生物间高阶相互作用研究进展

生物间的相互作用是物种共存和生物多样性维持的关键。传统的物种共存研究主要关注配对物种之间的直接相互作用, 而忽略了更为复杂的间接相互作用。本文首先介绍了两种间接相互作用: 链式相互作用(本质上仍是两两物种之间的相互作用)和高阶相互作用。在此基础上, 我们回顾了高阶相互作用定义的演变历史(包括狭义的高阶相互作用和广义的高阶相互作用)及其检验方法, 并介绍了高阶相互作用在多营养级之间和同一营养级内的研究概况。目前, 生态学家主要对多营养级之间(如食物网)的高阶相互作用的特征、发生机制、作用途径及实验证据等方面进行了详尽的研究。近年来, 同一营养级内的高阶相互作用也开始受到关注, 因此我们进一步介绍了同一营养级内个体水平高阶相互作用的重要意义和度量方法。从个体水平上研究高阶相互作用, 既能统一狭义和广义高阶相互作用在定义上的争议, 又可以将个体间的差异(如个体大小、个体的空间分布等信息)考虑进来。最后, 本文对高阶相互作用一些可能的重要研究方向进行了展望: 在自然群落中(尤其同一营养级内)检验高阶相互作用的普遍性与相对重要性, 探讨高阶相互作用的发生机制以及如何将高阶相互作用整合到现有的理论体系中等。高阶相互作用的研究有助于我们全面深刻地理解物种共存和生物多样性的维持机制, 丰富和完善群落生态学的理论框架, 为人类世背景下的生物多样性保护和生态系统功能维持与提升提供基础。     

生物多样性理论最新进展

生物多样性是生态系统复杂性的重要特征, 理解多样性的形成和维持机制一直是理论生态学研究的核心议题。本文从三方面概述了生物多样性理论的最新进展。一是物种共存和群落构建, 总结了现代共存理论和基于过程的群落构建理论的新进展。二是物种相互作用, 综述了利用经验数据推断物种相互作用关系和强度的最新方法。三是生态-进化动态, 介绍了生态-进化模型的一般框架及其在生物多样性研究中的应用。最后对生物多样性理论的发展趋势做了展望, 特别是多尺度整合理论和全球变化下的预测理论。     

植物群落稀有种维持机制与土壤反馈的研究进展

自Janzen-Connell (J-C)假说提出后半个世纪以来, 生态学家在热带及亚热带森林对该假说开展的大量实证研究表明, 由专性天敌导致的J-C效应所引起的负密度制约是维持森林多样性和决定群落组成的重要驱动力, 该假说成功地解释了热带及亚热带森林的丰富多样性。土壤病原真菌所引起的植物-土壤负反馈是J-C效应最主要的表现形式。然而, 对于植物-土壤负反馈是否能够维持森林群落中的大量稀有种仍然存在许多争议。基于当代物种共存理论的“稀有种优势”假说认为, 只有在满足“可入侵准则” (即物种在稀有时具有种群增加的趋势)的前提下, 稀有种才能在群落中与其他物种长期共存。然而, 当前基于土壤反馈的实验结果与该理论预测相悖, 因此在稀有种的维持机制方面仍存在较大的分歧。本文通过介绍植物-土壤反馈理论, 整合了可能对稀有种维持有较大影响的因素, 包括共生菌根真菌、土壤养分以及植物细根性状等在影响土壤负反馈方面的相关研究, 并对这些因素如何影响群落中物种多度和稀有种在群落中的维持进行了探讨。最后, 我们也从其他角度探讨了一些对稀有种维持的研究。我们认为在未来对稀有种的研究中, 探讨使其长期存续的“优势”和制约其种群扩大的“限制”同等重要, 将当代物种共存理论与新技术、新方法相结合对于探究稀有种的维持机制具有重要的意义, 可为稀有种保护提供理论依据。     

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。     

复杂性-稳定性研究: 数学模型的进展

对自然生态系统的观察给人们以复杂的群落更稳定的直观印象, 但数学模型却得出了截然相反的结论。这一“悖论”使得复杂性-稳定性研究自20世纪70年代以来成为长期的热点。本文对这一领域的数学模型研究进行简要综述。首先对这一论题进行概念剖析, 然后将各类模型分为线性和非线性两大类, 前者即群落矩阵法, 后者包括相互作用矩阵法、复杂网络数值模拟法和食物网构件动力学法。它们分别基于不同的群落构建方法和稳定性判断标准, 探求各物种是如何相互作用并实现共存的。总体而言, 在随机构建的群落模型中, 多样性和连接度的增长不利于系统稳定; 而在更接近真实自然群落的模型中, 相互作用方式、网络拓扑结构、相互作用强度分布等方面的机制提供了稳定效应, 按此组织的生态网络可达到很高的复杂度。然而, 复杂性-稳定性的研究还远未结束, 当前的模型仍不足以反映自然群落中的复杂相互作用, 稳定性的概念也有待拓展。对这一议题的深入研究在生态学理论和生态系统管理实践方面都具有重大价值。     

喜马拉雅山哺乳动物物种多样性垂直分布格局

生物多样性的空间分布及其相关机制一直是生态学、生物地理学和保护生物学研究的热点问题。山地生态系统生境异质性和生物多样性高, 适合研究生物多样性空间分布及其相关机制。喜马拉雅山脉位于青藏高原南缘, 是全球生态热点区域。其地形复杂, 海拔落差大(100-8,844 m), 具有明显的垂直气候带。本研究通过整合野外调查和文献资料, 系统地分析了10目23科160属313种喜马拉雅山地区哺乳动物物种多样性的垂直分布格局, 发现该区域哺乳动物总体及其子集的物种多样性垂直分布格局都为左偏倚的中峰格局, 物种多样性在海拔900-1,400 m之间最高, 不同物种子集的物种多样性垂直分布格局的模式有所不同。UPGMA聚类分析表明, 喜马拉雅山地区哺乳动物群落沿海拔梯度可以划分为5个聚类簇(海拔100-1,500 m、1,500-2,000 m、2,000-3,000 m、3,000-4,200 m以及4,200-6,000 m的地区), 大致与该地区植被的垂直带分布相吻合。喜马拉雅山地区哺乳动物物种多样性在中低海拔最为丰富, 可能跟东洋界与古北界生物群扩散后的交汇地带相关。喜马拉雅山区贯通南北的沟谷是生物扩散和迁移的通道, 沟谷内水热资源较好, 气候稳定性高, 为高山生态系统内各种生物创造了栖息条件。综上, 喜马拉雅山沟谷地区是生物多样性热点地区, 也是生物扩散和交流关键的“生态走廊”, 应加强对喜马拉雅山沟谷地区的保护, 以维系该区域较高的生物多样性。     

种间互作网络的结构、生态系统功能及稳定性机制研究

生态群落中不同物种间发生多样化的相互作用, 形成了复杂的种间互作网络。复杂生态网络的结构如何影响群落的生态系统功能及稳定性是群落生态学的核心问题之一。种间互作直接影响到物质和能量在生态系统不同组分之间的流动和循环以及群落构建过程, 使得网络结构与生态系统功能和群落稳定性密切相关。在群落及生态系统水平上开展种间互作网络研究将为群落的构建机制、生物多样性维持、生态系统稳定性、物种协同进化和性状分化等领域提供新的视野。当前生物多样性及生态系统功能受到全球变化的极大影响, 研究种间互作网络的拓扑结构、构建机制、稳定性和生态功能也可为生物多样性的保护和管理提供依据。该文从网络结构、构建机制、网络结构和稳定性关系、种间互作对生态系统功能的影响等4个方面综述当前种间网络研究进展, 并提出在今后的研究中利用机器学习和多层网络等来探究环境变化对种间互作网络结构和功能的影响, 并实现理论和实证研究的有效整合。     

中国森林生物多样性监测网络有力支撑生物群落维持机制研究

正物种共存机制是当前群落生态学的研究热点之一,生态学家提出了众多理论和假说来解释某一特定群落中的物种如何共存.大型森林动态样地途径为研究物种共存机制提供了独特的研究平台,实地观测、控制实验及建模则为生态学家提供了量化共存机制的工具.为了推动中国的森林生物多样性维持机制研究,中国科学院生物多样性委员会于2004年组织有关研究所的科研人员和院外相关单位的合作者,参照全球森     

次生常绿阔叶林的群落结构与物种组成:基于浙江乌岩岭9 ha森林动态样地

常绿阔叶林是我国亚热带地区的地带性植被, 但由于长期的人为干扰, 目前仅有少量分布且主要以次生常绿阔叶林的形式存在。因此, 了解次生常绿阔叶林的物种共存机制对于保护森林生物多样性非常重要。基于大型动态监测样地对森林的物种组成及群落结构进行研究, 是揭示生物多样性维持机制的重要手段。按照美国史密森研究院热带森林科学研究中心(Center for Tropical Forest Science, CTFS)大型森林动态样地建设标准, 于2011-2012年在浙江省乌岩岭国家级自然保护区内建立了1个9 ha森林动态监测样地。通过对样地内胸径≥ 1 cm木本植物的物种组成(如重要值)、群落结构(如区系组成和径级结构等)以及空间分布的分析发现: (1)样地有木本植物存活个体47科92属200种71,396株, 其中壳斗科、樟科、山茶科、冬青科、杜鹃花科和山矾科等占优势; (2)在区系组成上, 热带成分略多于温带成分; (3)群落径级结构接近倒“J”型, 表明群落更新良好, 其中优势种径级结构呈倒“J”型、偏常态型和波动型等各种形态, 且并未表现出种群衰退的趋势; (4)优势种表现出明显的生境偏好和聚集分布格局, 暗示生境异质性可能是影响亚热带次生常绿阔叶林物种空间分布的重要因素。     

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 m²)、中(约100 m²)、小(约50 m²)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 m²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。     

Habitat loss alters effects of intransitive higher-order competition on biodiversity: a new metapopulation framework

Recent studies have suggested that intransitive competition, as opposed to hierarchical competition, allows more species to coexist. Furthermore, it is recognized that the prevalent paradigm, which assumes that species interactions are exclusively pairwise, may be insufficient. More importantly, whether and how habitat loss, a key driver of biodiversity loss, can alter these complex competition structures (and therefore species coexistence) remain unclear. We thus present a new, simple yet comprehensive metapopulation framework that can account for any competition pattern and more complex higher-order interactions (HOIs) among species. We find that competitive intransitivity increases community diversity and that HOIs generally enhance this effect. Essentially, intransitivity promotes species richness by preventing the dominance of a few species, unlike the hierarchical competition, while HOIs facilitate species coexistence through stabilizing community fluctuations. However, variation in species’ vital rates and habitat loss can weaken or even reverse such higher-order effects, as their interaction can lead to a more rapid decline in competitive intransitivity under HOIs. Thus, it is essential to correctly identify the most appropriate interaction model for a given system before models are used to inform conservation efforts. Overall, our simple model framework provides a more parsimonious explanation for biodiversity maintenance than the existing theory.     

热带森林植物多样性及其维持机制

热带森林具有地球上最丰富的植物多样性。关于热带森林植物多样性的维持机制,虽有众多假说,但均未形成完善的理论体系。不同的学者从不同的角度出发得出了许多结论,但也引起了不少争议。本文主要简述了4种经常被用来解释热带森林高植物多样性的机制：生态位分化、取食压力、生活史负相关和随机竞争,每一种机制都有大量的证据支持。热带森林植物沿微环境梯度的非随机空间分布表明其生态位分化很明显,并对其多样性起重要作用。动物的取食降低母树周围同种幼苗的生长率和存活率,为其他物种存活提供了机会,这就是取食压力假说,这是一个极有前景但仍需大量实验验证的假说。生活史负相关使得热带森林的许多植物能够共存。冠层植物的抑制使得随机性在林下植物的建立过程中起决定作用。     

Competition and stoichiometry: coexistence of two predators on one prey

The competitive exclusion principle (CEP) states that no equilibrium is possible if n species exploit fewer than n resources. This principle does not appear to hold in nature, where high biodiversity is commonly observed, even in seemingly homogenous habitats. Although various mechanisms, such as spatial heterogeneity or chaotic fluctuations, have been proposed to explain this coexistence, none of them invalidates this principle. Here we evaluate whether principles of ecological stoichiometry can contribute to the stable maintenance of biodiverse communities. Stoichiometric analysis recognizes that each organism is a mixture of multiple chemical elements such as carbon (C), nitrogen (N), and phosphorus (P) that are present in various proportions in organisms. We incorporate these principles into a standard predator-prey model to analyze competition between two predators on one autotrophic prey. The model tracks two essential elements, C and P, in each species. We show that a stable equilibrium is possible with two predators on this single prey. At this equilibrium both predators can be limited by the P content of the prey. The analysis suggests that chemical heterogeneity within and among species provides new mechanisms that can support species coexistence and that may be important in maintaining biodiversity.     

Biodiversity in model ecosystems, II: species assembly and food web structure

This is the second of two papers dedicated to the relationship between population models of competition and biodiversity. Here, we consider species assembly models where the population dynamics is kept far from fixed points through the continuous introduction of new species, and generalize to such models the coexistence condition derived for systems at the fixed point. The ecological overlap between species and shared preys, that we define here, provides a quantitative measure of the effective interspecies competition and of the trophic network topology. We obtain distributions of the overlap from simulations of a new model based both on immigration and speciation, and show that they are in good agreement with those measured for three large natural food webs. As discussed in the first paper, rapid environmental fluctuations, interacting with the condition for coexistence of competing species, limit the maximal biodiversity that a trophic level can host. This horizontal limitation to biodiversity is here combined with either dissipation of energy or growth of fluctuations, which in our model limit the length of food webs in the vertical direction. These ingredients yield an effective model of food webs that produce a biodiversity profile with a maximum at an intermediate trophic level, in agreement with field studies.     

Temporal resource partitioning mediates vertebrate coexistence at carcasses: The role of competitive and facilitative interactions

Unravelling how biodiversity is maintained despite species competition for shared resources has been a central question in community ecology, and is gaining relevance amidst the current biodiversity crisis. Yet, we have still a poor understanding of the mechanisms that regulate species coexistence and shape the structure of assemblages in highly competitive environments such as carrion pulsed resources. Here, we study how large vertebrates coexist in scavenger assemblages by adapting their diel activity at large ungulate carcasses in NW Spain. We used camera traps to record vertebrate scavengers consuming 34 carcasses of livestock and hunted wild ungulates, which allowed us to assess also differences regarding carcass origin. To evaluate temporal resource partition among species, we estimated the overlap of diel activity patterns and the mean times of each scavenger at carcasses. We recorded 16 species of scavengers, 7 mammals and 9 birds, and found similar richness at both types of carcasses. Birds and mammals showed contrasting diel activity patterns, with birds using carcasses during daytime (mean= 11:38 h) and mammals mostly at night (23:09 h). The unimodal activity patterns of scavengers showed asynchronous peaks among species. Subordinate species modified their activity patterns at carcasses used by apex species to reduce temporal overlap. Also, diel activity patterns of vultures closely followed those of corvids, suggesting facilitation processes in which corvids would enhance carcass detection by vultures. Two mammal species (12.5%) increased nocturnality at carcasses of hunted ungulates, which could be a response to human disturbance. Our results suggest that both temporal segregation and coupling mediate the coexistence of large vertebrates at carcasses. These mechanisms might lead to richer scavenger assemblages and thereby more efficient ones in driving critical ecosystem functions related to carrion consumption, such as energy and nutrient recycling and biodiversity maintenance.     

Competitive intransitivity promotes species coexistence

Using a spatially explicit cellular automaton model with local competition, we investigate the potential for varied levels of competitive intransitivity (i.e., nonhierarchical competition) to promote species coexistence. As predicted, on average, increased levels of intransitivity result in more sustained coexistence within simulated communities, although the outcome of competition also becomes increasingly unpredictable. Interestingly, even a moderate degree of intransitivity within a community can promote coexistence, in terms of both the length of time until the first competitive exclusion and the number of species remaining in the community after 500 simulated generations. These results suggest that modest levels of intransitivity in nature, such as those that are thought to be characteristic of plant communities, can contribute to coexistence and, therefore, community-scale biodiversity. We explore a potential connection between competitive intransitivity and neutral theory, whereby competitive intransitivity may represent an important mechanism for "ecological equivalence."     

Heteromyopia and the spatial coexistence of similar competitors

Most spatial models of competing species assume symmetries in the spatial scales of dispersal and interactions. This makes analysis tractable, and has led to the conclusion that segregation of species in space does not promote coexistence. However, these symmetries leave parts of the parameter space uninvestigated. Using a moment‐approximation method, we present a spatial version of the Lotka–Volterra competition equations to investigate effects of removing symmetries in the distances over which individuals disperse and interact. Some spatial segregation of the species always comes about due to competition, and such segregation does not necessarily lead to coexistence. But, if interspecific competition occurs over shorter distances than intraspecific competition (heteromyopia), spatial segregation becomes strong enough to promote coexistence. Such coexistence is most likely when the species have similar dynamics, in contrast to the competition–colonization trade‐off that requires large competitive differences between species.