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。     

甘南高寒草甸退化对植物功能群多样性与物种多度分布的影响

为分析甘南高寒草甸植物功能群多样性及其物种多度分布对退化的响应,探讨其多样性的形成与维持机制,采用样地调查法收集数据,并使用物种多度模型对其进行拟合分析。结果表明:随着退化程度的加深,植被优势种以禾草科和莎草科植物为主,逐渐变为杂草功能群植物为主,杂草功能群在群落多样性分布中起着主导作用;全部物种多度分布随着退化程度的加深发生变化,其中,无退化草甸的最优拟合模型是VOLKOV,轻度退化草甸的最优拟合模型是GEO,中度退化草甸和重度退化草甸的最优拟合模型为BRO,资源分配模式由随机分配转向固定分配的分配模式;禾草功能群的最优拟合模型以生态位模型为主,资源分配方式由固定分配和随机分配共同主导,豆科功能群的最优拟合模型是BRO,资源分配方式以固定分配为主,杂草功能群的最优拟合模型从中性模型向生态位模型转变,与全部物种多度分布的最优模型基本一致,且资源分配由随机分配向固定分配转变,可以认为杂草功能群是影响群落物种多度分布的主要原因,但是禾草和豆科功能群的贡献也不可忽视。     

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]     

Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity

Neutral and niche theories give contrasting explanations for the maintenance of tropical tree species diversity. Both have some empirical support, but methods to disentangle their effects have not yet been developed. We applied a statistical measure of spatial structure to data from 14 large tropical forest plots to test a prediction of niche theory that is incompatible with neutral theory: that species in heterogeneous environments should separate out in space according to their niche preferences. We chose plots across a range of topographic heterogeneity, and tested whether pairwise spatial associations among species were more variable in more heterogeneous sites. We found strong support for this prediction, based on a strong positive relationship between variance in the spatial structure of species pairs and topographic heterogeneity across sites. We interpret this pattern as evidence of pervasive niche differentiation, which increases in importance with increasing environmental heterogeneity.     

物种分布模型理论研究进展

利用物种分布模型估计物种的真实和潜在分布区,已成为区域生态学与生物地理学中非常活跃的研究领域。然而,到目前为止,这项技术的理论基础仍然存在不足之处,一些关键的生态过程未能被有效纳入到物种分布模型的理论框架中,从而为解释物种分布模型预测的结果带来了诸多困惑。鉴于此,总结了物种分布模型的理论基础;系统探讨了物种分布模型与物种分布区的关系;特别指出了物种分布模型研究中存在的理论问题;重点阐述了物种分布模型未来的发展方向。研究认为,物种分布模型与生态位理论、源-库理论、种群动态理论、集合种群理论、进化理论等具有重要的联系;正确理解物种分布模型的预测结果与物种分布区的关系,有赖于对影响物种分布的3个主要因素(环境条件、物种相互作用与物种迁移能力)做出定量的分离;目前物种分布模型主要存在的问题是未能将物种的相互作用和物种的迁移能力有效纳入到模型的构建过程中;未来物种分布模型的发展应该加强模型背后理论框架的研究,并进一步加强整合物种相互作用过程、种群动态过程、迁移过程和物种进化过程等内容。研究还认为,从更高的理论层次模拟功能群和群落结构将是未来物种分布模型的重要发展方向。     

The quest for trait convergence and divergence in community assembly: are null‐models the magic wand?

The relevance of neutral versus niche‐based community assembly rules (i.e. the processes sorting species present in a larger geographical region into local communities) remains to be demonstrated in ecology and biogeography. To attempt to do this, a number of complex null models are increasingly being used that compare observed community functional diversity (FD, i.e. the extent of trait dissimilarity between coexisting species) with randomly simulated FD. However, little is known about the performance of these null models in detecting non‐neutral community assembly rules such as trait convergence and divergence of communities (supposedly revealing habitat selection and limiting similarity, respectively). Here, using both simulated and field communities, I show that assembly rule detection varies systematically with the magnitude of the observed FD, so that these null models do not really succeed in breaking down the observed functional relationships between species. This is a particular concern, making detection of community assembly dependent on: (1) the pool of samples considered, and (2) the capacity of observed FD to correctly discriminate these rules. Null models should be more thoroughly described and validated before being considered as a magic wand to reveal assembly patterns.     

The influence of stochasticity,landscape structure and species traits on abundant–centre relationships

Species have been commonly hypothesized to have high population densities in geographic areas which correspond to either the centre of the species geographic range or climatic niche (abundant–centre hypothesis). However, there is mixed empirical support for this relationship, and little theoretical underpinning. We simulate a species spreading across a set of replicated artificial landscapes to examine the expected level of support for abundant–centre relationships in geographic and niche space. Species niche constraints were modeled as a single axis which was related directly to population growth rates. We found strong evidence for abundant–centre relationships when populations follow deterministic growth, dispersal is high, environmental noise is absent and intraspecific competition is low. However, the incorporation of ecological realism reduced the detectability of abundant–centre relationships considerably. Our results suggest that even in carefully constructed artificial landscapes designed to demonstrate abundant–centre dynamics, the incorporation of small amounts of demographic stochasticity, environmental heterogeneity or landscape structure can strongly influence the relationship between species population density and distance to species geographic range or niche centre. While some simulated relationships were of comparable strength to common empirical support for abundant–centre relationships, our results suggest that these relationships are expected to be fairly variable and weak.