生物多样性与生态系统多功能性: 进展与展望

doi:10.17520/biods.2015091

摘要/Abstract

摘要：

全球变化和人类活动引起的生物多样性丧失将会对生态系统功能产生诸多不利影响, 如生产力下降、养分循环失衡等。因此, 始于20世纪90年代的生物多样性与生态系统功能(biodiversity and ecosystem functioning, BEF)研究一直是生态学界关注的热点。然而, 随着研究的深入, 人们逐步认识到生态系统并非仅仅提供单个生态系统功能, 而是能同时提供多个功能, 这一特性被称之为“生态系统多功能性” (ecosystem multifunctionality, EMF)。尽管有此认识, 但直到2007年, 研究者才开始定量描述生物多样性与生态系统多功能性(biodiversity and ecosystem multifunctionality, BEMF)的关系。目前, BEMF研究已成为生态学研究的一个重要议题, 但仍存在很多问题和争议, 如缺少公认的多功能性测度标准、生态系统不同功能之间的权衡问题等。本文概述了BEMF研究的发展历程、常用的量化方法、EMF的维持机制和不同研究视角下BEMF的关系。针对现有研究中的不足, 本文还总结了需要进一步深入研究的地方, 特别强调了优化EMF测度方法和研究不同维度生物多样性与EMF间关系的重要性, 以期对未来的BEMF研究有所帮助。

关键词: 生态系统功能, 物种丧失, 多功能性指数, 多功能

Abstract

As global biodiversity losses accelerate, there is increasing evidence shows that there may be negative impacts on ecosystem functioning, such as declines in plant primary productivity and imbalances in nutrient cycling. Thus, it is critical to understand the relationship between biodiversity and ecosystem functioning (BEF). However, ecosystems can provide multiple functions simultaneously (ecosystem multifunctionality, EMF). Since 2007, the quantification of relationships between biodiversity and ecosystem multifunctionality (BEMF) has generated additional questions and controversies, such as the lack of consensus in appropriate multifunctionality indices and uncertain trade-offs among ecosystem functions. In this review, we briefly summarize the history of BEMF studies and the methods of EMF quantification, then outline the mechanisms of EMF maintenance and current research progress. We emphasize the importance of optimizing EMF quantifications and investigating the relationship between different dimensions of biodiversity and EMF. We also provide suggestions and directions for future research on BEMF.

Key words: ecosystem functioning, species loss, multifunctionality indices, multiple functions

徐炜, 马志远, 井新, 贺金生 (2016) 生物多样性与生态系统多功能性: 进展与展望. 生物多样性, 24, 55. DOI: 10.17520/biods.2015091.

Wei Xu, Zhiyuan Ma, Xin Jing, Jin-Sheng He (2016) Biodiversity and ecosystem multifunctionality: advances and perspectives. Biodiversity Science, 24, 55. DOI: 10.17520/biods.2015091.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2015091

https://www.biodiversity-science.net/CN/Y2016/V24/I1/55

图/表 4

表1 生物多样性与生态系统多功能性(BEMF)热点文献

Table 1 Key references on biodiversity and ecosystem multifunctionality (BEMF)

序号 No.	文献 References	备注 Remarks
1	Sanderson et al, 2004	首次提出多功能性的概念。This paper firstly defined multifunctionality.
2	Hector & Bagchi, 2007	第一次定量分析了生物多样性同时对多个生态系统过程的作用, 引发BEMF研究热潮。This paper first quantified the effects of biodiversity on multiple ecosystem processes, and evoked research climax of BEMF.
3	Gamfeldt et al, 2008	提出一个概念模型来探讨物种丧失对生态系统多个功能的影响; 证明多功能冗余比单功能冗余程度低。This paper proposed a conceptual model to address how species loss affects overall ecosystem functioning and demonstrated that multifunctional redundancy was generally lower than single functional redundancy.
4	Zavaleta et al, 2010	第一次在多个时间尺度上探讨生物多样性与生态系统多功能性的关系。This paper studied the relationships between biodiversity and ecosystem multifunctionality at multiple temporal scales.
5	Isbell et al, 2011	评估生物多样性与生态系统多功能性的关系时, 第一次同时考虑了不同的时间、地点、功能和气候变化等因素。This study first considered multiple times, places, functions or climate change scenarios in the evaluations of the relationships between biodiversity and ecosystem services.
6	Maestre et al, 2012b	第一次研究全球干旱地的植物物种丰富度和生态系统多功能性的关系。This paper first investigated the relationships between plant species richness and ecosystem multifunctionality in drylands at global scale.
7	Maestre et al, 2012a	第一次通过实验探讨关键群落属性(物种丰富度、群落物种组成、均匀度和空间格局)同时改变对生态系统多功能性的影响。This paper first investigated the effects of simultaneously changes in key community traits (e.g., species richness, community components, evenness, and spatial patterns) on ecosystem multifucatioality.
8	Pasari et al, 2013	第一次研究了多个尺度的生物多样性对生态系统多功能性的影响。This paper first studied the relationships between several scales of BEMF.
9	Byrnes et al, 2014a	利用BIODEPTH实验数据和R软件的multifunc程序包系统综述了当前BEMF研究中存在的问题, 首次提出多阈值法。This paper systematically reviewed the challenges and solutions in current BEMF studies and first quantified EMF with multiple threshold approach using data from the BIODEPTH experiment and the multifunc packages in R.
10	Wagg et al, 2014	通过改变土壤群落多样性实验, 提出土壤生物多样性和土壤群落组成决定生态系统多功能性。This study demonstrated that soil biodiversity and soil community composition determine ecosystem multifunctionality by manipulating soil community biodiversity.
11	Perkins et al, 2015	第一次研究了变化环境下生物多样性与生态系统多功能性的关系。This paper first studied the relationships between biodiversity and ecosystem multifunctionality in a changing environmental scenario.
12	Valencia et al, 2015	首次研究了功能多样性与生态系统多功能性的关系。This paper first studied the relationship between functional diversity and ecosystem multifunctionality.
13	Lefcheck et al, 2015	第一次系统分析了不同分类群、营养级和生境条件下生物多样性对生态系统多功能性的作用。This study presented the first systematic investigation of biodiversity’s effect on ecosystem multifunctionality across multiple taxonomy, trophic levels, and habitats.

表1 生物多样性与生态系统多功能性(BEMF)热点文献

Table 1 Key references on biodiversity and ecosystem multifunctionality (BEMF)

序号 No.	文献 References	备注 Remarks
1	Sanderson et al, 2004	首次提出多功能性的概念。This paper firstly defined multifunctionality.
2	Hector & Bagchi, 2007	第一次定量分析了生物多样性同时对多个生态系统过程的作用, 引发BEMF研究热潮。This paper first quantified the effects of biodiversity on multiple ecosystem processes, and evoked research climax of BEMF.
3	Gamfeldt et al, 2008	提出一个概念模型来探讨物种丧失对生态系统多个功能的影响; 证明多功能冗余比单功能冗余程度低。This paper proposed a conceptual model to address how species loss affects overall ecosystem functioning and demonstrated that multifunctional redundancy was generally lower than single functional redundancy.
4	Zavaleta et al, 2010	第一次在多个时间尺度上探讨生物多样性与生态系统多功能性的关系。This paper studied the relationships between biodiversity and ecosystem multifunctionality at multiple temporal scales.
5	Isbell et al, 2011	评估生物多样性与生态系统多功能性的关系时, 第一次同时考虑了不同的时间、地点、功能和气候变化等因素。This study first considered multiple times, places, functions or climate change scenarios in the evaluations of the relationships between biodiversity and ecosystem services.
6	Maestre et al, 2012b	第一次研究全球干旱地的植物物种丰富度和生态系统多功能性的关系。This paper first investigated the relationships between plant species richness and ecosystem multifunctionality in drylands at global scale.
7	Maestre et al, 2012a	第一次通过实验探讨关键群落属性(物种丰富度、群落物种组成、均匀度和空间格局)同时改变对生态系统多功能性的影响。This paper first investigated the effects of simultaneously changes in key community traits (e.g., species richness, community components, evenness, and spatial patterns) on ecosystem multifucatioality.
8	Pasari et al, 2013	第一次研究了多个尺度的生物多样性对生态系统多功能性的影响。This paper first studied the relationships between several scales of BEMF.
9	Byrnes et al, 2014a	利用BIODEPTH实验数据和R软件的multifunc程序包系统综述了当前BEMF研究中存在的问题, 首次提出多阈值法。This paper systematically reviewed the challenges and solutions in current BEMF studies and first quantified EMF with multiple threshold approach using data from the BIODEPTH experiment and the multifunc packages in R.
10	Wagg et al, 2014	通过改变土壤群落多样性实验, 提出土壤生物多样性和土壤群落组成决定生态系统多功能性。This study demonstrated that soil biodiversity and soil community composition determine ecosystem multifunctionality by manipulating soil community biodiversity.
11	Perkins et al, 2015	第一次研究了变化环境下生物多样性与生态系统多功能性的关系。This paper first studied the relationships between biodiversity and ecosystem multifunctionality in a changing environmental scenario.
12	Valencia et al, 2015	首次研究了功能多样性与生态系统多功能性的关系。This paper first studied the relationship between functional diversity and ecosystem multifunctionality.
13	Lefcheck et al, 2015	第一次系统分析了不同分类群、营养级和生境条件下生物多样性对生态系统多功能性的作用。This study presented the first systematic investigation of biodiversity’s effect on ecosystem multifunctionality across multiple taxonomy, trophic levels, and habitats.

图1 物种丰富度与生态系统功能的关系随冗余度的变化。 (a)物种丧失对假想群落的影响。实线表示当丧失的每个物种都是群落中存在的最有效率的物种时, 功能丧失最大化(物种丰富度由高转低时)。箭头和虚线表示随着冗余的变化, 物种丧失对生态系统功能影响的变化。(b)此图表示概念上维持整体功能的概率如何依赖于功能的数量和物种间多功能冗余的程度。随着某些功能的特化, 多个功能将比单个功能更易受到物种丧失的影响, 这种敏感性将随着冗余度的降低而增加(修改自Gamfeldt et al, 2008)。

Fig. 1 The relationship between species richness and ecosystem functioning changes with redundancy. (a) The effect of species loss on hypothetical communities. The solid line represents maximum loss in function if each species lost is the most efficient species of the ones remaining in the community (as we move from high to low species richness). The arrow and the dashed line indicate how the influence of species loss on ecosystem function shifts with changing redundancy. (b) A conceptual model of how the probability of maintaining overall functioning depends on both the number of functions and the degree of multifunctional redundancy across species. With some functional specialization, multiple functions will always be more susceptible to species loss than single functions. This susceptibility will increase with decreasing redundancy (Adopted from Gamfeldt et al, 2008).

图2 以气候、土壤和生物多样性的分段结构方程模型作为生态系统多功能性的预测因素。(a)和(b)分别代表2种假设: (a)表示植物物种丰富度驱动地下生物多样性(土壤生物多样性)时的情况; (b)表示地下生物多样性驱动植物物种丰富度时的情况。黑色实线箭头表示正向路径(P < 0.05), 灰色实线箭头表示负向路径(P < 0.05), 灰色虚线箭头表示不显著的路径(P > 0.05)。图中的路径系数为标准化的效应值。通过Shipley检验的d分离法评估了分段结构方程模型的拟合效果: 分别是费希尔C统计(如果P > 0.05, 那么没有路径丢失, 模型拟合效果好)和AIC标准。MAP, 年平均降水量; MAT, 年平均温度; SM, 土壤湿度; Plant SR, 植物物种丰富度(修改自Jing et al, 2015)。

Fig. 2 Structural equation models (SEM) of climate, soil and biodiversity as predictors of ecosystem multifuncitonality (EMF), represented by two hypotheses: (a) plant species richness drives belowground biodiversity (soil biodiversity), and conversely, (b) belowground biodiversity drives plant species richness. Solid black arrows represent positive paths (P < 0.05, piecewise SEM), solid gray arrows represent negative paths (P < 0.05, piecewise SEM), and dotted gray arrows represent non-significant paths (P > 0.05, piecewise SEM). We report the path coefficients as standardized effect sizes. Overall fit of piecewise SEM is evaluated using Shipley’s test of d-separation: Fisher’s C statistic (if P > 0.05, then no paths are missing and the model is a good fit) and Akaike Information Criterion (AIC). MAP: Mean annual precipitation; MAT: Mean annual temperature; SM: Soil moisture; Plant SR: Plant species richness (Adopted from Jing et al, 2015).

图3 全球变化背景下不同维度多样性对生态系统多功能性的影响。紫色虚线框内为现阶段研究内容, 红色虚线框内为未来研究方向。

Fig. 3 The impacts of multiple biodiversity dimensionality on ecosystem multifunctionality under global change. The purple dashed box indicates current research focus while the red dashed box represents proposed future directions.

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