集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。     

中国湿地物种多样性与生境面积关系及其生态学机理的模拟研究

生物群落中的物种多样性是群落生态学研究的一个基本问题。大量的实验研究表明物种多样性和生境面积的常数次幂成比例,这说明它们的对数数量级呈线性正相关。我们研究了中国境内15块湿地的高等植物和32块湿地鸟类多样性,发现它们分别和湿地面积在对数尺度上呈线性正相关．进一步支持了种数与面积的幂指数关系。我们还借助计算机模拟系统地讨论了产生这种简单规律的生态学机理,包括中性理论、集合种群动态和物种分布的自相似性。中性理论假设了群落中物种的个体之间只有竞争关系,忽略了其它的种间关系。集合种群动态理论考虑的是由多个亚群落构成的集合群落,在研究种数和面积关系时也忽略了种间关系,所以也是中性的。尽管物种分布的自相似性可导致这种面积幂指数关系,但在自然界中自相似性也可能不成立。     

物种多样性与生态系统功能时间变异性的关系研究进展

近年来物种多样性的急剧丧失使得物种多样性与生态系统功能的时间变异性的关系及其机制问题的研究成为了生态学研究的一个热点。综述了物种多样性与群落集合性质变异性以及种群性质变异性的关系及其机制的最新研究成果：1、理论上探讨造成物种多样性与群落集合性质变异性负相关关系的机制包括：抽样效应、资源利用分化假说、统计平均效应、保险假说、种群变异性的均匀度效应等;但实验研究对理论预期的支持并不是普遍的;2．多样性与种群变异性之间的关系主要依赖于均值-方差尺度系数Z;理论上大部分自然群落是种群变异性应该随着多样性的增加而增加;但有研究表明：在变动环境中多样性对单个组分物种的种群水平有稳定性作用;而经验研究并不能得出多样性对种群变异性效应的清晰模式。讨论了目前的理论和实验研究中存在的和今后研究中需要认真思考的问题。     

应用中性理论分析局域群落中的物种多样性及稳定性

如何解释群落中物种的丰富与稀少,并对物种多样性和群落稳定性进行合理的量化评价是群落生态学研究中的一个热点问题。20世纪中期,MacArthur将影响自然群落稳定性的因素归结为物种数量多少以及物种间相互作用的大小,20世纪末Doak等学者提出群落的容纳能力和物种间的维持机制是决定群落稳定性的关键因素。同时对群落结构及物种间维持机制的研究也有了新的突破,Hubbell提出"生物多样性与生物地理学统一的中性理论(Unified Neutral Theory of Biodiversity and Biogeography)"为群落生态学研究提供了新的思路和方法。从群落中性理论的基本假设出发,对Hubbell中性理论中局域群落的物种多度动态模型进行分析,归纳得出群落中性理论中物种多样性与群落稳定性之间的量化关系。封闭的局域群落中,出现物种灭绝或单物种独占的时间与群落大小及物种相对多度成正比,物种多样性程度的增加可延长物种灭绝或独占的时间;开放的局域群落中,物种多度期望值与局域群落大小、物种在集合群落中的物种相对多度成正比,周围群落中物种的灭绝会引起局域群落中相应物种的灭绝,最终导致整个生态群落物种多样性的降低;群落中物种多度的方差与局域群落大小、迁移率、物种在集合群落中的物种相对多度相关,局域群落物种多度的波动幅度随着群落间生态隔离的减弱或物种多样性程度的增加而减小。由此,集合群落物种多样性是影响局域群落物种多样性的重要因素,生态隔离程度的减弱及物种多样性的增加都将增强群落的稳定性。     

物种保护的理论基础——从岛屿生物地理学理论到集合种群理论

全球面临着生境破碎化的危机,物种保护已成为人类面临的重大课题,并不是所有的人对岛屿生物地理学理论的产生及其关注的海洋岛屿都很熟悉,但是越来越多生物赖以生存的自然栖息地的丧失和破碎化都是有目共睹的,岛屿生物地理学和集合种群理论是目前物种保护的两个基本理论,物种迁入率和绝灭率的动态变化决策岛屿上的物种丰富度是岛屿生物地理学理论的核心内容,而集合种群理论关注的是局部种群之间个体迁移的动态以及物种的续存条件,在概述两个理论形成、发展及其核心内容的基础上,着重比较它们的异同点以及在生态学理论和实践中的应用,并论述物种保护理论范式从岛屿生物地理学向集合种群理论转变的基本背景和原因。     

阿拉善荒漠啮齿动物集合群落实证研究

当生态学家探求在破碎化的栖息地中,群落物种的共存机制、多样性、局域尺度的性质和过程被放到更广阔的时空框架内时,就出现了"集合群落"这一概念。Leibold提出了集合群落概念,他们将一个集合群落定义为局域群落集,这些群落由各个潜在的相互作用的物种的扩散连接在一起。集合群落理论描述了那些发生在集合群落尺度上的过程,并且提出思考关于物种相互作用的新方法。集合群落概念为群落生态学提供了一个新的革命性的范式,集合群落研究的最基本问题是同一系统中多物种共存的机理、多样性的形成原因与维持机制。该范式强调区域范围内群落中的综合变异,强调环境特证和栖息地之间通过扩散调节的生物相互作用和空间变化。Leibold等提出了解释集合群落结果理论上的4个生态范式,即(1)中性理论;(2)斑块动态理论;(3)物种分配理论;(4)集团效应理论。之后有大量有关检验这4种生态理论的研究,但是有关陆地脊椎动物系统的集合群落的研究较少。2010—2012年,通过在内蒙古阿拉善荒漠景观中的8个固定样地中,对啮齿动物、栖息地环境因子进行调查。利用冗余分析和偏冗余分析,评估环境特征和空间特征对物种组成的影响。结果表明,环境特征独自解释72.8%的啮齿动物物种组成变化,空间特征独自解释33.8%的物种组成变化,环境特征和空间特征共同解释86.5%的啮齿动物物种组成变化,结果显著(P=0.032);去除环境特征之后,空间特征解释13.7%的变化(P=0.246),结果不显著;去除空间特征之后,栖息地变化解释52.7%的变化(P=0.016);环境特征和空间特征的交互作用解释20.1%的物种组成的变化,该区域啮齿动物群落构成集合群落,物种共存中环境特征起着主导作用,由物种分配理论解释该集合群落结构。     

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。     

生境丧失对具有似Allee效应集合种群的影响及对策——以江苏盐城为例

似Allee效应对物种续存是潜在的扰动因素,稀有物种更易受其影响,可能增加生存于破碎化栖息地中的珍稀物种的死亡风险;但似Allee效应对多物种集合种群续存的影响及其在珍稀物种保护中的应用未能引起足够重视。将似Allee效应引入集合种群动力模式,建立了生境丧失下具有似Allee效应的n-珍稀物种的集合种群模式,并以江苏盐城滩涂湿地中的29种珍稀物种为研究实例。研究结果表明:(1)似Allee效应导致n-物种集合种群多度作长期变周期振荡,原本竞争共存物种可能无法继续共存,甚至灭绝。(2)似Allee效应增强对次强种及劣势种的生存极为不利,导致次强物种由强至弱灭绝,劣势物种由弱至强依次灭绝。(3)盐城天然湿地丧失29%后,11种劣势物种的集合种群由弱到强将最终依次灭绝,灭绝迟豫时间为304～890a,这些物种即Hanski所指的"活死者"。(4)适度增加栖息地面积是保护珍稀物种多样性的有效方法之一,在盐城现存3200km2的天然湿地基础上适度增加1801～2064km2左右栖息地面积,可以有效保护29种濒危物种的多样性,同时应注意结合针对具体物种的保护措施来提高濒危物种多度。研究结果对物种多样性保护及自然保护区建设具有重要的理论指导意义。     

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。     

气候变化情景下物种适宜生境预测研究进展

气候变化能够引起物种分布范围、生物物候等一系列生态现象和过程的变化,进而加速物种灭绝的速率。气候变化被认为是21世纪全球生物多样性面临的最主要威胁之一,将给未来的生物多样性保护工作带来严峻的挑战。利用物种分布模型预测气候变化情景下物种适宜生境的变化正成为当前的研究热点。本研究总结目前气候变化情景下物种适宜生境预测的最新方法及取得的主要成果。在研究方法上,多物种分布模型、多气候情景基础上的集合预测方法正成为目前研究采用的主要手段;在研究结果上,未来气候变化将有可能导致物种适宜生境面积减少,范围向高纬度、高海拔地区移动。最后本研究指出目前气候变化情景下物种适宜生境预测研究中存在的主要不足及今后的发展方向。     

Identifying the mechanisms that shape fungal community and metacommunity patterns in Yunnan,China

Fungi are key organisms in terrestrial ecosystems, functioning as decomposers, pathogens, and symbionts. Identifying the mechanisms that shape metacommunity patterns is likely to be critical for predicting how ecosystems will respond to global environmental change. Using fungal occurrence data and a hierarchical approach that combines three elements of metacommunity structure—coherence, turnover and boundary clumping—we identified the structures that best describe metacommunity patterns. We related these patterns to underlying environmental and spatial variables known to influence fungal distribution, and determined the relative importance of the environment and geographic distance in structuring fungal metacommunities. Fungal metacommunities had Clementsian and quasi-Clementsian structures, indicating that species distributions were compartmentalized along a dominant environmental gradient. This gradient was strongly associated with annual precipitation, precipitation seasonality and pH for the entire metacommunity. Variance partitioning revealed that the environment was relatively more important than geographic distance in explaining metacommunity patterns, indicating that niche-based processes are crucial in shaping species distributions among sites. However, the strength of the relationship between the latent gradient and environmental factors and the relative contributions of the environment and geographic distance to metacommunity structure varied across groups, suggesting that interactions among habitat, dispersal and life-history might be driving these differences.     

A neutral model of edge effects

In this paper a spatially implicit neutral model for explaining the edge effects between habitats is proposed. To analyze this model we use two different approaches: a discrete approach that is based on the Master equation for a one step jump process and a continuous approach based on the approximation of the discrete jump process with the Kolmogorov-Fokker-Planck forward and backward equations. The discrete and continuous approaches are applied to analyze the species abundance distributions and the time to species extinction. Moreover, with the aid of the continuous approach a realistic classification of the behavior of species in local communities is developed. The species abundance dynamics at the edge between two distinct habitats is compared with those located in the homogeneous interior habitats using species abundance distributions and the first time to species extinction. We show that the structure of the links between local community and the metacommunity plays an important role on species persistence. Specifically, species at the edge between two distinct metacommunities have higher extinction rate than those in the interior habitats connected only to one metacommunity. Moreover, the same species might be persistent in the homogeneous interior habitat, but its probability of extinction from the edge local community could be very high.     

Extinction debt in source-sink metacommunities

In an increasingly modified world, understanding and predicting the consequences of landscape alteration on biodiversity is a challenge for ecologists. To this end, metacommunity theory has developed to better understand the complexity of local and regional interactions that occur across larger landscapes. While metacommunity ecology has now provided several alternative models of species coexistence at different spatial scales, predictions regarding the consequences of landscape alteration have been done exclusively for the competition-colonization trade off model (CC). In this paper we investigate the effects of landscape perturbation on source-sink metacommunities. We show that habitat destruction perturbs the equilibria among species competitive effects within the metacommunity, driving both direct extinctions and an indirect extinction debt. As in CC models, we found a time lag for extinction following habitat destruction that varied in length depending upon the relative importance of direct and indirect effects. However, in contrast to CC models, we found that the less competitive species are more affected by habitat destruction. The best competitors can sometimes even be positively affected by habitat destruction, which corresponds well with the results of field studies. Our results are complementary to those results found in CC models of metacommunity dynamics. From a conservation perspective, our results illustrate that landscape alteration jeopardizes species coexistence in patchy landscapes through complex indirect effects and delayed extinctions patterns.     

Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.

     

Vertebrate metacommunity structure along an extensive elevational gradient in the tropics: a comparison of bats,rodents and birds

Aim We evaluated the structure of metacommunities for each of three vertebrate orders (Chiroptera, Rodentia and Passeriformes) along an extensive elevational gradient. Using elevation as a proxy for variation in abiotic characteristics and the known elevational distributions of habitat types, we assessed the extent to which variation in those factors may structure each metacommunity based on taxon‐specific characteristics. Location Manu Biosphere Reserve in the Peruvian Andes. Methods Metacommunity structure is an emergent property of a set of species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit structure for each metacommunity along an elevational gradient comprising 13 250‐m elevational intervals and 58 species of rodent, 92 species of bat or 586 species of passerine. Results For each taxon, the environmental gradient along which the metacommunity was structured was highly correlated with elevation. Clementsian structure (i.e. groups of species replacing other such groups along the gradient) characterized rodents, with a group of species that was characteristic of rain forests and a group of species that was characteristic of higher elevation habitats (i.e. above 1500 m). Distributions of bats were strongly nested, with more montane communities comprising subsets of species at lower elevations. The structure of the passerine metacommunity was complex and most consistent with a quasi‐Clementsian structure. Main conclusions Each metacommunity exhibited a different structure along the same elevational gradient, and each structure can be accounted for by taxon‐specific responses to local environmental factors that vary predictably with elevation. The structures of rodent and bird metacommunities suggest species sorting associated with habitat specializations, whereas structure of the bat metacommunity is probably moulded by a combination of species‐specific tolerances to increasingly cold, low‐productivity environs of higher elevations and the diversity and abundance of food resources associated with particular habitat types.     

Elements of metacommunity structure of Paraguayan bats: multiple gradients require analysis of multiple ordination axes

Techniques to evaluate elements of metacommunity structure (EMS; coherence, species turnover and range boundary clumping) have been available for several years. Such approaches are capable of determining which idealized pattern of species distribution best describes distributions in a metacommunity. Nonetheless, this approach rarely is employed and such aspects of metacommunity structure remain poorly understood. We expanded an extant method to better investigate metacommunity structure for systems that respond to multiple environmental gradients. We used data obtained from 26 sites throughout Paraguay as a model system to demonstrate application of this methodology. Using presence–absence data for bats, we evaluated coherence, species turnover and boundary clumping to distinguish among six idealized patterns of species distribution. Analyses were conducted for all bats as well as for each of three feeding ensembles (aerial insectivores, frugivores and molossid insectivores). For each group of bats, analyses were conducted separately for primary and secondary axes of ordination as defined by reciprocal averaging. The Paraguayan bat metacommunity evinced Clementsian distributions for primary and secondary ordination axes. Patterns of species distribution for aerial insectivores were dependent on ordination axis, showing Gleasonian distributions when ordinated according to the primary axis and Clementsian distributions when ordinated according to the secondary axis. Distribution patterns for frugivores and molossid insectivores were best described as random. Analysis of metacommunities using multiple ordination axes can provide a more complete picture of environmental variables that mold patterns of species distribution. Moreover, analysis of EMS along defined gradients (e.g., latitude, elevation and depth) or based on alternative ordination techniques may complement insights based on reciprocal averaging because the fundamental questions addressed in analyses are contingent on the ordination technique that is employed.     

Proximity of restored hedgerows interacts with local floral diversity and species' traits to shape long‐term pollinator metacommunity dynamics

Disconnected habitat fragments are poor at supporting population and community persistence; restoration ecologists, therefore, advocate for the establishment of habitat networks across landscapes. Few empirical studies, however, have considered how networks of restored habitat patches affect metacommunity dynamics. Here, using a 10‐year study on restored hedgerows and unrestored field margins within an intensive agricultural landscape, we integrate occupancy modelling with network theory to examine the interaction between local and landscape characteristics, habitat selection and dispersal in shaping pollinator metacommunity dynamics. We show that surrounding hedgerows and remnant habitat patches interact with the local floral diversity, bee diet breadth and bee body size to influence site occupancy, via colonisation and persistence dynamics. Florally diverse sites and generalist, small‐bodied species are most important for maintaining metacommunity connectivity. By providing the first in‐depth assessment of how a network of restored habitat influences long‐term population dynamics, we confirm the conservation benefit of hedgerows for pollinator populations and demonstrate the importance of restoring and maintaining habitat networks within an inhospitable matrix.     

The importance of landscape and habitat properties in explaining instantaneous and long‐term distributions of large branchiopods in subtropical temporary pans

1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long‐term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long‐term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.