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.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Ecological Equivalence: A Realistic Assumption for Niche Theory as a Testable Alternative to Neutral Theory

Background

Hubbell''s 2001 neutral theory unifies biodiversity and biogeography by modelling steady-state distributions of species richness and abundances across spatio-temporal scales. Accurate predictions have issued from its core premise that all species have identical vital rates. Yet no ecologist believes that species are identical in reality. Here I explain this paradox in terms of the ecological equivalence that species must achieve at their coexistence equilibrium, defined by zero net fitness for all regardless of intrinsic differences between them. I show that the distinction of realised from intrinsic vital rates is crucial to evaluating community resilience.

Principal Findings

An analysis of competitive interactions reveals how zero-sum patterns of abundance emerge for species with contrasting life-history traits as for identical species. I develop a stochastic model to simulate community assembly from a random drift of invasions sustaining the dynamics of recruitment following deaths and extinctions. Species are allocated identical intrinsic vital rates for neutral dynamics, or random intrinsic vital rates and competitive abilities for niche dynamics either on a continuous scale or between dominant-fugitive extremes. Resulting communities have steady-state distributions of the same type for more or less extremely differentiated species as for identical species. All produce negatively skewed log-normal distributions of species abundance, zero-sum relationships of total abundance to area, and Arrhenius relationships of species to area. Intrinsically identical species nevertheless support fewer total individuals, because their densities impact as strongly on each other as on themselves. Truly neutral communities have measurably lower abundance/area and higher species/abundance ratios.

Conclusions

Neutral scenarios can be parameterized as null hypotheses for testing competitive release, which is a sure signal of niche dynamics. Ignoring the true strength of interactions between and within species risks a substantial misrepresentation of community resilience to habitat loss.     

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.     

Beyond the continuum: a multi-dimensional phase space for neutral–niche community assembly

Neutral and niche processes are generally considered to interact in natural communities along a continuum, exhibiting community patterns bounded by pure neutral and pure niche processes. The continuum concept uses niche separation, an attribute of the community, to test the hypothesis that communities are bounded by pure niche or pure neutral conditions. It does not accommodate interactions via feedback between processes and the environment. By contrast, we introduce the Community Assembly Phase Space (CAPS), a multi-dimensional space that uses community processes (such as dispersal and niche selection) to define the limiting neutral and niche conditions and to test the continuum hypothesis. We compare the outputs of modelled communities in a heterogeneous landscape, assembled by pure neutral, pure niche and composite processes. Differences in patterns under different combinations of processes in CAPS reveal hidden complexity in neutral–niche community dynamics. The neutral–niche continuum only holds for strong dispersal limitation and niche separation. For weaker dispersal limitation and niche separation, neutral and niche processes amplify each other via feedback with the environment. This generates patterns that lie well beyond those predicted by a continuum. Inferences drawn from patterns about community assembly processes can therefore be misguided when based on the continuum perspective. CAPS also demonstrates the complementary information value of different patterns for inferring community processes and captures the complexity of community assembly. It provides a general tool for studying the processes structuring communities and can be applied to address a range of questions in community and metacommunity ecology.     

Relative contribution of neutral and deterministic processes in shaping fruit‐feeding butterfly assemblages in Afrotropical forests

The unified neutral theory of biodiversity and biogeography has gained the status of a quantitative null model for explaining patterns in ecological (meta)communities. The theory assumes that individuals of trophically similar species are functionally equivalent. We empirically evaluate the relative contribution of neutral and deterministic processes in shaping fruit‐feeding butterfly assemblages in three tropical forests in Africa, using both direct (confronting the neutral model with species abundance data) and indirect approaches (testing the predictions of neutral theory using data other than species abundance distributions). Abundance data were obtained by sampling butterflies using banana baited traps set at the forest canopy and understorey strata. Our results indicate a clear consistency in the kind of species or species groups observed at either the canopy or understorey in the three studied communities. Furthermore, we found significant correlation between some flight‐related morphological traits and species abundance at the forest canopy, but not at the understorey. Neutral theory's contribution to explaining our data lies largely in identifying dispersal limitation as a key process regulating fruit‐feeding butterfly community structure. Our study illustrates that using species abundance data alone in evaluating neutral theory can be informative, but is insufficient. Species‐level information such as habitat preference, host plants, geographical distribution, and phylogeny is essential in elucidating the processes that regulate biodiversity community structures and patterns.     

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]     

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.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

A truce with neutral theory: local deterministic factors, species traits and dispersal limitation together determine patterns of diversity in stream invertebrates

1. Studies seeking to explain local patterns of diversity have typically relied on niche explanations, reflected in correlations with local environmental conditions, or neutral theory, invoking dispersal processes and speciation. 2. We used macroinvertebrate community data from 10 streams that varied independently in local ecological conditions and spatial proximity. Neutral theory predicts that similarity in communities will be negatively associated with distance between sites, while niche theory suggests that community similarity will be positively associated with similarity in local ecological conditions. 3. Similarity in total invertebrate, grazer and predator assemblages showed negative relationships with distance and, for grazers and predators, positive relationships with local ecological conditions. However, the best model predicting community similarity in all three cases included aspects of both local ecological conditions and distance between sites. 4. When assemblages were analysed according to dispersal ability, high-dispersal species were shown to be freely accessing all sites and community similarity was not well predicted by either local ecology or spatial separation. Assemblages of species with low and moderate dispersal ability were best predicted by combined models, including distance between sites and local ecological factors. 5. The results suggest that the perceived dichotomy between neutral and local environmental processes in determining local patterns of diversity may not be useful. Neutral and niche processes structured these communities differentially depending on trophic level and species traits. 6. We emphasize the potential for both dispersal processes and local environmental conditions to explain local patterns of diversity.     

可利用资源波动对外来种入侵的抵抗性

生物入侵带来的生态和经济危害引起了人们的广泛关注。在入侵生态学研究方面,生物多样性与生物入侵之间的关系长久以来成为群落可入侵性探讨的焦点。Elton经典假说认为,物种多样性越高对外来种入侵的抵抗能力越强,许多模型或野外试验都支持这一假说。但现在越来越多的试验对此提出了异议,各种假说纷纷出现。究竟生物多样性会不会影响外来种的入侵?假设两种不同的群落结构(功能群),设计6种外来种入侵土著群落的情景分析不同多样性及相同多样性下外来种的入侵状况。结果发现,在多样性相同的情况下,两种群落对外来种入侵的抵抗力不同。外来种成功入侵等比群落,却被倍数群落排斥在系统之外。进一步分析表明这主要是由于可利用资源的波动引起的,即Davis提出的"资源机遇假说"。在相同的物种多样性下,由于倍数群落的特殊结构,整个群落所占有资源远远大于等比群落资源比率。因此,外来种在等比群落中更易找到合适的入侵机会。而在物种多样性不同的情况下,由于物种多样性与已占有资源的变化是成正比的,因此,混淆了多样性与剩余资源可利用性对外来种入侵的影响。     

植物群落的生物多样性及其可入侵性关系的实验研究

 生物入侵已经成为一个普遍性的环境问题,并为许多学者所关注。尽管一些理论研究和观察表明生物多样性丰富的群落不容易受到外来种的入侵,但后来有些实验研究并没能证实两者的负相关性,多样性 可入侵性假说仍然是入侵生态学领域争论比较多的一个焦点。人为构建不同物种多样性和物种功能群多样性（C3 禾本科植物、C4植物、非禾本科草本植物和豆科植物）梯度的小尺度群落,把其它影响可入侵性的外在因子和多样性效应隔离开来,研究入侵种喜旱莲子草(Alternanthera philoxeroides)在不同群落里的入侵过程来验证多样性 可入侵性及其相关假说。研究结果显示,物种功能群丰富的群落可入侵程度较低,功能群数目相同而物种多样性不同的群落可入侵性没有显著性差异,功能群特征不同的群落也表现出可入侵性的差异,生活史周期短的单一物种群落和有着生物固氮功能的豆科植物群落可入侵程度较高,与喜旱莲子草属于同一功能群且有着相似生态位的土著种莲子草(A. sessilis)对入侵的抵抗力最强。实验结果表明,物种多样性和群落可入侵性并没有很显著的负相关,而是与物种特性基础上的物种功能群多样性呈负相关,群落中留给入侵种生态位的机会很可能是决定群落可入侵性的一个关键因子。     

Niche separation and weak interactions in the high tidal zone of saltmarsh‐mangrove mixing communities

1. Saltmarsh‐mangrove ecotones occur at the boundary of the natural geographic distribution of mangroves and salt marshes. Climate warming and species invasion can also drive the formation of saltmarsh‐mangrove mixing communities. How these coastal species live together in a “new” mixed community is important in predicting the dynamic of saltmarsh‐mangrove ecosystems as affected by ongoing climate change or human activities. To date, the understanding of species interactions has been rare on adult species in these ecotones.
2. Two typical coastal wetlands were selected as cases to understand how mangrove and saltmarsh species living together in the ecotones. The leaves of seven species were sampled from these coastal wetlands based on their distribution patterns (living alone or coexisting) in the high tidal zone, and seven commonly used functional traits of these species were analyzed.
3. We found niche separation between saltmarsh and mangrove species, which is probably due to the different adaptive strategies they adopted to deal with intertidal environments.
4. Weak interactions between coexisting species were dominated in the high tidal zone of the two saltmarsh‐mangrove communities, which could be driven by both niche differentiation and neutral theory.
5. Synthesis. Our field study implies a potential opportunity to establish a multispecies community in the high tidal zone of saltmarsh‐mangrove ecotones, where the sediment was characterized by low salinity and high nitrogen.

     

Taxa-area relationship and neutral dynamics influence the diversity of fungal communities on senesced tree leaves

This study utilized individual senesced sugar maple and beech leaves as natural sampling units within which to quantify saprotrophic fungal diversity. Quantifying communities in individual leaves allowed us to determine if fungi display a classic taxa–area relationship (species richness increasing with area). We found a significant taxa–area relationship for sugar maple leaves, but not beech leaves, consistent with Wright's species‐energy theory. This suggests that energy availability as affected plant biochemistry is a key factor regulating the scaling relationships of fungal diversity. We also compared taxa rank abundance distributions to models associated with niche or neutral theories of community assembly, and tested the influence of leaf type as an environmental niche factor controlling fungal community composition. Among rank abundance distribution models, the zero‐sum model derived from neutral theory showed the best fit to our data. Leaf type explained only 5% of the variability in community composition. Habitat (vernal pool, upland or riparian forest floor) and site of collection explained > 40%, but could be attributed to either niche or neutral processes. Hence, although niche dynamics may regulate fungal communities at the habitat scale, our evidence points towards neutral assembly of saprotrophic fungi on individual leaves, with energy availability constraining the taxa–area relationship.     

On the balance between niche and neutral processes as drivers of community structure along a successional gradient: insights from alpine and sub-alpine meadow communities

BACKGROUND AND AIMS: Neutral theory predicts that the diversity and relative abundance of species in ecological communities do not depend on their specific traits. This prediction remains controversial, as many studies suggest that variations in the niches of species determine the structure of communities. The aim of this study was to test empirically the relative importance of niche and neutral processes as drivers of species abundance within plant communities along a successional gradient. METHODS: Information on the abundance (density and frequency) and traits (aboveground individual biomass and seed mass) of > 90 species was collected in alpine and sub-alpine meadows of the Tibet Plateau (China). A successional gradient (1, 3, 15 and 30 years after abandonment) was established in a sub-alpine meadow. The relationships between species traits and their abundance were evaluated using regression models. KEY RESULTS: Seed mass was negatively related to both species density (r = -0.6270, P < 0.001) and frequency (r = -0.5335, P = 0.005) in the 1-year meadow. Such relationships disappeared along the successional gradient evaluated (P > 0.07 in the 3-, 15- and 30-year meadows). Data gathered in all sites showed a significant negative relationship between the average individual biomass of a given species and its density within the community (r < -0.30, P < 0.025 in all cases). CONCLUSIONS: The results show that seed mass was a key driver of species abundance in early successional communities, and that niche forces may become more important as succession progresses. They also indicate that predictions from neutral theory, in its current form, do not hold for the meadow communities studied.     

The interpretation of biological surveys

Biological surveys provide the raw material for assembling ecological patterns. These include the properties of parameters such as range, abundance, dispersion, evenness and diversity; the relationships between these parameters; the relationship between geographical distributions and landscape structure; and the co-occurrence of species. These patterns have often been used in the past to evaluate the role of ecological processes in structuring natural communities. In this paper, I investigate the patterns produced by simple neutral community models (NCMs) and compare them with the output of systematic biological surveys. The NCM generates qualitatively, and in some cases quantitatively, the same patterns as the survey data. It therefore provides a satisfactory general theory of diversity and distribution, although what patterns can be used to distinguish neutral from adaptationist interpretations of communities, or even whether such patterns exist, remains unclear.     

Responses of species abundance distribution patterns to spatial scaling in subtropical secondary forests

To quantify and assess the processes underlying community assembly and driving tree species abundance distributions(SADs) with spatial scale variation in two typical subtropical secondary forests in Dashanchong state‐owned forest farm, two 1‐ha permanent study plots (100‐m × 100‐m) were established. We selected four diversity indices including species richness, Shannon–Wiener, Simpson and Pielou, and relative importance values to quantify community assembly and biodiversity. Empirical cumulative distribution and species accumulation curves were utilized to describe the SADs of two forests communities trees. Three types of models, including statistic model (lognormal and logseries model), niche model (broken‐stick, niche preemption, and Zipf‐Mandelbrodt model), and neutral theory model, were estimated by the fitted SADs. Simulation effects were tested by Akaike's information criterion (AIC) and Kolmogorov–Smirnov test. Results found that the Fagaceae and Anacardiaceae families were their respective dominance family in the evergreen broad‐leaved and deciduous mixed communities. According to original data and random sampling predictions, the SADs were hump‐shaped for intermediate abundance classes, peaking between 8 and 32 in the evergreen broad‐leaved community, but this maximum increased with size of total sampled area size in the deciduous mixed community. All niche models could only explain SADs patterns at smaller spatial scales. However, both the neutral theory and purely statistical models were suitable for explaining the SADs for secondary forest communities when the sampling plot exceeded 40 m. The results showed the SADs indicated a clear directional trend toward convergence and similar predominating ecological processes in two typical subtropical secondary forests. The neutral process gradually replaced the niche process in importance and become the main mechanism for determining SADs of forest trees as the sampling scale expanded. Thus, we can preliminarily conclude that neutral processes had a major effect on biodiversity patterns in these two subtropical secondary forests but exclude possible contributions of other processes.     

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.     

洲际入侵植物生态位稳定性研究进展

人类活动引起的大规模洲际物种交换与生物入侵, 改变了当地生态系统结构与功能, 使生物多样性受到日益严重的威胁。本文通过综合分析主要国家和地区入侵植物的地理起源, 发现洲际入侵主要包括东亚—北美、东亚—南美、欧洲—南非、欧洲—北美、欧洲—东亚、北美—大洋洲等, 这些洲际入侵造成的后果往往比陆内入侵更为严重。利用物种分布模型(SDMs)预测入侵物种潜在分布范围是有效管理和提早预防生物入侵的重要依据, 但这些模型的一个关键假定是: 入侵物种的生态位在空间和时间上是保守的、稳定的。然而, 对于远离原产地种群并能快速适应新生境的洲际入侵植物来说, 生态位可能发生显著的变化。入侵种能否在入侵地保持原有的生态位, 取决于制约其生态分布的限制因素和生态过程在不同地区间是否发生变化。本文中作者总结了洲际入侵与陆内入侵的生态与进化过程的异同点, 认为这些限制物种原产地分布的因素如扩散限制、种间互作、适应性进化、生态可塑性和种群遗传特性等均可能导致入侵物种生态位的改变。建议下一步的研究应该重视: (1)对生态位属性进行多尺度的研究, 包括时间、空间、环境或系统发育等几个方面; (2)对比生态位稳定与发生偏移的物种特性, 确定什么样的入侵物种更容易改变原有的生态位; (3)进行生态位时间动态格局研究, 探讨生态位变化的倾向、历史速率和偏移程度, 以便判定生态位变化趋势。这些研究结果将会进一步提高物种分布模型的预测能力, 有助于更为准确地揭示气候变化和物种入侵对生物多样性的影响。     

Slight differences among individuals and the unified neutral theory of biodiversity

The unified neutral theory of biodiversity provides a very simple and counterintuitive explanation of species diversity patterns. By specifying speciation, community size and dispersal, and completely ignoring differences among individual organisms and species, it generates biodiversity patterns that remarkably resemble natural ones. Here I show that adding even slight differences among organisms generates very different patterns and predictions. In large communities with widespread dispersal, heritable differences in viability among individual organisms lead to biodiversity patterns characterised by the overdominance of a single species comprising organisms with relatively high fitness. In communities with local dispersal, the same differences produce rapid community extinction. I conclude that the unified neutral theory is not robust to slight deviations from its most controversial assumption.