基于栖息地斑块尺度的青弋江河源溪流鱼类群落的时空格局

确定鱼类的栖息地利用格局是研究物种与环境关系的基础, 也是鱼类多样性保护和管理的必要前提。目前, 有关溪流鱼类群落的栖息地斑块利用格局尚存在争议。基于2012年9月至2013年8月对青弋江河源溪流的逐月调查数据, 初步研究了鱼类群落的栖息地斑块利用格局, 着重在栖息地斑块尺度上解析了鱼类群落的时空变化规律。主要研究结果显示, 深潭和急滩2类斑块间的底质、流速、水深、溶氧栖息地因子显著差异, 且深潭斑块的环境稳定性高于急滩。研究共采集鱼类15种, 其中鲤科鱼类8种, 占采集物种数50%以上。基于鱼类物种存在与否的不连续变量的分析结果显示, 鱼类物种组成的斑块间和月份间变化均不具显著性。但是, 基于鱼类物种多度的连续变量的分析结果显示, 鱼类群落结构存在有显著的斑块间变化和时间动态; 就斑块间变化而言, 原缨口鳅(Vanmanenia stenosoma)在急滩斑块中的多度更高, 而宽鳍 (Zacco platypus)、光唇鱼(Acrossocheilus fasciatus)和尖头 (Phoxinus oxycephalus)等其他关键物种则在深潭中具有更高多度。深潭斑块的鱼类物种数显著高于急滩, 但2类斑块间的个体数无显著差异。深潭斑块的鱼类物种数较稳定, 而个体数月变化显著, 可能与鱼类繁殖和群体补充以及越冬死亡等有关; 急滩鱼类物种数和个体数的月变化均显著, 除了与鱼类群体补充和越冬死亡有关以外, 还可能受越冬时栖息地斑块选择变化的影响。上述结果表明, 在栖息地斑块空间尺度上, 由于研究区域内大多数物种在栖息地斑块选择上无明显的特化性, 深潭和急滩斑块间鱼类的物种组成分布不符合前人所报道的生境-共位群格局, 但区域内常见种多度的变化可引起鱼类群落结构的斑块间差异和季节动态。       

集合群落(metacommunity)研究动态

由于人类活动等原因,自然生境在日益丧失和破碎化。因此,集合种群结构作为空间生态学的一个重要的研究途径越来越受到人们的重视,而斑块化环境中的生物群落即集合群落也成为生态学的一个研究热点。文中综述了集合群落动态,群落物种丰富度和物种分布,生境的丧失和破坏对集合群落的影响等的最新研究成果。     

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。     

集合群落及其在土壤动物群落构建中的应用

集合群落(metacommunity)指的是一系列由多个潜在的相互作用的物种通过扩散联系起来的局域群落。集合群落研究的基本问题是同一系统中多物种共存的机理、多样性的成因与维持机制。为了解决这些经典问题,Leibold提出了集合群落理论的4个经典范式,即斑块动态理论、物种分配理论、中性理论和集群效应理论。集合群落理论可以应用于土壤动物群落构建研究领域,解决多空间尺度土壤动物空间分布模式与驱动机制等问题。本文采用文献计量学方法梳理了集合群落自1992年提出到目前为止,集合群落理论、结构、驱动力以及基于功能性状的集合群落机制研究等四大核心方向与研究热点。同时,本文结合目前国内外土壤动物集合群落理论研究现状,提出进一步深入开展多尺度土壤动物多样性时空格局动态研究,旨在为土壤生物多样性管理提供对策措施。     

松嫩草地植物群落物种多度–分布关系及其解释

物种多度–分布关系是生物多样性科学的关键问题之一,但目前对其解释尚未有统一的观点,生态幅假说和集合种群动态理论是其中两个有代表性的假说。作者以松嫩平原羊草(Leymus chinensis)草甸的代表性群落羊草–杂类草群落斑块为研究对象,分别于2007年和2008年调查了其植物生长型组成和斑块距离,来检验上述两个假说对物种多度–分布关系的解释能力。作者提出以生长型适应指数(growth-form adaption index,GAI)作为生态幅的衡量指标,以最小斑块距离来反映植物的扩散和定居过程。结果表明:在生境尺度上,物种的多度和分布成显著正相关,且多度–分布关系在年际间相对稳定。物种的多度和分布均与植物生长型密切相关,分布广、多度高的物种以根茎型、直立型或分枝型为主,分布窄、多度低的物种以莲座型或刷状根型为主。GAI与物种的多度和分布均成显著线性正相关,少数物种的分布广,多数物种的分布窄,直接支持了生态幅假说。最近斑块距离与物种的多度和分布均成显著幂函数负相关,而群落相似性与斑块距离无显著相关性,因此本结果并未直接支持集合种群动态理论,但扩散过程可能是影响羊草草甸植物群落种多度–分布关系的重要潜在机制之一。     

多物种集合的种-多度关系模型研究进展

种 -多度关系是群落结构研究最基本的方面之一。Preston最早描述的物种“常见性和罕见性的分布”是群落中种 -多度关系的一般特性。自从 Motomura开创性地提出一个几何序列模型用于描述这种特性后 ,生态学家已建立了许多拟合种 -多度关系的经验模型和理论模型 ,主要包括对数序列分布、对数正态分布等统计模型 ,几何序列模型、分割线段模型等生态位模型以及反映群落动态和环境制约作用的动态模型 ,还有其中一些模型的扩展模型。近年来 ,种 -多度关系研究领域已大大拓宽 ,如“群落”的内涵已延伸为“多物种集合”,物种“多度”的测度推广到“广义多度”。尽管种 -多度关系研究已取得很大进展 ,仍有一些问题尚待解决 ,诸如多度指标选择、种 -多度分布时空变化规律及其机制以及种 -多度关系模型的选择、检验、解释与应用。     

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。     

溪流鱼类群落对低水头坝的大小及功能的响应——以皖南山区河源溪流为例

以皖南山区的河源溪流为研究区域, 根据2016年7月和11月两次的调查数据, 在区分土著物种和本地入侵物种的基础上, 初步比较研究了低水头坝的坝上蓄水区和冲刷区鱼类群落的季节动态及所受水坝大小和功能的影响。共采集鱼类24种, 隶属4目10科, 其中土著鱼类13种, 本地入侵鱼类11种。尽管坝下冲刷区的鱼类群落无显著季节变化, 但坝上蓄水区的鱼类群落季节动态显著, 其中11月的鱼类个体数显著高于7月。水坝大小对鱼类群落的影响相对较小, 仅冲刷区入侵鱼类群落结构随水坝大小显著变化; 水坝功能对鱼类群落的影响较大, 蓄水区土著鱼类群落及冲刷区入侵鱼类群落等都随水坝功能显著变化——同蓄水坝相比, 引渠坝的蓄水区中宽鳍鱲较少但中华花鳅和吻虾虎鱼较多, 且引渠坝的冲刷区中高体鳑鲏和泥鳅具有更高多度。研究表明, 在研究区域内, 低水头坝对鱼类群落分布的影响视水坝蓄水功能而不同, 而水坝大小的相对重要性较低。研究也进一步证实, 因亲流性土著鱼类与广布性入侵鱼类对低水头坝干扰的响应不同, 在开展有关人类活动对溪流鱼类多样性的影响评价时, 有必要区分土著物种和入侵物种来进行, 以确保研究结论的科学性。     

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。     

亚热带常绿阔叶林林分结构对物种组成变异的驱动作用:从局域到区域尺度

林分结构可以表征森林群落光的可利用性和光环境的异质性,对群落物种组成的变异具有重要驱动作用。然而,目前还鲜有研究将林分结构用于解释群落物种组成的变异。本研究以哀牢山亚热带中山湿性常绿阔叶林20 ha森林动态监测样地及其周边区域按公里网格设置的19个1 ha森林动态样地为研究对象,将林分结构参数、环境因子和空间结构变量共同作为解释变量,采用基于冗余分析的变差分解和层次分割方法,在局域和区域尺度上同时解析群落物种组成变异的驱动因素。结果表明,在局域和区域尺度上,纳入林分结构后均提高了对物种组成变异的解释率。在局域尺度上,加入林分结构作为解释变量后,单纯的空间结构的解释率明显下降,林分结构与环境因子累计贡献了41.0%的解释率。在区域尺度上,林分结构与环境因子累计贡献了23.0%的解释率。从局域到区域尺度,环境过滤的相对作用明显增强。林分结构指示的光的可利用性对林冠下方的树种组成具有较强的塑造作用,今后的研究应进一步探讨林分结构对亚热带常绿阔叶林物种组成变异的驱动机制。     

When can we distinguish between neutral and non‐neutral processes in community dynamics under ecological drift?

Determining whether the composition of ecological communities (species presence and abundance), can be predicted from species demographic traits, rather than being a result of neutral drift, is a key ecological question. Here we compare the similarity of community composition, from different community assembly models run under identical environmental conditions, where interspecific competition is assumed to be either neutral or niche-based. In both cases, species colonize a focal patch from a network of neighbouring patches in a metacommunity. We highlight the circumstances (rate and spatial scale of dispersal, and the relative importance of ecological drift) where commonly used community similarity metrics or species rank–abundance relationships are likely to give similar results, regardless of the underlying processes (neutral or non-neural) driving species' dynamics. As drift becomes more important in driving species abundances, deterministic niche structure has a smaller influence. Our ability to discriminate between different underlying processes driving community organization depends on the relative importance of different drift processes that operate on different spatial scales.     

Dispersal Ability Determines the Role of Environmental,Spatial and Temporal Drivers of Metacommunity Structure

Recently, community ecologists are focusing on the relative importance of local environmental factors and proxies to dispersal limitation to explain spatial variation in community structure. Albeit less explored, temporal processes may also be important in explaining species composition variation in metacommunities occupying dynamic systems. We aimed to evaluate the relative role of environmental, spatial and temporal variables on the metacommunity structure of different organism groups in the Upper Paraná River floodplain (Brazil). We used data on macrophytes, fish, benthic macroinvertebrates, zooplankton, periphyton, and phytoplankton collected in up to 36 habitats during a total of eight sampling campaigns over two years. According to variation partitioning results, the importance of predictors varied among biological groups. Spatial predictors were particularly important for organisms with comparatively lower dispersal ability, such as aquatic macrophytes and fish. On the other hand, environmental predictors were particularly important for organisms with high dispersal ability, such as microalgae, indicating the importance of species sorting processes in shaping the community structure of these organisms. The importance of watercourse distances increased when spatial variables were the main predictors of metacommunity structure. The contribution of temporal predictors was low. Our results emphasize the strength of a trait-based analysis and of better defining spatial variables. More importantly, they supported the view that “all-or- nothing” interpretations on the mechanisms structuring metacommunities are rather the exception than the rule.     

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.     

Do metacommunity theories explain spatial variation in fish assemblage structure in a pristine tropical river?

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Effect of landscape context on fish metacommunity structuring in stream networks

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Metacommunity of a host metapopulation: explaining patterns and structures of a fish parasite metacommunity in a Neotropical floodplain basin

Host-parasite metacommunities are influenced by a myriad of factors, although little is known about which processes affect this relationship at different scales. Here, we tested how local habitat characteristics and host traits explained the parasite metacommunity of a migratory fish in a large Brazilian river floodplain. The parasite metacommunity structure showed a Clementsian pattern, which indicates a more deterministic assembly pattern, in accordance with partial Redundancy Analysis results. Results indicated that species filtering is the predominant mechanism driving community assembly. Patterns were clearer in the dry season of the floodplain. Environmental determinism seems to explain ectoparasite metacommunities in the dry season, in contrast with endoparasites that were more correlated to host traits. Overall, our results indicated that ectoparasitism is an interaction marked by opportunity, whereas endoparasitism is likely related to host features. Thus, we argue that metacommunity structuring of parasites depends on the infection strategy. Our results show that floodplain dynamics are central not only for free-living animal organizations but also for symbiotic interactions. Here, we highlight the importance of understanding the factors influencing the distribution of parasites to predict their transmission, as well as the importance of floodplain dynamics and its hydrological regime on the maintenance of ecological interactions.

     

Temporal variability in the spatial and environmental determinants of functional metacommunity organization – stream fish in a human‐modified landscape

1. Quantifying the relative importance of environmental filtering versus regional spatial structuring has become an intensively studied area in the context of metacommunity ecology. However, most studies have evaluated the role of environmental and spatial processes using taxonomic data sets of single snapshot surveys. 2. Here, we examined temporal changes in patterns and possible processes behind the functional metacommunity organization of stream fishes in a human‐modified landscape. Specifically, we (i) studied general changes in the functional composition of fish assemblages among 40 wadeable stream sites during a 3‐year study period in the catchment area of Lake Balaton, Hungary, (ii) quantified the relative importance of spatial and environmental factors as determinants of metacommunity structure and (iii) examined temporal variability in the relative role of spatial and environmental processes for this metacommunity. 3. Partial triadic analysis showed that assemblages could be effectively ordered along a functional gradient from invertebrate consuming species dominated by the opportunistic life‐history strategy, to assemblages with a diverse array of functional attributes. The analysis also revealed that functional fish assemblage structure was moderately stable among the sites between the sampling periods. 4. Despite moderate stability, variance partitioning using redundancy analyses (RDA) showed considerable temporal variability in the contribution of environmental and spatial factors to this pattern. The analyses also showed that environmental variables were, in general, more important than spatial ones in determining metacommunity structure. Of these, natural environmental variables (e.g. altitude, velocity) proved to be more influential than human‐related effects (e.g. pond area, % inhabited area above the site, nutrient enrichment), even in this landscape with relatively low variation in altitude and stream size. 5. Pond area was, however, the most important human stressor variable that was positively associated with the abundance of non‐native species with diverse functional attributes. The temporal variability in the relative importance of environmental and spatial factors was probably shaped by the release of non‐native fish from fish ponds to the stream system during flood events. 6. To conclude, both spatial processes and environmental control shape the functional metacommunity organization of stream fish assemblages in human‐modified landscapes, but their importance can vary in time. We argue, therefore, that metacommunity studies should better consider temporal variability in the ecological mechanisms (e.g. dispersal limitation, species sorting) that determine the dynamics of landscape‐level community organization.     

Combining Niche and Dispersal in a Simple Model (NDM) of Species Distribution

Predicting the distribution of species has become a crucial issue in biodiversity research. Two kinds of model address this question: niche models, which are usually based on static approaches linking species distribution to habitat characteristics, and dispersal models, which are usually dynamic and process-based. We propose a model (NDM: niche and dispersal model) that considers the local presence of a species to result from a dynamic balance between extinction (based on the niche concept) and immigration (based on the dispersal concept), at a given moment in time, in a spatially explicit context. We show that NDM correctly predicts observed bird species and community distributions at different scales. NDM helps to reconcile the contrasting paradigms of metacommunity theory. It shows that sorting and mass effects are the factors determining bird species distribution. One of the most interesting features of NDM is its ability to predict well known properties of communities, such as decreasing species richness with decreasing patch size and increasing distance to the mainland, and the mid-domain effect at the regional scale, contrasting with predictions of much smaller effects at the local scale. NDM shows that habitat destruction in the matrix around patches of forest can affect the forest bird community, principally by decreasing the occurrence of typical matrix birds within the forest. This model could be used as the starting point for applied ecological studies on the management of species and community distributions.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

Molecular and morphological evidence for conspecificity of two common Indo-Pacific species of Palythoa (Cnidaria: Anthozoa)

Most studies characterize metacommunities based on a single snapshot of the spatial structure, which may be inadequate for taxa with high migratory behavior (e.g., fish). Here, we applied elements of metacommunity structure to examine variations in the spatial distributions of stream fishes over time and to explore possible structuring mechanisms. Although the major environmental gradients influencing species distributions remained largely the same in time, the best-fit pattern of metacommunity structure varied according to sampling occasion and whether or not we included non-native species in the analyses. Quasi-Clementsian and Clementsian structures were the predominant best-fit structures, indicating the importance of species turnover among sites and the existence of more or less discrete community boundaries. The environmental gradient most correlated with metacommunity structure was defined by altitude, area of artificial ponds in the catchment, and dissolved oxygen content. Our results suggest that the best-fit metacommunity structure of the native species can change in time in this catchment due to seasonal changes in distribution patterns. However, the distribution of non-native species throughout the landscape homogenizes the temporal variability in metacommunity structure of native species. Further studies are necessary from other regions to examine best-fit metacommunity structures of stream fishes within relatively short environmental gradients.