鄂西南亚热带山地常绿落叶阔叶混交林物种多度分布格局

基于七姊妹山自然保护区内6 hm~2监测样地多度数据,通过累计经验分布曲线(ECDF)表征该样地内不同生活型功能群的物种-多度分布格局,并采用6种模型对各功能群不同取样尺度物种等级-多度曲线进行拟合并检验其拟合效果,分析多度格局与模型拟合在不同尺度间的差异,探讨其背后的生态学过程与机制。结果表明:(1)各尺度下落叶种比常绿种的物种数多,物种多样性指数更大,但个体数相对较少;不同功能群稀有种比例排序为:落叶种所有种常绿种。(2)6种模型中的断棍模型的拟合效果较差;中大尺度(50 m×50 m、100 m×100 m)上不同生活型树种多度分布能接受的模型较少,除大尺度的常绿树种外,拟合最优模型均为对数正态分布模型,大尺度的常绿树种拟合最优模型为中性模型;小尺度上(20 m×20 m)常绿树种的最优模型为对数正态分布模型,落叶树种最优模型为生态位优先模型,所有树种在小尺度最优模型为Zipf-Mandelbrot模型。研究认为,随着尺度逐渐扩大,中性过程较生态位过程对物种-多度格局的解释力度更大,落叶树种物种多度格局的形成机制较常绿树种更接近于样地所有树种物种-多度格局的形成机制。     

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。     

亚热带常绿阔叶林群落物种多度分布格局对取样尺度的响应

为揭示物种多度格局随尺度的变化规律,探讨多度格局形成的机理及生态学过程,作者以古田山亚热带常绿阔叶林24 ha固定监测样地为背景.采用断棍模型(broken stick model)、对数正态模型(Iognormal distribution model)、生态位优先占领模型(preemption model)、Zipf模型(Zipf model)、Zipf-Mandelbrot模型(Zipf-Mandelbrot model)及中性理论模型(neutral model),对不同尺度下的物种多度分布格局进行拟合,并采用AIC检验和卡方检验选择最优拟合模型.结果表明,不同尺度上适合的物种一多度曲线模型不同;在取样边长为10 m和20 m时,除中性模型外的5个模型均不能被拒绝,它们均适合小尺度下的格局,这表明存小的尺度上生态位过程对物种一多度曲线的格局贡献较大;在取样边长为40 m时,最适合的模型为对数正态模型;取样边长为60 m和80 m时,Zipf-Mandelbrot模型为最优拟合模型;在取样边长为100 m时,尽管Zipf-Mandelbrot模型有最小的AIC值,但卡方检验拒绝了除中性模型外的5个模型;中性理论模型除了边长为10 m和20 m尺度以外,在其他尺度上均比前面5种模型的预测效果更好.因此在研究物种多度分布规律时必须注意空间尺度的影响.研究结果表明随着尺度的增加,中性过程成为决定物种一多度曲线格局的主要生态过程.     

长白山阔叶红松林草本层物种多度分布格局及其季节动态

草本层是森林生态系统的重要组成部分, 对维持森林生物多样性具有重要意义。本文以长白山阔叶红松(Pinus koraiensis)林25 ha固定监测样地为研究平台, 运用不同的统计模型(对数正态模型和对数级数模型)及机理模型(包括生态位模型: 断棍模型和生态位优先占领模型; 中性模型: 复合群落零和多项式模型和Volkov模型), 对不同季节草本物种多度分布进行拟合。采用Kolmogorov-Smirnov和AIC检验确定最优模型, 以揭示草本层物种多度分布格局随季节的变化规律, 探讨草本层物种组成与结构背后的生态学过程。结果表明: (1)草本层物种多度分布季节差异明显。春季各多度级物种数差异不大, 夏季中间种较多, 秋季则是稀有种较多; (2)模型拟合结果显示, 不同季节草本层物种多度分布的最优拟合模型相近。统计模型中对数级数模型表现最优, 机理模型中中性模型的拟合效果优于生态位模型。复合群落零和多项式模型较好地拟合了春夏季草本物种多度分布, Volkov模型较好地拟合了秋季草本物种多度分布。综上所述, 尽管长白山阔叶红松林草本植物不同季节的物种多度分布格局不尽一致, 但其背后的构建机制相似, 中性随机过程在草本层物种多样性维持过程中显得更为重要。     

浙江天童山区灌丛群落的物种多样性及其与演替的关系

根据浙江天童山区灌丛群落的样地调查数据,采用Shannon－Wiener多样性指数、Simpson指数和群落均匀度指数研究了灌丛群落的物种多样性,并与当地的常绿阔叶林进行了比较。结果表明：这3种指标能够有效地表征亚热带灌丛群落的组成结构特征。天童灌丛群落灌木层的物种多样性指数 SW 高于常绿阔叶林中乔木层,而低于常绿阔叶林中灌木层。灌丛群落灌木层的生态优势度 SN 低于常绿阔叶林中乔木层,但高于常绿阔叶林中灌木层。灌丛群落灌木层的群落均匀度 PW 比常绿阔叶林中乔木层的和灌水层的都低。此外,文中还就本区灌丛群落物种多样性的特点,讨论了加速其进展演替的恢复措施。     

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。     

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。     

太白山锐齿栎林物种-多度分布格局

以太白山1.5 hm²的锐齿栎原始林和次生林样地中环境因子和胸径≥1 cm的木本植物调查数据为基础,采用统计模型(对数正态模型)、生态位模型(Zipf模型、断棍模型、生态位优先模型)和中性模型,拟合了锐齿栎群落的物种多度分布。结果表明: 太白山锐齿栎林物种多度分布格局受到生境异质性的影响。其中,地形因子对原始林物种分布影响较大,在凹凸度较大的生境中,物种分布同时受到中性过程和生态位过程的影响,但中性过程发挥的作用较小;而在凹凸度较小的生境中,中性模型被拒绝,物种的多度分布符合生态位理论的假设。在群落坡度大的区域,群落中生态位过程和中性过程同等重要;而在坡度较小的平缓区域,生态位分化对群落物种分布的影响较大。在次生林中,影响物种分布的因素主要是土壤养分。在次生林土壤速效磷含量高的生境中,生态位过程是影响群落物种分布的主要生态学过程;而在土壤速效磷含量低的生境中,中性过程和生态位过程在群落物种分布中同时存在。太白山锐齿栎林物种多度分布格局存在明显的尺度效应。原始林在20 m×20 m尺度上,生态位模型和中性模型都能预测物种多度分布,而在40 m×40 m和70 m×70 m尺度上,生态位过程可解释物种多度分布格局。在次生林样地20 m×20 m、40 m×40 m、70 m×70 m尺度上,生态位过程和中性过程共同作用于物种的分布,但是生态位过程更为重要。可见,除了尺度和生境异质性外,原始林与受干扰的次生林中的物种多度分布也存在明显的差异。     

古田山常绿阔叶林萌生特征及其与群落物种多样性的关系

萌生更新是森林木本植物重要的生活史策略,能够促使萌生树种长时间占据原有空间,对物种共存、群落构建与生态系统稳定性有重要的潜在影响;同时,萌生更新的植株具有较强的资源获取与竞争能力。但萌生更新对群落物种共存的作用还远未被理解。本文基于古田山5 ha亚热带常绿阔叶林监测样地的调查数据,研究了萌生更新对群落物种多样性的影响及作用机制。结果表明:(1)古田山常绿阔叶林中萌生树种的丰富度比例较高(63.95%),已发生萌生的物种多度比例较高(38.53%),潜在萌生多度比例更高(59.51%);(2)萌生现象在各分类群普遍发生,壳斗科、杜鹃花科、金缕梅科、山茶科等类群的萌生能力较强;(3)萌生物种丰富度比例与群落物种多样性的关系不显著,萌生物种多度比例与群落物种多样性呈负相关关系。由此推测,作为古田山亚热带常绿阔叶林中一种占优势的更新方式,萌生更新在保证萌生树种持续存活从而增加群落物种多样性的同时,在一定阶段又能抑制群落物种多样性的发展,对群落物种共存、生物多样性维持起到了较重要的调节作用。     

湖北七姊妹山亚热带常绿落叶阔叶混交林的物种组成和群落结构

亚热带常绿落叶阔叶混交林是亚热带山地中重要的森林植被类型之一,是维持区域生态安全和保护生物多样性的重要屏障。为深入研究亚热带常绿落叶阔叶混交林中的生物多样性维持机理和动态过程,我们在湖北恩施七姊妹山国家级自然保护区6 ha亚热带常绿落叶阔叶混交林动态监测样地调查了所有胸径(DBH)≥1 cm的木本植物,分析了其群落组成、径级结构和优势种的空间结构特征。共记录到木本植物50,316株,其中独立个体有36,596株(去除分枝和萌生),隶属于57科124属245种,包括7种国家级保护植物和76种稀有种(每公顷个体数不超过1株)。样地内落叶物种分别占总物种数的61.2%(150种)和总个体数的52.6%(26,503株)。整个样地内植物个体胸径呈明显的倒"J"型分布,平均胸径为4.38 cm。样地小径级木本植物较多,胸径≥30 cm的木本植物仅175株。样地中重要值排名前4位的物种分别是川陕鹅耳枥(Carpinus fargesiana)、多脉青冈(Cyclobalanopsis multinervis)、交让木(Daphniphyllum macropodum)和缺萼枫香(Liquidambar acalycina)。这些优势种虽然在整个样地均有分布,但大径级个体(DBH≥10 cm)的空间分布差异较为明显。依据群落组成和径级结构特征,湖北七姊妹山森林动态样地属于人为干扰后处于自然恢复中期的森林群落。针对该样地开展长期的森林动态监测,将有助于深入了解亚热带常绿落叶阔叶混交林恢复过程中的群落构建规律和物种多样性维持机制。     

Responses of species abundance distribution patterns to spatial scaling in subtropical secondary forests

To quantify and assess the processes underlying community assembly and driving tree species abundance distributions(SADs) with spatial scale variation in two typical subtropical secondary forests in Dashanchong state‐owned forest farm, two 1‐ha permanent study plots (100‐m × 100‐m) were established. We selected four diversity indices including species richness, Shannon–Wiener, Simpson and Pielou, and relative importance values to quantify community assembly and biodiversity. Empirical cumulative distribution and species accumulation curves were utilized to describe the SADs of two forests communities trees. Three types of models, including statistic model (lognormal and logseries model), niche model (broken‐stick, niche preemption, and Zipf‐Mandelbrodt model), and neutral theory model, were estimated by the fitted SADs. Simulation effects were tested by Akaike's information criterion (AIC) and Kolmogorov–Smirnov test. Results found that the Fagaceae and Anacardiaceae families were their respective dominance family in the evergreen broad‐leaved and deciduous mixed communities. According to original data and random sampling predictions, the SADs were hump‐shaped for intermediate abundance classes, peaking between 8 and 32 in the evergreen broad‐leaved community, but this maximum increased with size of total sampled area size in the deciduous mixed community. All niche models could only explain SADs patterns at smaller spatial scales. However, both the neutral theory and purely statistical models were suitable for explaining the SADs for secondary forest communities when the sampling plot exceeded 40 m. The results showed the SADs indicated a clear directional trend toward convergence and similar predominating ecological processes in two typical subtropical secondary forests. The neutral process gradually replaced the niche process in importance and become the main mechanism for determining SADs of forest trees as the sampling scale expanded. Thus, we can preliminarily conclude that neutral processes had a major effect on biodiversity patterns in these two subtropical secondary forests but exclude possible contributions of other processes.     

基于地形因子的天童地区常绿树种和落叶树种共存机制研究

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种,那么它们与常绿树种的共存机制是什么?常绿树种和落叶树种生态习性差异较大,二者对生境的选择应有所不同,我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设,我们以天童20ha动态样地调查数据为依托,选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象,用典范对应分析(CCA)研究了地形因子对二者分布的影响差异,用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下:(1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%,对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明,与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%,成负关联的比例分别为40%和7%;与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%,成负关联的比例分别为10.9%和47.6%;与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种;两个植物类群对生境的选择多呈现相反格局,尤其是在沟谷生境和山脊生境,这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一,生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。     

Neutral theory and the species abundance distribution: recent developments and prospects for unifying niche and neutral perspectives

Published in 2001, The Unified Neutral Theory of Biodiversity and Biogeography (UNTB) emphasizes the importance of stochastic processes in ecological community structure, and has challenged the traditional niche‐based view of ecology. While neutral models have since been applied to a broad range of ecological and macroecological phenomena, the majority of research relating to neutral theory has focused exclusively on the species abundance distribution (SAD). Here, we synthesize the large body of work on neutral theory in the context of the species abundance distribution, with a particular focus on integrating ideas from neutral theory with traditional niche theory. First, we summarize the basic tenets of neutral theory; both in general and in the context of SADs. Second, we explore the issues associated with neutral theory and the SAD, such as complications with fitting and model comparison, the underlying assumptions of neutral models, and the difficultly of linking pattern to process. Third, we highlight the advances in understanding of SADs that have resulted from neutral theory and models. Finally, we focus consideration on recent developments aimed at unifying neutral‐ and niche‐based approaches to ecology, with a particular emphasis on what this means for SAD theory, embracing, for instance, ideas of emergent neutrality and stochastic niche theory. We put forward the argument that the prospect of the unification of niche and neutral perspectives represents one of the most promising future avenues of neutral theory research.     

The implicit assumption of symmetry and the species abundance distribution

Species abundance distributions (SADs) have played a historical role in the development of community ecology. They summarize information about the number and the relative abundance of the species encountered in a sample from a given community. For years ecologists have developed theory to characterize species abundance patterns, and the study of these patterns has received special attention in recent years. In particular, ecologists have developed statistical sampling theories to predict the SAD expected in a sample taken from a region. Here, we emphasize an important limitation of all current sampling theories: they ignore species identity. We present an alternative formulation of statistical sampling theory that incorporates species asymmetries in sampling and dynamics, and relate, in a general way, the community-level SAD to the distribution of population abundances of the species integrating the community. We illustrate the theory on a stochastic community model that can accommodate species asymmetry. Finally, we discuss the potentially important role of species asymmetries in shaping recently observed multi-humped SADs and in comparisons of the relative success of niche and neutral theories at predicting SADs.     

福建建溪流域常绿阔叶防护林物种多样性特征研究

从生物多样性保护原则出发,物种多样性应是评价防护林综合效益的重要指标之一。本文采用Weibull分布模型分析建溪流域防护林乔木层、灌木层的种－多度关系,用多种公式计算防护林各层次的物种多样性并与我国暖温带落叶阔叶林的物种多样性相比较,得出以下结论：1)该流域防护林乔木层的种－多度关系符合Weibull分布模型,说明个体数量较多的乔木仅限于少数几种主要的树种,群落的均匀度相对较小;2)该流域防护林主要群落内乔木层、灌木层和草本层的丰富度、均匀度和总多样性指数都较我国暖温带落叶阔叶林相对应的指数高。     

Evaluating the performance of neutrality tests of a local community using a niche‐structured simulation model

Understanding the processes that underlie species diversity and abundance in a community is a fundamental issue in community ecology. While the species abundance distributions (SADs) of various natural communities may be well explained by Hubbell's neutral model, it has been repeatedly pointed out that Hubbell's SAD‐fitting approach lacks the ability to detect the effects of non‐neutral factors such as niche differentiation; however, our understanding of its quantitative effect is limited. Herein, we conducted extensive simulations to quantitatively evaluate the performance of the SAD‐fitting method and other recently developed tests. For simulations, we developed a niche model that incorporates the random stochastic demography of individuals and the nonrandom replacements of those individuals, i.e. niche differentiation. It therefore allows us to explore situations with various degrees of niche differentiation. We found that niche differentiation has strong effects on SADs and the number of species in the community under this model. We then examined the performance of these neutrality tests, including Hubbell's SAD‐fitting method, using extensive simulations. It was demonstrated that all these tests have relatively poor performance except for the cases with very strong niche structure, which is in accordance with previous studies. This is likely because two important parameters in Hubbell's model are usually unknown and are commonly estimated from the data to be tested. To demonstrate this point, we showed that the precise estimation of the two parameters substantially improved the performance of these neutrality tests, indicating that poor performance can be owed to overfitting Hubbell's neutral model with unrealistic parameters. Our results therefore emphasize the importance of accurate parameter estimation, which should be obtained from data independent of the local community to be tested.