生态学的多样性指数:功能与系统发育

生物多样性是生态学的核心问题。传统的多样性指数仅包含物种数和相对多度的信息,这类基于分类学的多样性指数并不能很好地帮助理解群落构建和生态系统功能。不同物种对群落构建和生态系统功能所起到的作用类型和贡献也不完全相同,且物种在生态过程中的作用和贡献往往与性状密切相关,因此功能多样性已经成为反映物种群落构建、干扰以及环境因素对群落影响的重要指标。同时,由于亲缘关系相近的物种往往具有相似的性状,系统发育多样性也可以作为功能多样性的一个替代。功能多样性和系统发育多样性各自具有优缺点,但二者均比分类多样性更能揭示群落和生态系统的构建、维持与功能。     

基于功能性状的生态系统服务研究框架

功能性状通过影响生态系统的属性和过程及其维持来影响生态系统服务。功能多样性-生态系统功能关系的研究有助于深入探讨生态系统服务形成机制, 也为生态系统服务研究提供了一个切入点。该文对目前的功能性状和生态系统服务研究框架进行了介绍, 回顾了功能多样性-生态系统功能关系的研究现状, 总结了目前功能性状在生态系统服务研究中的应用, 提出了基于功能性状的生态系统服务研究框架。在这个研究框架中, 首先选取对生态系统功能影响显著的非生物因子和功能多样性指数, 然后量化非生物因子和功能多样性与生态系统功能, 以及生态系统功能-生态系统服务之间的关系, 进而构建功能多样性与生态系统服务的数量关系。与此同时, 利用群落构建理论和物种共存机制分析功能多样性-生态系统功能变化的机制联系, 以研究生态系统服务形成和变化机制, 为生态系统服务管理决策提供科学依据。     

植物功能性状研究进展

植物功能性状是指植物体具有的与其定植、存活、生长和死亡紧密相关的一系列核心植物属性,且这些属性能够显著影响生态系统功能,并能够反映植被对环境变化的响应.越来越多的研究表明,相比大多数基于植物分类和数量的研究,植物功能性状在种群、群落和生态系统尺度上,都已成为解决重要生态学问题的可靠途径.本文回顾了植物功能性状研究的发展历程,总结了近10年来基于植物功能性状研究的前沿科学问题,包括功能性状的全球分布格局和内在关联,沿环境梯度的变化规律,功能多样性的定义及应用,与群落物种共存机制和群落动态变化的关系,与系统发育的关系,对生态系统功能的影响以及对各类干扰的影响和响应.尽管功能性状研究已经延伸到生态学领域的各个方面,有力推动了各个前沿科学问题的研究发展,仍然有很多值得关注和着重研究的方向.本文也对未来基于植物功能性状的研究,从性状测量和选取、研究方法以及研究方向上提出了展望,并指出,在当前全球气候变化背景下,功能性状也可应用于指导生物多样性保护和生态系统管理政策的制定.     

刈割、围封、放牧三种利用方式下草原生态系统的多功能性与植物物种多样性之间的关系

随着全球变化对生物多样性的影响不断加剧, 生物多样性与生态系统功能之间相互关系(BEF)的研究显得极为重要。过去的20多年, BEF的研究大多集中在对物种多样性与单一或少数生态系统功能之间关系的探讨, 但生态系统最为重要的价值是同时维持多种服务和功能的能力, 基于此, 该文首次在国内引入近年来不断完善的生态系统多功能性(multifunctionality)的概念, 并对目前主流的评价方法进行了改进, 从而对内蒙古三种利用方式(刈割、围封、放牧)下的草地群落进行了多功能性评价, 并探讨了多功能性与物种多样性之间的关系。结果显示本研究改进的方法和目前主流方法评价得出的多功能性指数在样方和样地尺度上都有很高的相关性(R² = 0.6956, p < 0.0001; R² = 0.9231, p < 0.0001), 表明该文作者改进后的方法是可靠的。重度放牧的草地群落物种多样性水平最低, 绝大多数土壤功能指标较差, 表现出退化特征; 7年的围封和刈割群落均有较高的物种多样性水平和改善的土壤功能指标; 三者的多功能性指数为刈割(0.2178) >围封(0.0704) >放牧(-0.8031)。植被样方主要沿水肥梯度分布; 多样性指数中, 均匀度指数(Pielou index)和丰富度指数(Margelf index)对多功能性的影响作用最大, 均为样方尺度(R² = 0.1871, p < 0.0001; R² = 0.1601, p < 0.0001)小于样地尺度(R² = 0.5921, p = 0.0093; R² = 0.7499, p = 0.0007), 有尺度依赖性; 多功能性在样方和样地尺度上均与物种均匀度呈线性正相关关系, 而与物种丰富度呈单峰曲线关系。该文研究结果表明, 相对于重度放牧和围封, 刈割更有利于维持该地区生态系统的多功能性; 物种丰富度适中且物种分布均匀的生态系统可能有更好的多功能性。     

荒漠植物性状权衡策略及功能多样性研究进展

荒漠植物是干旱区具有独特功能性状与资源权衡表征的地带性植物。植物功能性状及其多样性格局与资源权衡策略对群落结构优化和生态系统功能改善起着关键作用。该综述主要从荒漠植物组织、器官功能性状特征、功能性状权衡策略、功能多样性组分及测度3个方面梳理了荒漠植物性状权衡策略与功能多样性研究的进展脉络：1）荒漠植物独特的根、茎、叶功能性状特征揭露了植被对环境变化的响应以及对生态系统功能的影响，基于植物功能性状的研究有助于解决许多生态学的关键性问题；2）作为植物功能性状之间存在的最普遍的联系，权衡策略是经过自然筛选后形成的性状组合，关键性状已经被发掘并创造性的提出了"经济谱"概念。荒漠植物研究过程中，应分析其根、茎、叶的特征属性筛选关键性状，着眼于关键性状间及整株植物性状间的权衡策略；3）功能多样性是影响生态系统运行和发挥作用的生物多样性的重要组成部分，荒漠植物功能多样性能预测和指示群落中物种对于荒漠生态系统功能发挥和过程变化的影响。功能多样性的组分可以从不同角度反映群落的生态位占据状况和资源利用程度，指数的选择要体现在群落内部物种的功能特征之间的差异程度，同时要考虑这些物种自身在群落内的优势程度。本研究为未来荒漠植物功能性状及多样性研究梳理了一些新的研究方向和内容，期望为荒漠植物生理生态学研究的选题和发展提供一些新的思路。     

山西五鹿山森林群落木本植物功能多样性

通过选取群落中木本植物种子的扩散方式、传粉方式、植株高度和盖度等13个功能性状,计算出群落的6个功能多样性指数:功能性状距离、功能性状平均距离、功能体积、功能均匀度、功能分散指数和Rao二次熵指数,结合群落物种丰富度指数、Shannon-Wiener指数和物种均匀度指数对山西五鹿山森林群落木本植物功能多样性进行研究。结果表明:(1)功能性状距离、功能性状平均距离、功能体积与物种丰富度、Shannon-Wiener指数显著正相关;功能均匀度与Shannon-Wiener指数、物种均匀度指数显著正相关;功能分散指数、Rao二次熵指数与物种均匀度指数、Shannon-Wiener指数显著正相关;(2)功能多样性的差异很大程度上是由于物种差异所引起的;(3)6个功能多样性指数可分为三类:功能性状距离、功能性状平均距离、功能体积为功能丰富度指数;功能均匀度为功能均匀度指数;功能分散指数和Rao二次熵指数为功能离散度指数。该分类结果符合指数的计算方法和生态学意义,以及相互独立的标准。     

陆地植物群落物种多样性研究进展

生物多样性是当前生态学研究的热点之一,物种多样性层次是最直接、最易观察和最适合研究生物多样性的层次。总结了与群落动态、生境因子、取样尺度及生态系统相关的陆地植物物种多样性研究。同时,根据目前的趋势提出了多样性动态研究的发展动向。     

功能多样性及其研究方法

功能多样性是联系生物多样性和生态系统功能的关键性因素。植物群落内功能多样性的量化描述对于研究生物多样性响应环境变化及其对生物多样性-生态系统功能关系的影响至关重要。由于功能多样性的概念含混不清,使得对其范围的确定、定量描述及其对生态系统功能的作用的评价存在较大的难度。在查阅大量文献的基础上,以植物群落为例,介绍了功能多样性的定义、计算方法及研究中应注意的问题。在植物群落功能多样性指数的选择和测定过程中,对获取相关物种必要的功能特征信息、确定其权重、确定不同物种在群落中的相对丰富度、利用所得的功能多样性指数解释和预测生态系统过程的变化等问题,进行了研究。     

植物功能性状、功能多样性与生态系统功能: 进展与展望

植物功能性状与生态系统功能是生态学研究的一个重要领域和热点问题。开展植物功能性状与生态系统功能的研究不仅有助于人类更好地应对全球变化情景下生物多样性丧失的生态学后果,而且能为生态恢复实践提供理论基础。近二十年来,该领域的研究迅速发展,并取得了一系列的重要研究成果,增强了人们对植物功能性状-生态系统功能关系的认识和理解。本文首先明确了植物功能性状的概念, 评述了近年来植物功能性状-生态系统功能关系领域的重要研究结果, 尤其是植物功能性状多样性-生态系统功能关系研究现状; 提出了未来植物功能性状与生态系统功能关系研究中应加强植物地上和地下性状之间关系及其与生态系统功能、植物功能性状与生态系统多功能性、不同时空尺度上植物功能性状与生态系统功能, 以及全球变化和消费者的影响等方面。     

高寒草地植物物种多样性与功能多样性的关系

物种多样性与功能多样性的关系是生态学当前研究的热点问题之一,不同区域典型生态系统物种多样性和功能多样性的关系研究有利于生物多样性保护理论的全面发展。以青藏高原地区的主要草地生态系统—高寒草甸和高寒草原为研究对象,采用4个物种多样性指数(Patrick丰富度指数、Shannon-Weiner多样性指数、Pielou均匀度指数和Simpson优势度指数)和9个功能多样性指数(FAD功能性状距离指数、MFAD功能性状平均距离指数、基于样地的FDp和基于群落的FDc功能树状图指数、FRic功能体积指数、FEve功能均匀度指数、Rao功能离散度常二次熵指数、FDiv功能离散指数、FDis功能分散指数),分析了高寒草地植物物种多样性、功能多样性关系及其与初级生产力的关系,以期阐明3个科学问题:不同草地类型的高寒草地生态系统植物物种多样性和功能多样性有何差异?高寒草地生态系统的植物物种多样性和功能多样性有何关系?高寒草地生态系统物种多样性、功能多样性对生态系统功能的影响有何异同?研究结果表明:(1)与高寒草原相比,高寒草甸具有更高的物种多样性、功能丰富度和功能离散度;(2)高寒草甸中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc)和功能离散度指数(FDiv)的具有较强的相关性,最优拟合方程分别为幂函数和二次多项式函数;(3)高寒草原中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc、FRic)、Shannon指数和Simpson指数与FEve指数的相关性较强,最优拟合方程为二次多项式函数,Pielou指数与FEve指数的相关性较强,最优拟合方程为指数函数;(4)高寒草甸的初级生产力分别与物种丰富度指数Patrick、功能离散指数FDiv具有较强的相关性;而高寒草原的初级生产力与4个物种多样性指数间均具有较强的相关性,与功能离散指数FDiv具有较强的相关性,最佳拟合方程均为二次多项式函数。研究的总体结论为:物种多样性、功能多样性、二者之间的关系以及二者与生态系统服务功能(以初级生产力为例)之间的关系在高寒草甸和高寒草原群落中表现迥异,因此在研究青藏高原高寒草地的生态功能时,不能仅仅测度传统的物种多样性,还应测度与物种多样性、生态功能密切相关的功能多样性。     

Dominance determines fish community biomass in a temperate seagrass ecosystem

Biodiversity and ecosystem function are often correlated, but there are multiple hypotheses about the mechanisms underlying this relationship. Ecosystem functions such as primary or secondary production may be maximized by species richness, evenness in species abundances, or the presence or dominance of species with certain traits. Here, we combine surveys of natural fish communities (conducted in July and August 2016) with morphological trait data to examine relationships between biodiversity and ecosystem function (quantified as fish community biomass) across 14 subtidal eelgrass meadows in the Northeast Pacific (54°N, 130°W). We employ both taxonomic and functional trait measures of diversity to investigate whether ecosystem function is best predicted by species diversity (complementarity hypothesis) or by the presence or dominance of species with particular trait values (selection or dominance hypotheses). After controlling for environmental variation, we find that fish community biomass is maximized when taxonomic richness and functional evenness are low, and in communities dominated by species with particular trait values, specifically those associated with benthic habitats and prey capture. While previous work on fish communities has found that species richness is often positively correlated with ecosystem function, our results instead highlight the capacity for regionally prevalent and locally dominant species to drive ecosystem function in moderately diverse communities. We discuss these alternate links between community composition and ecosystem function and consider their divergent implications for ecosystem valuation and conservation prioritization.     

生态系统性状对宏生态研究的启示与挑战

功能性状在器官-物种-种群-群落-生态系统水平都具有其特定的适应或功能优化的意义,但目前对功能性状的测定和研究大都局限于器官或物种水平。然而,当前高速发展的宏生态新研究技术和方法(如遥感观测、通量观测、模型模拟)的研究对象都是在生态系统或区域尺度上,如何将传统功能性状与其相连结并服务于生态环境问题和全球变化问题是科学界的一大难题。为了解决传统性状与宏生态研究"尺度不统一"和"量纲不统一"的难题,研究人员最新发展了"生态系统性状(Ecosystem traits, ESTs)"概念体系,并从"理念-数据源-推导方法-应用"等多角度为后续研究提供了可借鉴案例。生态系统性状将传统性状研究从器官水平拓展到了群落和生态系统水平,以单位土地面积为基础构建了传统性状与宏生态研究(或地学研究)的桥梁,开启了性状研究从"器官到群落"、从"经典理论验证到宏观应用"的美好愿景,为多学科交叉提供了新思路。然而,它在方法学和数据源等方面还存在诸多问题与挑战;在此,我们呼吁相关专家从研究方法、概念体系和应用实践上赋予"生态系统性状"更强大的生命力,尤其从动物群落性状和微生物群落性状等角度。本文在深入解读先前生态性状概念体系、理论意义和潜在挑战的基础上,结合最新进展进行了补充,希望通过广泛讨论,完善生态系统性状概念体系,逐步形成"以性状为基础的生态系统生态学"新研究框架,切实推动宏生态研究和区域生态环境问题的解决。     

Trait structure and redundancy determine sensitivity to disturbance in marine fish communities

Trait diversity is believed to influence ecosystem dynamics through links between organismal traits and ecosystem processes. Theory predicts that key traits and high trait redundancy—large species richness and abundance supporting the same traits—can buffer communities against environmental disturbances. While experiments and data from simple ecological systems lend support, large‐scale evidence from diverse, natural systems under major disturbance is lacking. Here, using long‐term data from both temperate (English Channel) and tropical (Seychelles Islands) fishes, we show that sensitivity to disturbance depends on communities’ initial trait structure and initial trait redundancy. In both ecosystems, we found that increasing dominance by climatically vulnerable traits (e.g., small, fast‐growing pelagics/corallivores) rendered fish communities more sensitive to environmental change, while communities with higher trait redundancy were more resistant. To our knowledge, this is the first study demonstrating the influence of trait structure and redundancy on community sensitivity over large temporal and spatial scales in natural systems. Our results exemplify a consistent link between biological structure and community sensitivity that may be transferable across ecosystems and taxa and could help anticipate future disturbance impacts on biodiversity and ecosystem functioning.     

Revisiting the Holy Grail: using plant functional traits to understand ecological processes

One of ecology's grand challenges is developing general rules to explain and predict highly complex systems. Understanding and predicting ecological processes from species' traits has been considered a ‘Holy Grail’ in ecology. Plant functional traits are increasingly being used to develop mechanistic models that can predict how ecological communities will respond to abiotic and biotic perturbations and how species will affect ecosystem function and services in a rapidly changing world; however, significant challenges remain. In this review, we highlight recent work and outstanding questions in three areas: (i) selecting relevant traits; (ii) describing intraspecific trait variation and incorporating this variation into models; and (iii) scaling trait data to community‐ and ecosystem‐level processes. Over the past decade, there have been significant advances in the characterization of plant strategies based on traits and trait relationships, and the integration of traits into multivariate indices and models of community and ecosystem function. However, the utility of trait‐based approaches in ecology will benefit from efforts that demonstrate how these traits and indices influence organismal, community, and ecosystem processes across vegetation types, which may be achieved through meta‐analysis and enhancement of trait databases. Additionally, intraspecific trait variation and species interactions need to be incorporated into predictive models using tools such as Bayesian hierarchical modelling. Finally, existing models linking traits to community and ecosystem processes need to be empirically tested for their applicability to be realized.     

Functional traits explain ecosystem function through opposing mechanisms

The ability to explain why multispecies assemblages produce greater biomass compared to monocultures, has been a central goal in the quest to understand biodiversity effects on ecosystem function. Species contributions to ecosystem function can be driven by two processes: niche complementarity and a selection effect that is influenced by fitness (competitive) differences, and both can be approximated with measures of species’ traits. It has been hypothesised that fitness differences are associated with few, singular traits while complementarity requires multidimensional trait measures. Here, using experimental data from plant assemblages, I show that the selection effect was strongest when trait dissimilarity was low, while complementarity was greatest with high trait dissimilarity. Selection effects were best explained by a single trait, plant height. Complementarity was correlated with dissimilarity across multiple traits, representing above and below ground processes. By identifying the relevant traits linked to ecosystem function, we obtain the ability to predict combinations of species that will maximise ecosystem function.     

What does species richness tell us about functional trait diversity? Predictions and evidence for responses of species and functional trait diversity to land‐use change

In the conservation literature on land‐use change, it is often assumed that land‐use intensification drives species loss, driving a loss of functional trait diversity and ecosystem function. Modern research, however, does not support this cascade of loss for all natural systems. In this paper we explore the errors in this assumption and present a conceptual model taking a more mechanistic approach to the species–functional trait association in a context of land‐use change. We provide empirical support for our model's predictions demonstrating that the association of species and functional trait diversity follows various trajectories in response to land‐use change. The central premise of our model is that land‐use change impacts upon processes of community assembly, not species per se. From the model, it is clear that community context (i.e. type of disturbance, species pool size) will affect the response trajectory of the relationship between species and functional trait diversity in communities undergoing land‐use change. The maintenance of ecosystem function and of species diversity in the face of increasing land‐use change are complementary goals. The use of a more ecologically realistic model of responses of species and functional traits will improve our ability to make wise management decisions to achieve both aims in specific at‐risk systems.     

Functional regularity: a neglected aspect of functional diversity

Functional diversity has been identified as a key to understanding ecosystem and community functioning. However, due to the lack of a sound definition its nature and measurement are still poorly understood. In the same way that species diversity can be split into species richness and species evenness, so functional diversity can be split into functional richness (i.e. the amount of functional trait/character/attribute space filled) and functional evenness (i.e. the evenness of abundance distribution in functional trait space). We propose a functional regularity index (FRO) as a measure of functional evenness for situations where species are represented only by a single functional trait value (e.g. mean, median or mode), and species abundances are known. This new index is based on the Bulla O index of species evenness. When dealing with functional types or categorical functional traits, the Bulla O or any other accepted species evenness index may be used directly to measure functional evenness. The advantage of FRO is that it supplies a measure of functional evenness for continuous trait data. The FRO index presented in this paper fulfils all the a priori criteria required. We demonstrate with two example datasets that a range of FRO values may be obtained for both plant and animal communities. Moreover, FRO was strongly related to ecosystem function as seen in photosynthetic biomass in plant communities, and was able to differentiate sampling stations in a lagoon based on the functional traits of fish. Thus, the FRO index is potentially a highly useful tool for measuring functional diversity in a variety of ecological situations.     

Which components of plant diversity are most correlated with ecosystem properties? A case study in a restored wetland in northern China

The relationship between plant diversity and ecosystem services is a controversial topic in ecology that may be due, at least in part, to the variety of methods used to define and quantify diversity. This study examined the relationship between plant diversity and 11 ecosystem properties of a restored wetland in northern China by considering four primary components of diversity (dominance, richness, evenness, and divergence). Each diversity component was expressed by eight taxonomic and functional diversity indices respectively. Results showed that trait-based functional diversity had a stronger correlation with ecosystem processes than non-trait taxonomic diversity did. Among the four components of diversity, dominance (in terms of mean trait value index) was the best in explaining the variation in ecosystem processing. Richness and divergence also had significant correlations with ecosystem properties in some instances. By contrast, evenness had no significant correlation with most of the studied ecosystem properties. Our results indicated that wetland ecosystem properties are significantly related to certain traits of the dominant species. Thus, the dominant species and functional traits should be considered before the number of species in managing diversity and enhancing certain ecosystem functions of wetlands, especially in the case of conservation.