局域与区域过程对生物多样性的影响: 群落构建的饱和与开放

EFFECTS OF LOCAL AND REGIONAL PROCESSES ON BIOLOGICAL DIVERSITY: SATURATION AND OPENNESS OF COMMUNITY ASSEMBLY

  • 摘要: 自MacArthur和Wilson建立岛屿生物地理学的平衡模型以来, 以生物多样性为核心的群落生态学进入了范式转变的阶段, 从更为广阔的视角重新审视群落构建这一决定生物多样性的核心机制。与经典群落生态学不同, 新范式的群落生态学将局域群落视为一个开放的单元, 认为区域群落和集合群落内部的物种扩散与迁移是平衡局域群落中物种灭绝的关键力量。局域群落的物种组成与多样性是局域过程与区域过程共同作用的产物, 局域过程包括生物过滤和非生物过滤, 而区域过程包含了影响物种形成的地理、历史和进化过程。局域过程与区域过程在局域群落构建中的相对重要性具有时空依赖性。文章回顾了半个世纪以来群落生态学研究范式的变化, 从群落构建、物种库与群落饱和性等概念入手, 系统地介绍了基于局域-区域物种丰富度关系分析区域与局域过程相对重要性的方法及其局限性, 并介绍了克服这些局限性的辅助分析方法。物种库是生物地理学的基础概念, 物种库的物种数量与组成记录了地理与区域尺度上历史上发生的生态与进化过程, 物种库为表述区域过程的作用提供了可定量化的变量。局域群落是区域物种库所有物种的子集, 局域群落构建是这一子集形成的过程与机制。局域-区域物种丰富度关系是用于比较群落构建中区域与局域过程的相对重要性的直观方法, 这种关系的非线性反映了群落的饱和性。由于这种关系受到空间尺度、环境条件和群落中物种特性的影响, 在判读群落是否饱和时需要结合其他理论与经验方法。对于物种均处于同一营养级的群落而言, 当局域尺度足够小时, 容易检测到群落的饱和性。而对于包括多个营养级的群落, 局域-区域物种丰富度关系的解读需要考虑跨营养级的相互作用。环境选择与种间相互作用都是通过性状实现的, 实际观测的局域群落与随机群落的功能多样性比较可以辅助判断局域过程的性质与强度。准确定义物种库是理解局域群落构建与生物多样性维持机制的前提, 同时应考虑物种库的大小与功能属性, 这将促进我们对群落结构与生物多样性格局的理解与认识。

     

    Abstract: The publication of the equilibrium model of island biogeography by MacArthur and Wilson (1963) initiated a paradigm shift in community ecology that focuses on biodiversity. Community assembly as the core mechanism of determining biodiversity has been re-examined from a larger-scale perspective by linking to biogeography. Being different from classical community ecology, the new paradigm-based community ecology regards local communities as an open unit, and species dispersal and migration within regional- and meta- communities acting as the key forces to balance local species extinction. The species composition and diversity of local communities result from the interaction between local and regional processes. Regional processes include geographical, historical, and evolutionary factors that affect speciation and diversification of species. The relative importance of local versus regional processes in community assembly is temporal and spatial scale dependent. This review first briefly retrospect the shift in community ecology paradigms over the past half century, and then systematically introduces the concepts of community assembly, species pools, and community saturation, followed by methods for analyzing the relative importance of regional and local processes based on local-regional relationships for species richness. We also discuss potential issues and introduce further analytical methods for these issues. Species pools are a basic concept of biogeography. The number and composition of species in a species pool record the ecological and evolutionary processes that have occurred historically in at geographical or regional scales. The species pool concept allows us to quantitatively describe the role of regional processes. A local community is a subset of all species in a regional species pool, and local community assembly refers to the processes and mechanisms of the formation of this subset. The local-regional relationship for species richness provides an empirical approach to determine the relative importance of regional and local processes in community assembly. As the curvilinearity of this relationship spatial scale, environmental conditions, and species characteristics, additional theoretical and empirical methods are needed to interpret the community saturation. When the local scale is small enough, community saturation in species at the same trophic level can be detected with greater sensitively. For communities that consist of species at multiple trophic levels, the interactions between different trophic levels must be carefully considered to interpret the local-regional relationships for species richness. Environmental selection and interspecific interactions play on traits rather than species. The comparison of functional diversity between observed and random communities can reveal the type and intensity of local processes. We emphasize that, to reveal community assembly and the maintenance of biodiversity, it is necessary to accurately define the species pool and incorporate both the size and functional attributes.

     

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