生物多样性丧失已成为全球关注的问题, 并对人类赖以生存的生态系统服务产生了负面影响(Wagg et al, 2014)。1992年在巴西里约热内卢举行的地球峰会之后, 人们对了解物种的丧失如何影响生态系统功能的兴趣不断增加(Schulze & Mooney, 1993), 涌现出一大批关于生物多样性和生态系统功能(biodiversity and ecosystem function, BEF)关系的研究成果(Hazard et al, 2017)。这些研究促进了生态学思维的重大转变, 因为在此之前, 生物多样性仅被认为是对环境变化和生态系统功能的响应变量, 而不是生态系统功能的驱动变量(Naeem, 2002; Tilman et al, 2014)。1994年, Tilman和Downing (1994)在美国明尼苏达州雪松溪生态系统科学保护区利用超过200个草地样方的数据得出物种数量的增加提高了草地抗旱能力的结果; Naeem等(1995)利用盆栽实验发现种植更多物种会拥有更高的初级生产力。这些成果均是BEF理论的有力佐证。
BEF关系的研究大多只关注生物量的贮存和生产、凋落物的分解、营养循环、土壤有机碳贮存、生物量稳定性、病原体和草食性动物的损害以及授粉等单一的生态系统功能(van der Plas, 2019)。然而, Hector和Bagchi (2007)的草地生物多样性实验发现, 不同的物种影响不同的生态系统功能, 高的生物多样性水平可能提供多种多样的功能; 当同时考虑多种功能时, 生物多样性的影响比只考虑单一功能时更加重要(Gamfeldt et al, 2008)。这让人们意识到一个生态系统具有同时提供多种功能的能力, 这个概念被称为生态系统多功能性(ecosystem multifunctionality, EMF)。近10年来, 生物多样性与生态系统多功能性(biodiversity-ecosystem multifunctionality, BEMF)关系的研究已迅速成为生态学研究的热点(Manning et al, 2018; van der Plas, 2019; Jing et al, 2020)。
目前, 关于BEMF关系的研究主要以草地、森林、干旱地、农业生态系统为对象(Garland et al, 2021)。在控制性实验研究中, 生物多样性对EMF的显著正效应较为普遍(Lefcheck et al, 2015); 但在自然生态系统中, BEMF还存在显著负相关和不显著相关关系(van der Plas, 2019)。在国内, 徐炜等(2016a, b)详细介绍了BEMF关系研究的发展历程和测度方法, 提出了需要深入研究的领域。目前, 已有科学家意识到并试图解决功能和服务指标选择的问题。例如, Hölting等(2019)总结得出当前的研究中功能和服务指标的平均个数为8个左右; Manning等(2018)以及井新和贺金生(2021)指出将功能和服务指标区别对待已成为今后研究的趋势; Garland等(2021)对现有研究的功能和服务指标进行总结归类, 提出了可参考的建议。绝大多数BEMF关系的研究都集中在分类多样性中的物种丰富度(Hector & Bagchi, 2007; Maestre et al, 2012b; Zavaleta et al, 2010), 也有评估物种均匀度(Maestre et al, 2012a; Soliveres et al, 2014; Dooley et al, 2015)、物种优势度(Lohbeck et al, 2016; Li et al, 2017)等对EMF的影响。
功能多样性也是EMF的关键驱动因子, 因为共存种间存在差异的功能性状增加了整体的资源利用, 从而促进了生态系统功能或者优势种的性状较强地影响着生态系统功能(Le Bagousse-Pinguet et al, 2019)。目前这方面的研究开始涌现(Valencia et al, 2015; Finney & Kaye, 2017; Huang et al, 2019; Mensah et al, 2020; Liu et al, 2021)。物种亲缘关系的远近可表现为功能上的差异, 因此系统发育多样性被认为是EMF的一个重要预测者(Oka et al, 2019), 但这方面的研究还不多(Zirbel et al, 2019; Liu et al, 2021)。
1 功能指标的选择
生态系统功能和服务指标的选择是EMF研究的第一步(Hölting et al, 2019)。现有研究中, 生态系统功能和服务指标的数量差异很大, 从2个到82个不等(Minden & Kleyer, 2015; Meyer et al, 2018; Garland et al, 2020), 功能指标的数量是否影响生物多样性对EMF的效应目前还存在争议(Gamfeldt & Roger, 2017; Fanin et al, 2018; Meyer et al, 2018)。涉及的生态系统功能指标主要有土壤有机碳(Maestre et al, 2012b; Soliveres et al, 2014; Jing et al, 2015; Birkhofer et al, 2018)、氮和磷的营养循环(Jing et al, 2015; Wang et al, 2020)、凋落物分解(Mouillot et al, 2011; Lohbeck et al, 2016; Mori et al, 2016; Lucas-Borja & Delgado-Baquerizo, 2019)、土壤性质与肥力(Finney & Kaye, 2017; Chandregowda et al, 2018)等; 生态系统服务指标主要有生产力(Mouillot et al, 2011; Pasari et al, 2013; Lohbeck et al, 2016)、传粉(Soliveres et al, 2016; Birkhofer et al, 2018)、抵抗力(van der Plas et al, 2016a; Ratcliffe et al, 2017; Hautier et al, 2018)等。Garland等(2021)根据EMF研究中已发表的775个功能指标, 将其归为24个功能范畴和4个生态系统服务功能类型, 可作为指标选取的参考。
此外, 研究者也针对不同的生态系统和研究目标选择某些特别的功能指标。如Mensah等(2020)在非洲山地森林研究中, 选择栖息地质量和落石保护功能等作为评价指标; Schuldt等(2018)在针对不同营养级水平的BEMF研究中, 选择寄生状况等22个指标来评价EMF。在BEF-China实验中, Trogisch等(2017)认为需要积极发展标准的BEF方法来评估森林生态系统的EMF, 并提出了包括植物生物量生产和树木生长、树木多样性、地上多营养级关联、地下微生物交互作用、营养循环、土壤流失控制相关的17个功能指标及测定的标准方法。这些实用、便捷、可重复和低成本的方法有利于森林生态系统BEMF关系的综合评价。因此, 在开展BEMF关系研究之前, 可立足于需要解决的科学问题, 选择合适的备用指标并注意区分生态系统功能和服务指标;在代替不同功能和服务的备用指标里尽可能选择涉及面广且容易获得的指标。
在功能指标的选择过程中, 生物多样性对EMF的效应是否依赖于功能的数量还存在争议(Jing et al, 2020)。评估EMF需要对功能间的权衡和协同效应作整体分析(Hölting et al, 2019)。如果所选功能间存在正相关关系并且依赖于生物多样性, 最大化EMF则需要高的生物多样性水平, 那么研究时应考虑增加多种功能; 如果功能间呈负相关关系, 即功能间存在权衡, 最大化一种功能必然导致其他功能的下降, BEMF的关系将依赖于特定的功能以及它们与生物多样性之间的关系(Meyer et al, 2018)。在EMF研究中, 容易把生态系统属性指标(如地形因子和土壤的pH值、厚度、含水量、孔隙度等)混淆为生态系统功能指标(Sacchelli et al, 2013; Li et al, 2017; Garibotti et al, 2018; Meyer et al, 2018)。事实上, 绝大多数研究都是用生态系统功能指标来替代生态系统功能(Isbell et al, 2011; Maestre et al, 2012b; Bowker et al, 2013; Lefcheck et al, 2015), 并把随生物活性、时间和空间改变的变量称之为直接变量(Reiss et al, 2009), 其他间接变量为受生态系统功能影响的关键生态系统属性(Jax, 2010)。Manning等(2018)和Garland等(2021)建议通过直接测量过程速率(process rates)来评估生态系统功能; 如果过程速率太慢而无法直接测量, 那么对某些营养库的测量可以作为较慢过程的替代。该观点尽可能地包含了所有已公布的涉及过程速率和营养库的功能指标, 同时也跨越了多个时间尺度和研究焦点(Garland et al, 2021)。
长期以来, 研究者并没有意识到所选功能指标代表的是生态系统功能还是生态系统服务。Manning等(2018)根据研究对象是基础问题还是应用问题, 提出将EMF分为“生态系统功能多功能性” (ecosystem function multifunctionality, EFM)和“生态系统服务-多功能性” (ecosystem service multifunctionality, ESM)两个水平来定义。前者侧重于生态系统中的一系列生物、物理和地球化学过程; 后者则指生态系统提供的与人类利益需求相关的多种生态系统服务。事实上, 生态系统服务-多功能性这一概念被优化为生态系统多服务性(ecosystem multiserviceability或multiservicing ability, EMS)更为贴切(贺金生等, 2000; 井新和贺金生, 2021)。Allan等(2015)和Manning等(2018)指出, 在评估ESM时需要利益相关者对涉及的每个生态系统服务赋予不同的权重, 但问题是他们并没有利用实际调查数据来研究不同利益相关群体的权重。目前, 已有利用生态系统服务功能指标来评估ESM的研究。Peura等(2018)利用8个生态系统服务功能指标比较了芬兰近自然经营和轮伐期经营森林的ESM, 发现近自然经营森林的ESM要高于轮伐期经营的森林, 为可持续林业管理提供了很好的依据。Jönsson和Snäll (2020)利用瑞典全国范围内2,000多个样地的4类生态系统服务数据, 评估了低产林的ESM, 指出低产林的ESM随林龄的增加而增加, 针叶和落叶混交林的老龄林能支持更多的生态系统服务功能, 对当地制定森林保护和管理政策具有直接的指导意义。为了以最低成本获得最大的整体效益来满足不同利益相关者, Zeng等(2021)对7个林龄序列的杉木(Cunninghamia lanceolata)人工林的ESM进行了量化, 发现林龄对供应服务的影响大于对支持服务的影响, ESM在不同管理场景下的变异可分为两个不同的阶段, 结果可为杉木人工林乃至其他人工林的管理提供具体的建议。Linders等(2021)利用访谈的方式调查了不同生态系统服务在8个利益相关群体中的权重来评估入侵树种对不同利益相关者ESM的影响, 发现随着入侵树木的增加, 大多数利益相关群体的净ESM下降, 或总体上没有显著变化; ESM随入侵树木增加的只涉及两个利益相关群体, 即木炭生产者和涉及区域发展的非政府组织, 结果有助于制定经营决策以及土地管理方案。生态系统多服务性这一概念因其强调涉及利益相关者的生态系统服务指标, 在生态系统的可持续管理以及生物多样性保护等领域应深入研究。
2 生物多样性的多重维度与生态系统多功能性
2.1 物种多样性
但是, Gamfeldt和Roger (2017)指出, 物种丰富度在大多数情况下(特别是野外调查而不是控制实验时)可能并不适用于BEMF研究; 它被广泛应用只因为是最简单的度量标准, 并可扩展到其他多样性度量标准。基于多度的Shannon-Wiener指数和Simpson指数等多样性参数因能赋予稀有种和常见种不同的权重而更能代表真实的物种多样性水平(Jing et al, 2020)。Hertzog等(2019)指出, 在树木丰富度程度较低的温带森林里, 树木的特性和优势度对生态系统功能的影响往往比物种多样性要强; 除了探究需要多少物种来维持某种程度的EMF外, 更要了解哪些树种以及树种组成的效应(Baeten et al, 2019)。
不同尺度的物种多样性(α、β以及γ多样性)在各生态系统中与EMF的关系也不尽相同。在欧洲森林中, α和β多样性均是EMF的重要驱动者, 生物同质化可能会对大尺度的EMF产生尚未被认识的负面影响(van der Plas et al, 2016b)。在北美大草原, β多样性的增加比α多样性更能够提升EMF, 群落的异质性对草原恢复期间重建EMF具有重要作用(Grman et al, 2018)。但同样是草地生态系统, Pasari等(2013)却发现, α多样性与绝大多数的单一生态系统功能和EMF均有强烈的正效应, 而β和γ多样性只与EMF呈显著正相关; 高的β多样性降低了EMF的变异, 说明群落的异质性是维持EMF的前提。
2.2 功能多样性
功能多样性是基于物种功能性状来定义的, 指影响生态系统功能的物种所具有的功能性状的大小、范围及分布(Dı́az & Cabido, 2001; Petchey & Gaston, 2002)。植物功能性状是指一系列与其个体定植、存活、生长以及死亡紧密相关的属性, 且这些属性能够显著影响生态系统功能, 并能反映植被对环境影响的响应(刘晓娟和马克平, 2015)。功能性状与个体扩散、生长、养分循环、能量利用、生态策略等密切相关, 是研究功能多样性与生态系统功能关系的重要纽带(Pérez-Harguindeguy et al, 2013)。理论和实证表明, 与物种多样性相比, 功能多样性能够更好地反映生态系统功能(Steudel et al, 2016)。究其原因, 是因为物种多样性尤其是物种丰富度并不直接提供性状的信息, 物种丰富度与生态系统功能间的正相关关系可能是因为物种数量的增加导致了性状多样性的增加; 而功能性状与物种的资源利用方式(包括选择效应和互补效应)有关(Dı́az & Cabido, 2001), 可能是生态系统功能较好的预测者(Zirbel et al, 2019)。由于性状间存在差异的共存物种能够利用不同时空中的相同资源而增加整体的资源利用效率, 所以高水平的性状多样性能够提高生态系统功能(Hooper et al, 2005; Gross et al, 2007; de Bello et al, 2010)。
研究表明, 在多种不同的生态系统(草地、农业、森林)中, 功能多样性均是EMF的主要驱动者(Mouillot et al, 2011; Finney & Kaye, 2017; Huang et al, 2019)。Valencia等(2015)指出, 在地中海干旱生态系统中, 功能多样性是干旱和灌木入侵响应下的EMF的重要驱动者, 能够增进生态系统对干旱的抵抗力。Gross等(2017)根据全球124个干旱区植物群落的数据研究发现, 功能性状多样性对EMF拥有最多的解释率, 因而能够与EMF的最大化联系起来, 性状的分布可以用来预测陆地生态系统生物多样性丧失的功能后果。最新研究表明, 功能性状多样性比物种丰富度和功能优势值更能解释EMF的变异; 在多个物种组成的森林中, 拥有不同性状的不同物种比功能上相似的不同物种对EMF的贡献要大, 这在农林生态系统的管理上具有重要意义(Mensah et al, 2020)。在云南松(Pinus yunnanensis)林的经营管理中, 择伐强度主要通过增加功能多样性来间接提高EMF (Huang et al, 2020)。Yan等(2020)指出, 功能多样性的两个尺度α和β多样性在维持草原生态系统土壤EMF方面均发挥着重要作用, β多样性是减缓干旱对土壤EMF不利影响的重要调节者。
2.3 系统发育多样性
系统发育多样性指一个群落中物种谱系距离的总和, 它受平均种间亲缘关系和群落中物种数量的影响(Srivastava et al, 2012), 可以反映分类群的亲缘关系和进化信息。由于亲缘关系相近的物种具有相似的性状, 系统发育多样性往往可以作为功能多样性的替代(Wiens & Graham, 2005)。因为一般难以测量所有与功能相关的性状, 当系统发育多样性有效地包括了与生态系统功能相关的生物学性状时, 它将是影响生态系统功能的一个关键生物多样性属性, 比功能多样性更能揭示生态系统功能(Flynn et al, 2011; Venail et al, 2015)。近年来, 已有学者开始综合考虑分类学多样性、功能多样性和系统发育多样性对EMF的影响(Le Bagousse-Pinguet et al, 2019; Luo et al, 2019; Zirbel et al, 2019; Huang et al, 2020)。但很多研究表明, 相比功能多样性, 系统发育多样性并不能解释更多的EMF变异(Zirbel et al, 2019; Huang et al, 2020; Liu et al, 2021), 可能的解释是一些功能性状丧失了演化上的保守模式(Srivastava et al, 2012)。
3 微生物多样性与生态系统多功能性
土壤微生物是地球上最丰富多样的有机体(Locey & Lennon, 2016), 占整个地球生物多样性的1/4 (Wagg et al, 2019)。在不同的生态系统中, 土壤微生物通过促进凋落物分解、养分循环以及资源可利用性, 在维持EMF方面也扮演着关键角色(Delgado-Baquerizo et al, 2016b, 2020; Wagg et al, 2019)。诸多研究表明, 气候变化、土地利用变化等因素导致的土壤微生物多样性丧失将威胁土壤微生物支持EMF的能力(Jing et al, 2015; Ren et al, 2018; Delgado-Baquerizo et al, 2017, 2020)。
真菌和细菌多样性对EMF的影响并不一致, 这种差异可能也反映了不同类群微生物(比如真菌和细菌)生活史的差异(Jing et al, 2015)。在农田生态系统中, 土壤细菌和真菌多样性均与EMF呈显著正相关(Luo et al, 2018)。在青藏高原草地生态系统中, 细菌多样性对EMF的调节就比真菌重要(Jing et al, 2015)。在寒带森林中, 真菌多样性对EMF贡献最大(Li et al, 2019)。在温带森林中, 土壤微生物对多种功能的均值没有直接影响, 但其是维持高水平EMF的关键驱动者(Yuan et al, 2020)。也有研究指出, 微生物是有结构的, 相互之间会形成复杂的微生物网络, 不同生态系统类型的微生物网络的复杂性也并不相同, 因此不能绝对地将真菌和细菌割裂开来(de Vries et al, 2018; Banerjee et al, 2019)。Wagg等(2019)首次将微生物网络的复杂性与EMF联系起来, 揭示了真菌和细菌群落内部的相互作用对提高EMF的重要性, 并证明了地下微生物群落复杂生态联系的消失将会损害生态系统功能。
4 生态系统多功能性的其他驱动因子
4.1 土地利用变化
Allan等(2015)研究发现, 在草地生态系统中, 土地利用强度对EMF整体上有很强的效应, 除直接效应外, 还通过生物多样性和功能结构的变化间接改变EMF。Wen等(2020)揭示, 土地利用强度的加剧可通过降低植物多样性及其与土壤细菌多样性的交互作用降低EMF。近年来, 放牧强度如何影响EMF的研究也逐渐增多, 并发现EMF随放牧强度的增加呈下降趋势(Ren et al, 2018), 中低强度的放牧有助于提高EMF, 且能提高土壤肥力和稳定性以及土壤碳贮量(Peco et al, 2017)。Luo等(2018)发现, 长期的施肥管理措施能够增加微生物多样性, 促进农业生态系统功能。Birkhofer等(2018)分析了农业生态系统中由土地利用变化引起的景观复杂性与EMF间的关系, 并没有发现二者显著性相关。Fu等(2018)针对森林生态系统的研究表明, 土地覆盖变化通过影响群落的功能结构而影响EMF。总的来说, 关于土地利用变化影响EMF的研究开展的时间并不长, 在不同生态系统土地利用快速变化的背景下, 土地利用变化与EMF关系的研究还有待于进一步加强。
4.2 气候变化
气候变化导致的干旱正在全球范围内蔓延, 干旱生态系统已占地球表面积的41%, 承载着38%的人口(MEA, 2005)。干旱导致生物多样性降低, 严重影响生态系统结构和功能, 因此保护植物和微生物等生物多样性对减缓干旱对生态系统功能的消极影响至关重要(Maestre et al, 2012a; Valencia et al, 2015; Delgado-Baquerizo et al, 2016a; Berdugo et al, 2017, 2020)。在泥炭沼泽地中, 干旱对EMF造成了严重威胁(Robroek et al, 2017)。关于温度和降水对EMF的影响目前尚未得到一致的结论。在青藏高原草地生态系统中, 年均降水量对EMF没有显著影响, 但年均温度有显著正影响(Jing et al, 2015); Pan等(2017)的研究结果表明年均温度对EMF有直接和间接的消极影响。在全球旱地生态系统中, 年均降水量对EMF有显著正影响, 年均温度却没有影响(Delgado-Baquerizo et al, 2016a)。同样在干旱地区, Maestre等(2012b)则发现年均温度的升高会降低EMF。因此, 还需要加强对不同生态系统类型及不同尺度的研究, 积累更丰富的数据资料。
4.3 其他驱动因子
近年来, 科学家针对不同生态系统的研究发现, 地质过程、景观变化、生境多样性、土壤的化学计量比、森林破碎化以及林分复杂性等因子均是EMF重要的驱动因子。
Hu等(2020)量化了地质过程(母岩、风化等)和当代环境对植物、微生物群落和生态系统功能的相对贡献, 表明植物、微生物多样性和EMF受地质过程和当代环境共同驱动。在大空间尺度上, 样地的环境条件可能抑制或增加多样性对EMF的影响(van der Plas et al, 2016b; Hautier et al, 2018)。van der Plas等(2016b)和Hautier等(2018)都发现, 在大的空间范围内(欧洲森林和世界范围的草原), 环境变化对EMF的影响有时比生物多样性的影响更大。Zirbel等(2019)指出, 尽管BIODEPTH实验(泛欧洲生物多样性-生态系统功能实验)没有明确包括环境变量, 但样地分布在8个欧洲国家, 很可能包括相对大的环境变化。
在温带森林演替阶段EMF变化的研究中, 土壤的化学计量比(C : N)是EMF变化的主要驱动因子; 近100年来, 森林有机质含量(较低的C : N)的增加促进了EMF的提高(Lucas-Borja & Delgado-Baquerizo, 2019)。
由于人类活动, 世界范围内的生态系统正面临着生境同质化的危险, Alsterberg等(2017)通过对生境多样性如何影响EMF的研究发现, 生境多样性对EMF有直接和间接的效应, 且表现为季节间的差异。
5 展望
5.1 功能指标的选择和量化方法
缺乏界定功能和服务指标的统一标准限制着BEMF关系的深入研究, 同时ESM研究也必然涉及功能和服务指标的选择(井新和贺金生, 2021)。未来的研究中应明确涵盖的功能指标, 如常用的涉及生产力、营养循环、凋落物分解、气候调节等方面的指标, 因为某个功能可能有很多合适的功能指标, 所以必须解释为什么要用某个特定的指标来评估某一项功能, 以便在具体解决某一科学问题的背景下恰当地解释EMF指数。ESM概念的提出为不同利益相关群体对生态系统服务的关切找到了一种平衡方式(Manning et al, 2018; 井新和贺金生, 2021)。目前, 基于基础研究的EFM和基于应用研究的ESM还仅仅是一种概念上的分化(Hölting et al, 2019), 绝大部分的研究兼顾了功能和服务价值的评估(Allan et al, 2015; van der Plas et al, 2016b)。归根到底, ESM取决于单一功能的价值, 价值的估算应结合全部利益相关者(森林管理者、自然资源保护者以及公众等)赋予的不同权重(Hertzog et al, 2019)。这种不同功能和服务交互的研究更需要注重概念和方法的发展。比如在多功能性基础上, 生态系统多服务性概念的提出就是一种概念上的发展, 新概念需要新理论和新方法的支持, 这在未来依然是一个挑战。
目前量化EMF的方法都有其优缺点(Byrnes et al, 2014; Manning et al, 2018; Hölting et al, 2019), 基于不同统计学原理的方法会得到不同的结果(Jing et al, 2020; 井新和贺金生, 2021), 这严重阻碍了BEMF关系研究的发展。鉴于生态系统功能可通过相互作用的网络关联而共享驱动因子, Manning等(2018)提出为避免个别功能类别权重过大, 可通过聚类分析将功能指标分成n类, 在分析中对每一类赋予相同的权重。例如, 如果某类功能包含了4个功能指标, 则对其每个指标赋予权重0.25, 以避免相似的功能指标比例过高。Jing等(2020)提出了一种基于变量数值范围的标准化方法(scaling standardization method), 可将平均或加和参数转变为改进的标准化多功能性指数, 以便对不同的研究结果进行比较, 并可应用到阈值法中解决真正的生态学问题, 避免数理统计上的一些假象。这些新方法在今后的研究中应得到更多的应用。
5.2 生物多样性与生态系统多功能性关系的驱动机制
阐明BEMF的驱动机制是管理生态系统及其提供的服务的前提和研究的重点, 生物多样性驱动生态系统功能的选择性和互补性机制可能也是驱动EMF的潜在机制(Gamfeldt & Roger, 2017)。除了生物多样性, 非生物因子机制也已被发现, 并越来越受到重视(Constán-Nava et al, 2015; Zhang et al, 2016; Chandregowda et al, 2018; Luo et al, 2018)。通过线性模型、结构方程模型、随机森林模型等统计方法可以区分生物因子和非生物因子与EMF之间的复杂关系(Jing et al, 2015; Delgado-Baquerizo et al, 2017; Luo et al, 2019; Zirbel et al, 2019; Hu et al, 2020)。为了促进支撑人类福祉的生态系统功能, 不仅要保护生物多样性本身, 也要促进有利于物种具有适当性状组合的非生物条件(van der Plas, 2019)。
BEMF具有多种驱动机制, 并且常常交织在一起, 给研究带来了挑战(Giling et al, 2019)。这也说明了揭示驱动BEMF关系潜在机制的重要性。例如Meyer等(2018)的研究表明, 当考虑更多的功能时, 生物多样性的效应会变得更强, 且强度取决于功能的特性; 而Gamfeldt和Roger (2017)的研究表明, BEMF的关系并不因功能数量的改变而改变。Jing等(2020)通过对统计学原理的解释揭示了量化EMF指数的方法不同会得出截然不同的结果, 提升了对BEMF关系的认识。Slade等(2019)指出, 群落内的竞争关系降低了物种的个体多度及其对生态系统功能的贡献, 充分了解环境条件、功能间的权衡和物种间的竞争这三者间的关系是了解BEMF复杂关系背后维持机制的必要条件。大量研究表明, 地上和地下生物多样性以及生物多样性的不同尺度或维度的综合影响能够解释EMF更多的变异(Yan et al, 2020; Yuan et al, 2020)。因此, 未来的研究应包括地上和地下不同尺度或维度的生物多样性指标, 尽可能全面地分析EMF的生物驱动因子。
5.3 生物多样性与生态系统多功能性关系的研究对象
综上, 我们建议未来BEMF关系的研究应在区分功能和服务的基础上并行不悖地发展。对于功能或服务指标的选择应制定指南性质的标准以供参考。未来的研究可侧重于不同尺度上的典型森林BEMF关系对全球变化的响应; 加强不同生物多样性维度、微生物多样性以及非生物因子对EMF的综合影响以及多营养级水平的生物多样性对EMF影响机制的研究; 尽快在相关研究中应用提出的新概念(比如生态系统多服务性)以及开发的新方法(比如基于变量数值范围的标准化方法)。
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Functional trait diversity is a stronger predictor of multifunctionality than dominance: Evidence from an Afromontane forest in South Africa
DOI:10.1016/j.ecolind.2020.106415 URL [本文引用: 4]
Biodiversity-multifunctionality relationships depend on identity and number of measured functions
Ecosystem multifunctionality of coastal marshes is determined by key plant traits
DOI:10.1111/jvs.12276 URL [本文引用: 1]
Low multifunctional redundancy of soil fungal diversity at multiple scales
DOI:10.1111/ele.2016.19.issue-3 URL [本文引用: 1]
Functional structure of biological communities predicts ecosystem multifunctionality
DOI:10.1371/journal.pone.0017476 URL [本文引用: 3]
Ecosystem consequences of biodiversity loss: The evolution of a paradigm
DOI:10.1890/0012-9658(2002)083[1537:ECOBLT]2.0.CO;2 URL [本文引用: 1]
Empirical evidence that declining species diversity may alter the performance of terrestrial ecosystems
DOI:10.1098/rstb.1995.0025 URL [本文引用: 1]
Phylogenetic clustering in beneficial attributes of tree species directly linked to provisioning, regulating and cultural ecosystem services
DOI:10.1016/j.ecolind.2018.09.035 URL [本文引用: 1]
Climatic and geographic factors affect ecosystem multifunctionality through biodiversity in the Tibetan alpine grasslands
DOI:10.1007/s11629-016-4242-6 URL [本文引用: 1]
Several scales of biodiversity affect ecosystem multifunctionality
Effects of grazing abandonment on soil multifunctionality: The role of plant functional traits
DOI:10.1016/j.agee.2017.08.013 URL [本文引用: 1]
New handbook for standardised measurement of plant functional traits worldwide
DOI:10.1071/BT12225 URL [本文引用: 1]
Functional diversity (FD), species richness and community composition
DOI:10.1046/j.1461-0248.2002.00339.x URL [本文引用: 1]
Continuous cover forestry is a cost-efficient tool to increase multifunctionality of boreal production forests in Fennoscandia
DOI:10.1016/j.biocon.2017.10.018 URL [本文引用: 1]
Biodiversity and ecosystem functioning relations in European forests depend on environmental context
DOI:10.1111/ele.12849
PMID:28925074
[本文引用: 1]
The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests.© 2017 John Wiley & Sons Ltd/CNRS.
Emerging horizons in biodiversity and ecosystem functioning research
DOI:10.1016/j.tree.2009.03.018 URL [本文引用: 1]
Livestock grazing regulates ecosystem multifunctionality in semi-arid grassland
DOI:10.1111/fec.2018.32.issue-12 URL [本文引用: 2]
Diverse fen plant communities enhance carbon-related multifunctionality, but do not mitigate negative effects of drought
DOI:10.1098/rsos.170449
PMID:29134063
[本文引用: 1]
Global change, like droughts, can destabilize the carbon sink function of peatlands, either directly or indirectly through changes in plant community composition. While the effects of drought and plant community composition on individual carbon (C) related processes are well understood, their effect on multiple C-related processes simultaneously-multifunctionality-is poorly known. We studied the effect of drought on four C-related processes (net and gross CO exchange, methane fluxes, and dissolved organic carbon content) in a plant removal experiment. Plant functional type (PFT) removal (graminoids, herbs, spp., incl. combinations) negatively affected multifunctionality; most markedly when all PFTs were removed. Our results corroborate a negative drought effect on C-related multifunctionality. Drought reduced multifunctionality, and this reduction was again largest when all PFTs were removed. Our data further indicate that much of these negative drought effects were carried over and maintained from the initial removal treatment. These results suggest that while a high diversity in plant functional types is associated to high C-related multifunctionality, plant community assembly does not drive the ability of peatlands to withstand the negative impacts of drought on multifunctionality. Hence, to safeguard the carbon cycling function in intact peatlands, the effects of climate change on the functional composition of the peatland plant community needs to be minimized.
Bioenergy production and forest multifunctionality: A trade-off analysis using multiscale GIS model in a case study in Italy
DOI:10.1016/j.apenergy.2012.11.038 URL [本文引用: 1]
Context- dependency of tree species diversity, trait composition and stand structural attributes regulate temperate forest multifunctionality
DOI:10.1016/j.scitotenv.2020.143724 URL [本文引用: 1]
Biodiversity across trophic levels drives multifunctionality in highly diverse forests
DOI:10.1038/s41467-018-05421-z URL [本文引用: 6]
Biodiversity and ecosystem function
When do more species maximize more ecosystem services?
DOI:10.1016/j.tplants.2019.06.014 URL [本文引用: 1]
Plant diversity and ecosystem multifunctionality peak at intermediate levels of woody cover in global drylands
The global spread of woody plants into grasslands is predicted to increase over the coming century. While there is general agreement regarding the anthropogenic causes of this phenomenon, its ecological consequences are less certain. We analyzed how woody vegetation of differing cover affects plant diversity (richness and evenness) and multiple ecosystem functions (multifunctionality) in global drylands, and how this changes with aridity.224 dryland sites from all continents except Antarctica widely differing in their environmental conditions (from arid to dry-subhumid sites) and woody covers (from 0 to 100%).Using a standardized field survey, we measured the cover, richness and evenness of perennial vegetation. At each site, we measured 14 ecosystem functions related to soil fertility and the build-up of nutrient pools. These functions are critical for maintaining ecosystem function in drylands.Species richness and ecosystem multifunctionality were strongly influenced by woody vegetation, with both variables peaking at relative woody covers (RWC) of 41-60%. This relationship shifted with aridity. We observed linear positive effects of RWC in dry-subhumid sites. These positive trends shifted to hump-shaped RWC-diversity and multifunctionality relationships under semiarid environments. Finally, hump-shaped (richness, evenness) or linear negative (multifunctionality) effects of RWC were found under the most arid conditions.Plant diversity and multifunctionality peaked at intermediate levels of woody cover, although this relationship became increasingly positive under wetter environments. This comprehensive study accounts for multiple ecosystem attributes across a range of woody covers and environmental conditions. Our results help us to reconcile contrasting views of woody encroachment found in current literature and can be used to improve predictions of the likely effects of encroachment on biodiversity and ecosystem services.
Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality
Phylogenetic diversity and the functioning of ecosystems
DOI:10.1111/j.1461-0248.2012.01795.x
PMID:22583836
[本文引用: 2]
Phylogenetic diversity (PD) describes the total amount of phylogenetic distance among species in a community. Although there has been substantial research on the factors that determine community PD, exploration of the consequences of PD for ecosystem functioning is just beginning. We argue that PD may be useful in predicting ecosystem functions in a range of communities, from single-trophic to complex networks. Many traits show a phylogenetic signal, suggesting that PD can estimate the functional trait space of a community, and thus ecosystem functioning. Phylogeny also determines interactions among species, and so could help predict how extinctions cascade through ecological networks and thus impact ecosystem functions. Although the initial evidence available suggests patterns consistent with these predictions, we caution that the utility of PD depends critically on the strength of phylogenetic signals to both traits and interactions. We advocate for a synthetic approach that incorporates a deeper understanding of how traits and interactions are shaped by evolution, and outline key areas for future research. If these complexities can be incorporated into future studies, relationships between PD and ecosystem function bear promise in conceptually unifying evolutionary biology with ecosystem ecology.© 2012 Blackwell Publishing Ltd/CNRS.
Contrasting biodiversity-ecosystem functioning relationships in phylogenetic and functional diversity
DOI:10.1111/nph.2016.212.issue-2 URL [本文引用: 1]
Biodiversity and stability in grasslands
DOI:10.1038/367363a0 URL [本文引用: 1]
Biodiversity and ecosystem functioning
DOI:10.1146/ecolsys.2014.45.issue-1 URL [本文引用: 1]
Toward a methodical framework for comprehensively assessing forest multifunctionality
DOI:10.1002/ece3.2017.7.issue-24 URL [本文引用: 1]
Functional diversity enhances the resistance of ecosystem multifunctionality to aridity in Mediterranean drylands
DOI:10.1111/nph.13268
PMID:25615801
[本文引用: 4]
We used a functional trait-based approach to assess the impacts of aridity and shrub encroachment on the functional structure of Mediterranean dryland communities (functional diversity (FD) and community-weighted mean trait values (CWM)), and to evaluate how these functional attributes ultimately affect multifunctionality (i.e. the provision of several ecosystem functions simultaneously). Shrub encroachment (the increase in the abundance/cover of shrubs) is a major land cover change that is taking place in grasslands worldwide. Studies conducted on drylands have reported positive or negative impacts of shrub encroachment depending on the functions and the traits of the sprouting or nonsprouting shrub species considered. FD and CWM were equally important as drivers of multifunctionality responses to both aridity and shrub encroachment. Size traits (e.g. vegetative height or lateral spread) and leaf traits (e.g. specific leaf area and leaf dry matter content) captured the effect of shrub encroachment on multifunctionality with a relative high accuracy (r(2) = 0.63). FD also improved the resistance of multifunctionality along the aridity gradient studied. Maintaining and enhancing FD in plant communities may help to buffer negative effects of ongoing global environmental change on dryland multifunctionality. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
Biodiversity and ecosystem functioning in naturally assembled communities
Jack-of-all-trades effects drive biodiversity-ecosystem multifunctionality relationships in European forests
DOI:10.1038/ncomms11109 URL [本文引用: 2]
Biotic homogenization can decrease landscape- scale forest multifunctionality
Species richness, but not phylogenetic diversity, influences community biomass production and temporal stability in a re-examination of 16 grassland biodiversity studies
DOI:10.1111/fec.2015.29.issue-5 URL [本文引用: 1]
Soil biodiversity and soil community composition determine ecosystem multifunctionality
Fungal-bacterial diversity and microbiome complexity predict ecosystem functioning
DOI:10.1038/s41467-019-12798-y URL [本文引用: 3]
High ecosystem multifunctionality under moderate grazing is associated with high plant but low bacterial diversity in a semi-arid steppe grassland
DOI:10.1007/s11104-020-04430-6 URL [本文引用: 2]
Land-use intensity indirectly affects soil multifunctionality via a cascade effect of plant diversity on soil bacterial diversity
DOI:10.1016/j.gecco.2020.e01061 URL [本文引用: 1]
Niche conservatism: Integrating evolution, ecology, and conservation biology
DOI:10.1146/ecolsys.2005.36.issue-1 URL [本文引用: 1]
A review on the measurement of ecosystem multifunctionality
DOI:10.17520/biods.2015170 URL [本文引用: 1]
生态系统多功能性的测度方法
Biodiversity and ecosystem multifunctionality: Advances and perspectives
DOI:10.17520/biods.2015091 URL [本文引用: 1]
生物多样性与生态系统多功能性: 进展与展望
Plant functional β diversity is an important mediator of effects of aridity on soil multifunctionality
DOI:10.1016/j.scitotenv.2020.138529 URL [本文引用: 2]
Above- and below-ground biodiversity jointly regulate temperate forest multifunctionality along a local-scale environmental gradient
DOI:10.1111/jec.v108.5 URL [本文引用: 3]
Sustaining multiple ecosystem functions in grassland communities requires higher biodiversity
Ecosystem service multifunctionality of Chinese fir plantations differing in stand age and implications for sustainable management
DOI:10.1016/j.scitotenv.2021.147791 URL [本文引用: 1]
Biotic communities cannot mitigate the negative effects of grazing on multiple ecosystem functions and services in an arid shrubland
DOI:10.1007/s11104-015-2754-4 URL [本文引用: 1]
Landscape context explains ecosystem multifunctionality in restored grasslands better than plant diversity
