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.

Experimental evidence that soil heterogeneity enhances plant diversity during community assembly

Aims Environmental heterogeneity is a primary mechanism explain-ing species coexistence and extant patterns of diversity.Despite strong theoretical support and ample observational evidence,few experimental studies in plant communities have been able to demonstrate a causal link between environmental heterogene-ity and plant diversity.This lack of experimental evidence sug-gests that either fine-scale heterogeneity has weak effects on plant diversity or previous experiments have been unable to effectively manipulate heterogeneity.Here,we utilize a unique soil manipu-lation to test whether fine-scale soil heterogeneity will increase plant richness through species sorting among experimental patch types.Methods This experiment was conducted in the tallgrass prairie region of south-central Kansas,USA.We utilized the inherent variation found in the vertical soil profile,which varied in both biotic and abiotic characteristics,and redistributed these strata into either homoge-neous or heterogeneous spatial arrangements in 2.4×2.4 m plots.After the soil manipulation,34 native prairie species were sown into all plots.We conducted annual censuses at peak biomass to quantify species composition and plant density by species within the experimental communities.Important Findings After 2 years,species richness was significantly higher in heteroge-neous relative to homogeneous plots and this pattern was independ-ent of total plant density.In the heterogeneous plots,13 species had higher establishment in a specific patch type representing one of the three soil strata.Conversely,no species had greater estab-lishment in the mixed stratum,which comprised the homogene-ous plots,relative to the heterogeneous strata.These species sorting patterns suggest that fine-scale heterogeneity creates opportunities for plant establishment due to niche differences,which translates into increased plant diversity at the plot scale.Species richness was more strongly related to plant density among patches comprising homogenous plots-where fine-scale heterogeneity was minimized,but weak in heterogeneous plots.This pattern is consistent with the idea that richness-density relationships dominate when neutral pro-cesses are important but are weak when niche processes operate.Unlike many previous attempts,our results provide clear,experi-mental evidence that fine-scale soil heterogeneity increases species richness through species sorting during community assembly.

Diversity patterns and phylogenetic structure of vascular plants along elevational gradients in a mountain ecosystem, South Korea

Recently, a phylogenetic diversity and community structure analysis as complementary to species-centric approaches in biodiversity studies provides new insights into the processes of community assembly. In this study, we analyzed species and phylogenetic diversity and community structures for woody and herbaceous plants along two elevational transects on Mt. Baekhwa, South Korea. The species richness and phylogenetic diversity of woody plants showed monotonic declining patterns with increasing elevation along all transects, whereas herbaceous plants showed different patterns, such as no relationship and a reversed unimodal pattern, between the study transects. The main drivers of these patterns were climate and habitat variables for woody and herbaceous plants, respectively. In addition, the phylogenetic community structure primarily showed phylogenetic clustering regulated by deterministic processes, especially environmental filtering, such as climate or habitat factors, along the two transects, although herbaceous plants along a transect depicted phylogenetic randomness as a result of a neutral process. Our findings suggest that deterministic and neutral processes may simultaneously control the community structures along small-scale elevational gradients such as local transects, although the deterministic process may be the predominant type.

Dispersal and recruitment limitation contribute differently to community assembly

Aims The neutral theory of biodiversity has been criticized for being fragile with even slight deviations from its basic assumption of equal fitness among species.In response to this criticism,Hubbell((2001)The Unified Neutral Theory of Biodiversity and Biogeography.Princeton,NJ:Princeton University Press)proposed that competitive exclusion can be infinitely delayed by dispersal and recruitment limitation,thus making species effectively neutral.But the theoretical foundation for this claim still remains unclear and controversial,and the effects of dispersal and recruitment limitation are often confounded,especially in field studies.This study aims to provide an affirmative theoretical answer to the question of whether dispersal limitation and recruitment limitation can separately or jointly overwhelm the effects of fitness differences among species and lead to neutral community dynamics.Methods Computer simulations were used to investigate the effects of dispersal and recruitment limitation on delaying competitive exclusion in a homogeneous habitat in a spatially explicit context.Important Findings We found that even a slight competitive asymmetry would require extremely strong dispersal and recruitment limitation for neutrality to emerge.Most importantly,when the effects of dispersal and recruitment limitation were set apart,it is found that recruitment limitation is more effective in delaying competitive exclusion,whereas dispersal limitation tends to have a stronger impact on the general shape of both species abundance distributions and species–area relationships.

群落生态学的中性理论

生物多样性的分布格局和维持机制一直是群落生态学研究的核心问题,其中的关键是物种的共存机制。长期以来,生态位分化的思想在这一研究领域占据着主导地位。然而这一理论在解释热带雨林很高的物种多样性时遇到了困难。而以Hubbell为代表提出的群落中性漂变理论则假定在同一营养级物种构成的群落中不同物种的不同个体在生态学上可看成是完全等同的;物种的多度随机游走,群落中的物种数取决于物种灭绝和物种迁入/新物种形成之间的动态平衡。在这一假定之下,该理论预言了两种统计分布。一种是集合群落在点突变形成新物种的模式下其各个物种相对多度服从对数级数分布,而受扩散限制的局域群落以及按照随机分裂为新物种模式形成的集合群落则服从零和多项式分布。与生态位理论相反,中性理论不以种间生态位差异作为研究群落结构的出发点,而是以物种间在个体水平上的对等性作为前提。该理论第一次从基本生态学过程(出生、死亡、迁移、物种分化)出发,给出了群落物种多度分布的机理性解释,同时其预测的物种多度分布格局在实际群落中也得到了广泛的印证。因此,中性理论自诞生以来便在生态学界引发了极大的反响,也包括一些反对的声音。该文重点综述了关于中性理论的假设、预测和物种形成模式等方面的最新研究进展,包括中性理论本身的发展、关于中性理论的假设和预测的合理性检验以及在集合群落尺度上物种分化模式的讨论;并指出未来发展方向可能是在生态位理论和中性理论之间架起一座桥梁,同时发展包含随机性的群落生态位模型,以及允许种间差异的近中性模型。

Point process models, the dimensions of biodiversity and the importance of small-scale biotic interactions

Aims Recent mechanistic explanations for community assembly focus on the debates surrounding niche-based deterministic and dispersalbased stochastic models.This body of work has emphasized the importance of both habitat filtering and dispersal limitation,and many of these works have utilized the assumption of species spatial independence to simplify the complexity of the spatial modeling in natural communities when given dispersal limitation and/or habitat filtering.One potential drawback of this simplification is that it does not consider species interactions and how they may influence the spatial distribution of species,phylogenetic and functional diversity.Here,we assess the validity of the assumption of species spatial independence using data from a subtropical forest plot in southeastern China.Methods We use the four most commonly employed spatial statistical models—the homogeneous Poisson process representing pure random effect,the heterogeneous Poisson process for the effect of habitat heterogeneity,the homogenous Thomas process for sole dispersal limitation and the heterogeneous Thomas process for joint effect of habitat heterogeneity and dispersal limitation—to investigate the contribution of different mechanisms in shaping the species,phylogenetic and functional structures of communities.Important Findings Our evidence from species,phylogenetic and functional diversity demonstrates that the habitat filtering and/or dispersal-based models perform well and the assumption of species spatial independence is relatively valid at larger scales(50×50 m).Conversely,at local scales(10×10 and 20×20 m),the models often fail to predict the species,phylogenetic and functional diversity,suggesting that the assumption of species spatial independence is invalid and that biotic interactions are increasingly important at these spatial scales.

山东半岛潮上带沙草地的物种多度格局及其对人为干扰的响应

沿海岸线分布的潮上带沙滩是受风暴潮影响的独特生境,蕴含着独特的植物群落与植物资源。很多沙滩已受到不同程度的人为干扰。人们尚未了解潮上带沙滩植物群落的构建方式,也忽视了各种人为干扰对它的影响。该文以几何级数模型、分割线段模型、重叠生态位模型和中性理论模型,剖析了山东半岛潮上带3处典型沙草地的物种多度格局,同时研究了滩涂养殖和旅游践踏两种最重要的干扰方式对潮上带沙滩植物群落的影响。对于这3处植被,中性理论模型的拟合效果最好,其次是几何级数模型,而分割线段模型和重叠生态位模型的拟合效果不够理想。旅游践踏和沙滩养殖使潮上带沙草地的植被盖度降低,不利于珍稀特有植物的生存。在潮上带沙滩上很多植物具有不少相似的特征,导致它们在个体水平上的空间替代有强烈的随机性,服从群落中性理论。然而,它们的性状毕竟存在或多或少的差异,这些差异难免对各种植物的资源分配或竞争能力产生微弱的影响,经过很多年的积累,这种影响必然在群落水平上逐渐显现出来,以至于用几何级数模型也可拟合物种的多度格局。作者建议应控制人为干扰对潮上带沙滩珍稀植物的不利影响。