片段化森林群落构建的生态过程及其检验方法

鉴于全球森林均呈现片段化(破碎化)的分布状态,理解片段化森林群落构建的过程很有必要。该文通过综述群落构建的主要生态过程如生态漂变、扩散、选择和物种形成等在片段化森林群落构建中的相对作用,发现因片段化森林形成方式的不同,重构群落(片段化生境中通过次生演替重新形成的森林群落)和解构群落(原有森林被片段化后形成的森林群落)在不同演替阶段所受到的主要生态过程的相对作用有所不同。虽然利用基于群落内物种分布格局推测构建过程(如物种多度分布、零模型结合β多样性的方法、功能特征的收敛和发散等)、人工控制实验、群落结构动态分析等方法对片段化森林中群落构建的过程进行了有效的检验,但是针对片段化森林群落构建过程的实验性研究仍然不足。未来有待在理论模型、群落构建过程的检验以及理论与物种保护相结合等方面继续开展深入的研究。     

长白山不同演替阶段针阔混交林群落物种多度分布格局

为解释长白山温带森林群落构建和物种多度格局的形成过程, 该文以不同演替阶段的针阔混交林监测样地数据为基础, 采用中性理论模型、生物统计模型(对数正态分布模型)和生态位模型(Zifp模型、分割线段模型、生态位优先模型)拟合森林群落物种多度分布, 并用χ ²检验、Kolmogorov-Smirnov (K-S)检验和赤池信息准则(AIC)选择最佳拟合模型。结果显示: 中性模型能很好地预测长白山温带森林不同演替阶段植物群落的物种多度分布。在10 m × 10 m尺度上, 5种模型均可被χ ²检验和K-S检验接受, 但中性模型拟合效果不如对数正态分布模型、Zifp模型、分割线段模型和生态位优先模型, 表明小尺度上中性过程和生态位过程均能解释群落物种多度分布, 但生态位过程的解释能力相对较大。而在中大尺度上(30 m × 30 m、60 m × 60 m和90 m × 90 m), 中性模型为最优拟合模型, 并且随着研究尺度增加, 生态位模型和生物统计模型逐渐被χ ²检验拒绝, 表明中性过程在长白山针阔混交林群落物种多度分布格局形成中的作用随着研究尺度增加而逐渐增大。该文证实了中性过程在长白山温带针阔混交林群落结构形成中具有重要作用, 但未否认生态位机制在群落构建中的贡献。因此, 温带森林群落构建过程中中性理论和生态位理论并非相互矛盾, 而是相互融合的。在研究森林群落物种多度分布时, 应重视取样尺度和演替阶段的影响, 并采用多种模型进行拟合。     

黄土高原森林群落物种多度的分布格局

选用物种多度统计模型（对数级数分布模型、对数正态分布模型）和物种多度生态位模型（分割线段模型、生态位重叠模型、随机分配模型）研究了黄土高原子午岭马栏林区不同演替阶段3种森林群落物种多度分布格局的变化。结果表明：（1）物种分布统计模型拟合显示,演替初期的白桦（Betula platyphylla）林,物种丰富,但分布不均匀;演替中期的天然油松（Pinus tabulaeformis）林,油松优势地位明显,林下灌木和草本植物数量少但分布集中;演替顶极的辽东栎（Quercus liaotungensis）林,物种分布均匀且集中,枯立木形成的林隙较多。（2）在物种多度生态位模型拟合中,3种群落均能被较好地拟合。说明在演替过程中,影响物种分布的自然和人为因素较多,不能简单地选择一个生态位模型来解释。因此,研究森林群落物种分布时,最好采用多个模型进行拟合。     

石灰岩山地淡竹林演替序列的群落物种多度分布格局

利用断棍模型(BSM)、生态位优先占领模型(NPM)、优势优先模型(DPM)、随机分配模型(RAM)和生态位重叠模型(ONM),对石灰岩山地淡竹林演替序列3类群落15个样地的种 多度关系进行拟合,并利用卡方(x²)和赤池信息量准则(AIC)检验.结果表明: 淡竹纯林、竹阔混交林和阔叶林最优物种多度分布格局模型分别为：DPM(x²=35.86,AIC=-69.77)、NPM(x²=1.60,AIC=-94.68)和NPM(x²=0.35,AIC=-364.61);BSM对混交林和阔叶林的拟合效果较好,对淡竹纯林的拟合欠佳;RAM和ONM对3类群落的拟合均不能接受;在淡竹纯林向阔叶林演替过程中,物种数逐渐增加,多度分布均匀,物种多度分布格局由DPM向NPM转变.由生境过滤作用主导转换成种间竞争作用主导是淡竹林演替序列物种多度格局变化的主要原因.采用多种模型和检验方法综合分析群落演替内、外因素变化,将有助于深入理解群落演替的生态过程.     

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。     

塔里木荒漠河岸林不同生境群落物种多度分布格局

为解释塔里木荒漠河岸林群落构建和物种多度分布格局形成的机理, 本文以塔里木荒漠河岸林2个不同生境(沙地、河漫滩) 4 ha固定监测样地为研究对象, 基于两样地物种调查数据, 采用统计模型(对数级数模型、对数正态模型、泊松对数正态分布模型、Weibull分布模型)、生态位模型(生态位优先占领模型、断棍模型)和中性理论模型(复合群落零和多项式模型、Volkov模型)拟合荒漠河岸林群落物种多度分布, 并用K-S检验与赤池信息准则(AIC)筛选最优拟合模型。结果表明: (1)随生境恶化(土壤水分降低), 植物物种多度分布曲线变化减小, 群落物种多样性、多度和群落盖度降低, 常见种数减少。(2)选用的3类模型均可拟合荒漠河岸林不同生境群落物种多度分布格局, 统计模型和中性理论模型拟合效果均优于生态位模型。复合群落零和多项式模型对远离河岸的干旱沙地生境拟合效果最好; 对数正态模型和泊松对数正态模型对洪水漫溢的河漫滩生境拟合效果最优; 中性理论模型与统计模型无显著差异。初步推断中性过程在荒漠河岸林群落构建中发挥着主导作用, 但模型拟合结果只能作为推断群落构建过程的必要非充分条件, 不能排除生态位过程的潜在作用。     

长白山温带森林不同演替阶段群落功能性状的空间变化

群落构建机制研究是生态学研究的热点。长白山自然保护区拥有完整的原始阔叶红松林生态系统, 近年来随着物种多样性丧失愈发严重, 对该地区开展群落构建机制研究显得尤为重要。该研究以长白山不同演替阶段的3块5.2 hm ²固定监测样地(次生杨桦林、次生针阔混交林、原始椴树红松林)为研究对象, 通过采集样地内主要树种的6个关键功能性状(叶面积、比叶面积、叶片厚度、叶氮含量、叶磷含量、最大树高), 分析不同空间尺度下(5 m × 5 m, 10 m × 10 m, 20 m × 20 m, 30 m × 30 m, 40 m × 40 m, 50 m × 50 m和60 m × 60 m)及不同演替阶段群落性状空间值的变化, 结合零模型的模拟结果对长白山温带森林演替过程中的群落构建机制进行讨论。结果表明: 种库大小对于研究结果具有重要影响。在较大的种库下, 环境过滤作用影响显著。而在样地水平进行研究时, 演替早期和中期, 群落性状空间值与零模型模拟值无显著差异, 在演替的晚期, 群落性状空间值显著高于零模型模拟值。结合多个群落功能多样性指数分析发现, 环境过滤和竞争作用共同决定该地区顶级群落的物种组成。在演替早期大量物种迁入, 群落内物种间存在强烈的资源竞争, 而随着演替进行, 部分物种逐渐被竞争排除出群落, 群落中的物种呈现明显的生态位分化, 竞争作用是维持物种共存的主要机制。     

长白山温带森林不同演替阶段群落功能性状的空间变化

群落构建机制研究是生态学研究的热点。长白山自然保护区拥有完整的原始阔叶红松林生态系统,近年来随着物种多样性丧失愈发严重,对该地区开展群落构建机制研究显得尤为重要。该研究以长白山不同演替阶段的3块5.2hm2固定监测样地(次生杨桦林、次生针阔混交林、原始椴树红松林)为研究对象,通过采集样地内主要树种的6个关键功能性状(叶面积、比叶面积、叶片厚度、叶氮含量、叶磷含量、最大树高),分析不同空间尺度下(5 m×5 m, 10 m×10 m, 20 m×20 m, 30 m×30m, 40 m×40 m, 50 m×50 m和60 m×60 m)及不同演替阶段群落性状空间值的变化,结合零模型的模拟结果对长白山温带森林演替过程中的群落构建机制进行讨论。结果表明:种库大小对于研究结果具有重要影响。在较大的种库下,环境过滤作用影响显著。而在样地水平进行研究时,演替早期和中期,群落性状空间值与零模型模拟值无显著差异,在演替的晚期,群落性状空间值显著高于零模型模拟值。结合多个群落功能多样性指数分析发现,环境过滤和竞争作用共同决定该地区顶级群落的物种组成。在演替早期大量物种迁入,群落内物种间存在强烈的资源竞争,而随着演替进行,部分物种逐渐被竞争排除出群落,群落中的物种呈现明显的生态位分化,竞争作用是维持物种共存的主要机制。     

亚热带喀斯特地区不同地形植物群落物种多度分布格局

为探究亚热带喀斯特地区不同地形下植物群落物种多度分布格局,揭示不同地形下群落的物种多度格局形成的作用机制,丰富该地区植物群落构建理论,该文以贵州茂兰喀斯特地区山脊、槽谷、鞍部、洼地4种典型地形下植物群落的乔木层与灌木层为对象,统计物种多度,采用累计经验分布曲线(ECDF)表征多度分布格局,采取Wilcoxon秩和检验探究不同地形之间物种多度的差异性。采用不同生态学模型进行多度拟合,利用Kolmogorov-Smirnov(K-S)检验与赤池信息量准则(AIC)检验模型接受与拟合优度。结果表明:(1)不同地形下植物群落的个体数量与物种数存在差异,鞍部个体数最多,洼地的物种数最多,山脊的个体数、物种数均最少。(2)不同地形下植物群落的乔木层物种多度格局无显著差异,灌木层之间出现显著差异,山脊与鞍部、洼部,鞍部与槽谷、洼部都存在显著差异。(3)不同地形下乔木层物种多度对中性模型接受较好,其中山脊拟合最优,对生态位模型接受较差,仅山脊与鞍部通过两种生态位模型,拟合优度不及中性模型。灌木层对中性模型接受也较好,鞍部拟合最优,对生态位模型接受较差,仅洼地通过断棍模型。整体而言,乔木层比灌木层能更好地接受两种生态学模型,可能乔木层物种多度格局有更明显生态过程的印记,但不同地形下灌木层拟合优度差异更大,可能与灌木层物种对环境变化更剧烈有关。不同地形会引起群落构建不同程度的生态学过程,物种多度分布格局会逐渐适应地形。     

群落构建研究的新进展:进化和生态相结合的群落谱系结构研究

群落如何构建足群落生态学中的重要问题.群落谱系结构研究将物种间的亲缘进化关系运用到群落生态学研究中,利用物种的系统发育状况推测历史因素对现有群落的影响,为推断影响群落组成的生态学机制提供了有效方法.群落谱系结构的研究方法是首先建立可代表群落物种库的超级系统进化树,然后计算群落内物种间的谱系距离,最后通过统计方法检测其与随机模型下的谱系距离是否有显著差异来获得谱系结构(如谱系聚集、谱系发散),从而揭示群落构建中的关键生态过程(如生境过滤、竞争作用).群落谱系结构与空间尺度、分类群尺度、时间尺度等不同研究尺度有关.在小的空间尺度下,随着分类群尺度降低、树木年龄级增大,群落谱系结构从聚集逐渐转为发散;而随群落空间尺度的增大,谱系趋向于聚集.谱系结构受到环境因素影响,因此分析集合群落下的谱系可以揭示区域生态过程的影响.另外,群落谱系结构研究还有助于探讨中性理论、密度制约假说等生态学理论,并预测干扰作用下的群落演化趋势.在利用谱系结构深入探讨群落构建成因时,需要基于生态特征和环境变量共同分析,同时考虑小尺度局域过程(群落的微环境或群落内种间相互作用等)和大尺度区域过程(地史过程和物种形成等),并可结合生态控制实验,以确认群落构建的关键因素.在研究方法和手段上,今后需要注重通过选择合适的基因片段建立系统树,然后通过生态特征来加以校正,以更准确地反映物种间的亲缘距离.另外,获得谱系树后还需要寻找更加合理的统计模型和指数,增加统计分析和解决问题的能力.     

The composition of arbuscular mycorrhizal fungal communities in the roots of a ruderal forb is not related to the forest fragmentation process

Land‐use changes and forest fragmentation have strong impact on biodiversity. However, little is known about the influence of new landscape configurations on arbuscular mycorrhizal fungal (AMF) community composition. We used 454 pyrosequencing to assess AMF diversity in plant roots from a fragmented forest. We detected 59 virtual taxa (VT; phylogenetically defined operational taxonomic units) of AMF – including 10 new VT – in the roots of Euphorbia acerensis. AMF communities were mainly composed of members of family Glomeraceae and were similar throughout the fragmented landscape, despite variation in forest fragment size (i.e. small, medium and large) and isolation (i.e. varying pairwise distances). AMF communities in forest fragments were phylogenetically clustered compared with the global, but not regional and local AMF taxon pools. This indicates that non‐random community assembly processes possibly related to dispersal limitation at a large scale, rather than habitat filtering or biotic interactions, may be important in structuring the AMF communities. In this system, forest fragmentation did not appear to influence AMF community composition in the roots of the ruderal plant. Whether this is true for AMF communities in soil and the roots of other ecological groups of host plants or in other habitats deserves further study.     

High Conservation Value of Forest Fragments for Plant and Frugivore Communities in a Fragmented Forest Landscape in South Africa

Fragmentation is a major threat factor for plant–frugivore communities in tropical and subtropical forests. Resulting changes in the distribution of traits within these communities, e.g., a loss in large‐bodied frugivores, may lead to strong changes in plant–frugivore interactions in fragmented forests. Yet, we still lack a thorough understanding of the interplay between forest fragmentation, the trait‐composition of communities and resulting plant–frugivore interactions on a community‐scale. In a fragmented South African landscape comprising different forest categories—i.e., continuous natural forest, forest fragments surrounded by natural grassland, and forest fragments surrounded by sugarcane—we investigated the relationship between communities of fruiting plants and their frugivore visitors in response to forest fragmentation, as well as the interactive effects of forest fragmentation and fruit size of the plants on the number of frugivore visitors and their body size. Neither the fruit size of plant nor the body mass of frugivore communities differed between natural forest sites and forest fragments. Moreover, in‐depth analyses of frugivore assemblages visiting plant species revealed no effect of forest category on the number of frugivore visits or their mean body mass. The number of visits and body mass of frugivores were merely determined by the crop and fruit size of the focal plant species. Overall, our results suggest that frugivory of plant species with differently sized fruits was not reduced in forest fragments. Thus, fragments with high fruit availability may be key elements maintaining the functional connectivity of a heterogeneous forest landscape.     

Elevation‐dependent effects of forest fragmentation on plant–bird interaction networks in the tropical Andes

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human‐induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species‐poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.     

Ecological filtering or random extinction? Beta‐diversity patterns and the importance of niche‐based and neutral processes following habitat loss

Although both niche‐based and neutral processes are involved in community assembly, most models on the effects of habitat loss are stochastic, assuming neutral communities mainly affected by ecological drift and random extinction. Given that habitat loss is considered the most important driver of the current biodiversity crisis, unraveling the processes underlying the effects of habitat loss is critical from both a theoretical and an applied perspective. Here we unveil the importance of niche‐based and neutral processes to species extinction and community assembly across a gradient of habitat loss, challenging the predictions of neutral models. We draw on a large dataset containing the distribution of 3653 individuals of 42 species, representing 35% of the small mammal species of the Atlantic Forest hotspot, obtained in 68 sites across three continuously‐forested landscapes and three adjacent 10 000‐ha fragmented landscapes differing in the amount of remaining forest (50%, 30% and 10%). By applying a null‐model approach, we investigated β‐diversity patterns by detecting deviations of observed community similarity from the similarity between randomly assembled communities. Species extinction following habitat loss was decidedly non‐random, in contrast to the notion that fragmented communities are mainly driven by ecological drift. Instead, habitat loss led to a strong biotic homogenization. Moreover, species composition changed abruptly at the same level of landscape‐scale habitat loss that has already been associated with a drastic decline in species richness. Habitat loss, as other anthropogenic disturbances, can thus be seen as a strong ecological filter that increases (rather than decreases) the importance of deterministic processes in community assembly. As such, critical advances for the development of conservation science lie on the incorporation of the relevant niche traits associated with extinction proneness into models of habitat loss. The results also underscore the fundamental importance of pro‐active measures to prevent human‐modified landscapes surpassing critical ecological thresholds.     

华北区域环境梯度上阔叶林构建模式及分布成因

森林群落的构建即多样性维持机制是当今生态学研究的热点问题。然而, 当前群落构建和群落多样性的研究多在间接梯度上进行, 而在水、热等影响物种在区域内定植的关键且直接的环境梯度上研究群落构建和多样性模式则鲜有报道。结合环境因子, 基于物种组成和谱系方法探讨不同群落的分布成因, 有助于解释群落构建过程中的关键问题。该研究基于华北森林群落调查数据和环境数据, 涉及7个省市区的29个以壳斗科、桦木科为优势种的群落, 探讨了直接环境梯度上的群落构建和多样性模式, 同时用典范对应分析研究了不同群落分布的环境解释。结果发现, 相似的群落具有相似的生境偏好, 相似的生境条件会形成物种组成相同或相似的群落。环境热量主导了本区域的谱系关系, 在年平均气温较低的地区, 群落构建主要表现为生境过滤的模式。此外, 随着年降水量的增加, 生境过滤作用逐渐增加。在温度梯度上, 谱系多样性表现为钟形模式, 而降水量的增加能导致谱系多样性的增加。     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

Exploring tree-habitat associations in a Chinese subtropical forest plot using a molecular phylogeny generated from DNA barcode loci

Elucidating the ecological mechanisms underlying community assembly in subtropical forests remains a central challenge for ecologists. The assembly of species into communities can be due to interspecific differences in habitat associations, and there is increasing evidence that these associations may have an underlying phylogenetic structure in contemporary terrestrial communities. In other words, by examining the degree to which closely related species prefer similar habitats and the degree to which they co-occur, ecologists are able to infer the mechanisms underlying community assembly. Here we implement this approach in a diverse subtropical tree community in China using a long-term forest dynamics plot and a molecular phylogeny generated from three DNA barcode loci. We find that there is phylogenetic signal in plant-habitat associations (i.e. closely related species tend to prefer similar habitats) and that patterns of co-occurrence within habitats are typically non-random with respect to phylogeny. In particular, we found phylogenetic clustering in valley and low-slope habitats in this forest, indicating a filtering of lineages plays a dominant role in structuring communities in these habitats and we found evidence of phylogenetic overdispersion in high-slope, ridge-top and high-gully habitats, indicating that distantly related species tended to co-occur in these high elevation habitats and that lineage filtering is less important in structuring these communities. Thus we infer that non-neutral niche-based processes acting upon evolutionarily conserved habitat preferences explain the assembly of local scale communities in the forest studied.     

Hyperdynamism in fragmented habitats

Abstract. Are the dynamics of most ecological processes fundamentally increased in frequency or magnitude in fragmented habitats? Hyperdynamism could alter a wide range of population, community, and landscape phenomena, and appears to be evident in fragmented tropical, temperate, and boreal communities. I suggest some potential causes and consequences of hyperdynamism, and argue that the responses of many species and ecological processes to habitat fragmentation can be understood in this context.