应用中性理论分析局域群落中的物种多样性及稳定性

如何解释群落中物种的丰富与稀少,并对物种多样性和群落稳定性进行合理的量化评价是群落生态学研究中的一个热点问题。20世纪中期,MacArthur将影响自然群落稳定性的因素归结为物种数量多少以及物种间相互作用的大小,20世纪末Doak等学者提出群落的容纳能力和物种间的维持机制是决定群落稳定性的关键因素。同时对群落结构及物种间维持机制的研究也有了新的突破,Hubbell提出"生物多样性与生物地理学统一的中性理论(Unified Neutral Theory of Biodiversity and Biogeography)"为群落生态学研究提供了新的思路和方法。从群落中性理论的基本假设出发,对Hubbell中性理论中局域群落的物种多度动态模型进行分析,归纳得出群落中性理论中物种多样性与群落稳定性之间的量化关系。封闭的局域群落中,出现物种灭绝或单物种独占的时间与群落大小及物种相对多度成正比,物种多样性程度的增加可延长物种灭绝或独占的时间;开放的局域群落中,物种多度期望值与局域群落大小、物种在集合群落中的物种相对多度成正比,周围群落中物种的灭绝会引起局域群落中相应物种的灭绝,最终导致整个生态群落物种多样性的降低;群落中物种多度的方差与局域群落大小、迁移率、物种在集合群落中的物种相对多度相关,局域群落物种多度的波动幅度随着群落间生态隔离的减弱或物种多样性程度的增加而减小。由此,集合群落物种多样性是影响局域群落物种多样性的重要因素,生态隔离程度的减弱及物种多样性的增加都将增强群落的稳定性。     

太白山锐齿栎林物种-多度分布格局

以太白山1.5 hm²的锐齿栎原始林和次生林样地中环境因子和胸径≥1 cm的木本植物调查数据为基础,采用统计模型(对数正态模型)、生态位模型(Zipf模型、断棍模型、生态位优先模型)和中性模型,拟合了锐齿栎群落的物种多度分布。结果表明: 太白山锐齿栎林物种多度分布格局受到生境异质性的影响。其中,地形因子对原始林物种分布影响较大,在凹凸度较大的生境中,物种分布同时受到中性过程和生态位过程的影响,但中性过程发挥的作用较小;而在凹凸度较小的生境中,中性模型被拒绝,物种的多度分布符合生态位理论的假设。在群落坡度大的区域,群落中生态位过程和中性过程同等重要;而在坡度较小的平缓区域,生态位分化对群落物种分布的影响较大。在次生林中,影响物种分布的因素主要是土壤养分。在次生林土壤速效磷含量高的生境中,生态位过程是影响群落物种分布的主要生态学过程;而在土壤速效磷含量低的生境中,中性过程和生态位过程在群落物种分布中同时存在。太白山锐齿栎林物种多度分布格局存在明显的尺度效应。原始林在20 m×20 m尺度上,生态位模型和中性模型都能预测物种多度分布,而在40 m×40 m和70 m×70 m尺度上,生态位过程可解释物种多度分布格局。在次生林样地20 m×20 m、40 m×40 m、70 m×70 m尺度上,生态位过程和中性过程共同作用于物种的分布,但是生态位过程更为重要。可见,除了尺度和生境异质性外,原始林与受干扰的次生林中的物种多度分布也存在明显的差异。     

群落构建的中性理论和生态位理论

物种共存和生物多样性维持一直是生态学研究的中心论题。基于物种生态位分化的群落构建理论已经发展了近一个世纪, 但我们对群落构建和生物多样性维持的机理仍然不清楚。近年来, 群落中性理论以其简约性和预测能力成为群落生态学研究的焦点, 但由于其“物种在生态功能上等价”的假设与大量研究结果相悖, 同时对自然群落结构的准确预测也只限于少数的生态系统, 因而饱受质疑。如今, 越来越多的生态学家认为群落构建的生态位理论与中性理论之争的最终归宿应该是二者的整合。 在本文中, 我们在简要回顾生态位理论和群落中性理论发展的基础上, 分析二者之间的主要分歧和互补性, 试图梳理二者整合的途径。我们认为, 尽管中性理论的发展极大地丰富了群落构建理论, 但二者的整合尚处于初级阶段; 群落构建零模型假说、中性—生态位连续体假说、随机生态位假说等都不失为有价值的尝试, 今后需要在其他类型的生态系统中进行实验验证, 以更好地理解确定性过程和随机过程在决定群落构建和生物多样性维持中的作用。     

中国湿地物种多样性与生境面积关系及其生态学机理的模拟研究

生物群落中的物种多样性是群落生态学研究的一个基本问题。大量的实验研究表明物种多样性和生境面积的常数次幂成比例,这说明它们的对数数量级呈线性正相关。我们研究了中国境内15块湿地的高等植物和32块湿地鸟类多样性,发现它们分别和湿地面积在对数尺度上呈线性正相关．进一步支持了种数与面积的幂指数关系。我们还借助计算机模拟系统地讨论了产生这种简单规律的生态学机理,包括中性理论、集合种群动态和物种分布的自相似性。中性理论假设了群落中物种的个体之间只有竞争关系,忽略了其它的种间关系。集合种群动态理论考虑的是由多个亚群落构成的集合群落,在研究种数和面积关系时也忽略了种间关系,所以也是中性的。尽管物种分布的自相似性可导致这种面积幂指数关系,但在自然界中自相似性也可能不成立。     

天童常绿阔叶林中常绿与落叶物种的物种多度分布格局

物种多度分布是对群落内不同物种多度情况的数量描述, 作为理解群落性质的基石, 其形成机制受到广泛关注。常绿与落叶物种是两类有着不同物候性状与生长策略的物种集合, 它们普遍共存于常绿阔叶林中。在天童20 ha常绿阔叶林动态监测样地内, 虽然常绿物种在物种多度和胸高断面积等指标上占有绝对优势, 但其在物种丰富度上却不及落叶物种。分析两者在常绿阔叶林中的物种多度分布特征, 能够为理解常绿阔叶林内物种多样性的维持机制提供一个全新的视角。为此, 我们基于天童样地的植被调查数据, 一方面利用累积经验分布函数对两类生活型植物的物种多度分布进行描述, 使用Kolmogorov-Smirnov检验(K-S检验)判断其差异性; 另一方面, 采用纯统计模型、生态位模型和中性理论模型对二者的物种多度分布曲线进行拟合, 并基于K-S检验的结果以及AIC值进行最优模型的筛选。结果显示: (1)常绿与落叶物种的物种多度分布曲线间并无显著差异。(2)在选用的3类模型中, 中性理论模型对于两类物种多度分布曲线的拟合效果都最好, 而生态位模型的拟合效果则一般。从上述结果可以看出, 尽管常绿与落叶物种在物种数量和多度等方面均存在差异, 但它们却有着近似的物种多度分布格局以及相近的多样性维持机制。然而, 鉴于模型拟合的结果只能作为理解群落多样性构建机制的必要非充分条件, 故而只能初步判定中性过程对于常绿与落叶物种的物种多样性格局影响更大, 却不能排除或衡量诸如生态位分化等其他过程在两类生活型多样性格局形成中的贡献。     

物种多度格局研究进展

物种多度格局研究始于20世纪30年代,是种群生态学和群落生态学研究的起点。物种多度格局研究主要在两个水平上进行：1）初期研究主要集中于群落水平,希望在不同群落之间发现一个共同的整体格局来描述群落的组织结构。常用模型包括几何级数、对数级数、对数正态和断棍模型,不同模型代表了不同的生态学过程。2）目前转向重视物种水平,并以物种多度的区域分布规律及其生态学机制研究为主。物种分布区多度关系有正相关、无相关和负相关3种形式。局部多度高的物种一般趋于广布,而局部多度低的物种趋于受限分布。物种多度区域分布的生态位模型预测为单峰型,还经常会出现“热点地区”;而异质种群模型预测为双峰型。物种多度的区域分布主要由环境资源特性、物种生态位和扩散过程等因素决定。3）物种多度格局的时间变化与空间变异类似,代表了这些生态学过程的时间异质性。4）物种多度格局的尺度变化经常表现出自相似性,但该规律并非一直存在,因为生物多样性由不同尺度上的不同生态学过程决定。5）多度（稀有度）是物种保护的基本依据,而群落多度模型能够指示生态学和干扰过程变化对群落结构的影响。物种多度格局的模型手段仍需改进,机制研究尚不系统,应用研究亟待扩展,对于物种多度格局的深入理解将为揭示生物多样性分布机制和有效保护提供帮助。     

生态位分化与森林群落物种多样性维持研究展望

一直以来,生态学家和进化生物学家对森林群落物种多样格局及其形成机制持有不同的观点。虽然Robert Ricklefs将进化和生态过程整合的观点已经被群落生态学家广泛接受,但是区域物种进化历史以及局域群落微进化过程是否能够影响群落生态学过程以及这些过程如何影响群落结构和动态还有待商榷。经典的生态位理论同时强调了种间和种内生态位分化对群落多样性维持的影响。但是生态学家普遍认为种间差异足以代表群落内个体间的相互作用关系,并且由于进化过程导致的种内分化往往涉及较长的时间尺度,因此,虽然种内差异是自然选择的重要材料,物种对环境的适应性进化过程所导致的种内分化对群落构建的影响往往被生态学家所忽视。为此,通过回顾种间和个体生态位分化的研究历史,对两类研究分别进行简要阐述,强调在今后的群落生态学研究中需要考虑个体分化对局域群落构建的影响。     

亚热带喀斯特地区不同地形植物群落物种多度分布格局

为探究亚热带喀斯特地区不同地形下植物群落物种多度分布格局,揭示不同地形下群落的物种多度格局形成的作用机制,丰富该地区植物群落构建理论,该文以贵州茂兰喀斯特地区山脊、槽谷、鞍部、洼地4种典型地形下植物群落的乔木层与灌木层为对象,统计物种多度,采用累计经验分布曲线(ECDF)表征多度分布格局,采取Wilcoxon秩和检验探究不同地形之间物种多度的差异性。采用不同生态学模型进行多度拟合,利用Kolmogorov-Smirnov(K-S)检验与赤池信息量准则(AIC)检验模型接受与拟合优度。结果表明:(1)不同地形下植物群落的个体数量与物种数存在差异,鞍部个体数最多,洼地的物种数最多,山脊的个体数、物种数均最少。(2)不同地形下植物群落的乔木层物种多度格局无显著差异,灌木层之间出现显著差异,山脊与鞍部、洼部,鞍部与槽谷、洼部都存在显著差异。(3)不同地形下乔木层物种多度对中性模型接受较好,其中山脊拟合最优,对生态位模型接受较差,仅山脊与鞍部通过两种生态位模型,拟合优度不及中性模型。灌木层对中性模型接受也较好,鞍部拟合最优,对生态位模型接受较差,仅洼地通过断棍模型。整体而言,乔木层比灌木层能更好地接受两种生态学模型,可能乔木层物种多度格局有更明显生态过程的印记,但不同地形下灌木层拟合优度差异更大,可能与灌木层物种对环境变化更剧烈有关。不同地形会引起群落构建不同程度的生态学过程,物种多度分布格局会逐渐适应地形。     

长白山不同演替阶段针阔混交林群落物种多度分布格局

为解释长白山温带森林群落构建和物种多度格局的形成过程, 该文以不同演替阶段的针阔混交林监测样地数据为基础, 采用中性理论模型、生物统计模型(对数正态分布模型)和生态位模型(Zifp模型、分割线段模型、生态位优先模型)拟合森林群落物种多度分布, 并用χ ²检验、Kolmogorov-Smirnov (K-S)检验和赤池信息准则(AIC)选择最佳拟合模型。结果显示: 中性模型能很好地预测长白山温带森林不同演替阶段植物群落的物种多度分布。在10 m × 10 m尺度上, 5种模型均可被χ ²检验和K-S检验接受, 但中性模型拟合效果不如对数正态分布模型、Zifp模型、分割线段模型和生态位优先模型, 表明小尺度上中性过程和生态位过程均能解释群落物种多度分布, 但生态位过程的解释能力相对较大。而在中大尺度上(30 m × 30 m、60 m × 60 m和90 m × 90 m), 中性模型为最优拟合模型, 并且随着研究尺度增加, 生态位模型和生物统计模型逐渐被χ ²检验拒绝, 表明中性过程在长白山针阔混交林群落物种多度分布格局形成中的作用随着研究尺度增加而逐渐增大。该文证实了中性过程在长白山温带针阔混交林群落结构形成中具有重要作用, 但未否认生态位机制在群落构建中的贡献。因此, 温带森林群落构建过程中中性理论和生态位理论并非相互矛盾, 而是相互融合的。在研究森林群落物种多度分布时, 应重视取样尺度和演替阶段的影响, 并采用多种模型进行拟合。     

甘肃祁连山脉雪豹及其同域分布大型食肉动物时间生态位关系

<正>群落内多物种如何共存是群落生态学和生物多样性研究的核心内容之一。经典物种共存理论强调物种之间的生态位分化,侧重于物种对环境的需求,Hutchinson (1957)提出超体积生态位概念,认为物种适合度是由多个因素共同决定,即物种只有在满足其生态位需求的多维空间,     

Neutral theory in community ecology

One of the central goals of community ecology is to understand the forces that maintain species diversity within communities. The traditional niche-assembly theory asserts that species live together in a community only when they differ from one another in resource uses. But this theory has some difficulties in explaining the diversity often observed in specie-rich communities such as tropical forests. As an alternative to the niche theory, Hubbell and other ecologists introduced a neutral model. Hubbell argues that the number of species in a community is controlled by species extinction and immigration or speciation of new species. Assuming that all individuals of all species in a trophically similar com-munity are ecologically equivalent, Hubbell's neutral theory predicts two important statistical distributions. One is the asymptotic log-series distribution for the metacommunities under point mutation speciation, and the other is the zero-sum multinomial distribution for both local communities under dispersal limitation and metacommunities under random fission speciation. Unlike the niche-assembly theory, the neutral theory takes similarity in species and individuals as a starting point for investigating species diversity. Based on the fundamental processes of birth, death, dispersal and spe-ciation, the neutral theory provided the first mechanistic explanation of species abundance distribution commonly observed in natural communities. Since the publication of the neutral theory, there has been much discussion about it, pro and con. In this paper, we summarize recent progress in the assumption, prediction and speciation mode of the neutral theory, including progress in the theory itself, tests about the assumption of the theory, prediction and speciation mode at the metacommunity level. We also suggest that the most important task in the future is to bridge the niche-assembly theory and the neutral theory, and to add species differences to the neutral theory and more stochasticity to the niche theory.     

Neutral theory in community ecology

One of the central goals of community ecology is to understand the forces that maintain species diversity within communities. The traditional niche-assembly theory asserts that species live together in a community only when they differ from one another in resource uses. But this theory has some difficulties in explaining the diversity often observed in specie-rich communities such as tropical forests. As an alternative to the niche theory, Hubbell and other ecologists introduced a neutral model. Hubbell argues that the number of species in a community is controlled by species extinction and immigration or speciation of new species. Assuming that all individuals of all species in a trophically similar community are ecologically equivalent, Hubbell’s neutral theory predicts two important statistical distributions. One is the asymptotic log-series distribution for the metacommunities under point mutation speciation, and the other is the zero-sum multinomial distribution for both local communities under dispersal limitation and metacommunities under random fission speciation. Unlike the niche-assembly theory, the neutral theory takes similarity in species and individuals as a starting point for investigating species diversity. Based on the fundamental processes of birth, death, dispersal and speciation, the neutral theory provided the first mechanistic explanation of species abundance distribution commonly observed in natural communities. Since the publication of the neutral theory, there has been much discussion about it, pro and con. In this paper, we summarize recent progress in the assumption, prediction and speciation mode of the neutral theory, including progress in the theory itself, tests about the assumption of the theory, prediction and speciation mode at the metacommunity level. We also suggest that the most important task in the future is to bridge the niche-assembly theory and the neutral theory, and to add species differences to the neutral theory and more stochasticity to the niche theory. __________ Translated from Journal of Plant Ecology, 2006, 30(5): 868–877 [译自:植物生态学报]     

Departures from neutrality induced by niche and relative fitness differences

Breaking the core assumption of ecological equivalence in Hubbell’s “neutral theory of biodiversity” requires a theory of species differences. In one framework for characterizing differences between competing species, non-neutral interactions are said to involve both niche differences, which promote stable coexistence, and relative fitness differences, which promote competitive exclusion. We include both in a stochastic community model in order to determine if relative fitness differences compensate for changes in community structure and dynamics induced by niche differences, possibly explaining neutral theory’s apparent success. We show that species abundance distributions are sensitive to both niche and relative fitness differences, but that certain combinations of differences result in abundance distributions that are indistinguishable from the neutral case. In contrast, the distribution of species’ lifetimes, or the time between speciation and extinction, differs under all combinations of niche and relative fitness differences. The results from our model experiment are inconsistent with the hypothesis of “emergent neutrality” and support instead a hypothesis that relative fitness differences counteract effects of niche differences on distributions of abundance. However, an even more developed theory of interspecific variation appears necessary to explain the diversity and structure of non-neutral communities.     

Dispersal, edaphic fidelity and speciation in species-rich Western Australian shrublands: evaluating a neutral model of biodiversity

Over evolutionary time, the number of species in a community reflects the balance between the rate of speciation and the rate of extinction. Over shorter time‐scales local species richness is also affected by how often species move into and out of the local community. These processes are at the heart of Hubbell's ‘unified neutral theory of biodiversity’ ( Hubbell 2001 ). Hubbell's spatially implicit, dispersal‐limited neutral model is the most widely used of the many implementations of neutral theory and it provides an estimate of the rate of speciation in a metacommunity (if metacommunity size is known) and the rate at which species migrate into the local community from the wider metacommunity. Recently, this neutral model has been used to compare rates of speciation and migration in the species‐rich fynbos of South Africa and in neotropical forests. Here we use new analytical methods for estimating the neutral model's parameters to infer speciation and dispersal rates for three sites in species‐rich sclerophyll shrublands (equivalent to fynbos) in Western Australia (WA). Our estimates suggest that WA shrublands are intermediate between fynbos and tropical rainforest in terms of speciation and dispersal. Although a weak test, the model predicts species abundance distributions and species accumulation curves similar to those observed at the three sites. The neutral model's predictions also remain plausible when confronted with independent data describing: (1) known edaphic relationships between sites, (2) estimates of metacommunity species richness and (3) rates of speciation among resprouters and nonsprouters. Two of the site pairs, however, show species turnovers significantly different from those predicted by the spatially implicit form of the neutral model that we use. This suggests that non‐neutral processes, in this case probably edaphic specialisation, are important in the WA shrubland metacommunity. The neutral model predicts similar rates of speciation in resprouter and sprouter taxa, a finding supported by recent molecular phylogenies. Finally, when converted into temporally scaled speciation rates and species longevities, the estimates produced by the neutral model seem implausible. The apparent departure from neutrality in the turnover of species between some sites and the implausible temporal dynamics may be due to the particular model chosen and does not reduce the significance of our other results, which confirm that local dispersal limitation, coupled with broader scale edaphic fidelity, combine to structure this biodiverse metacommunity.     

Community differentiation on landscapes: drift, migration and speciation

Theories of the differentiation of ecological communities on landscapes have typically not considered evolutionary dynamics. Here we analytically study the expected differentiation among local communities in a large metacommunity, undergoing speciation, ecological drift and intercommunity dispersal, in the context of neutral theory. We demonstrate that heterogeneity in species diversity and abundance arises among communities when local communities are small and intercommunity migration is infrequent. We propose a new measure to describe community differentiation, defined as the average correlation or the average probability (C_st) that two randomly sampled individuals of the same species within local communities are from the same ancestor. The effects of driving forces (migration, mutation, and ecological drift) are incorporated into the two-level hierarchical community structure in a finite island model of neutral communities. Community differentiation can increase the effective metacommunity size or the Hubbell's fundamental species diversity in the metacommunity by a factor (1−C_st)⁻¹. Significant community differentiation arises when C_st≠0. Intercommunity migration promotes species diversity in local communities but reduce species diversity in the metacommunity. In either the finite or infinite island case, one can estimate the number of intercommunity migrants by using multiple local community datasets when the speciation is negligible in the neutral local communities, or by using the metacommunity dataset when the speciation is included in the local neutral communities. These results highlight the significance of the evolutionary mechanisms in generating heterogeneous communities in the absence of complicated ecological processes on large landscapes.     

Metacommunity speciation models and their implications for diversification theory

The emergence of new frameworks combining evolutionary and ecological dynamics in communities opens new perspectives on the study of speciation. By acknowledging the relative contribution of local and regional dynamics in shaping the complexity of ecological communities, metacommunity theory sheds a new light on the mechanisms underlying the emergence of species. Three integrative frameworks have been proposed, involving neutral dynamics, niche theory, and life history trade‐offs respectively. Here, we review these frameworks of metacommunity theory to emphasise that: (1) studies on speciation and community ecology have converged towards similar general principles by acknowledging the central role of dispersal in metacommunities dynamics, (2) considering the conditions of emergence and maintenance of new species in communities has given rise to new models of speciation embedded in the metacommunity theory, (3) studies of diversification have shifted from relating phylogenetic patterns to landscapes spatial and ecological characteristics towards integrative approaches that explicitly consider speciation in a mechanistic ecological framework. We highlight several challenges, in particular the need for a better integration of the eco‐evolutionary consequences of dispersal and the need to increase our understanding on the relative rates of evolutionary and ecological changes in communities.     

The zero-sum assumption in neutral biodiversity theory

The neutral theory of biodiversity as put forward by Hubbell in his 2001 monograph has received much criticism for its unrealistic simplifying assumptions. These are the assumptions of functional equivalence among different species (neutrality), the assumption of point mutation speciation, and the assumption that resources are continuously saturated, such that constant resource availability implies constant community size (zero-sum assumption). Here we focus on the zero-sum assumption. We present a general theory for calculating the probability of observing a particular species-abundance distribution (sampling formula) and show that zero-sum and non-zero-sum formulations of neutral theory have exactly the same sampling formula when the community is in equilibrium. Moreover, for the non-zero-sum community the sampling formula has this same form, even out of equilibrium. Therefore, the term "zero-sum multinomial (ZSM)" to describe species abundance patterns, as coined by Hubbell [2001. The Unified Neutral Theory of Biodiversity and Biogeography, Princeton University Press, Princeton, NJ], is not really appropriate, as it also applies to non-zero-sum communities. Instead we propose the term "dispersal-limited multinomial (DLM)", thus making explicit one of the most important contributions of neutral community theory, the emphasis on dispersal limitation as a dominant factor in determining species abundances.     

Beta diversity in spatially implicit neutral models: a new way to assess species migration

The spatially implicit neutral model (SINM) of S. P. Hubbell predicts species' abundance distributions at two levels: local communities where extinction balances immigration, characterized by the immigration number I, and the metacommunity, a source pool of migrants where speciation balances extinction. Previously, a plot's I was estimated from its species abundance distribution. Here, we relate neutral theory to the additive partitioning of species diversity and calculate the immigration rate into different plots from the variation in species composition among them. We revisit the G(ST) statistic of population genetics to introduce a new version, G(ST)(k), conditional on each community sample k. We derive an analytical expectation of G(ST)(k) as a function of the local immigration number, I(k), under a generalized version of the SINM, which allows the pool of migrants to deviate from the large-scale speciation-extinction balance. Simulations and field data suggest that G(ST)(k) provides reasonable estimates of immigration numbers, which were compared with the results from alternative likelihood-based estimations.     

Do fitness-equalizing tradeoffs lead to neutral communities?

Neutral theory in ecology is aimed at describing communities where species coexist due to similarities rather than the classically posited niche differences. It assumes that all individuals, regardless of species identity, are demographically equivalent. However, Hubbell suggested that neutral theory may describe even niche communities because tradeoffs equalize fitness across species which differ in their traits. In fact, tradeoffs can involve stabilization as well as fitness equalization, and stabilization involves different dynamics and can lead to different community patterns than neutral theory. Yet the important question remains if neutral theory provides a robust picture of all fitness-equalized communities, of which communities with demographic equivalence are one special case. Here, I examine Hubbell’s suggestion for a purely fitness-equalizing interspecific birth–death tradeoff, expanding neutral theory to a theory describing this broader class of fitness-equalized communities. In particular, I use a flexible framework allowing examination of the influence of speciation dynamics. I find that the scaling of speciation rates with birth and death rates, which is poorly known, has large impacts on community structure. In most cases, the departure from the predictions of current neutral models is substantial. This work suggests that demographic and speciation complexities present a challenge to the future development and use of neutral theory in ecology as null model. The framework presented here will provide a starting point for meeting that challenge, and may also be useful in the development of stochastic niche models with speciation dynamics.