植物生态学报, 2022, 46(11): 1317-1320 doi: 10.17521/cjpe.2022.0431

编者评述

中国典型灌丛研究: 格局与过程

刘庆,,*, 陈冬东

中国科学院成都生物研究所, 中国科学院山地生态恢复与生物资源利用重点实验室, 生态恢复与生物多样性保育四川省重点实验室, 成都 610041

Studies on typical shrubland in China: pattern and process

LIU Qing,,*, CHEN Dong-Dong

CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China

通讯作者: *刘庆, E-mail:liuqing@cib.ac.cn

编委: 郭柯

责任编辑: 乔鲜果

收稿日期: 2022-10-31   接受日期: 2022-11-11  

基金资助: 国家科技基础性工作专项(2015FY110300)
国家自然科学基金(32101234)

Received: 2022-10-31   Accepted: 2022-11-11  

Fund supported: The Special Foundation for National Science and Technology Basic Research Program of China(2015FY110300)
The National Natural Science Foundation of China(32101234)

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引用本文

刘庆, 陈冬东. 中国典型灌丛研究: 格局与过程. 植物生态学报, 2022, 46(11): 1317-1320. DOI: 10.17521/cjpe.2022.0431

LIU Qing, CHEN Dong-Dong. Studies on typical shrubland in China: pattern and process. Chinese Journal of Plant Ecology, 2022, 46(11): 1317-1320. DOI: 10.17521/cjpe.2022.0431

灌丛在我国山地及干旱半干旱地区广泛分布, 是我国陆地生态系统的重要组成部分(Tang et al., 2018), 在水源涵养、生物固碳、土壤保持及防风固沙等方面起到了重要的作用(Piao et al., 2009; 胡雄蛟等, 2021)。近年来, 我国开始重视灌丛研究, 先后开展了中国科学院先导专项课题“中国灌丛生态系统的固碳现状、变化和机制研究”及国家科技基础性工作专项项目“我国主要灌丛植物群落调查”等工作。通过十余年的研究, 对我国灌丛植物群落的类型、分布格局、物种组成以及碳收支等有了较系统的认知(谢宗强等, 2019)。

近年来, 国内外学者针对灌丛群落格局及过程开展了大量的研究工作, 并取得了一系列的重要进展。近期的相关植被调查初步掌握了我国灌丛分布格局, 明确了我国灌丛可分为常绿针叶灌丛、落叶阔叶灌丛、常绿阔叶灌丛、肉质刺灌丛及竹丛共5个植被型(郭柯等, 2020)。群落多样性格局的量化从单一描述群落物种丰富度, 逐渐发展到整合群落分类多样性、谱系多样性及功能多样性的多维度研究(Pistón et al., 2016)。丰富的群落调查数据为研究灌丛群落多样性格局的形成机制, 即群落构建机制, 奠定了良好的数据基础。通常, 群落构建受到区域物种进化及谱系地理历史、区域种库效应、环境选择、随机扩散、种间关系等多个过程的共同作用(Stokes & Archer, 2010; HilleRisLambers et al., 2012)。对于灌丛群落而言, 其构建过程具有较强的特异性, 其可能是原生演替形成, 也可能是森林群落退化形成或者退化群落的重组。当前, 群落生态学研究中兴起以零模型为代表的以格局推断过程的研究方法(Miller et al., 2017)。这类方法通过检测群落谱系相似性或功能性状差异性与零模型之间是否存在显著差异来揭示确定性过程或随机过程对于群落结构的影响(Pavoine & Bonsall, 2011)。尽管不同的生态过程组合可能会产生相同的群落格局而导致零模型的适用范围存在一定的争议, 但其对于揭示自然生态系统中多样性格局形成的过程仍具有一定的指导意义(Molina & Stone, 2020)。

灌丛独特的生态位促使其占据了大量森林和草地无法发育的生境, 针对灌丛群落多样性格局及其形成过程的研究对于了解灌丛生态系统功能具有积极作用, 对其进行生态系统服务及价值评估可为灌丛的有效保护及科学利用提供重要依据。例如, 研究发现灌木物种的生长能够对微环境起到改善作用, 庇护不同物种在其冠层下存活, 提升冠层下物种丰富度(Pistón et al., 2016)。为此, 灌丛对冠层下植物的促进作用已作为受损生态系统中物种多样性与维持的重要驱动因素, 成为植被修复的重要技术支撑(Gómez-Aparicio, 2009)。在环境胁迫下, 灌丛可以通过资源分配来调节各个器官的生物量以缓解环境胁迫, 对其生物量的分布格局及分配策略的研究有助于深化环境选择对灌丛群落影响的认识(杨弦等, 2017)。

基于以上论述的意义和目的, 我们组织了“中国典型灌丛研究: 格局与过程”专辑, 以期推进我国灌丛生态学的研究, 促进灌丛格局及其变化机制的前沿研究。本专辑收录了10篇文章, 内容涉及典型区域的灌丛植物群落特征及植被分类方案, 环境因素及干扰对灌丛群落特征的影响, 从群落功能性状的角度探讨了植物株高、叶性状及茎性状之间及其与气候和土壤化学计量特征之间的关系, 并基于不同时期植株群落叶性状的动态特征, 评估了胁迫环境中灌丛物种在资源权衡策略上的变化趋势。同时对灌丛群落内木本植物间、木本-草本植物间的竞争及促进作用进行了系统的梳理, 突出了种间关系在塑造灌丛群落结构中的重要作用。

在灌丛植物群落方面, 研究人员基于灌丛群落调查数据, 对我国典型灌丛植物群落组成与结构特征进行了研究, 系统展示了四川巴塘海子山高寒灌丛群落(刘秋蓉等, 2022)与内蒙古地区虎榛子(Ostryopsis davidiana)灌丛群系(要振宇等, 2022)的基本特征, 对区域灌丛群落的物种组成、生态环境特征及其相互关系进行了详细的描述, 并根据《中国植被志》研编内容与规范(王国宏等, 2020)对群落进行了数量分类。在放牧的影响下, 亚热带喀斯特山地灌丛群落的物种组成和群落结构发生了显著变化(黄侩侩等, 2022), 放牧干扰强度的增加会驱使灌草丛物种组成与群落结构简单化、稀疏化和矮小化, 从而显著降低地上生物量并增加喀斯特山地灌丛植被退化的潜在风险。

在群落功能性状方面, 基于选取的9种锦鸡儿属(Caragana)植物以及41个锦鸡儿属样地的群落调查数据, 研究人员综合植物的株高、叶性状和茎性状, 分析了不同功能性状之间及功能性状与气候和土壤特征之间的关系(罗源林等, 2022), 发现植物的功能性状沿着气候梯度, 尤其是干旱程度与土壤氮含量梯度, 呈现出明显的变化趋势, 表明植物功能性状在一定程度上能够适应环境变化。代远萌等(2022)在毛乌素沙地研究了黑沙蒿(Artemisia ordosica)叶性状在沙丘不同固定阶段的动态特征, 发现在沙丘固定前期黑沙蒿倾向于增强光合作用来提高生存能力, 而中后期对叶片光合能力的投资会随物理防御结构的增加而降低, 在沙丘的固定过程中, 黑沙蒿在资源权衡策略上存在从“快速投资-收益型”向“缓慢投资-收益型”变化的趋势, 以此来适应环境变化。此外, 祁鲁玉等(2022)通过室内盆栽实验, 对中国暖温带3种常见本地灌木和2种外来灌木的叶性状及生物量等指标进行测量, 分析了不同物种幼苗的生长策略与生态位分化机制, 表明了本地种表现出比较保守的资源利用能力, 而外来物种偏向于较高的光合速率。

在种间关系及生态适应方面, 张零念等(2022)在滇中云南杨梅灌丛中发现木本植物物种的生态位重叠指数普遍偏小, 主要物种之间对环境资源的需求相似性不大, 种间竞争较弱; 而主要物种的种间联结呈显著正相关关系, 群落各物种之间可以协调共存, 促进群落结构稳定。在干旱半干旱区灌丛群落中, 灌木物种对草本植物的促进作用随水分的减少并不总是线性增强, 而是受到多种群落生态特征的共同影响, 这对于系统认识灌丛促进作用在退化生态系统植被恢复中的贡献具有重要意义(崔光帅等, 2022)。此外, 高璐鑫等(2022)在区域尺度上对中亚热带北部灌丛群落中物种的空间周转规律及其驱动因素进行了分析, 发现气候因子对物种周转产生显著影响, 海拔、坡度、土壤总氮含量等因素与物种周转速率呈现正相关关系, 土壤因子为驱动物种周转的关键因素。

针对山地灌丛群落的土壤化学计量与分异规律, 牟文博等(2022)在祁连山地区开展了灌丛土壤养分化学计量特征的空间分布格局研究, 表明了灌丛群落土壤养分含量存在显著的海拔格局, 有机碳、全氮的含量随着海拔升高呈现先增后减的趋势, 且与灌丛地上生物量呈正相关关系, 而全磷含量随海拔升高持续增加, 高寒灌丛植物生产力受氮制约严重, 适量增施氮肥有助于缓解土壤氮限制性作用对高寒地区灌丛群落稳定性的影响。

本专辑未覆盖中国所有灌丛群落的多样性格局及空间分布规律, 但针对典型灌丛群落类型的物种组成及其变化特征、形成机制、生态功能及化学计量特征等方面对灌丛群落进行了研究, 这对于了解我国灌丛植物群落的格局与过程具有重要意义。

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Mou WB, Xu DH, Wang XJ, Jing WM, Zhang RY, Gu YL, Yao GQ, Qi SH, Zhong L, Gou YF (2022).

Soil carbon, nitrogen, and phosphorus stoichiometry along an altitude gradient in shrublands in Pailugou watershed, China

Chinese Journal of Plant Ecology, 46, 1422-1431.

[本文引用: 1]

[ 牟文博, 徐当会, 王谢军, 敬文茂, 张瑞英, 顾玉玲, 姚广前, 祁世华, 张龙, 苟亚飞 (2022).

排露沟流域不同海拔灌丛土壤碳氮磷化学计量特征

植物生态学报, 46, 1422-1431.]

[本文引用: 1]

Pavoine S, Bonsall MB (2011).

Measuring biodiversity to explain community assembly: a unified approach

Biological Reviews, 86, 792-812.

DOI:10.1111/j.1469-185X.2010.00171.x      URL     [本文引用: 1]

Piao SL, Fang JY, Ciais P, Peylin P, Huang Y, Sitch S, Wang T (2009).

The carbon balance of terrestrial ecosystems in China

Nature, 458, 1009-1013.

DOI:10.1038/nature07944      URL     [本文引用: 1]

Pistón N, Schöb C, Armas C, Prieto I, Pugnaire FI (2016).

Contribution of co-occurring shrub species to community richness and phylogenetic diversity along an environmental gradient

Perspectives in Plant Ecology, Evolution and Systematics, 19, 30-39.

DOI:10.1016/j.ppees.2016.02.002      URL     [本文引用: 2]

Qi LY, Chen HN, Sairebieli K, Ji TY, Meng GD, Qin HY, Wang N, Song YX, Liu CY, Du N, Guo WH (2022).

Growth strategies of five shrub seedlings in warm temperate zone based on plant functional traits

Chinese Journal of Plant Ecology, 46, 1388-1399.

[本文引用: 1]

[ 祁鲁玉, 陈浩楠,库丽洪•赛热别力, 籍天宇, 孟高德, 秦慧颖, 王宁, 宋逸欣, 刘春雨, 杜宁, 郭卫华 (2022).

基于植物功能性状的暖温带5种灌木幼苗生长策略

植物生态学报, 46, 1388-1399.]

[本文引用: 1]

Stokes CJ, Archer SR (2010).

Niche differentiation and neutral theory: an integrated perspective on shrub assemblages in a parkland savanna

Ecology, 91, 1152-1162.

PMID:20462129      [本文引用: 1]

Investigations of structure in ecological communities need to move beyond the dichotomy between niche and neutral theory to broader conceptual frameworks that accommodate both neutral stochastic and biological structuring processes in organizing species assemblages. We propose and test a framework that integrates niche and neutral-assembly perspectives and determines their relative contributions in structuring diverse shrub species assemblages in a parkland savanna. Our approach proposes that stochastic dispersal processes initially govern the assemblage of species in discrete shrub clusters developing in grassland, but that community structure subsequently develops through the progressive action of first positive, then negative interactions among species. A comparison of observed patterns of occurrence and niche models for 12 shrub cluster species against neutral predictions revealed that neutral stochastic, island biogeographic processes accounted for most patterns of species occurrence. One species showed strong evidence of successional differentiation, whereas evidence of slight recruitment biases for five others was equivocal. Our results demonstrate the usefulness of an approach that accommodates contributions of both neutral and niche assembly rather than assuming either process alone is sufficient to account for community structure. Further development and testing of robust and falsifiable neutral theory will allow ecologists to critically evaluate the relative roles of niche differentiation and neutral, stochastic processes in structuring communities.

Tang XL, Zhao X, Bai YF, Tang ZY, Wang WT, Zhao YC, Wan HW, Xie ZQ, Shi XZ, Wu BF, Wang GX, Yan JH, Ma KP, Du S, Li SG, et al. (2018).

Carbon pools in China’s terrestrial ecosystems: new estimates based on an intensive field survey

Proceedings of the National Academy of Sciences of the United States of America, 115, 4021-4026.

[本文引用: 1]

Wang GH, Fang JY, Guo K, Xie ZQ, Tang ZY, Shen ZH, Wang RQ, Wang XP, Wang DL, Qiang S, Yu D, Peng SL, Da LJ, Liu Q, Liang CZ (2020).

Contents and protocols for the classification and description of Vegetation Formations, Alliances and Associations of Vegetation of China

Chinese Journal of Plant Ecology, 44, 128-178.

DOI:10.17521/cjpe.2019.0272      [本文引用: 1]

Vegegraphy, a compound word of prefix “vege-” of “vegetation” and suffix of “-graphy” (description), is a series of monographs that describe species composition, structures, functions, distribution and environmental settings of a set of plant communities and/or their combinations. The Vegegraphy of China, which will be composed of 48 volumes and about 110 issues, is the first version to describe the medium-level units (Alliance Group, Alliance) and the lower-level units (Association Group, Association), and summarizes the higher-level units (Vegetation Formation Group, Vegetation Formation, Vegetation Subformation) in the Chinese Vegetation Classification System. The description of the higher-level units is highly generalized, mainly based on the data and evidences from the classification and description of the medium- and the lower-level classification units of the Chinese vegetation classification system, focusing on the geographic distribution, natural environment, community physiognomy, vegetation type and diversity, dominant species, values for biodiversity conservation, and the status of current vegetation resources at the scales of both worldwide and China. It is an important revision and expansion of the relevant content of Vegetation of China (1980) and will be the most authoritative and accurate description of the basic characteristics of Chinese vegetation. Alliance Group in each issue is briefly described on geographical distribution, natural environment, vegetation types, significance and conservation etc. Alliance as a key medium- level unit is to be described on several aspects, i.e., geographical distribution, natural environment, ecological characteristics, vegetation composition, vegetation structure, vegetation types and characteristics, biological characteristics of dominant species, biomass and productivity, vegetation dynamics and succession, and significance and conservation. As a core content of the above mentioned aspects, vegetation classification and description are conducted under the guidance of the “80 scheme” of Vegetation of China, using vegetation survey data as credentials to quantitatively differentiate vegetation types accordingly. Specifically, a vegetation classification scheme is determined based on supraterraneous stratification, and a set of diagnostic species while vegetation descriptions focus on physiognomy, community structure, species composition, including quantitative features of dominant species, companion species, constant species and accidental species, as well as their environmental preferences. In this study, we first put forward a code of vegetation nomenclature for the medium-level units (Alliance Group, Alliance) and the lower-level units (Association Group, Association) in the Chinese Vegetation Classification System. According to this vegetation nomenclature, the name of a vegetation type is composed of the name of a diagnostic species (may also be a dominant species) of each layer of the vegetation type and the name of the higher-level units (Vegetation Formation or Vegetation Formation Group) to which it belongs, which guarantees both scientific soundness and practical sense of the vegetation nomenclature. Contents and protocols for the research and editing of “Vegegraphy of China”, including vegetation survey and sample collection, literature compilation, climate, soil and topography data collection and compilation, vegetation classification, vegetation nomenclature, vegetation description outlines, are elaborated or exemplified in detail.

[ 王国宏, 方精云, 郭柯, 谢宗强, 唐志尧, 沈泽昊, 王仁卿, 王襄平, 王德利, 强胜, 于丹, 彭少麟, 达良俊, 刘庆, 梁存柱 (2020).

《中国植被志》研编内容与规范

植物生态学报, 44, 128-178.]

DOI:10.17521/cjpe.2019.0272      [本文引用: 1]

植被志是基于植被(或植物群落)调查资料, 全面记叙植被的外貌、物种组成、结构和功能, 以及地理分布和生境条件等特征, 并对同类植被进行归纳和总结的志书。《中国植被志》是第一部对中国植被进行全面记述的志书, 预计完成约48卷110册。在借鉴《中国植被》(1980)(简称“80方案”)植被分类基本原则的基础上,《中国植被志》将对中国植被分类系统的高级分类单位(植被型组、植被型和植被亚型)进行归纳和总结, 对中级和低级分类单位(群系组、群系、亚群系, 群丛组、群丛)进行详细描述。植被高级分类单位的描述具有概括性质, 是在中国植被分类系统中级和低级分类单位描述的基础上, 对其在全球和中国境内的地理分布、自然环境、群落外貌、植被类型及多样性、优势种或共优势种、生物多样性保育价值以及资源现状等进行概述, 并对“80方案”相关内容进行修订和拓展, 将提供对中国植被基本特征客观、准确的记述。在植被中级和低级分类单位中, 群系组描述的内容包括地理分布、自然环境、群落外貌、植被类型以及价值与保育等内容; 群系描述的主要内容包括地理分布、自然环境、生态特征、物种组成、群落结构、群丛组和群丛的分类与描述、优势种的生物学特性、生物量与生产力、植被动态与演替以及价值与保育等方面。作为植被志研编的核心内容, 群丛组和群丛的分类与描述主要基于植被调查资料, 采用数量分类方法, 根据群落结构和物种组成的差异划分出不同的植被类型, 并对其基本特征进行定量描述和归纳。其中, 群落的层片结构、特定植被分类单元的特征种或特征种组的筛选与甄别是植被类型划分的关键环节; 而群落外貌, 群落结构, 物种组成, 各类物种的生长习性、生境的偏适性等是群丛组和群丛描述与归纳的重点内容。该文提出了中国植被中级和低级分类单位的命名方案, 其特点在于植被类型的科学名称中同时体现了植被分类单元特征种或优势种的名称及其所属的高级植被分类单位(植被型组或植被型)的名称, 兼顾了植被名称的规范性与实用性。《中国植被志》的研编工作由文献整编、群落调查、数据分析与整理、文本撰写等环节组成。该文对植被样方的调查与收集, 文献收集与整编, 气候、土壤、地形等相关数据的来源及其整理方法, 植被分类方法, 植被命名, 植被分类单元描述的内容, 植被志章节编写大纲、体裁及撰写等多个规范进行了详细的阐述或示例。

Xie ZQ, Tang ZY, Liu Q, Xu WT (2019). Carbon Budget of Shrub Ecosystem in China. Science Press, Beijing.

[本文引用: 1]

[ 谢宗强, 唐志尧, 刘庆, 徐文婷 (2019). 中国灌丛生态系统碳收支研究. 科学出版社, 北京.]

[本文引用: 1]

Yang X, Guo YP, Mohhamot A, Liu HY, Ma WH, Yu SL, Tang ZY (2017).

Distribution of biomass in relation to environments in shrublands of temperate China

Chinese Journal of Plant Ecology, 41, 22-30.

DOI:10.17521/cjpe.2016.0199      [本文引用: 1]

Aims Shrubland is one of the most widely distributed vegetation types in northern China. Previous studies on pattern and dynamics of plant biomass have been focused on forest and grassland ecosystems, while relevant knowledge on shrubland ecosystems is lacking. It is important to include shrublands in northern China to improve the accuracy in estimating the terrestrial ecosystem biomass in China. Methods Based on investigations and samplings from 433 shrubland sites, we explored the distribution and allocation patterns of biomass in relation to climatic and soil nutrient factors of shrublands of temperate China. Important findings The average shrubland biomass density in northern China is 12.5 t·hm-2. It decreases significantly from temperate deciduous shrubland in northeast to desert shrubland in northwest. The average biomass density of temperate deciduous shrubland, alpine shrubland, and desert shrubland is 14.4, 28.8, and 5.0 t·hm-2, respectively. Within temperate deciduous shrublands, plant biomass is lower in North China than in Northeast China. The average aboveground and belowground biomass density of shrub layer is 4.5 and 5.4 t·hm-2, respectively; while that of grass layer is 0.8 and 1.8 t·hm-2, respectively. Environmental factors affect biomass allocation across different plant organs. The belowground-aboveground biomass ratio of shrub exhibits no significant changes with environmental variables. The leaf-stem ratio increases with annual precipitation, and leaf biomass is low in arid region.

[ 杨弦, 郭焱培,安尼瓦尔•买买提, 刘鸿雁, 马文红, 于顺利, 唐志尧 (2017).

中国北方温带灌丛生物量的分布及其与环境的关系

植物生态学报, 41, 22-30.]

DOI:10.17521/cjpe.2016.0199      [本文引用: 1]

目前对植物生物量分布格局和分配的研究多集中在森林和草地生态系统, 对灌丛的相关研究较少。灌丛是中国北方广泛分布的植被。研究灌丛生物量分布格局及其分配是对估算我国陆地生态系统碳库的重要补充。该文通过对中国北方温带灌丛的大范围野外调查和采样, 计算中国北方433个典型灌丛样地的生物量及其在各器官间的分配, 并研究它们与气候和土壤营养等环境因子的关系。结果表明: 中国北方温带灌丛平均生物量为12.5 t&#x000b7;hm<sup>-2</sup>, 其中灌木层地上、地下生物量分别为4.5和5.4 t&#x000b7;hm<sup>-2</sup>, 草本层地上、地下生物量分别为0.8和1.8 t&#x000b7;hm<sup>-2</sup>; 凋落物量为2.5 t&#x000b7;hm<sup>-2</sup>。不同类型中, 温带落叶灌丛、亚高山落叶阔叶灌丛、荒漠灌丛平均生物量分别为14.4、28.8和5.0 t&#x000b7;hm<sup>-2</sup>。东西部生物量分布差异较大, 东部温带落叶灌丛总生物量高于西部的荒漠灌丛。东部温带落叶灌丛中, 东北地区的灌丛生物量稍低于华北地区。灌木的地下-地上生物量比不随水分和土壤养分变化, 而叶-枝生物量比受水分影响, 在干旱区域叶-枝生物量比较低。

Yao ZY, Shi YB, Ma WH, Zhao LQ, Juhua, Haoren T, Su C, Zhang XY, Liang CZ (2022).

Community characteristics of Ostryopsis davidiana alliance in Nei Mongol, China

Chinese Journal of Plant Ecology, 46, 1342-1349.

[本文引用: 1]

[ 要振宇, 史亚博, 马文红, 赵利清, 菊花, 浩仁塔本, 苏闯, 张芯毓, 梁存柱 (2022).

内蒙古地区虎榛子群系的群落特征

植物生态学报, 46, 1342-1349.]

[本文引用: 1]

Zhang LN, Zhu GQ, Yang K, Liu XY, Gong HD, Zheng L (2022).

Niche and interspecific association of main woody plants in Myrica nana shrubland in central Yunnan, China

Chinese Journal of Plant Ecology, 46, 1400-1410.

[本文引用: 1]

[ 张零念, 朱贵青, 杨宽, 刘星月, 巩合德, 郑丽 (2022).

滇中云南杨梅灌丛主要木本植物生态位与种间联结

植物生态学报, 46, 1400-1410.]

[本文引用: 1]

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