草地群落物种多样性维持机制的研究（Ⅱ）：物种实现生态位

尽管为解释种类丰富的植物群落物种共存和多样性维持机制,生态学家们做了大量的努力并提出了许多假说和模型,但这一问题仍处在争议之中,需要更多的证据支持他们的观点或提出新的看法,使这一生物多样性难题不断地向前推进。以松嫩平原物种丰富度较高的羊草—杂类草群落为对象,在土壤C、N、P、K和H2O等5个资源轴上,探讨了物种多样性与实现生态位的关系。结果表明：尽管物种生态位存在一定程度的分化,但多数物种的生态位是高度重叠的,物种生态位的分化在草地群落物种共存和多样性维持中,不是唯一的途径,认为应更加重视物种在长期协同进化中所形成的生物学特性。     

物种共存理论研究进展

群落内的多物种如何共存是群落生态学的核心研究内容之一。经典的物种共存理论强调物种之间的生态位分化,注重具体共存机制的研究。这种以具体共存机制为研究对象的方法一定程度上促进了当代物种共存理论框架的形成。在当代物种共存理论框架下,物种间的差异被划分为两类综合性的抽象差异——生态位差异和平均适合度差异,前者促进物种共存,对应稳定化机制;后者导致竞争排除,对应均等化机制。本文在简要回顾经典物种共存理论的基础上,介绍了当代物种共存理论的框架(包括理论的形成和定义)、基于该理论的部分实验验证工作及其在一些重要生态学问题中的应用。当代物种共存理论不仅揭示了群落内物种是如何共存的这一基本理论问题,更重要的是在全球变化的背景下该理论对生物多样性的保护和管理具有重要的应用价值。期望本文的介绍有助于国内生态学和生物多样性工作者了解当代物种共存理论,并将其应用于群落构建和生物多样性维持机制等方面的研究。     

群落构建的中性理论和生态位理论

物种共存和生物多样性维持一直是生态学研究的中心论题。基于物种生态位分化的群落构建理论已经发展了近一个世纪, 但我们对群落构建和生物多样性维持的机理仍然不清楚。近年来, 群落中性理论以其简约性和预测能力成为群落生态学研究的焦点, 但由于其“物种在生态功能上等价”的假设与大量研究结果相悖, 同时对自然群落结构的准确预测也只限于少数的生态系统, 因而饱受质疑。如今, 越来越多的生态学家认为群落构建的生态位理论与中性理论之争的最终归宿应该是二者的整合。 在本文中, 我们在简要回顾生态位理论和群落中性理论发展的基础上, 分析二者之间的主要分歧和互补性, 试图梳理二者整合的途径。我们认为, 尽管中性理论的发展极大地丰富了群落构建理论, 但二者的整合尚处于初级阶段; 群落构建零模型假说、中性—生态位连续体假说、随机生态位假说等都不失为有价值的尝试, 今后需要在其他类型的生态系统中进行实验验证, 以更好地理解确定性过程和随机过程在决定群落构建和生物多样性维持中的作用。     

天童常绿阔叶林中常绿与落叶物种的物种多度分布格局

物种多度分布是对群落内不同物种多度情况的数量描述, 作为理解群落性质的基石, 其形成机制受到广泛关注。常绿与落叶物种是两类有着不同物候性状与生长策略的物种集合, 它们普遍共存于常绿阔叶林中。在天童20 ha常绿阔叶林动态监测样地内, 虽然常绿物种在物种多度和胸高断面积等指标上占有绝对优势, 但其在物种丰富度上却不及落叶物种。分析两者在常绿阔叶林中的物种多度分布特征, 能够为理解常绿阔叶林内物种多样性的维持机制提供一个全新的视角。为此, 我们基于天童样地的植被调查数据, 一方面利用累积经验分布函数对两类生活型植物的物种多度分布进行描述, 使用Kolmogorov-Smirnov检验(K-S检验)判断其差异性; 另一方面, 采用纯统计模型、生态位模型和中性理论模型对二者的物种多度分布曲线进行拟合, 并基于K-S检验的结果以及AIC值进行最优模型的筛选。结果显示: (1)常绿与落叶物种的物种多度分布曲线间并无显著差异。(2)在选用的3类模型中, 中性理论模型对于两类物种多度分布曲线的拟合效果都最好, 而生态位模型的拟合效果则一般。从上述结果可以看出, 尽管常绿与落叶物种在物种数量和多度等方面均存在差异, 但它们却有着近似的物种多度分布格局以及相近的多样性维持机制。然而, 鉴于模型拟合的结果只能作为理解群落多样性构建机制的必要非充分条件, 故而只能初步判定中性过程对于常绿与落叶物种的物种多样性格局影响更大, 却不能排除或衡量诸如生态位分化等其他过程在两类生活型多样性格局形成中的贡献。     

物种多样性与植物群落的维持机制

物种多样性的形成与维持是生物多样性研究中的核心问题之一,形成了许多假说。剖析了群落物种多样性维持具有增加、减少和稳定共存三方面内容,着重阐明物种稳定共存的内因是物种的生物学与生态学特性差异,外因是生境具有小尺度的差异;群落生境异质性是其物种多样性维持的基础,并影响着群落的组织过程;群落的斑块镶嵌结构源于林隙形成的生境异质性,导致了物种多样性;群落斑块镶嵌结构是群落的基本属性,使群落的结构和功能得以长期维持等论点。最后,还以鼎湖山季风常绿阔叶林为例,对群落斑块镶嵌结构的形成及其对物种多样性维持和群落生产力构成与维持等方面的意义予以说明。     

荒漠草原“土岛”生境群落物种共存机制

经过长期破碎化,荒漠草原原生硬质灰钙土斑块散布在广大沙化土地中,形成类似"土岛"的土被结构。为揭示土岛生境的群落物种共存机制,2016年在宁夏盐池县皖记沟村选取大(200–300 m~2)、中(约100 m~2)、小(约50 m~2)土岛各3个开展调查,采用Jaccard相异系数、物种生态位宽度和生态位重叠度、零模型、Meta分析,综合计算和分析土岛内部与外部植物群落相似性、物种生态位宽度和生态位重叠、物种共存格局及其影响因子。研究发现,(1)随着破碎化加剧,土岛内部植物多样性整体呈现下降趋势,群落优势种从短花针茅(Stipa breviflora)转变为猪毛蒿(Artemisia scoparia)和短花针茅共优种,土岛内外群落相似性增加。(2)土岛内外绝大多数物种生态位重叠较小,生态位重叠在土岛内呈集中分布,而土岛外则呈均匀发散分布。(3)环境过滤为主的生态过程决定了土岛生境群落物种的共存格局,随着土岛面积减小,环境因子对群落物种共存的调控强度降低,关键性环境因子由土壤细砂粒和黏粒转变为粗砂粒,显著性竞争物种共存格局在小岛出现。综上所述,土岛生境对于维持草原物种具有重要作用,环境过滤主导了荒漠草原物种共存格局。随着生境破碎化加剧或土岛面积减小,物种共存格局及其调控因子发生转变。保护面积在200 m~2以上的大土岛对于恢复荒漠草原区草原成分种和其物种多样性机制都十分必要。     

甘肃祁连山脉雪豹及其同域分布大型食肉动物时间生态位关系

<正>群落内多物种如何共存是群落生态学和生物多样性研究的核心内容之一。经典物种共存理论强调物种之间的生态位分化,侧重于物种对环境的需求,Hutchinson (1957)提出超体积生态位概念,认为物种适合度是由多个因素共同决定,即物种只有在满足其生态位需求的多维空间,     

群落内物种多样性发生与维持的一个假说

本文根据作者对竞争排除法则的研究而提出了一个新的群落多样性假说。按照作者的观点,占用相同生态位的物种可以稳定共存;这样,群落内物种多样性将受到４个基本因子所控制。它们分别是：（Ⅰ）生态位的数量;（Ⅱ）区域物种库的大小;（Ⅲ）物种迁入速率,以及（Ⅳ）物种灭绝速率。该假说强调区域生物地理过程与局域生态过程共同决定了群落内种多样性的大小及分布模式。     

米槁群落主要物种资源利用特征分析

开展米槁群落物种生态位研究,可了解米槁群落物种对资源的利用特征,为米槁的保护与管理提供科学依据。基于贵州省米槁天然林群落的实际调查数据,通过生态位宽度、生态位重叠指数,分析该群落主要物种的资源利用特征。群落中米槁(Cinnamomum migao)、杜茎山(Maesa japonica)、菝葜(Smilax china)的生态位宽度最大,在群落中处于明显优势地位,具有较强竞争力和资源利用能力;生态位宽度越大的物种间生态位重叠越大,但在相同或相似环境条件下,生态位宽度值小的种对也会拥有较高的生态位重叠;米槁群落主要物种间的生态位重叠指数偏低,说明物种间对资源利用的相似性较低,物种的生态位分化明显,竞争不激烈,群落结构较为稳定,物种间能够较好地相互共存。     

水淹干扰下水库水滨带植物群落谱系结构与物种多样性格局

群落谱系结构将物种系统发育信息整合到群落中，给生物多样性研究提供了全新视角。为探讨水淹干扰下水库水滨带植物群落物种多样性维持机制，以丹江口水库水滨带为研究区域，以南水北调中线一期工程运行前的原有水滨带植被为研究对象，分析不同水淹干扰强度下群落谱系结构和物种多样性及其与环境因素的关系。结果表明：(1)随着水淹干扰强度增加，群落物种丰富度、物种多样性和谱系多样性显著降低。谱系结构指数虽无显著差异，但随着水淹干扰强度增加呈现由发散到聚集的变化趋势。表明水滨带植物群落物种多样性维持机制是生境过滤和生态位分化共同驱动的结果，并且随着水淹干扰强度增加呈现生境过滤作用增加、物种间竞争排斥作用减弱的生态过程影响规律。(2)坡度、土壤质地、速效氮和全氮含量对整个水滨带植物群落物种丰富度、物种多样性和谱系多样性影响显著。此外，土层厚度和速效磷对物种丰富度和土层厚度对谱系多样性也影响显著。而仅有速效氮含量对谱系结构指数影响显著；轻度水淹干扰下坡向、速效磷含量与多样性指数显著相关；中度水淹干扰下坡度、土层厚度、土壤质地、土壤总氮含量与多样性指数显著相关，土壤pH、全氮和全磷含量与谱系结构指数显著相关；重度水淹...     

Coexistence and emergent neutrality generate synchrony among competitors in fluctuating environments

Many competitive communities exhibit a puzzling amount of species diversity. In this study, we model a community of symmetric competitors in a fluctuating environment. We use biologically realistic temperature-dependent growth curves with a widely hypothesized trade-off between maximum growth and nice breadth to control the shapes of the curves of different species. We perform three analyses of the community dynamics to investigate the role of environmental fluctuations in community composition and species diversity. We initiate communities with equal abundances of all species and randomize the temperature fluctuations so that there is no correlation between species responses, only noise. We initiate single populations and allow other species to randomly invade the community. We also knock out extant species one by one from an established community and allow them to reinvade after the remaining species have adjusted. We find that competitors with sufficiently different temperature niches coexist via temporal niche differentiation. We also find long-term persistence of species that are very similar to a dominant competitor. This creates communities with species clumped along a temperature niche axis, with stable coexistence between groups and near neutrality within groups. The near neutrality results in interspecific synchrony within the groups, providing an explanation for the maintenance of high diversity in competitive communities where synchrony is commonly observed.     

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.     

Dietary and Temporal Niche Differentiation in Tropical Ants—Can They Explain Local Ant Coexistence?

How species with similar ecological requirements avoid competitive exclusion remains contentious, especially in the species‐rich tropics. Niche differentiation has been proposed as a major mechanism for species coexistence. However, different niche dimensions must be studied simultaneously to assess their combined effects on diversity and composition of a community. In most terrestrial ecosystems, ants are among the most abundant and ubiquitous animals. Since they display direct, aggressive competition and often competitively displace subordinate species from resources, niche differentiation may be especially relevant among ants. We studied temporal and trophic niche differentiation in a ground ant community in a forest fragment in French Guiana. Different baits were presented during day and night to assess the temporal and dietary niches of the local species. They represented natural food resources such as sugars, carrion, excrements, seeds, and live prey. In addition, pitfalls provided a background measure of ant diversity. The communities attracted to the different baits significantly differed from each other, and even less attractive baits yielded additional species. We detected species specialized on living grasshoppers, sucrose, seeds, or dead insects. Community‐level differences between day and night were larger than those between baits, and many species were temporally specialized. In contrast to commonness, foraging efficiency of species was correlated to food specialization. We conclude that many ant species occupy different temporal or dietary niches. However, for many generalized species, the dietary, and temporal niche differentiation brought forward through our sampling effort, cannot alone explain their coexistence.     

Evidence that niche specialization explains species-energy relationships in lake fish communities

1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.     

Climatic niche partitioning following successive invasions by fruit flies in La Réunion

1. Biological invasions have profound effects on community structure. The community composition following invasions can be influenced by the habitat diversity and the species' responses to abiotic factors. 2. We evaluated the tolerance to climatic factors and analysed the field distribution of four polyphagous fruit flies (Diptera: Tephritidae) of La Réunion Island (three exotic species that successively invaded the island and the endemic species Ceratitis catoirii) in order to evaluate the opportunities of coexistence by niche differentiation. 3. Atmospheric humidity and immersion in water in the laboratory greatly influence the survival of fruit fly pupae. While C. catoirii and C. rosa are very sensitive to desiccation, C. capitata and especially Bactrocera zonata are relatively tolerant. B. zonata also tolerated immersion in water much longer than did C. rosa and C. catoirii, that in turn were more resistant than C. capitata. Overall, field distributions agree with the predictions based on this study of humidity combined with previous data on the effects of temperature. 4. Climatic niche partitioning promotes coexistence between some but not all pairs of invasive species. Thus, C. rosa can coexist with both C. capitata and B. zonata at the regional scale, while climatic niches are not different enough to promote coexistence of the latter two species. The endemic species has no private climatic niche either and this now very rare species could be in the process of extinction. 5. By promoting coexistence or not, climatic diversity in invaded areas can directly affect the community composition following invasions.     

Stable isotope analyses reveal dense trophic species packing and clear niche differentiation in a malagasy primate community

Understanding the mechanisms maintaining local species richness is a major topic in tropical ecology. In ecological communities of Madagascar, primates represent a major part of mammalian diversity and, thus, are a suitable taxon to study these mechanisms. Previous research suggested that ecological niche differentiation facilitates the coexistence of lemurs. However, detailed data on all species making up diverse local primate assemblages is rarely available, hampering community‐wide tests of niche differentiation among Malagasy mammals. Here, we took an indirect approach and used stable isotopes as long‐term indicators of individuals' diets to answer the question of whether trophic patterns and food‐related mechanisms stabilize coexistence in a species‐rich lemur community. We analyzed stable carbon and nitrogen isotopes in hair collected from eight syntopic lemurs in Kirindy Forest. We found that lemur species were well separated into trophic niches and ranged over two trophic levels. Furthermore, species were densely packed in isotopic space suggesting that past competitive interactions between species are a major structuring force of this dry forest lemur community. Results of other comparative studies on primates and our findings underline that—in contrast to communities worldwide—the structure and composition of lemur communities follow predictions of ecological niche theory. Patterns of competitive interactions might be more clearly revealed in Malagasy primate communities than elsewhere because lemurs represent a large fraction of ecologically interacting species in these communities. The pronounced trophic niche differentiation among lemurs is most likely due to intense competition in the past as is characteristic for adaptive radiations. Am J Phys Anthropol 153:249–259, 2014. © 2013 Wiley Periodicals, Inc.     

How variation between individuals affects species coexistence

Although the effects of variation between individuals within species are traditionally ignored in studies of species coexistence, the magnitude of intraspecific variation in nature is forcing ecologists to reconsider. Compelling intuitive arguments suggest that individual variation may provide a previously unrecognised route to diversity maintenance by blurring species‐level competitive differences or substituting for species‐level niche differences. These arguments, which are motivating a large body of empirical work, have rarely been evaluated with quantitative theory. Here we incorporate intraspecific variation into a common model of competition and identify three pathways by which this variation affects coexistence: (1) changes in competitive dynamics because of nonlinear averaging, (2) changes in species’ mean interaction strengths because of variation in underlying traits (also via nonlinear averaging) and (3) effects on stochastic demography. As a consequence of the first two mechanisms, we find that intraspecific variation in competitive ability increases the dominance of superior competitors, and intraspecific niche variation reduces species‐level niche differentiation, both of which make coexistence more difficult. In addition, individual variation can exacerbate the effects of demographic stochasticity, and this further destabilises coexistence. Our work provides a theoretical foundation for emerging empirical interests in the effects of intraspecific variation on species diversity.     

Shade tolerance,canopy gaps and mechanisms of coexistence of forest trees

The belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche differentiation in response to gaps has focused on evidence for an interspecific tradeoff between low light survival and high light growth. The objectives of this study are first to distinguish the possible mechanisms allowing coexistence based on this tradeoff, and second, to explore their limitations. We present a theory of forest dynamics driven by small‐scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non‐linearity’, ‘the successional niche’ and the ‘storage effect’. We find that temporal fluctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. The rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.     

The importance of niche differentiation for coexistence on large scales

It is widely accepted that niche differentiation plays a key role in coexistence on relatively small scales. With regard to a large community scale, the recently propounded neutral theory suggests that species abundances are more influenced by history and chance than they are by interspecies competition. This inference is mainly based on the probability that competitive exclusion is largely slowed by recruitment limitation, which may be common in species rich communities. In this respect, a theoretical study conducted by Hurtt and Pacala (1995) for a niche differentiated community has been frequently cited to support neutral coexistence. In this paper, we focused on the effect of symmetric recruitment limitation on delaying species competitive exclusion caused by both symmetric and asymmetric competition in a large homogeneous habitat. By removing niche differentiation in space, we found that recruitment limitation could delay competitive exclusion to some extent, but the effect was rather limited compared to that predicted by Hurtt and Pacala's model for a niche differentiated community. Our results imply that niche differentiation may be important for species coexistence even on large scales and this has already been confirmed in some species rich communities.     

Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.