Species diversity in local neutral communities

We extend the neutral theory of macroecology by deriving biodiversity models (relative species abundance and species-area relationships) in a local community-metacommunity system in which the local community is embedded within the metacommunity. We first demonstrate that the local species diversity patterns converge to that of the metacommunity as the size (scale) of the embedded local community increases. This result shows that in continuous landscapes no sharp boundaries dividing the communities at the two scales exist; they are an artificial distinction made by the current spatially implicit neutral theory. Second, we remove the artificial restriction that speciation cannot occur in a local community, even if the effects of local speciation are small. Third, we introduce stochasticity into the immigration rate, previously treated as constant, and demonstrate that local species diversity is a function not only of the mean but also of the variance in immigration rate. High variance in immigration rates reduces species diversity in local communities. Finally, we show that a simple relationship exists between the fundamental diversity parameter of neutral theory and Simpson's index for local communities. Derivation of this relationship extends recent work on diversity indices and provides a means of evaluating the effect of immigration on estimates of the fundamental diversity parameter derived from relative species abundance data on local communities.     

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.     

Sampling Hubbell's neutral theory of biodiversity

In the context of neutral theories of community ecology, a novel genealogy‐based framework has recently furnished an analytic extension of Ewens’ sampling multivariate abundance distribution, which also applies to a random sample from a local community. Here, instead of taking a multivariate approach, we further develop the sampling theory of Hubbell's neutral spatially implicit theory and derive simple abundance distributions for a random sample both from a local community and a metacommunity. Our result is given in terms of the average number of species with a given abundance in any randomly extracted sample. Contrary to what has been widely assumed, a random sample from a metacommunity is not fully described by the Fisher log‐series, but by a new distribution. This new sample distribution matches the log‐series expectation at high biodiversity values (θ > 1) but clearly departs from it for species‐poor metacommunities (θ < 1). Our theoretical framework should be helpful in the better assessment of diversity and testing of the neutral theory by using abundance data.     

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.     

A sampling theory for asymmetric communities

We introduce the first analytical model of asymmetric community dynamics to yield Hubbell's neutral theory in the limit of functional equivalence among all species. Our focus centers on an asymmetric extension of Hubbell's local community dynamics, while an analogous extension of Hubbell's metacommunity dynamics is deferred to an appendix. We find that mass-effects may facilitate coexistence in asymmetric local communities and generate unimodal species abundance distributions indistinguishable from those of symmetric communities. Multiple modes, however, only arise from asymmetric processes and provide a strong indication of non-neutral dynamics. Although the exact stationary distributions of fully asymmetric communities must be calculated numerically, we derive approximate sampling distributions for the general case and for nearly neutral communities where symmetry is broken by a single species distinct from all others in ecological fitness and dispersal ability. In the latter case, our approximate distributions are fully normalized, and novel asymptotic expansions of the required hypergeometric functions are provided to make evaluations tractable for large communities. Employing these results in a Bayesian analysis may provide a novel statistical test to assess the consistency of species abundance data with the neutral hypothesis.     

Asymmetry in species regional dispersal ability and the neutral theory

The neutral assumption that individuals of either the same or different species share exactly the same birth, death, migration, and speciation probabilities is fundamental yet controversial to the neutral theory. Several theoretical studies have demonstrated that a slight difference in species per capita birth or death rates can have a profound consequence on species coexistence and community structure. Whether asymmetry in migration, a vital demographic parameter in the neutral model, plays an important role in community assembly still remains unknown. In this paper, we relaxed the ecological equivalence assumption of the neutral model by introducing differences into species regional dispersal ability. We investigated the effect of asymmetric dispersal on the neutral local community structure. We found that per capita asymmetric dispersal among species could reduce species richness of the local community and result in deviations of species abundance distributions from those predicted by the neutral model. But the effect was moderate compared with that of asymmetries in birth or death rates, unless very large asymmetries in dispersal were assumed. A large difference in species dispersal ability, if there is, can overwhelm the role of random drift and make local community dynamics deterministic. In this case, species with higher regional dispersal abilities tended to dominate in the local community. However, the species abundance distribution of the local community under asymmetric dispersal could be well fitted by the neutral model, but the neutral model generally underestimated the fundamental biodiversity number but overestimated the migration rate in such communities.     

Reconciling niche and neutrality: the continuum hypothesis

In this study, we ask if instead of being fundamentally opposed, niche and neutral theories could simply be located at the extremes of a continuum. First, we present a model of recruitment probabilities that combines both niche and neutral processes. From this model, we predict and test whether the relative importance of niche vs. neutral processes in controlling community dynamics will vary depending on community species richness, niche overlap and dispersal capabilities of species (both local and long distance). Results demonstrate that niche and neutrality form ends of a continuum from competitive to stochastic exclusion. In the absence of immigration, competitive exclusion tends to create a regular spacing of niches. However, immigration prevents the establishment of a limiting similarity. The equilibrium community consists of a set of complementary and redundant species, with their abundance determined, respectively, by the distribution of environmental conditions and the amount of immigration.     

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]     

Reconciling neutral community models and environmental filtering: theory and an empirical test

It is widely believed that the neutral theory of biodiversity cannot be used for parameter inference if the assumption of neutrality is not met. The goal of this work is to extend this neutral framework to quantify the intensity of recruitment limitation (limited dispersal plus environmental filtering) in natural species assemblages. We model several local communities as part of a larger metacommunity, and we assume that neutrality holds in each local community, but not in the metacommunity. The immigration rate m does not only reflect dispersal limitation into a given local community, but also the intensity of environmental filtering. We develop a novel statistical method to infer the immigration parameter m in each local community. Using simulated datasets, we show that m indeed depends on both dispersal limitation and on the intensity of environmental filtering. We then apply this method to a network of tropical tree plots in central Panama. Inferred recruitment rates m were positively correlated with the fraction of trees dispersed by mammals, and with annual rainfall, possibly due to a weaker environmental filtering as rainfall increases. Finally, m, as estimated from trees greater than 1 cm trunk diameter, were significantly larger than an estimation based on trees greater than 10 cm trunk diameter. This suggests a cumulative effect of environmental filtering upon trees throughout their ontogeny.     

A new sampling formula for neutral biodiversity

The neutral model of biodiversity, proposed by Hubbell (The Unified Neutral Theory of Biodiversity and Biogeography, Princeton University Press, Princeton, NJ, 2001) to explain the diversity of functionally equivalent species, has been subject of hot debate in community ecology. Whereas Hubbell studied the model mostly by simulations, recently analytical treatments have yielded expressions of the expected number of species of a particular abundance in a local community with dispersal limitation. Moreover, a formula has been offered for the joint likelihood of observing a given species‐abundance dataset in a local community with dispersal limitation, but this formula is too complicated to allow practical applications. Here, I present a much simplified expression that can be regarded as an enhanced version of the famous Ewens sampling formula. It can be used in maximum likelihood methods for quick estimation of the model parameters, using all information in the data, and for model comparison. I also show how to rapidly generate examples of species‐abundance distributions for a given set of model parameters and how to calculate Simpson's diversity index.     

Temporal dynamics of a local fish community are strongly affected by immigration from the surrounding metacommunity

A 5‐year time series of annual censuses was collected from a large floodplain lake to determine how dynamics of the local fish community were affected by changes in hydrological connectivity with the surrounding metacommunity. The lake was disconnected from the metacommunity for 1 year prior to our study and remained disconnected until 3 months before our third annual census, when a flood reconnected the lake to the metacommunity. We determined how changes in connectivity affected temporal dynamics of (1) local community composition and (2) the population composition, condition, and growth of catfish, to shed light on how immigration of other species might affect local population dynamics. Before reconnection, the community was likely shaped by interactions between the local environment and species traits. The reconnection caused significant immigration and change in community composition and correlated with a significant and abrupt decline in catfish condition, growth, and abundance; effects likely due to the immigration of a competitor with a similar trophic niche: carp. The community was slow to return to its preconnection state, which may be due to dispersal traits of the fishes, and a time‐lag in the recovery of the local catfish population following transient intensification of species interactions. The dynamics observed were concordant with the species sorting and mass‐effects perspectives of metacommunity theory. Floods cause episodic dispersal in floodplain fish metacommunities, and so, flood frequency determines the relative importance of regional and local processes. Local processes may be particularly important to certain species, but these species may need sufficient time between floods for population increase, before the next flood‐induced dispersal episode brings competitors and predators that might cause population decline. Accordingly, species coexistence in these metacommunities may be facilitated by spatiotemporal storage effects, which may in turn be regulated by flood frequency.     

Time-dependent solutions of the spatially implicit neutral model of biodiversity

Previous research into the neutral theory of biodiversity has focused mainly on equilibrium solutions rather than time-dependent solutions. Understanding the time-dependent solutions is essential for applying neutral theory to ecosystems in which time-dependent processes, such as succession and invasion, are driving the dynamics. Time-dependent solutions also facilitate tests against data that are stronger than those based on static equilibrium patterns. Here I investigate the time-dependent solutions of the classic spatially implicit neutral model, in which a small local community is coupled to a much larger metacommunity through immigration. I present explicit general formulas for the eigenvalues, left eigenvectors and right eigenvectors of the models’s transition matrix. The time-dependent solutions can then be expressed in terms of these eigenvalues and eigenvectors. Some of these results are translated directly from existing results for the classic Moran model of population genetics (the Moran model is equivalent to the spatially implicit neutral model after a reparameterization); others of the results are new. I demonstrate that the asymptotic time-dependent solution corresponding to just these first two eigenvectors can be a good approximation to the full time-dependent solution. I also demonstrate the feasibility of a partial eigendecomposition of the transition matrix, which facilitates direct application of the results to a biologically relevant example in which a newly invading species is initially present in the metacommunity but absent from the local community.     

Immigration,Local Dispersal Limitation,and the Repeatability of Community Composition under Neutral and Niche Dynamics

Repeatability of community composition has been a critical aspect for community structure, which is closely associated with community stability, predictability, conservation biology and ecological restoration. It has been shown that both immigration and local dispersal limitation can affect the community composition in both neutral and niche model. Hence, we use a spatially explicit individual-based model to investigate the potential influence of immigration rate and strength of local dispersal limitation on repeatability in both neutral and niche models. Similarity measures are used to quantify repeatability. We examine the repeatability of community composition among replicate communities (which means the same community repeats many times), and between niche and neutral replicate communities. We find the correlation between repeatability and immigration rate is positive in the neutral model and an inverted unimodal in the niche model. The correlation between repeatability and local dispersal distance is positive in the niche model and negative in the neutral model. High repeatability between niche communities and neutral communities is observed with high immigration rates or when high local dispersal distance appears in the niche model or low local dispersal distance in the neutral model. Our results show that repeatability of community composition is not only dependent on the types of community models (niche vs. neutrality) but also strongly determined by immigration rates and local dispersal limitation.     

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。     

Hubbell's fundamental biodiversity parameter and the Simpson diversity index

Central to Hubbell's neutral theory of biodiversity is a universal, dimensionless fundamental biodiversity parameter that is the product of community size and speciation rate. One of the most important discoveries of Hubbell's theory is that the species‐abundance distribution and the species–area relationship of the neutral metacommunity is completely determined by this fundamental biodiversity parameter, although the diversity patterns of the local community are collectively determined by the biodiversity parameter and migration. Using the relative abundance of species and following the concept of heterozygosity of population genetics, here we developed an analytical relationship between this biodiversity parameter and the well‐known Simpson diversity index. This relationship helps bridge the evolutionary aspect of biodiversity to the ecological and statistical aspect of the diversity. The relationship between these two parameters suggests that diversity patterns of the metacommunity can also be equally described by the Simpson index. This relationship provides an alternative approach to interpret and estimate the fundamental biodiversity parameter for the metacommunity.     

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.     

Do Spatially-Implicit Estimates of Neutral Migration Comply with Seed Dispersal Data in Tropical Forests?

Neutral community models have shown that limited migration can have a pervasive influence on the taxonomic composition of local communities even when all individuals are assumed of equivalent ecological fitness. Notably, the spatially implicit neutral theory yields a single parameter I for the immigration-drift equilibrium in a local community. In the case of plants, seed dispersal is considered as a defining moment of the immigration process and has attracted empirical and theoretical work. In this paper, we consider a version of the immigration parameter I depending on dispersal limitation from the neighbourhood of a community. Seed dispersal distance is alternatively modelled using a distribution that decreases quickly in the tails (thin-tailed Gaussian kernel) and another that enhances the chance of dispersal events over very long distances (heavily fat-tailed Cauchy kernel). Our analysis highlights two contrasting situations, where I is either mainly sensitive to community size (related to ecological drift) under the heavily fat-tailed kernel or mainly sensitive to dispersal distance under the thin-tailed kernel. We review dispersal distances of rainforest trees from field studies and assess the consistency between published estimates of I based on spatially-implicit models and the predictions of the kernel-based model in tropical forest plots. Most estimates of I were derived from large plots (10–50 ha) and were too large to be accounted for by a Cauchy kernel. Conversely, a fraction of the estimates based on multiple smaller plots (1 ha) appeared too small to be consistent with reported ranges of dispersal distances in tropical forests. Very large estimates may reflect within-plot habitat heterogeneity or estimation problems, while the smallest estimates likely imply other factors inhibiting migration beyond dispersal limitation. Our study underscores the need for interpreting I as an integrative index of migration limitation which, besides the limited seed dispersal, possibly includes habitat filtering or fragmentation.     

Universal scaling of species‐abundance distributions across multiple scales

The scale‐dependent species abundance distribution (SAD) is fundamental in ecology, but few spatially explicit models of this pattern have thus far been studied. Here we show spatially explicit neutral model predictions for SADs over a wide range of spatial scales, which appear to match empirical patterns qualitatively. We find that the assumption of a log‐series SAD in the metacommunity made by spatially implicit neutral models can be justified with a spatially explicit model in the large area limit. Furthermore, our model predicts that SADs on multiple scales are characterized by a single, compound parameter that represents the ratio of the survey area to the species’ average biogeographic range (which is in turn set by the speciation rate and the dispersal distance). This intriguing prediction is in line with recent empirical evidence for a universal scaling of the species‐area curve. Hence we hypothesize that empirical SAD patterns will show a similar universal scaling for many different taxa and across multiple spatial scales.