosystem processes (short‐term effects of biodiversity)? (2) How does biodiversity contribute to the stability and maintenance of ecosystem processes in the face of perturbations (long‐term effects of biodiversity)?
Positive short‐term effects of species diversity on ecosystem processes, such as primary productivity and nutrient retention, have been explained by two major types of mechanisms: (1) functional niche complementarity (the complementarity effect), and (2) selection of extreme trait values (the selection effect). In both cases, biodiversity provides a range of phenotypic trait variation. In the complementarity effect, trait variation then forms the basis for a permanent association of species that enhances collective performance. In the selection effect, trait variation comes into play only as an initial condition, and a selective process then promotes dominance by species with extreme trait values. Major differences between within‐site effects of biodiversity and across‐site productivity–diversity patterns have also been clarified. The local effects of diversity on ecosystem processes are expected to be masked by the effects of varying environmental parameters in across‐site comparisons.
A major reappraisal of the paradigm that has dominated during the last decades seems necessary if we are to account for long‐term effects of biodiversity on ecosystem functioning. The classical deterministic, equilibrium approaches to stability do not explain the reduced temporal variability of aggregate ecosystem properties that has been observed in more diverse systems. On the other hand, stochastic, nonequilibrium approaches do show two types of biodiversity effects on ecosystem productivity in a fluctuating environment: (1) a buffering effect, i.e., a reduction in the temporal variance; and (2) a performance‐enhancing effect, i.e., an increase in the temporal mean. The basic mechanisms involved in these long‐term insurance effects are very similar to those that operate in short‐term biodiversity effects: temporal niche complementarity, and selection of extreme trait values. The ability of species diversity to provide an insurance against environmental fluctuations and a reservoir of variation allowing adaptation to changing conditions may be critical in a long‐term perspective.
These recent theoretical developments in the area of biodiversity and ecosystem functioning suggest that linking community and ecosystem ecology is a fruitful avenue, which paves the way for a new ecological synthesis.     

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.     

Quantifying the evidence for biodiversity effects on ecosystem functioning and services

Concern is growing about the consequences of biodiversity loss for ecosystem functioning, for the provision of ecosystem services, and for human well being. Experimental evidence for a relationship between biodiversity and ecosystem process rates is compelling, but the issue remains contentious. Here, we present the first rigorous quantitative assessment of this relationship through meta-analysis of experimental work spanning 50 years to June 2004. We analysed 446 measures of biodiversity effects (252 in grasslands), 319 of which involved primary producer manipulations or measurements. Our analyses show that: biodiversity effects are weaker if biodiversity manipulations are less well controlled; effects of biodiversity change on processes are weaker at the ecosystem compared with the community level and are negative at the population level; productivity-related effects decline with increasing number of trophic links between those elements manipulated and those measured; biodiversity effects on stability measures ('insurance' effects) are not stronger than biodiversity effects on performance measures. For those ecosystem services which could be assessed here, there is clear evidence that biodiversity has positive effects on most. Whilst such patterns should be further confirmed, a precautionary approach to biodiversity management would seem prudent in the meantime.     

Conservation implications of the link between biodiversity and ecosystem functioning

The relationship between biodiversity and individual ecosystem processes is often asymptotic, saturating at relatively low levels, with some species contributing more strongly than others. This has cast doubt on arguments for conservation based on maintenance of the functioning of ecosystems. However, we argue that the link between biodiversity and ecosystem functioning is an important additional argument for conservation for several reasons. (1) Although species differ in importance to ecosystem processes, we do not believe that this argues for preservation of just a few species for two reasons: first, it is nearly impossible to identify all species important to the numerous systems and processes on which humans depend; second, the important species themselves may depend on an unknown number of other species in their communities. (2) Arguments for conservation based on ecosystem functioning are complementary to other utilitarian, ethical and aesthetic justifications. No single reason will convince all people or protect all species, however the combination produces a strong case for conservation of biodiversity. (3) Even if the relationship between biodiversity and ecosystem functioning is asymptotic at local spatial scales and in the short term, effects of biodiversity loss are likely to be important at larger temporal and spatial scales. (4) Initial arguments for the importance of biodiversity for ecosystem functioning were largely based on a precautionary approach (points 1-3). However, we are now moving to a scientific position based on accumulating experimental evidence. The future challenge is the integration of this scientific research with policy.     

Context dependency of relationships between biodiversity and ecosystem functioning is different for multiple ecosystem functions

Increasing concern over the loss of biodiversity has led to attempts to quantify relationships between biodiversity and ecosystem functioning. While manipulative investigations have accumulated substantial evidence to support the notion that decreasing biodiversity can be detrimental to the functioning of ecosystems, recent investigations have identified the potential importance of physical processes in moderating biodiversity – ecosystem function relationships at larger geographical scales. In this study, the relationship between the genus richness of benthic macro‐invertebrates and five measures of ecosystem functioning (macrofaunal biomass, depth of the apparent redox discontinuity, fluxes of ammonium and NO_x and the abundance of nematodes) was determined over a large scale wave‐induced bed shear stress gradient on the seabed of the northern Irish Sea. Ecosystem functioning was significantly correlated to genus richness for four out of five ecosystem functions. However, wave stress moderated the genus richness – ecosystem functioning relationship for only one of the ecosystem functions; genus richness had a positive effect on the depth of the apparent redox discontinuity in the sediment at high wave stress but not at low wave stress. These results indicate that the effects of biodiversity on some ecosystem functions may be sufficiently strong to generate patterns in ecosystems where other factors are also affecting ecosystem processes, but that the biodiversity–ecosystem function relationship for can be dependent on environmental conditions for specific ecosystem functions.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.