Niches, rather than neutrality, structure a grassland pioneer guild

Pioneer species are fast-growing, short-lived gap exploiters. They are prime candidates for neutral dynamics because they contain ecologically similar species whose low adult density is likely to cause widespread recruitment limitation, which slows competitive dynamics. However, many pioneer guilds appear to be differentiated according to seed size. In this paper, we compare predictions from a neutral model of community structure with three niche-based models in which trade-offs involving seed size form the basis of niche differentiation. We test these predictions using sowing experiments with a guild of seven pioneer species from chalk grassland. We find strong evidence for niche structure based on seed size: specifically large-seeded species produce fewer seeds but have a greater chance of establishing on a per-seed basis. Their advantage in establishment arises because there are more microsites suitable for their germination and early establishment and not directly through competition with other seedlings. In fact, seedling densities of all species were equally suppressed by the addition of competitors' seeds. By the adult stage, despite using very high sowing densities, there were no detectable effects of interspecific competition on any species. The lack of interspecific effects indicates that niche differentiation, rather than neutrality, prevails.     

Niches versus neutrality: uncovering the drivers of diversity in a species-rich community

Ecological models suggest that high diversity can be generated by purely niche-based, purely neutral or by a mixture of niche-based and neutral ecological processes. Here, we compare the degree to which four contrasting hypotheses for coexistence, ranging from niche-based to neutral, explain species richness along a body mass niche axis. We derive predictions from these hypotheses and confront them with species body-mass patterns in a highly sampled marine phytoplankton community. We find that these patterns are consistent only with a mechanism that combines niche and neutral processes, such as the emergent neutrality mechanism. In this work, we provide the first empirical evidence that a niche-neutral model can explain niche space occupancy pattern in a natural species-rich community. We suggest this class of model may be a useful hypothesis for the generation and maintenance of species diversity in other size-structured communities.     

A mathematical synthesis of niche and neutral theories in community ecology

The debate between niche-based and neutral community theories centers around the question of which forces shape predominantly ecological communities. Niche theory attributes a central role to niche differences between species, which generate a difference between the strength of intra- and interspecific interactions. Neutral theory attributes a central role to migration processes and demographic stochasticity. One possibility to bridge these two theories is to combine them in a common mathematical framework. Here we propose a mathematical model that integrates the two perspectives. From a niche-based perspective, our model can be interpreted as a Lotka-Volterra model with symmetric interactions in which we introduce immigration and demographic stochasticity. From a neutral perspective, it can be interpreted as Hubbell's local community model in which we introduce a difference between intra- and interspecific interactions. We investigate the stationary species abundance distribution and other community properties as functions of the interaction coefficient, the immigration rate and the strength of demographic stochasticity.     

Niche partitioning at multiple scales facilitates coexistence among mosquito larvae

A theoretical dichotomy in community ecology distinguishes between mechanisms that stabilize species coexistence and those that cause neutral drift. Stable coexistence is predicted to occur in communities where competing species have niche-partitioning mechanisms that reduce interspecific competition. Neutral communities are predicted to be structured by stochastic processes that are not influenced by species identity, but that may be influenced by priority effects and dispersal limitation. Recent developments have suggested that neutral interactions may be more common at local scales, while niche structuring may be more common at larger scales. We tested for mechanisms that could promote either stable coexistence or neutral drift in a bromeliad-dwelling mosquito community by evaluating A) if a hypothesized within-bromeliad niche partitioning mechanism occurs in the community, B) if this mechanism correlates with local species co-occurrence patterns, and C) if patterns of coexistence at the larger (metacommunity) scale were consistent with those at the local scale. We found that mosquitoes in this community do partition space within containers, and that species with the strongest potential for competition co-occurred least. Species with overlapping spatial niches minimized co-occurrence by specialising in bromeliads of differing sizes, effectively changing the scale at which they coexist. In contrast, we found no evidence to support neutral dynamics in mosquito communities at either scale. In this community, a niche-based mechanism that is predicted to stabilize species coexistence explains co-occurrence patterns within and among bromeliads.     

When can we distinguish between neutral and non‐neutral processes in community dynamics under ecological drift?

Determining whether the composition of ecological communities (species presence and abundance), can be predicted from species demographic traits, rather than being a result of neutral drift, is a key ecological question. Here we compare the similarity of community composition, from different community assembly models run under identical environmental conditions, where interspecific competition is assumed to be either neutral or niche-based. In both cases, species colonize a focal patch from a network of neighbouring patches in a metacommunity. We highlight the circumstances (rate and spatial scale of dispersal, and the relative importance of ecological drift) where commonly used community similarity metrics or species rank–abundance relationships are likely to give similar results, regardless of the underlying processes (neutral or non-neural) driving species' dynamics. As drift becomes more important in driving species abundances, deterministic niche structure has a smaller influence. Our ability to discriminate between different underlying processes driving community organization depends on the relative importance of different drift processes that operate on different spatial scales.     

Analyzing tropical forest tree species abundance distributions using a nonneutral model and through approximate Bayesian inference

The neutral theory of biodiversity challenges the classical niche-based view of ecological communities, where species attributes and environmental conditions jointly determine community composition. Functional equivalence among species, as assumed by neutral ecological theory, has been recurrently falsified, yet many patterns of tropical tree communities appear consistent with neutral predictions. This may mean that neutral theory is a good first-approximation theory or that species abundance data sets contain too little information to reject neutrality. Here we present a simple test of neutrality based on species abundance distributions in ecological communities. Based on this test, we show that deviations from neutrality are more frequent than previously thought in tropical forest trees, especially at small spatial scales. We then develop a nonneutral model that generalizes Hubbell's dispersal-limited neutral model in a simple way by including one additional parameter of frequency dependence. We also develop a statistical method to infer the parameters of this model from empirical data by approximate Bayesian computation. In more than half of the permanent tree plots, we show that our new model fits the data better than does the neutral model. Finally, we discuss whether observed deviations from neutrality may be interpreted as the signature of environmental filtering on tropical tree species abundance distributions.     

Soil phosphorus heterogeneity promotes tree species diversity and phylogenetic clustering in a tropical seasonal rainforest

The niche theory predicts that environmental heterogeneity and species diversity are positively correlated in tropical forests, whereas the neutral theory suggests that stochastic processes are more important in determining species diversity. This study sought to investigate the effects of soil nutrient (nitrogen and phosphorus) heterogeneity on tree species diversity in the Xishuangbanna tropical seasonal rainforest in southwestern China. Thirty‐nine plots of 400 m² (20 × 20 m) were randomly located in the Xishuangbanna tropical seasonal rainforest. Within each plot, soil nutrient (nitrogen and phosphorus) availability and heterogeneity, tree species diversity, and community phylogenetic structure were measured. Soil phosphorus heterogeneity and tree species diversity in each plot were positively correlated, while phosphorus availability and tree species diversity were not. The trees in plots with low soil phosphorus heterogeneity were phylogenetically overdispersed, while the phylogenetic structure of trees within the plots became clustered as heterogeneity increased. Neither nitrogen availability nor its heterogeneity was correlated to tree species diversity or the phylogenetic structure of trees within the plots. The interspecific competition in the forest plots with low soil phosphorus heterogeneity could lead to an overdispersed community. However, as heterogeneity increase, more closely related species may be able to coexist together and lead to a clustered community. Our results indicate that soil phosphorus heterogeneity significantly affects tree diversity in the Xishuangbanna tropical seasonal rainforest, suggesting that deterministic processes are dominant in this tropical forest assembly.     

Complementarity effects are strengthened by competition intensity and global environmental change in the central boreal forests of Canada

Increases in niche complementarity have been hypothesised to reduce the intensity of interspecific competition within natural forests. In regions currently experiencing potentially enhanced growth under global environmental change, niche complementarity may become even more beneficial. However, few studies have provided direct evidence of this mechanism. Here, we use data from 180 permanent sample plots in Manitoba, Canada, with a full spatial mapping of all stems, to show that complementarity effects on average increased with neighbourhood competition intensity and temporally rising CO₂, warming and water availability. Importantly, complementarity effects increased with both shade tolerance and phylogenetic dissimilarity between the focal tree and its neighbours. Our results provide further evidence that increasing stand functional and phylogenetic diversity can improve individual tree productivity, especially for individuals experiencing intense competition and may offer an avenue to maintain productivity under global environmental change.     

Temporal niche dynamics, relative abundance and phylogenetic signal of coexisting species

The coexistence of similar species accounts for some 30% of diversity within communities, yet the coexistence and relative abundance of similar species is a continuing ecological conundrum. Using close phylogenetic relatedness as a measure of similarity, we previously demonstrated that neither classic niche theory nor neutral theory can explain the relative abundances of co-occurring pairs of similar tree species in a diverse tropical forest. Here, we show that the stable, focused competition of a temporal niche dynamic fits the distribution of observed fractional abundances (pairwise relative abundances). Previously published, independent evidence of temporal dynamics in this community supports our results; our model identifies additional criteria for field tests of differential sensitivity (DS) temporal dynamics. The success of temporal dynamics at explaining the observed distribution—and the failure of alternative hypotheses to do so—indicates that current diagnostics of community structure and assembly needs general re-examination. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Simulated responses to hypothetical fundamental niches

Abstract. Two graphical models of plant competition, the ‘ghost of competition past’ and the ‘hierarchical’ model are compared using a greatly simplified individual-based forest dynamics simulation. Assumed fundamental niche shapes are used to determine the basic growth responses of the species, but competition alters the realized niche. Differences in the two models, amount of niche overlap, disturbance, and removal and invasion of species are examined in simulation experiments. Without disturbance, the realized niche responses reveal abrupt boundaries between species in all cases, and thus the responses are generally platykurtic to rectangular with little overlap. In some cases overlap through the extension of abundance of weaker competitors into the area of greater resources — dominated by better competitors but still within their fundamental niche — creates skewed distributions, as have been observed and simulated in the past. When species are removed or invade, the abrupt boundaries and the lack of difference in final response shape indicate that past removals may be difficult to detect. This difficulty may be important because former species may be influencing the responses that are observed in common non-equilibrium distributions. Even when assumptions favor the illustration of a competitive hierarchy, actual distributions and dynamics do not differentiate between it and a model of the ghost of competition past.     

塔里木荒漠河岸林不同生境群落物种多度分布格局

为解释塔里木荒漠河岸林群落构建和物种多度分布格局形成的机理, 本文以塔里木荒漠河岸林2个不同生境(沙地、河漫滩) 4 ha固定监测样地为研究对象, 基于两样地物种调查数据, 采用统计模型(对数级数模型、对数正态模型、泊松对数正态分布模型、Weibull分布模型)、生态位模型(生态位优先占领模型、断棍模型)和中性理论模型(复合群落零和多项式模型、Volkov模型)拟合荒漠河岸林群落物种多度分布, 并用K-S检验与赤池信息准则(AIC)筛选最优拟合模型。结果表明: (1)随生境恶化(土壤水分降低), 植物物种多度分布曲线变化减小, 群落物种多样性、多度和群落盖度降低, 常见种数减少。(2)选用的3类模型均可拟合荒漠河岸林不同生境群落物种多度分布格局, 统计模型和中性理论模型拟合效果均优于生态位模型。复合群落零和多项式模型对远离河岸的干旱沙地生境拟合效果最好; 对数正态模型和泊松对数正态模型对洪水漫溢的河漫滩生境拟合效果最优; 中性理论模型与统计模型无显著差异。初步推断中性过程在荒漠河岸林群落构建中发挥着主导作用, 但模型拟合结果只能作为推断群落构建过程的必要非充分条件, 不能排除生态位过程的潜在作用。     

Competition theory,evolution, and the concept of an ecological niche

This article examines some of the main tenets of competition theory in light of the theory of evolution and the concept of an ecological niche. The principle of competitive exclusion and the related assumption that communities exist at competitive equilibrium - fundamental parts of many competition theories and models - may be violated if non-equilibrium conditions exist in natural communities or are incorporated into competition models. Furthermore, these two basic tenets of competition theory are not compatible with the theory of evolution. Variation in ecologically significant environmental factors and non-equilibrium in population numbers should occur in most natural communities, and such changes have important effects on community relations, niche overlap, and the evolution of ecosystems. Ecologists should view competition as a process occurring within a complexdynamic system, and should be wary of theoretical positions built upon simple laboratory experiments or simplistic mathematical models.In considering the relationship between niche overlap and competition, niche overlap should not be taken as a sufficient condition for competition; many factors may prevent or diminish competition between populations with similar resource utilization patterns. The typically opposing forces of intraspecific and interspecific competition need to be simultaneously considered, for it is the balance between them that in large part determines niche boundaries.     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

Maintenance of forest species diversity and latitudinal gradient

The species diversity of trees maintained in tropical rain forests is much higher than in temperate, boreal, or seasonally dry tropical forests. Many hypotheses have been proposed for higher diversity in tropical rain forests, including: (i) higher specialization of resource use, (ii) different mode of disturbance, (iii) smaller opportunity for competition on oligotrophic soil, (iv) higher productivity, (v) more active specific herbivores and pathogens, (vi) evolutionary/ecological history. In this paper we report mathematical models for tree-by-tree replacement. First the analysis of random drift model shows that the effect of gap size to species diversity is not very strong. Second we study phenological segregation model, which has the following assumptions: Basic mechanism for many species to coexist in the community is assumed given by the storage effect of lottery model, as species differ in seasonality in peak fruit production and in the subsequent period of high regeneration ability. Gaps formed during unfavorable season accumulate and become available for regeneration in the beginning of the growing season. The resulting synchronization of regeneration opportunity jeopardizes the coexistence of many similar species in seasonal environments. Analysis of a mathematical model shows: (1) the existence of unfavorable season can greatly reduce the diversity of coexisting species. (2) Diversity in the equilibrium community can be high when niche width of each species is broad and resource use is strongly overlapped. (3) Equilibrium community may include several distinct groups of species differing in phenology of regeneration. Effect of unequal niche width and frequency dependent regeneration are also examined.     

Local community size mediates ecological drift and competition in metacommunities

The outcome of competitive interactions is likely to be influenced by both competitive dominance (i.e. niche-based dynamics) and ecological drift (i.e. neutral dynamics governed by demographic stochasticity). However, spatial models of competition rarely consider the joint operation of these two processes. We develop a model based on the original competition-colonization trade-off model that incorporates niche and neutral processes and several realistic facets of ecological dynamics: it allows local competition (i.e. competition within a patch) to occur within communities of a finite size, it allows competitors to vary in the degree of competitive asymmetry, and it includes the role of local migration (i.e. propagule pressure). The model highlights the role of community size, i.e. the number of competitors in the local community, in mediating the relative importance of stochastic and deterministic forces. In metacommunities where local communities are small, ecological drift is substantial enough that strong competitors become effectively neutral, creating abrupt changes in the outcome of competition not predicted by the standard competition-colonization trade-off. Importantly, the model illustrates that, even when other aspects of species interactions (e.g. migration ability, competitive ability) are unchanged, local community size can alter the dynamics of metacommunity persistence. Our work demonstrates that activities which reduce the size of local communities, such as habitat destruction and degradation, effectively compound the extinction debt.     

Concurrent niche and neutral processes in the competition-colonization model of species coexistence

The importance of neutral dynamics is contentiously debated in the ecological literature. This debate focuses on neutral theory's assumption of fitness equivalency among individuals, which conflicts with stabilizing fitness that promotes coexistence through niche differentiation. I take advantage of competition-colonization trade-offs between species of aquatic micro-organisms (protozoans and rotifers) to show that equalizing and stabilizing mechanisms can operate simultaneously. Competition trials between species with similar colonization abilities were less likely to result in competitive exclusion than for species further apart. While the stabilizing mechanism (colonization differences) facilitates coexistence at large spatial scales, species with similar colonization abilities also exhibited local coexistence probably due to fitness similarities allowing weak stabilizing mechanisms to operate. These results suggest that neutral- and niche-based mechanisms of coexistence can simultaneously operate at differing temporal and spatial scales, and such a spatially explicit view of coexistence may be one way to reconcile niche and neutral dynamics.     

Predicted distributions,niche comparisons,and conservation status of the spotted linsang (<Emphasis Type="Italic">Prionodon pardicolor</Emphasis>) and banded linsang (<Emphasis Type="Italic">Prionodon linsang</Emphasis>)

Knowledge of a species’ distribution and habitat preferences is of paramount importance when assessing its conservation status. We used accurately recorded occurrence records and ecological niche modeling to predict the distribution of two poorly known small carnivore species that occur in Asia, the spotted linsang (Prionodon pardicolor) and banded linsang (Prionodon linsang), and analyzed their niche overlaps for habitat and elevation. We then assessed the loss of their suitable habitat and estimated the proportion of predicted presence under protected areas. We identified and assessed possible anthropogenic threats, and used our modeling predictions to recommend surveying priorities. Our study confirmed that these two linsangs are geographically separated, with no known overlap of their distributions. Our results indicated that the habitat and elevation niches of these two linsangs are similar: they both occur primarily in evergreen forest and can be found at all elevations up to around 3,000 m. Although these two linsangs might be allopatric due to intense interspecific competition between them, other explanations could explain their distribution patterns, such as biogeographical and ecological barriers. Our findings suggest that these two linsangs might be threatened by the loss, degradation, and fragmentation of tropical evergreen forests. Urgent field studies are thus needed to learn more about their habitat requirements and the impact of anthropogenic threats, including tropical forest disturbance and hunting.     

The contribution of rare species to community phylogenetic diversity across a global network of forest plots

Niche differentiation has been proposed as an explanation for rarity in species assemblages. To test this hypothesis requires quantifying the ecological similarity of species. This similarity can potentially be estimated by using phylogenetic relatedness. In this study, we predicted that if niche differentiation does explain the co-occurrence of rare and common species, then rare species should contribute greatly to the overall community phylogenetic diversity (PD), abundance will have phylogenetic signal, and common and rare species will be phylogenetically dissimilar. We tested these predictions by developing a novel method that integrates species rank abundance distributions with phylogenetic trees and trend analyses, to examine the relative contribution of individual species to the overall community PD. We then supplement this approach with analyses of phylogenetic signal in abundances and measures of phylogenetic similarity within and between rare and common species groups. We applied this analytical approach to 15 long-term temperate and tropical forest dynamics plots from around the world. We show that the niche differentiation hypothesis is supported in six of the nine gap-dominated forests but is rejected in the six disturbance-dominated and three gap-dominated forests. We also show that the three metrics utilized in this study each provide unique but corroborating information regarding the phylogenetic distribution of rarity in communities.     

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.