牦牛粪沉积下高寒草甸植物群落种间关联研究

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孔杨云, 于应文, 侯扶江.牦牛粪沉积下高寒草甸植物群落种间关联研究. 草业学报, 2017,26(3): 44-52
KONG Yang-Yun, YU Ying-Wen, HOU Fu-Jiang.Interspecific associations in plant communities under yak dung depositions in an alpine meadow. Acta Prataculturae Sinica,2017,26(3): 44-52 复制到剪切板

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牦牛粪沉积下高寒草甸植物群落种间关联研究

孔杨云, 于应文^*, 侯扶江

草地农业生态系统国家重点实验室,兰州大学草地农业科技学院,甘肃兰州 730020

*通信作者Corresponding author. E-mail: yuyw@lzu.edu.cn

作者简介:孔杨云(1992-),女,甘肃永靖人,在读硕士。E-mail:kongyy15@lzu.edu.cn

收稿日期: 2016-04-05

基金项目:国家重点研发计划课题(2016YFC0501902),草地农业生态系统国家重点实验室2015年度开放课题和云南省高端科技人才引进项目(2012HA012)资助

摘要

植物群落构成和种间关联,反映物种在空间上的分布及相互依存和制约关系。为探究牦牛粪沉积和植被斑块互作下草地植物群落演变趋势及稳定性机制,以青藏高原东北缘高寒草甸禾草/杂类草斑块和嵩草/杂类草斑块为对象,分析牦牛粪沉积下两种斑块中主要植物种的重要值( IV)、Jaccard种间关联指数和Spearman秩相关系数的变化规律。结果表明,1)粪沉积使禾草/杂类草斑块的建群种扁穗冰草的 IV增加和西北针茅的 IV降低,嵩草/杂类草斑块的建群种线叶嵩草的 IV降低;2)禾草/杂类草斑块中,粪沉积(DP)和对照(CK)的正负关联种对比为0.61和0.63,显著联结种对为6.6%和6.4%;嵩草/杂类草斑块中,DP和CK的正负关联种对比为0.90和0.72,显著联结种对为2.8%和14.5%;且二者显著联结的种对多存在于优势种或亚优势种与伴生种之间;3)两种斑块中多数种对间联结性弱,群落处于不稳定阶段;粪沉积使嵩草/杂类草斑块的稳定性增强,利于禾草/杂类草斑块的演变;4)依据优势种对环境的适应方式和主导生态因子,将两种斑块的群落植物种划分为喜阳耐旱植物、喜阴喜湿植物和喜阳喜湿植物三大生态种组。

关键词: 粪斑; 禾草/杂类草斑块; 嵩草/杂类草斑块; 重要值; Jaccard指数; Spearman秩相关

Interspecific associations in plant communities under yak dung depositions in an alpine meadow

KONG Yang-Yun, YU Ying-Wen^*, HOU Fu-Jiang

State Key Laboratory of Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China

Abstract

Plant community composition and interspecific associations reflect inter-dependence and competition between species, influencing their spatial distribution. To explore the evolutionary trends and stability mechanisms of grassland communities influenced by the interaction between yak dung depositions and vegetation we analyzed the importance values ( IV) of different species, Jaccard association indices and Spearman rank correlation coefficients of the main species present in patches of grass/forb and Kobresia/forb swards under yak dung deposition in an alpine meadow on the northeast Qinghai-Tibetan Plateau. The results indicated that: 1) dung deposition (DP) increased the Agropyron cristatum IV and decreased the Stipa krylovii IV in grass/forb patches, while decreasing the Kobresia capillifolia IV for Kobresia/forb parches; 2) in grass/forb patches, the species-pairs ratios of positive and negative associations under DP and control (CK) were 0.61 and 0.63, respectively and their respective associated species-pairs were 6.6% and 6.4%. For Kobresia/forb patches, the species-pairs ratio of positive and negative associations in DP and CK were 0.90 and 0.72, respectively and their respective associated species-pairs were 2.8% in DP and 14.5% in CK, exhibited mainly between the dominant or sub-dominant species and the companion species; 3) the interspecific association of most species-pairs was weak in both vegetation types; yak dung strengthened the stability of Kobresia/forb patches benefiting the generation of grass/forb patches; 4) according to their adaptability to environment and the key ecological factors affecting plants, the main plant species of two patches were divided into three ecological groups: shade-intolerant and drought-tolerant plants, shade-tolerant and drought-intolerant plants, and, shade-intolerant and drought-intolerant plants.

Keyword: dung patch; grass/forb patches; Kobresia/forb patches; important value; Jaccard association index; Spearman rank correlation

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植物种间关联通常由群落生境差异影响物种分布而引起, 是群落基本特征之一^[1] 。根据群落内种对对生境的选择及其相互促进和抑制关系, 将群落植物种间关系分为正相关、负相关和不相关^[2] 。种对间表现为正相关, 是由于物种具有相近的生物学特性和对环境具有相似的生态适应性以及一个种群的存在可以为另一个种群的生存提供条件; 种对间表现为负相关, 是由于物种生物学特性和对环境喜好的不同及相互竞争^[3] 。植物种间关联既是一种对物种存在与否二元数据的定性分析, 也是对物种数量特征数据的定量分析^[2] 。研究草地植物种间关联, 有助于深入了解草地植物物种间的相互依存和制约关系及群落结构和演替动态变化。采食、践踏和排泄物沉积作为放牧系统中家畜作用于草地的3种方式, 均是草地植物种间关系和植被构成发生变化的主要影响因素, 在草地植被演替中起重要作用^[4]。

青藏高原高寒草甸放牧历史悠久, 拥有世界独特的面积为1.3亿hm²高寒草地及1400万头牦牛, 从而牦牛排泄物沉积量大, 分布广^[5] 。由于垫状牛粪沉积对草地植物具有窒息作用, 使粪斑下植物枯萎死亡^[6] ; 且粪沉积后期或粪斑破碎后因粪中养分的施肥作用, 促进粪块及其毗邻区域植物生长; 二者均通过影响草地植被构成的改变而引起小尺度植被异质性的形成。因此, 研究牦牛粪沉积下, 高寒草甸植被构成及种间关联, 对于高寒草甸植被演替动态和植物种间共存竞争及群落稳定性机制的揭示具有重要意义^{[7, 8, 9]}。

诸多学者对群落类型^[10]和不同管理方式(封育和放牧)^{[11, 12]}及鼠丘密度^[13]下, 青藏高原高寒草甸植物群落构成及种间关联进行了分析。研究表明, 高寒草甸植物种间关联与封育年限有关, 短期封育表现为微弱负关联, 5~6年的封育时间无关联^[11]; 自由放牧较划区轮牧降低高寒草甸主要植物种间的关联程度和群落稳定性^[12]; 线叶嵩草(Kobresia capillifolia)草地群落总体关联性表现为显著负相关且处于不稳定阶段^[10]; 高山嵩草(Kobresia pygmaea)草地群落的总体联结性与高原鼠兔有效洞口密度有关, 由低洞口密度(≤ 224 个/hm²)下的正关联转变为高洞口密度(> 224 个/hm²)下的负关联^[13]。但家畜排泄物沉积下, 高寒草甸植物构成及种间关系的研究, 限于牦牛粪沉积下草地植物种生态位^[14]及植物种子库构成特征^{[15, 16]}分析。由于草地植被异质性和家畜采食的非均匀分布, 使粪斑和草地植被斑块产生叠加效应, 而植被构成对粪沉积的响应因群落植物种构成不同^[17]。截至目前, 牦牛粪沉积与草地植被构成的整合研究尚未见报道。

本研究采用Jaccard种间关联度指数及Spearman秩相关方法, 对牦牛粪沉积下青藏高原东北缘高寒草甸禾草/杂类草斑块和嵩草/杂类草斑块植物群落构成和种间关联进行定量分析, 揭示高寒草甸植物物种共存竞争及群落稳定性机制, 为高寒草甸的管理提供理论基础和实践依据。

1 材料与方法

1.1 试验地概况

研究区位于青藏高原东缘的甘肃省天祝县抓喜秀龙乡的高寒草甸上, 地理坐标为E 102° 40'-102° 47', N 37° 11'-37° 14', 海拔2960 m。年均气温-0.1 ℃, 1月均温-18.3 ℃, 7月均温12.7 ℃, > 0 ℃年积温 1380 ℃; 年降水量416 mm, 无绝对无霜期, 仅分冷热两季, 土壤类型为亚高山草甸土。草地植被斑块主要有禾草/杂类草、嵩草/杂类草、珠芽蓼(Polygonum viviparum)斑块等。草地植物种主要有矮嵩草(Kobresia humilis)、线叶嵩草、扁穗冰草(Agropyron cristatum)、西北针茅(Stipa krylovii)、洽草(Koeloria cristata)、早熟禾(Poa annua)、垂穗披碱草(Elymus nutans)、珠芽蓼和球花蒿(Artemisia smithii)等。

1.2 样地设置和粪斑标记

2012年5月末, 在研究区多年放牧牦牛的禾草/杂类草斑块与嵩草/杂类草斑块普遍出现的高寒草甸冷季牧场上, 设置面积为0.3~0.5 hm²的样地3块。在设置的3块样地的两类斑块上, 用木桩和彩色绳子分别标记沉积约3个月内的牦牛粪斑及其配对的对照样方各20~30个, 粪斑之间间距10 m以上, 每类斑块共标记60~90个粪斑。各样地设置后仍作为牦牛冬春放牧地, 除每年6-10月休牧外。草地利用率为80%~90%。

在研究区牦牛粪斑大小观测为0.02~0.03 m²的前期基础上, 考虑牛粪对草地植物的影响效应通常约为粪斑大小的2倍^[18], 将本研究牦牛粪斑大小设置为0.1 m², 以客观反映牦牛粪斑对草地植物的作用效应。

1.3 植物种生物量和盖度测定及重要值计算

依据牦牛粪斑对植物作用效应的前期观测结果, 即牦牛粪沉积对草地植物的作用效应一般在1年多后才表现出来。故在2013年7-8月(牦牛粪沉积约1年多后), 进行草地植物特性的测定。具体植被特征观测时, 以标记的粪斑为中心, 设置0.1 m²的正方形样方, 作为粪斑处理(dung patch, DP); 再在离标记的粪斑周围50~60 cm处草地上, 设置无粪尿排泄物沉积的与各标记牦牛粪斑配对的0.1 m²的正方形对照样方(control, CK)。分别测定各标记粪斑及其配对对照样方内的所有植物种的地上生物量和分种盖度, 地上生物量以干重计。因标记牦牛粪斑的部分破坏等, 用于本研究群落植物种重要值和种间关联分析的数据, 每类斑块为38对。

以分种生物量和盖度数据为基础, 计算两种群落的粪斑及其配对对照区域的植物种重要值。植物种重要值(importance values, IV)计算公式为:IV=(RC+RB)/2。式中:RC和RB分别表示植物种的相对盖度(relative coverage, RC)和相对生物量(relative biomass, RB), 样方中某种的相对盖度(或相对生物量)等于该种盖度(或生物量)除以所有植物种的盖度(或生物量)之和。

1.4 植物种种间关联度Jaccard指数

JI指数反映的是物种共同出现的百分率。本研究采用Jaccard指数^[2](简称JI指数)测定物种间关联程度, 它们以2× 2列关联表为基础计算, 其公式为:JI=a/(a+b+c)。式中:a为所有样方中种A和种B共同出现的样方数, b、c分别为所有样方中种A或种B单独出现的样方数。JI指数满足当a=0时, 取值为0, 表示种A和种B不同时出现在同一样方内, 即种A和种B完全独立或相异; 当a=S(样方数)时, b=c=0, 取值为1, 表示物种A、B同时出现在所有样方中, 即种A与种B绝对关联; 当c=0时, 表示完全关联, 即种B依赖于种A, 种B随种A出现而出现, 而种A可以单独出现。在借鉴张金屯^[2]划分方法, 将JI指数划分为以下4个等级, 0.75< JI≤ 1为强联结, 0.5< JI≤ 0.75为次强联结, 0.25< JI≤ 0.5为弱联结, 0≤ JI≤ 0.25为无联结, 以对嵩草/杂类草和禾草/杂类草草地群落主要植物种种间关联进行分析。

1.5 植物种种间Spearman秩相关系数计算

以植物种重要值数据为依据, 采用SPSS 16.0中Correlate中的Bivariate模块, 计算两种群落粪斑和对照区域主要植物种间Spearman秩相关系数及显著性检验^[19]。

2 结果与分析

2.1 植物种重要值

由表1可知, 禾草/杂类草斑块中, DP和CK的优势种和亚优势种均为扁穗冰草、西北针茅和洽草, 粪沉积使扁穗冰草的重要值增加26%, 使西北针茅的重要值降低19%; DP主要伴生种有早熟禾、大籽蒿、矮嵩草、紫花针茅、天蓝韭、阴山扁蓿豆、线叶嵩草、垂穗披碱草、冷蒿、藨草、二裂委陵菜和高山紫菀; CK主要伴生种有矮嵩草、早熟禾、大籽蒿、紫花针茅、天蓝韭、阴山扁蓿豆、二裂委陵菜、赖草、藨草和线叶嵩草; 且DP的线叶嵩草重要值约为CK的2.0倍。嵩草/杂类草斑块中, DP和CK的优势种和亚优势种分别为线叶嵩草和矮嵩草, 粪沉积使线叶嵩草重要值降低10%; DP主要伴生种有紫花针茅、珠芽蓼、垂穗披碱草、苔草、洽草、球花蒿和瑞香狼毒, CK主要伴生种有紫花针茅、珠芽蓼、洽草、高山嵩草、球花蒿、瑞香狼毒、垂穗披碱草、甘肃棘豆和乳白香青; 且斑处垂穗披碱草和洽草的重要值分别约为对照的3.6和0.7倍。说明, 粪沉积对禾草/杂类草斑块中植物种重要值构成的影响比嵩草/杂类草斑块的大, 利于禾草/杂类草群落植物种的演变。

表1 禾草/杂类草和嵩草/杂类草斑块中主要植物种重要值 Table 1 The importance values (IV) of main species in grass/forb and Kobresia/forb patches (IV≥ 0.01)

禾草/杂类草斑块Grass/forb patches						嵩草/杂类草斑块Kobresia/forb patches
粪斑Dung patch				对照CK		粪斑Dung patch				对照CK
植物名Plant name			重要值IV	植物名Plant name		重要值IV	植物名Plant name	重要值IV		植物名Plant name	重要值IV
扁穗冰草Agropyron cristatum (Acr)	0.216± 0.041	西北针茅 S. krylovii		0.259± 0.027		线叶嵩草 K. capillifolia		0.466± 0.087	线叶嵩草 K. capillifolia			0.514± 0.042
西北针茅Stipa krylovii (Sk)	0.211± 0.007	洽草K. cristata		0.183± 0.022	矮嵩草 K. humilis		0.150± 0.024	矮嵩草 K. humilis			0.159± 0.024
洽草Koeloria cristata (Kcr)	0.188± 0.012	扁穗冰草 A. cristatum		0.172± 0.014	紫花针茅 S.purpurea		0.072± 0.016	紫花针茅 S. purpurea			0.074± 0.018
早熟禾Poa annua (Pa)	0.070± 0.007	矮嵩草 K. humilis		0.072± 0.015	珠芽蓼 P. viviparum (Pv)		0.062± 0.021	珠芽蓼 P. viviparum			0.067± 0.017
大籽蒿Artemisia sieversiana (Asi)	0.050± 0.012	早熟禾 P. annua		0.060± 0.009	垂穗披碱草 E. nutans		0.045± 0.033	洽草K. cristata			0.038± 0.008
矮嵩草Kobresia humilis (Kh)	0.062± 0.003	大籽蒿A. sieversiana		0.046± 0.010	苔草Carex sp. (Csp)		0.041± 0.017	高山嵩草Kobresia pygmaea (Kp)			0.031± 0.017
紫花针茅Stipa purpurea (Sp)	0.037± 0.019	紫花针茅 S.purpurea		0.042± 0.036	洽草K. cristata		0.027± 0.003	球花蒿A. smithii			0.022± 0.002
天蓝韭Allium cyaneum (Acy)	0.029± 0.007	天蓝韭 A. cyaneum		0.034± 0.006	球花蒿 A. smithii (Asm)		0.027± 0.002	瑞香狼毒 S. chamaejasma			0.013± 0.005
阴山扁蓿豆Melilotoids ruthenica var. inschanica (Mri)	0.022± 0.002	阴山扁蓿豆M. ruthenica var. inschanica		0.026± 0.010	瑞香狼毒Stellera chamaejasma (Sc)		0.013± 0.002	垂穗披碱草 E. nutans			0.013± 0.009
线叶嵩草Kobresia capillifolia (Kca)	0.020± 0.006	二裂委陵菜 P. bifurca		0.019± 0.005				甘肃棘豆Oxytropis kansuensis (Ok)			0.012± 0.005
垂穗披碱草Elymus nutans (En)	0.020± 0.014	赖草Leymus secalinus (Ls)		0.018± 0.014				乳白香青Anaphalis lactea (Al)			0.012± 0.006
冷蒿Artemisia frigida (Af)	0.016± 0.006	藨草 S. triqueter		0.012± 0.006
藨草Scirpus triqueter (St)	0.012± 0.011	线叶嵩草 K. capillifolia		0.010± 0.007
二裂委陵菜Potentilla bifurca (Pb)	0.010± 0.004
高山紫菀Aster alpinus (Aa)	0.010± 0.008

表1 禾草/杂类草和嵩草/杂类草斑块中主要植物种重要值 Table 1 The importance values (IV) of main species in grass/forb and Kobresia/forb patches (IV≥ 0.01)

2.2 植物种种间关联度JI指数

主要植物种种间Jaccard指数显示, 禾草/杂类草斑块中的DP和CK中, 联结性较强的种对为11.4%和14.0%; 嵩草/杂类草斑块的DP和CK中, 联结性较强的种对为38.9%和38.2%(表2)。说明, 两种斑块中, 较多种对联结性弱, 种间独立性较强, 且禾草/杂类草斑块比嵩草/杂类草斑块的植物种对的联结更为松散。

表2 禾草/杂类草斑块和嵩草/杂类草斑块主要植物种种间Jaccard指数 Table 2 The interspecific Jaccard indices of main plant species in grass/forb and Kobresia/forb patches

2.3 植物种种间Spearman秩相关分析

Spearman秩相关分析显示, 禾草/杂类草斑块的正相关和负相关的种对数, DP为34和56对, CK为30和48对, DP和CK的正负关联种对比为0.61和0.63; 显著相关和不相关种对数, DP为6和84对, CK为5和73对, 显著联结种对为6.6%和6.4%(图1)。嵩草/杂类草斑块正相关和负相关的种对数, DP为17和19对, CK为23和32对, DP和CK的正负关联种对比为0.90和0.72; 显著相关和不相关种对数, DP为1和35对, CK为8和47对, 显著联结种对为2.8%和14.5%(图2)。说明, 两种斑块中多数种对间联结性弱, 群落处于不稳定阶段, 粪沉积使嵩草/杂类草斑块的稳定性增强。

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	图1 禾草/杂类草斑块主要植物种Spearman秩相关分析半矩阵图Fig.1 Semi-matrix of Spearman rank correlation analysis of main species in grass/forb patches * P< 0.05, * * P< 0.01, the same below.

在禾草/杂类草斑块中, DP为扁穗冰草-天蓝韭和西北针茅-藨草种对间显著正联结, 扁穗冰草-阴山扁蓿豆、扁穗冰草-垂穗披碱草、扁穗冰草-藨草和西北针茅-紫花针茅种对间显著负联结; CK为洽草-天蓝韭、洽草-大籽蒿、扁穗冰草-二裂委陵菜种对间极显著正联结, 西北针茅-紫花针茅、西北针茅-藨草、洽草-矮嵩草、洽草-大籽蒿、扁穗冰草-赖草、矮嵩草-天蓝韭和天蓝韭-赖草种对间显著负联结(图1)。在嵩草/杂类草斑块中, DP仅苔草-洽草种对间显著负联结, CK为矮嵩草-垂穗披碱草和瑞香狼毒-乳白香青种对间显著正联结, 珠芽蓼-洽草和瑞香狼毒-垂穗披碱草、线叶嵩草-矮嵩草、线叶嵩草-珠芽蓼、线叶嵩草-垂穗披碱草和垂穗披碱草-乳白香青种对间显著负联结(图2)。说明, 显著联结的种对多存在于优势种或亚优势种与伴生种之间。

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	图2 嵩草/杂类草斑块主要植物种Spearman秩相关分析半矩阵图Fig.2 Semi-matrix of Spearman rank correlation analysis of main species in Kobresia/forb patches

2.4 生态种组划分

高寒草甸植物群落植物种的生态习性不同, 依据群落主要植物种对主导环境因子, 如光照和水分的适应方式, 以及植物种间关联性特征, 将两种斑块群落植物种划分为三大生态种组(表3)。其中, 第一生态种组为多数禾本科、豆科和菊科等喜阳耐旱植物, 主要包括扁穗冰草、西北针茅、紫花针茅、赖草、高山嵩草、甘肃棘豆、阴山扁蓿豆和大籽蒿等13种处于草层的中上层, 具有喜光、耐旱、耐土壤贫瘠等趋同特征的植物, 草甸退化标志植物瑞香狼毒亦出现在该组, 组内多物种间呈显著相关, 群落处于不稳定状态。第二生态种组禾本科的早熟禾、莎草科的藨草和苔草及蓼科的珠芽蓼4种喜阴喜湿植物, 因本研究区高寒草甸水分不足, 第二生态种组植物种少且组内各物种间无显著关联, 植物种间独立性较强。第三生态种组为禾本科的垂穗披碱草和洽草, 莎草科的矮嵩草和线叶嵩草, 菊科的球花蒿和百合科的天蓝韭等喜阳喜湿的植物, 高寒草甸很难满足其生境要求而仅由6种植物种构成, 其中线叶嵩草与垂穗披碱草和矮嵩草呈极显著负关联而垂穗披碱草和矮嵩草呈显著正关联, 且线叶嵩草是嵩草/杂类草斑块的优势种, 种组内竞争激烈, 种组有演替的趋势。

表3 禾草/杂类草和嵩草/杂类草斑块群落中主要植物种生态分组 Table 3 Ecological species groups of main species in grass/forb and Kobresia/forb patches

3 讨论

本研究主要植物种Jaccard指数和种间Spearman秩相关分析均显示, 无论粪沉积与否, 禾草/杂类草斑块和嵩草/杂类草斑块中, 多数种对间联结性弱, 群落成熟度较低, 结构简单, 整体上处于不稳定状态。这是由于草地在长期演替过程中, 植物种的生态位进一步分化, 生境异质性增加^[8], 各物种都占据有利的位置, 和谐共处, 相互依赖和相互竞争均大大降低, 多数种对联结程度不强, 独立性强^[9], 从而使两种斑块的群落结构不稳定, 具有明显的次生性或正处于演替前期阶段。

本研究嵩草/杂类草斑块和禾草/杂类草斑块的植物种对数的正负关联比均< 1, 但前者更接近于1, 从而前者的群落稳定性比后者强。其主要原因是嵩草/杂类草斑块中的优势植物为嵩草属的线叶嵩草和矮嵩草, 其在长期适应寒冷气候的进化过程中, 形成特殊稳定的适应外界不良环境的维持性特征, 如叶片少, 植株矮化、细绒化和草毡表层加厚等极度发育等, 从而具有较强的抗外界干扰和自我调控能力^[20]; 而禾草/杂类草斑块中的优势植物为扁穗冰草、西北针茅和洽草等, 因其叶片宽而多, 茎秆直立高大, 根系较深, 具较多须根, 对外界干扰敏感。这与房飞等^[21]对新疆天山南坡高寒草原夏季牧场羊茅(Festuca ovina)+紫花针茅群落, 以及王伟等^[10]和郭倩等^[22]报道的青海省称多县藏嵩草(Kobresia tibetica)群落和高山嵩草+异针茅(Stipa aliena)/杂类草群落及河南县线叶嵩草草地群落, 植物种对的正负关联比均≥ 1, 且群落整体上处于稳定发展阶段的研究结果存在分异。这可能与二者取样大小不同有关, 本研究为测定粪斑效应, 将样方大小设为0.1 m², 而这些相关研究的样方大小为0.25 m², 一般样方大则正关联的种对数多, 反之亦然; 也可能是二者样方数量、群落自身物种数的多寡和群落类型等差异造成^{[11, 21, 22]}。

本研究粪沉积使嵩草/杂类草斑块群落稳定性增强, 而对禾草/杂类草斑块群落稳定性影响小。类似施肥研究表明, 施肥降低群落稳定性^[23], 施N肥及N、P、K单施或联施或施羊粪, 均增加青藏高原矮嵩草群落和退化高寒草甸草地中禾草比例, 降低杂类草比例^{[24, 25]}。本研究与后者结果存在分异, 其主要原因是二者所施肥料类型不同。本研究牛粪不但富含氮、磷、钾及多种有机养分, 且为垫状形态, 其沉积后首先在草地上形成致密粪垫层, 进而对粪垫下的草地植物造成窒息而使其枯黄死亡, 因群落结构和植物建群种构成不同, 嵩草/杂类草斑块比禾草/杂类草斑块中处于上层的高大禾草和杂类草比例少, 从而前者比后者因粪窒息造成的枯黄死亡比例少。同时, 由于禾草相对莎草对牛粪和施肥反应更敏感, 更易吸收肥料中的养分^[14], 粪沉积后期, 禾草/杂类草斑块中的禾草因在短时间易吸收利用牦牛粪中的养分, 生长增强, 迅速占领群落上层空间, 进而遮蔽其他矮小物种, 以此弥补了粪沉积前期因粪斑覆盖草地而造成的损失, 从而使其在与其他功能群植物的竞争中取胜, 而嵩草/杂类草斑块中的莎草吸收牛粪中的养分少且慢。因此, 粪沉积后, 嵩草/杂类草斑块向趋于嵩草群落的方向发展, 更趋于稳定; 而禾草/杂类草群落向趋于禾草的方向发展, 更趋于不稳定。同时, 由于粪斑具有特殊气味及寄生虫, 粪沉积初期-中期, 家畜弃食粪斑及其毗邻区域植物^[26]; 随粪块沉积时间的延长及粪中养分的释放, 粪斑及其周围植物通过土-草系统吸收粪中养分后形成高营养斑块, 反过来又吸引家畜采食^{[26, 27]}, 从而家畜粪沉积和采食互作对草地植被的复杂效应, 使粪块及其周围区域植物群落构成和种间关系发生改变^[6]。另一方面, 家畜具有在食源值与食源报酬之间的空间判断和权衡能力, 其也频繁采食草地上数量多、面积大的中、低营养植被斑块^[28]。可见, 草地群落稳定性和发展演替受放牧干扰复杂效应的调控。本研究仅阐释了家畜粪沉积下草地植物群落构成及种间关系变化, 而牛粪沉积对草地植被的作用过程和机制, 还需进一步研究。

植物种间关联或物种联结性是指不同物种在空间上的相互联结性, 体现物种间的依赖性与排斥性, 正联结种对对环境或资源有相互依存的相似需求, 负联结种对生境或资源具有相互排斥的不同需求^[29]。植物群落中优势种和亚优势种对群落的结构和稳定性起重要作用^[30]。本研究禾草/杂类草斑块中, 粪沉积使西北针茅-藨草的种间关联由负显著变为正显著, 西北针茅-紫花针茅由显著负关联变为极显著负关联。这是由于西北针茅属密丛型浅根禾草, 具抗旱耐寒特征, 藨草属莎草科挺拔直立型根状茎植物, 具抗寒耐湿特征, 二者因生态特征差异对生境和资源具不同要求而互相排斥, 但随着牦牛粪的沉积, 两者吸收粪中养分资源而具有相似的资源需求所导致; 西北针茅和紫花针茅均为密丛型浅根同属禾草, 在长期进化和不同微环境适应过程中, 对生境具有不同的生态适应性或相互竞争而具显著负关联, 是其对不同空间资源利用的结果, 而粪沉积下二者对粪中养分资源利用程度的竞争加剧而转为极显著负关联。

本研究分析了高寒草甸植物种间关联及群落构成特征, 推测了群落的稳定性和演替动态, 其结果有助于认识草地群落的基本特征和物种替代机制, 为放牧家畜排泄物的深入研究奠定了基础。

4 结论

无论粪沉积与否, 禾草或嵩草/杂类草斑块总体上群落植物种间关系均表现为负联结, 多数种对间联结性弱, 禾草/杂类草斑块比嵩草/杂类草斑块的植物种对的联结更为松散, 群落处于不稳定阶段。粪沉积对禾草/杂类草斑块中建群种重要值的影响比嵩草/杂类草斑块的大, 利于禾草/杂类草斑块的演变, 使嵩草/杂类草斑块的稳定性增强。依据两种斑块群落优势种对环境的适应方式和主导生态因子, 将植物种划为喜阳耐旱植物、喜阴喜湿植物和喜阳喜湿生植物三大生态种组。

The authors have declared that no competing interests exist.

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[19]	Guo S L, Yu J, Chen G Q. Ecological Data Analyses-Methods, Programs and Software[M]. Beijing: Science Press, 2015: 56. 郭水良, 于晶, 陈国奇. 生态学数据分析——方法、程序和软件[M]. 北京: 科学出版社, 2015: 56. [本文引用:1]
[20]	Cao G M, Long R J. *System stability and its self: maintaining mechanism by grazing in Alpine Kobresia* Meadow*. Chinese Journal of Agrometeorology, 2009, 30(4): 553-559. 曹广民, 龙瑞军. 放牧高寒嵩草草甸的稳定性及自我维持机制*. 中国农业气象, 2009, 30(4): 553-559. [本文引用:1]
[21]	Fang F, Hu Y K, Zhang W, et al. Numerical analysis of interspecific relationships in alpine steppe community in Bayanbulak. Acta Ecologica Sinica, 2012, 32(6): 1898-1906. 房飞, 胡玉昆, 张伟, 等. 高寒草原植物群落种间关系的数量分析. 生态学报, 2012, 32(6): 1898-1906. [本文引用:2]
[22]	Guo Q, Wang X L, Zhang R X, et al. *Analysis on the vegetation composition and interspecific association of Kobresia* meadow in southern pastoral area of southern Qinghai Province*. Acta Agrestia Sinica, 2016, 24(2): 24-31. 郭倩, 王旭丽, 张润霞, 等. 青南牧区南部嵩草草甸群落植被构成及种间关联*. 草地学报, 2016, 24(2): 24-31. [本文引用:2]
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[24]	Shen Z X, Zhou X M, Chen Z Z, et al. *Response of plant groups to simulated rainfall and nitrogen supply in Alpine Kobresia humilis* meadow*. Acta Phytoecologica Sinica, 2002, 26(3): 288-296. 沈振西, 周兴民, 陈佐忠, 等. 高寒矮嵩草草甸植物类群对模拟降水和施氮的响应*. 植物生态学报, 2002, 26(3): 288-296. [本文引用:1]
[25]	Chen W Y, Qi D C, Li G Y, et al. Effects of fertilization on grass community diversity and productivity of degraded alpine grassland at Maqu, in South of Gansu Province. Journal of China Agricultural University, 2009, 14(6): 31-36. 陈文业, 戚登臣, 李广宇, 等. 施肥对甘南高寒草甸退化草地植物群落多样性和生产力的影响. 中国农业大学学报, 2009, 14(6): 31-36. [本文引用:1]
[26]	Hutchings M R, Gordon I J, Kyriazkis I, et al. Sheep avoidance of faeces-contaminated patches leads to a trade-off between intake rate of forage and parasitism in subsequent foraging decisions. Animal Behaviour, 2001, 62: 955-964. [本文引用:2]
[27]	Sun H, Yu Y W, Ma X L, et al. *Effects animal dung deposition on grassland nutrients and botanical composition in mixed Lolium perenne* and Trifolium repens pasture in Guizhou plateau*. Pratacultural Science, 2014, 31(3): 488-498. 孙红, 于应文, 马向丽, 等. 畜粪沉积对贵州高原黑麦草+白三叶草地养分和植被构成的影响*. 草业学报, 2014, 31(3): 488-498. [本文引用:1]
[28]	Edwards G R, Newman J A, Parsons A J, et al. The use of spatial memory by grazing animals to locate food patches in spatially heterogeneous environments: an example with sheep. Applied Animal Behaviour Science, 1996, 50: 147-160. [本文引用:1]
[29]	Zhu G L, Yang H Q, Wei Z J, et al. *Interspecific associations in Stipa breviflora* desert steppe community*. Acta Prataculturae Sinica, 2004, 13(5): 33-38. 朱桂林, 杨洪琴, 卫智军, 等. 短花针茅草原群落种间联结研究*. 草业学报, 2004, 13(5): 33-38. [本文引用:1]
[30]	Qin L. Statistical Ecology[M]. Beijing: China Forestry Press, 2009: 81-97. 覃林. 统计生态学[M]. 北京: 中国林业出版社, 2009: 81-97. [本文引用:1]

2015

0.0

. 2015, 39(12):1136-1145 DOI:doi:10.17521/cjpe.2015.0110

Species association and correlation between vertical layers in the Liquidambar formosana community in Tiantong Region, Zhejiang Province

浙江天童枫香树群落不同垂直层次物种间的联结性与相关性

Zhou L L , Zhang Q Q , Zhao Y T

周刘丽, 张晴晴, 赵延涛

Aims Interspecific relationship is one of the most important properties in plant community. Analyzing species association and correlation between vertical layers in plant community is ecologically crucial for understanding community structure, dynamics and classification. The objective of this study was to test how plant species in contrasting vertical layers of plant community associate and correlate interspecifically. Methods The study sites are located in Tiantong region in Zhejiang Province. Community structure and species composition were measured in an early succession community of Liquidambar formosana. Interspecific relationships were examined between 3 tree species and 3 sub-tree species, between 3 tree species and 28 shrub species, and between 3 sub-tree species and 28 shrub species. Based on 2×2 contingency tables of species presence/absence, variance ratioanalysis, association coefficient, χ^2 test, Pearson correlation and Spearman rank correlation were conducted simultaneously to determine species association and correlation. Important findings With respect to tree and sub-tree layers, significantly positive association and correlation were not observed for overall species. Species were independent each other for the most species pairs, with few species pairs showing significantly negative association and correlation only. Regarding tree and shrub layers, there were significantly positive association and correlation for each of overall species and specific species pairs. Species between tree and shrub pair were more dependent each other. Similarly, significantly positive association and correlation were also found for overall species and specific species pairs between sub-tree and shrub layers, indicating a close interspecific relationship. These results suggest that differences in convergences of species behavior and habitat acclimatization, and complementary differences in habitat requirement over species between vertical layers might be the main driver affecting patterns of species association and correlation in the early successional community.

种间关系是植物群落重要的数量和结构特征之一，分析群落不同垂直层次物种的种间联结和相关性对于理解群落结构、动态和分类等具有重要的生态意义。本研究选取浙江省天童处于演替前期枫香树群落乔木层的3个物种、亚乔木层3个物种以及灌木层28个物种为研究对象，基于物种存在与否的2×2联列表，运用方差比率法、χ^2统计量、联结系数、Pearson 相关系数和 Spearman 秩相关系数检验了三个垂直层次间物种的联结性与相关性。结果发现：(1) 乔木层与亚乔木层物种总体间不存在显著正联结，乔木层与亚乔木层主要物种间相互独立的种对较多，较少种对显著联结或相关；(2)乔木层与灌木层物种总体显著正联结，显著关联或相关的种对较多，种对间依赖性较强；(3)亚乔木层与灌木层物种总体同样存在显著正联结，种对间关系较为紧密。该研究结果表明，群落垂直层次间物种生态习性和对生境适应的趋同性，以及在垂直方向上对生境要求的互补性差异是决定演替前期其种对关系的主要因素。

... 植物种间关联通常由群落生境差异影响物种分布而引起,是群落基本特征之一^[1] ...

2011

0.0

... 根据群落内种对对生境的选择及其相互促进和抑制关系,将群落植物种间关系分为正相关、负相关和不相关^[2] ...

... 植物种间关联既是一种对物种存在与否二元数据的定性分析,也是对物种数量特征数据的定量分析^[2] ...

... 本研究采用Jaccard指数^[2](简称JI指数)测定物种间关联程度,它们以2#cod#x000D7 ...

... 在借鉴张金屯^[2]划分方法,将JI指数划分为以下4个等级,0 ...

2011

0.0

. 2011, 29(3):278-287 DOI:doi:10.3724/SP.J.1142.2011.30278

Interspecific associations among dominant plant populations in Keteleeria hainanensis communities in Bawangling, Hainan Island

海南霸王岭海南油杉群落优势种群的种间联结性研究

Chen Y K , Yang X B , Li D H

陈玉凯, 杨小波, 李东海

Based on 2×2 contingency tables,a series of parameters including X 2 test,percentage of co-occurrence(PC),association coefficient(AC) and variance ratio(VR) were used to analyze the characteristics of interspecific associations among nineteen main plant populations in Keteleeria hainanensis communities.The results showed the association was nonsignificant positive correlation among the nineteen main populations.Among the 171 species pairs,87 pairs were positively correlated,82 pairs were negatively correlated,and 2 pairs showed no correlation.The X 2 test indicated that six positively correlated pairs and five negatively correlated pairs were significant or very significant.The percentage co-occurrence(PC) value for 144 species pairs was less than 40 percent,for 21 species pairs was equal or more than 40 percent,and for 6 species pairs was 0,indicating that a relatively low correlation existed among these species pairs.The more intense the positive association,the larger the value of niche overlap.The more intense the negative association,the smaller the value of niche overlap.In different dominant tree populations,the interspecific associations were different,but in the dominant species of Lithocarpus fenzelianus and K.hainanensis,the community showed high interspecific association and was more stable.Overall,K.hainanensis had the largest positive association with Psychotria rubra.

基于2×2联列表,应用 X 2 统计量、共同出现百分率、联结系数和多物种关联指数为参数,研究分析了海南油杉( Keteleeria hainanensis )所在群落19个主要种群的种间联结性特征。结果表明,19个主要种群在总体水平上表现为不显著的正联结。19个种群组成的171个种对中,有87对表现为正关联,82对为负关联,2对无关联。 X 2 检验有6对表现显著或极显著正关联,有5对表现显著或极显著负关联。有144个种对0 PC PC ≥40%,另有6个种对PC=0,表明171对种对中,只有少数种对存在显著的相关性,大多数种对联结关系未达到显著水平,种对间相关性较弱。种间正联结性越强,其生态位重叠值越大;种间负联结性越强,其生态位重叠值越小。不同优势种群下的种间联结性存在一定的差异,而以红椆(Lithocarpus fenzelianus)、海南油杉等为优势种群的群落正负关联比例较高,种间联结性较高,群落结构较稳定。总体上,海南油杉与九节( Psychot riarubra )存在最大的正联结关系。

... 种对间表现为负相关,是由于物种生物学特性和对环境喜好的不同及相互竞争^[3] ...

2004

0.0

. 2004, 24(4):784-789 DOI:doi:10.3321/j.issn:1000-0933.2004.04.020

A review on trampling by grazed livestock

放牧家畜的践踏作用研究评述

Hou F J , Chang S H , Yu Y W

侯扶江, 常生华, 于应文

Trampling is one of three ways for livestock to impact on grazing land is recognized to have longer active time and more permanent effect than intaking and excrement. It is therefore important to study the mechanism of trampling on grazing land in order to discover the mechanism of grassland degeneration and properly manage grazing ecosystems.(1) Trampling on vegetationWhile the trampling of livestock physically damaged aboveground herbage and buried the seeds, it was determined to be beneficial to the germ...

放牧家畜的践踏作用是家畜作用于草地的 3个途径之一 ,相对于采食和排泄物的作用具有作用时间长、直接作用的草地组分多和效果持久的特点。家畜践踏损伤牧草、埋实种子、促进萌发 ,减少凋落物的现存量 ,增加土壤的紧实度、容重 ,降低土壤孔隙度、水稳性团聚体、透水性和透气性 ,导致雨后水涝和植物根区缺氧 ,引发水土流失。国内外使用的 4种践踏强度指标分别存在不确定性大、忽视践踏强度的时间依赖性和畜种间差异、没有考虑放牧行为及其时间分配模式等问题 ,分析认为践踏强度应为家畜放牧行为、不同行为的持续时间、家畜体重、放牧地面积和坡度的函数。家畜的践踏作用因草地的健康状况差异而与之存在正或负的反馈机制以及践踏强度阈值 ,可能在草地退化和健康维护中起主导作用。今后需建立严格的研究方法和科学的践踏强度指标 ,重视草地植被对践踏的耐受阈限、践踏对牧草更新的调控机理及其对凋落物产生和分解的影响 ,以及土壤水分与践踏作用的关系 ,确定放牧系统的践踏强度阈值。

... 采食、践踏和排泄物沉积作为放牧系统中家畜作用于草地的3种方式,均是草地植物种间关系和植被构成发生变化的主要影响因素,在草地植被演替中起重要作用^[4] ...

2010

0.0

. 2010, :- DOI:doi:10.7666/d.y1740248

Yak Dung is the Mechanism of Action of the Qinghai-Tibet Plateau Alpine Grassland Health

牦牛粪维系青藏高原高寒草地健康的作用机制

Yu X J

鱼小军

青藏高原高寒草地是我国重要的牧区之一,也是广大藏族同胞赖以生存的基础。其生态系统的稳定不仅对我国东部和西南部气候产生巨大影响,而且对北半球甚至全球的气候产生显著影响。然而,日趋严重的草地退化问题已给青藏高原及其周边地区的生态环境和社会经济的可持续发展构成了严重威胁。在青藏高原,牦牛粪常作为牧民生活最依赖的燃料从草地上被捡走后,燃烧牦牛粪使灌丛草

... 3亿hm²高寒草地及1400万头牦牛,从而牦牛排泄物沉积量大,分布广^[5] ...

2010

0.0

... 由于垫状牛粪沉积对草地植物具有窒息作用,使粪斑下植物枯萎死亡^[6] ...

... 随粪块沉积时间的延长及粪中养分的释放,粪斑及其周围植物通过土-草系统吸收粪中养分后形成高营养斑块,反过来又吸引家畜采食^[26,27],从而家畜粪沉积和采食互作对草地植被的复杂效应,使粪块及其周围区域植物群落构成和种间关系发生改变^[6] ...

2010

0.0

. 2010, 24(5):569-574 DOI:doi:10.1088/0256-307X/17/9/008

Dynamics of associations between tree species in a secondary tropical montane rain forest at Jianfengling on Hainan Island

海南岛尖峰岭次生热带山地雨林树种间的联结动态

Huang S N , Li Y D , Luo T S

黄世能, 李意德, 骆土寿

应用方差检验法和基于２×２联列表的分析手段，对海南岛尖峰岭热带山地雨林采伐后２５～３５年次生群落２４００ｍ２固定样地中胸径≥７．０ｃｍ林木的种间联结动态进行了研究，结果如下：１）群落内的树种总体呈正联结，但未达到显著水平。联结指数Ｖ界于１．００～１．２１之间，接近无联结假设条件下的Ｖ值（Ｖ＝１），证实了作者关于 “种的分布相对独立”的假说；２）种间联结的比例和强度随演替进程而下降，群落朝种间无联结方向发展的趋势较为明显；３）群落早期建群种与后期入侵种种间没有负联结，两类树种之间有２６％的种对呈正联结，但均不显著，说明他们不仅可通过分割资源而和谐共处，而且更趋于独立存在。后期

... 因此,研究牦牛粪沉积下,高寒草甸植被构成及种间关联,对于高寒草甸植被演替动态和植物种间共存竞争及群落稳定性机制的揭示具有重要意义^[7,8,9] ...

2004

0.0

. 2004, 24(8):1435-1440 DOI:doi:10.3321/j.issn:1000-4025.2004.08.014

Interspecific association and correlation of dominant species of Lishan Mountain meadow in

历山山地草甸优势种的种间关联和相关分析

Wang L , Zhang J T

王琳, 张金屯

选取山西历山山地草甸中的22个优势种,运用χ2检验、3个关联指数以及Pearson相关和Spearman秩相关系数进行种间关联和种间相关分析.优势种对间大多呈正关联和正相关,但呈显著和极显著的种对不是很多,各种间相互依存,群落稳定发展.相关分析表明优势种对间大多呈正相关,Spearman秩相关要优于Pearson相关.Pearson相关星座图将22个优势种分为4个生态种组,各生态种组内的种有相同的资源利用方式和生态要求,各种组间有互相转化和演替的趋势.

... 这是由于草地在长期演替过程中,植物种的生态位进一步分化,生境异质性增加^[8],各物种都占据有利的位置,和谐共处,相互依赖和相互竞争均大大降低,多数种对联结程度不强,独立性强^[9],从而使两种斑块的群落结构不稳定,具有明显的次生性或正处于演替前期阶段 ...

2008

0.0

. 2008, 27(4):509-513

Analysis of interspecific association of Carex lasiocarpa community in recent 30-year succession in Sanjiang Plain

三江平原毛苔草群落近 30 年演替过程中的种间联结性分析

Lou Y J , Zhao K Y

娄彦景, 赵魁义

采用2×2列联表,通过方差分析、X2检验和Ochiai指数对三江平原毛苔草群落1973年、1983年、2003年3个阶段主要物种的种间联结进行了定量分析。结果表明:随着演替进展,群落的总体种间关联性均表现为显著正关联,关联性排列为1983年1973年2003年;成对物种间的联结性质即正、负联结的比例未发生变化;成对物种间达显著相关的种对所占比例呈逐渐增加趋势;物种间的联结程度逐渐增强;相同种对的联结性质或关联程度在不同时期存在差异。

2015

0.0

. 2015, 35(10):2096-2102 DOI:doi:10.7606/j.issn.1000-4025.2015.10.2096

Community composition and interspecific association analysis of Kobresia capillifolia grassland

线叶嵩草草地群落构成及种间关联分析

Wang W , Guo Q , Kang H J

王伟, 郭倩, 康海军

We firstly analyzed that the overall association characteristics of non-degraded and degraded Kobresia capillifolia communities by the variance ratio method.Then basing on 2×2 contingency table and important value of plant species,we investigated the composition of communities,interspecific relationship of plants and the stages of community succession,by using Chi square test,Jaccard index and Spearman rank correlation analysis.The results suggest that:(1)the differences in species composition between two communities are small;(2)the overall association of two communities shows significant negative correlation,and the interspecific relationship of each community are close on the whole;(3)most of the species of two communities have a weak interspecific association,namely,great independence,and the species pairs showing a negative or a positive association respectively occupies about half of total species pairs.It is studied that both non-degraded and degraded K.capillifolia communities are in the unstable stage.

以线叶嵩草群落和退化的线叶嵩草群落为对象，采用方差比率法分析群落总体关联性；基于2×2列联表及植物种重要值，运用χ 2 检验和Jaccard指数及Spearman秩相关分析，探讨了群落构成、植物种间关系及群落演替的阶段。结果表明：(1)2个群落物种组成差异小；（2)群落总体关联性均为显著负相关，群落种间关系整体上比较紧密；（3)群落内多数物种间联结性较弱，独立性较强，呈正相关和负相关的种对数约各占一半。研究认为，2个线叶嵩草群落均处于不稳定阶段。

... 诸多学者对群落类型^[10]和不同管理方式(封育和放牧)^[11,12]及鼠丘密度^[13]下,青藏高原高寒草甸植物群落构成及种间关联进行了分析 ...

... 线叶嵩草(Kobresia capillifolia)草地群落总体关联性表现为显著负相关且处于不稳定阶段^[10] ...

... 这与房飞等^[21]对新疆天山南坡高寒草原夏季牧场羊茅(Festuca ovina)+紫花针茅群落,以及王伟等^[10]和郭倩等^[22]报道的青海省称多县藏嵩草(Kobresia tibetica)群落和高山嵩草+异针茅(Stipa aliena)/杂类草群落及河南县线叶嵩草草地群落,植物种对的正负关联比均#cod#x02265 ...

2009

0.0

. 2009, 26(6):18-24

Study on interspecific association of spring-autumn pasture under enclosure and recovering conditions in Yili River valley

伊犁河谷春秋草地围封恢复过程中种间关联性研究

Li J B , Liu H L , Zhu J Z

李军保, 刘洪来, 朱进忠

The interspecific association in the major population of grazing communities and enclosed communities for 1, 5 and 6 years (springautumn pasture) in Yili River valley were studied and the results showed that the species in the communities enclosed for 5 and 6 years unassociated, and those enclosed for 1 year and unenclosed slightly associated. Along with the vegetation restoration, the species with positive association remarkably increased and those with negative association remarkably reduced. The positive and negative association rate performed a decreasing and then increasing pattern. The distribution of dominant species showed a relatively stable status and neutrally associated with other populations, the other populations were easily affected by environmental factors. The association coefficient reflected the relationships among species and with the environmental factors to a certain degree and it was strongly affected by environmental factors at the beginning of enclosure and the biological affection gradually reduced along with the improvement of habitat.

采用样方调查法，通过对方差分析、x2检验及种间关联度OI值的计算，对伊犁河谷春秋草地围封6年、5年、1年和未围封的草地群落中主要种群间的联结性进行了比较。结果表明，围封6年和5年的总体表现无关联,种间关联松散，围封1年和未围封表现轻微负关联。随着围封植被的恢复正关联显著的种对增加，负关联显著的种对明显减少，正负关联率表现为先减小后增大的趋势。优势种分布相对稳定，与其他种群表现中性联结，其他种群易受物种和环境的影响。种间联结测定值在一定程度上反映了物种间的相互关系以及它们与环境因子的关系，在未围封和围封初期种间关系受环境因素影响较大，随着群落生境的改善，生物因素对种间关联影响逐渐增强。

... 诸多学者对群落类型^[10]和不同管理方式(封育和放牧)^[11,12]及鼠丘密度^[13]下,青藏高原高寒草甸植物群落构成及种间关联进行了分析 ...

... 研究表明,高寒草甸植物种间关联与封育年限有关,短期封育表现为微弱负关联,5~6年的封育时间无关联^[11] ...

... 也可能是二者样方数量、群落自身物种数的多寡和群落类型等差异造成^[11,21,22] ...

2013

0.0

. 2013, 22(6):976-982 DOI:doi:10.3969/j.issn.1674-5906.2013.06.011

. Analysis of the role of Stipa breviflora desert grassland plant populations and the interspecific relationships under different grazing systems

不同放牧制度下短花针茅荒漠草原植物种群作用和种间关系分析

Lv S J , Wei Z J

吕世杰, 卫智军

摘　要：以短花针茅荒漠草原植物种群为研究对象,采用2×2列联表法和灰色关联分析方法,探讨植物种群地位和作用及种间关系在不同放牧制度下的变化。结果表明,放牧会使建群种短花针茅（Stipa breviflora Griseb.）重要值增大,使碱韭（Alliumpolyrhizum Turcz.ex Regel）的重要值减小;轮牧有利于无芒隐子草（Cleistogenes songorica（Roshev.）Ohwi）在群落中的地位和作用增加,自由放牧则相反。植物群落种间关系正联结强度表现为禁牧区（CK）〉划区轮牧区（RG）〉自由放牧区（CG）,负联结种对数的多少表现为CK

... 诸多学者对群落类型^[10]和不同管理方式(封育和放牧)^[11,12]及鼠丘密度^[13]下,青藏高原高寒草甸植物群落构成及种间关联进行了分析 ...

... 自由放牧较划区轮牧降低高寒草甸主要植物种间的关联程度和群落稳定性^[12] ...

2015

0.0

. 2015, 24(5):224-230 DOI:doi:10.11686/cyxb20150526

Effect of burrow entrance densities of plateau pike (Ochotona curzoniae) on interspecific association in Kobresia pygmaea meadow

高原鼠兔有效洞口数密度对高山嵩草草甸植物种间联结性的影响

Pang X P , Wang Q , Jia T T

庞晓攀, 王倩, 贾婷婷

Changes in the active burrow entrance densities of plateau pika ( Ochotona curzoniae ) can result in changes in the botanical composition in Kobresia pygmaea meadow which can influence interspecific relationships in the meadow. A field survey was carried out to determine the effect of active plateau pika active burrow entrance densities on the interspecific association between the major plant species in K. pygmaea meadow. This study showed that the dominance index for K. pygmaea (dominant species), decreased, while the dominance index of associated species including Gentiana dahurica and Anemone rivularis var. flore-minors increased as burrow density increased. The overall interspecific association among the main plant species of the K. pygmaea alpine meadow transformed from significantly positive association at 128 and 224 burrow entrances per ha to a negative association at 368 and 544 burrow entrances per ha. The relationship between K. pygmaea and Carex moorcroftii transformed from positive association at 128 and 224 active burrow entrances per ha to negative association at 368 and 544 active burrow entrances per ha, while associations between Euphorbia esula and Elsholtzia densa , and between Oxytropis ochrocephala and E. esula were negative at 128 and 224 active burrow entrances per ha but positive at 368 and 544 burrow entrances per ha. These results suggested that the active burrow entrance densities regulated the overall interspecific association among the main plant species in K. pygmaea alpine meadow, and the interspecific association among the main plant species was strongest and the stability of the alpine meadow community was greatest at 224 active burrow entrances per ha.

高原鼠兔有效洞口数密度变化会引起高山嵩草草甸植物组分变化,而且引起种间关系的改变。研究了高原鼠兔有效洞口数密度对高山嵩草草甸植物种间联结性的影响。结果表明,随高原鼠兔有效洞口数密度增加,原优势种高山嵩草优势度降低,而原伴生种达乌里秦艽和小花草玉梅等植物的优势度却逐渐增加;植物群落内主要物种间的总体联结性从有效洞口数密度小于或等于224个/hm 2 时的正关联,转变为有效洞口数密度超过224个/hm 2 时的负关联;高山嵩草和青藏苔草的联结性从有效洞口数密度≤224个/hm 2 时的正关联,转变为有效洞口数密度超过224个/hm 2 时的负关联,而密花香薷和乳浆大戟、乳浆大戟和黄花棘豆的联结性则相反。这说明高原鼠兔有效洞口数密度变化改变了高山嵩草草甸植物种间的总体联结性,有效洞口数密度为224个/hm 2 时种间联结性最强,群落最稳定。

... 诸多学者对群落类型^[10]和不同管理方式(封育和放牧)^[11,12]及鼠丘密度^[13]下,青藏高原高寒草甸植物群落构成及种间关联进行了分析 ...

... 224 个/hm²)下的负关联^[13] ...

2013

0.0

. 2013, 30(10):1594-1601

Effects of yak dung deposition on community characteristics and niche parameters alpine meadow

牦牛粪对高寒草甸植被群落特征和生态位参数的影响

Mu X M , Yu Y W , Zhang H M

牟晓明, 于应文, 张红梅

To provide foundation for researches on vegetation succession and mechanism of heterogeneity formation in alpine meadow, changes of vegetation composition, forage nutrients, plant importance value, plant niche breadth and plant niche overlap in Tibet Plateau alpine meadow with yak dung depositing were quantitative analyzed in this study. The results showed that, yak dung deposition significantly increased herbage biomass, green matter weight, plant height, coverage and K content of herbage, but had no significant influence on dead matter biomass and number of species. For dominant species, yak dung deposition decreased importance values of Stipa purpurea, Koeleria cristatav and Kobresia humilis , increased niche breadth values of these three species, and increased importance values of K.capillifolia and Agropyron cristatum . So yak dung deposition made these dominant species become to generalization species. For accompanying species, yak dung deposition decreased importance value of S.aliena , increased importance vale of S.krylovii , and decreased importance value and niche breadth value of Polygonum viviparum . For few species, yak dung deposition increased niche breadth values of Elymus nutans, Aremisia Smithii, A. capillaries, A. frigida, Allium sikkimense and Stellera chamaejasme . Beside, yak dung increased the whole niche overlap value of plants, increased species pairs, whose overlap values were >0.85, by increased 13.8%, while decreased species pairs, whose overlap values were<0.45, by 42.9%. In a word, yak dung deposition increased the proportion of grasses, and promoted the meadow transformation from sedge+grasses community to grasses community or grasses+sedges community.

本研究定量分析了青藏高原高寒草甸牦牛粪斑与对照处植被构成、牧草养分、植物种重要值、生态位宽度和生态位重叠值的变化,为高寒草甸植被演替和异质性形成机制的揭示提供科学依据。结果表明,牦牛粪沉积增加植物总生物量和活体质量,使草层高度、盖度和牧草K含量增加,而对死物质质量和物种数无影响。粪沉积降低优势植物紫花针茅( Stipa purpurea)、艹洽草(Koeleria cristata)和矮嵩草(Kobresia humilis)的重要值,增加其生态位宽度,并提高线叶嵩草(Kobresia capillifolia)和扁穗冰草(Agropyron cristatum)的重要值,使优势种植物更倾向于泛化种。粪沉积使伴生种异针茅(S.aliena)的重要值降低,西北针茅(S.krylovii)的重要值增加,珠芽蓼(Polygonum viviparum)的重要值和生态位宽度均降低,使少量种垂穗披碱草(Elymus nutans)、球花蒿(Artemsia pratensis)、茵陈蒿(A.capillaries)、冷蒿(A.frigida)、高山韭(Allium sikkimense)和狼毒(Stellera chamaejasme )的生态位宽度增加。粪沉积使草地植物的整体生态位重叠值增大,使生态位重叠值>0.85的种对增加13.8%,生态位重叠值<0.45的种对降低42.9%。总之,粪沉积使草地禾草比例增加,促进高寒草甸草地由莎草+禾草群落向禾草群落或禾草+莎草群落转化。

... 但家畜排泄物沉积下,高寒草甸植物构成及种间关系的研究,限于牦牛粪沉积下草地植物种生态位^[14]及植物种子库构成特征^[15,16]分析 ...

... 同时,由于禾草相对莎草对牛粪和施肥反应更敏感,更易吸收肥料中的养分^[14],粪沉积后期,禾草/杂类草斑块中的禾草因在短时间易吸收利用牦牛粪中的养分,生长增强,迅速占领群落上层空间,进而遮蔽其他矮小物种,以此弥补了粪沉积前期因粪斑覆盖草地而造成的损失,从而使其在与其他功能群植物的竞争中取胜,而嵩草/杂类草斑块中的莎草吸收牛粪中的养分少且慢 ...

2013

0.0

... 但家畜排泄物沉积下,高寒草甸植物构成及种间关系的研究,限于牦牛粪沉积下草地植物种生态位^[14]及植物种子库构成特征^[15,16]分析 ...

2014

0.0

. 2014, 33(10):2603-2609

Density and diversity of plant seeds in yak and sheep dung in cold season pasture meadow

高寒草甸冷季牧场牦牛和藏羊粪中植物种子密度和多样性

Jing Y Y , Xu C L , Chen L J

景媛媛, 徐长林, 陈陆军

In order to identify the features of plant seeds in dung of yak and Tibetan sheep grazing at different grazing stages of winter pasture (alpine meadow) on Eastern Qilian Mountain, the number of species, the seed diversity index, the seed number and density were studied. The results showed that in the same grazing stage, the number of seed species in yak dung was higher than in Tibetan sheep dung. The seed density in yak dung was lower than in the Tibetan sheep dung, but the difference was significant only at the middle grazing stage. At the initial and final grazing stages, the seed diversity index in yak dung was significantly higher than that in Tibetan sheep dung. The seed species number and seed density (grains·kg -1 ) in yak dung at the different grazing stages were both in order of final grazing stage > initial grazing stage > middle grazing stage, and those in Tibetan sheep dung were in order of initial grazing stage > final grazing stage > middle grazing stage and of middle grazing stage > final grazing stage > initial grazing stage, respectively. The seed number of sedge was the highest in yak dung, while the seed number of forbs was the highest in Tibetan sheep dung. The different conditions of winter pasture at the different grazing stages made the seed distribution different in each synusia of plant community, together with the differences in grazing habit, forage intake per bite and digestibility of yak and Tibetan sheep, causing the differences in seed density, seed species and seed diversity of their dung.

为明确高寒草甸冷季牧场牦牛粪和藏羊粪中种子含量，研究了天祝东祁连山高寒草甸冷季牧场不同放牧时期牦牛粪和藏羊粪中可萌发植物种子的密度数量和多样性。结果表明，高寒草甸冷季牧场的同一放牧时期，牦牛粪中种子的物种数高于藏羊粪；种子密度牦牛粪低于藏羊粪，放牧初期和末期差异不显著，放牧中期差异显著。放牧初期和末期牦牛粪中种子的Shannon指数显著高于藏羊粪，放牧中期差异不显著。牦牛粪中的种子物种数为：末期>初期>中期；藏羊粪为：初期>末期>中期。单位重量牦牛粪中的种子数量变化规律是：末期>初期>中期，藏羊粪为：中期>末期>初期。牦牛粪中所含莎草科的植物种子最多，而藏羊粪中则以杂类草为主。高寒草甸冷季牧场不同放牧时期草地状况的不同，使得草地贮草量和种子在草层内的分布不同，加之牦牛和藏羊各自采食习性、单口采食量的差异，以及牦牛和藏羊消化道对种子的消化率不同，最终导致牦牛和藏羊粪便中的种子密度、种类和多样性存在差异。

... 但家畜排泄物沉积下,高寒草甸植物构成及种间关系的研究,限于牦牛粪沉积下草地植物种生态位^[14]及植物种子库构成特征^[15,16]分析 ...

2010

0.0

... 由于草地植被异质性和家畜采食的非均匀分布,使粪斑和草地植被斑块产生叠加效应,而植被构成对粪沉积的响应因群落植物种构成不同^[17] ...

2006

0.0

. 2006, 15(4):30-35 DOI:doi:10.3321/j.issn:1004-5759.2006.04.005

The impact of cattle dung deposition on grasslands in the Songnen grassland

牛粪堆积对草地影响的研究

Jiang S C , Zhou D W

姜世成, 周道玮

初步研究了松嫩草地牛粪堆积对草地的影响.结果表明,草地牛粪堆积能提高土壤水分、有机质、全氮、全磷、全钾含量,降低土壤pH值,因而改善土壤理化性质,利于其周围植物生长,但只能延及到牛粪堆外围3～5 cm处,牛粪堆积造成其影响范围内的草地生产力下降11.2%.在草地牛粪中共调查到大型节肢动物3目8科15种,粪食性型种类10种,直蜉金龟在4,5,6和9月为优势种,而肖秋家蝇和东方角蝇幼虫在7,8月为优势种.牛粪中种子种类较少,数量较多,主要为稗草种子,其发芽率较低.牛粪中植物种子对碱斑地植被建植有重要意义.

... 03 m²的前期基础上,考虑牛粪对草地植物的影响效应通常约为粪斑大小的2倍^[18],将本研究牦牛粪斑大小设置为0 ...

2015

0.0

... 0中Correlate中的Bivariate模块,计算两种群落粪斑和对照区域主要植物种间Spearman秩相关系数及显著性检验^[19] ...

2009

0.0

. 2009, 30(4):553-559 DOI:doi:10.3969/j.issn.1000-6362.2009.04.016

System stability and its self: maintaining mechanism by grazing in Alpine Kobresia Meadow

放牧高寒嵩草草甸的稳定性及自我维持机制

Cao G M , Long R J

曹广民, 龙瑞军

The Alpine Kobresia Meadow Ecosystems at the different stage of degradation in the Haibei Alpine Meadow Ecosystems Research Station and the Three Rivers Source Region were studied to reveal the plant communities,surface conditions,mattic peipedon thickness,roots and soil ratio,and moisture saturation ratio,so as to distinct the system stability and its self-maintaining mechanism in the course of the degradation in alpine meadow.The results showed that the alpine Kobresia meadow system had performed special mechanism to maintain its stability in the long-term evolution process although it had a simple structure.So it could bear the man-made interferences and climate fluctuations to some extents,and had higher ability of controlling and resuming system,as it maintained its system stability through lowing,villiform and mattic growth.But the alpine Kobresia meadow system had a very bad recovering ability after the system destroyed.Nowadays,the reason for the degeneration of alpine meadow in large areas was that the carrying capacity endowed by human had exceeded the capability threshold.

以空间代时间,在"三江源"和中国科学院海北高寒草甸生态系统研究站地区,将处于不同退化阶段的高寒嵩草草甸作为研究对象,进行了其植物群落、地表状况、草毡表层厚度、根土比和水分渗透速率的演替过程与规律研究,以明晰放牧高寒嵩草草甸退化过程中其系统稳定性及自我维持机制。结果表明,高寒嵩草草甸虽然结构简单,但在长期适应寒冷气候进化过程中形成了低矮化、细绒化和草毡表层加厚、极度发育等一系列特殊的稳定性维持机制,可以承受一定范围内的人为干扰和气候波动,具有较高的系统稳定性与自我调控能力,但系统遭到破坏后的恢复能力极差。今日高寒草甸的大面积退化,是人类所赋加于草地的承载力远超过其承载力阈值而导致系统稳定性崩溃的结果。

... 其主要原因是嵩草/杂类草斑块中的优势植物为嵩草属的线叶嵩草和矮嵩草,其在长期适应寒冷气候的进化过程中,形成特殊稳定的适应外界不良环境的维持性特征,如叶片少,植株矮化、细绒化和草毡表层加厚等极度发育等,从而具有较强的抗外界干扰和自我调控能力^[20] ...

2012

0.0

. 2012, 32(6):1898-1906 DOI:doi:10.5846/stxb201102170182

Numerical analysis of interspecific relationships in alpine steppe community in Bayanbulak

高寒草原植物群落种间关系的数量分析

Fang F , Hu Y K , Zhang W

房飞, 胡玉昆, 张伟

Highland ecosystems are believed to be of vital importance in ecological study, partly due to the rapid climate change worldwide and the highest sensitive feature of this special system, not only its productivity and biodiversity but also its inter-specific relationship, to environmental alterations. The Bayabulak Steppe distributes in the middle part of Tianshan Mountains, and is situated at 42°18'-43°34' N, 82°27'-86°17' E with altitude between 2460-2760 m. Climate here belongs to typical alpine cold climate with yearly mean temperature of -4.8 ℃, yearly mean precipitation here is 276.2 mm, yearly evaporation is (1 135.2±112.3) mm, the snow period in Bayabulak is about 150-180d, and there is no frost-free time in this region. Stipa purpurea and Festuca ovina grassland are the dominant vegetation types in our study site. In order to clarify the species composition and inter-specific association of the alpine grassland community of Bayanbulak in Xinjiang, China, variance ratio ( VR ) of the overall association, association coefficient ( AC ), Ochiai index ( OI ) and Spearman rank correlation coefficient, based on the 2 × 2 contingency table, were used in our quantitative analysis. Field data were collected in July 2009. Three parallel transect were firstly set up and the distance between each of them was 50 m, then 20 plots (0.50 m × 0.50 m for each unit) were randomly set within each transect, in so doing total of 60 plots were involved in the present work. Our results showed that: 1) There are 36 species with 14 common species of plants in Bayinbulak alpine steppe, and they belong to 26 Genus of 16 Family. W = 61.52, a statistic index to test whether the VR is significant or not, indicated that the overall correlation of all plant species was positive but not significant. 2) Inter-specific associations of keystone species with others were neutral or not significant. Keystone species showed greater independence and occupied prior ecological niche in the community, and their distributions were not susceptible to other species. Values of OI were basically identical to those of association coefficient; Based on the measuring of OI , 19.79% of total 91species-pairs composed by 14 common species was significant or very significant, and 72.21% of them showed positive correlation while 27.79% showed negative correlation for those significantly related species-pairs; 3) Inter-specific associations of accompanying species or weeds with others were significant (negative or positive), suggesting higher dependency of accompanying species or weeds on other species, namely they were susceptible to other species and only occupied inferior ecological niche in the community. Furthermore, populations of these plant species are instability in the process of ecosystem development; 4) This study provides new dimensions for the theory of species association. According to their adaptability to the environment and the relationships between plants and environment, the 14 common species were divided into three ecological species groups. Positive correlations occurred among the species in the same ecological species groups, while the negative correlations were found in the different ecological species groups. It mainly due to their different biological features induced by different adaptability to habitats or separated niches; 5) Several suggestions were proposed to promote the ecological restoration of degraded steppe systems in the study region.

为了探明新疆巴音布鲁克高寒草原植物群落的物种组成和种间关系,在对其草原群落进行样方调查的基础上,利用方差比率法,χ 2 检验以及种间关联度指数对物种关联性和相关性进行了分析。结果表明:1)巴音布鲁克高寒草原植物群落高等植物分属16科,26属,36种;主要有14个常见植物种;2)用来检验方差比率( VR )偏离1是否显著的统计量 W = 61.52,说明该草原群落内物种整体关联度表现为不显著相关;表征种间关联程度的Ochiai指数( OI 值),与联结系数 AC 值基本吻合;3)伴生种或杂类草与其它物种之间联结显著,正向或负向,在群落中表现出很强的依赖性,分布易受其它物种的影响,在群落中不能稳定存在且只占据劣势生态位;4)根据14个优势种群对环境的适应方式和主导生态因素,可将它们划分为3个生态种组,同一生态种组内,各种间具有较强的正联结,而不同组间联结较为松散,主要是由于他们具有不同的生物学特性,对生境具有不同的生态适应性和相互分离的生态位所致;5)对退化草地的恢复提出了建议。

... 也可能是二者样方数量、群落自身物种数的多寡和群落类型等差异造成^[11,21,22] ...

2016

0.0

. 2016, 24(2):24-31 DOI:doi:10.11733/j.issn.1007-0435.2016.02.008

Analysis on the vegetation composition and interspecific association of Kobresia meadow in southern pastoral area of southern Qinghai Province

青南牧区南部嵩草草甸群落植被构成及种间关联

Guo Q , Wang X L , Zhang R X

郭倩, 王旭丽, 张润霞

摘　要：植物群落构成和植物种间联结,反映植物物种空间上相互关系,对于草地演替动态及稳定性的揭示具有重要理论意义。以青南牧区南部3个嵩草群落为对象,分析了群落生物量构成变化;并基于2×2列联表及植物种多度,采用Ochiai种间关联度指数及Spearman秩相关分析,探讨了不同嵩草群落的植物种间关系及群落演替特征。结果表明：藏嵩草群落的生物量显著高于高山嵩草＋异针茅/杂类草群落,群落生物量主要由莎草科植物构成,其占总生物量的45%-80%,而禾草、杂类草和不可食植物则分别约占总生物量的8.1%,15.2%和16.4%;3个嵩草群落内的植物种对一般存在完全关联、绝对关联和完全独立3种关系,藏嵩草群落和高山嵩草＋杂类草群落中完全独立的种对较多,为38.1%和40.0%;而高山嵩草＋异针茅群落中完全关联种对较多,为36.3%;3个群落中较多植物种对间存在显著联结性,且显著正联结种对数多于显著负联结种对数,同一种对在不同群落中的种间关系不同;各嵩草群落均处于稳定发展阶段。

... 也可能是二者样方数量、群落自身物种数的多寡和群落类型等差异造成^[11,21,22] ...

2010

0.0

... 类似施肥研究表明,施肥降低群落稳定性^[23],施N肥及N、P、K单施或联施或施羊粪,均增加青藏高原矮嵩草群落和退化高寒草甸草地中禾草比例,降低杂类草比例^[24,25] ...

2002

0.0

. 2002, 26(3):288-296 DOI:doi:doi:http://ir.nwipb.ac.cn/handle/363003/815

Response of plant groups to simulated rainfall and nitrogen supply in Alpine Kobresia humilis meadow

高寒矮嵩草草甸植物类群对模拟降水和施氮的响应

Shen Z X , Zhou X M , Chen Z Z

沈振西, 周兴民, 陈佐忠

The response of different plant groups to simulated rainfall changes in summer, increased snow in winter and spring, and nitrogen fertilization application in late June were studied in an alpine Kobresia humilis meadow in Qingzang plateau. The results indicated that the response of summed dominated ratio (SDR) and above-ground biomass in different plant groups were not significantly different with 20%-40% rainfall reduction or 20%-40% increased rainfall in 1999. A 100% increase in snow in winter and spring was more favorable to the growth of plant groups than were other rainfall treatments. Both a single late June application of 150 N kg·hm-2 and 300 N kg·hm-2 in soil increased the coverage ratio, height ratio and SDR of grasses. The coverage ratio, height ratio and SDR were significantly decreased at the strong N application (300 N kg·hm-2). There were obvious compensatory effects between grasses and forbs at the application of 150 N kg·hm-2. The sensitivity of the three plant groups was quite different. Grasses were most, forbs were less and sedges were least sensitive to simulated rainfall and available N in the improved disturbing condition. Forbs were most, grasses were less, and sedges were least sensitive to simulated rainfall and available N in the undesirable disturbing condition. It seemed that sedges were most tolerant to fluctuation in environmental resources. Simulated rainfall increasing by 20%-40% could moderate the effect of lower precipitation in a water deficient year (1999), while rainfall decreased 20%-40% could intensify the effect of drought.

研究了青藏高原高寒矮嵩草（Kobresia humilis）草甸植物类群对模拟夏季增减雨量、冬春增雪以及增施氮肥下的响应。结果表明：1999年模拟减少降雨20%～40%和增加雨量20%～40%下禾草类、杂类草和莎草类的综合优势比（SDR）和地上生物量变化均不显著。冬春增雪100%有利于禾草类夏季的生长，冬春增雪对植物类群的影响大于夏季雨量的增加。夏季增施氮150 kg·hm－2和增施氮300 kg·hm－2禾草类的盖度比、高度比、SDR和地上生物量明显增大，而杂类草的盖度比和高度比、SDR及地上生物量在施氮300 kg·hm－2下显著减低，在施氮150 kg·hm－2水平上禾草类的生物量的增加与杂类草生物量的降低存在相互补偿的作用机制。在水分资源不利的（如减少雨量）的干扰下，其敏感性表现为杂类草大于禾草类，莎草类最小。莎草类植物对各种处理下响应不敏感，也说明它对资源环境的波动有很强的适应性。缺水年（1999年）模拟增加雨量20%～40%的条件下，可缓解降水量减少的影响，相反模拟减少雨量20%～40%会增强干旱的影响程度。

2009

0.0

. 2009, 14(6):31-36

Effects of fertilization on grass community diversity and productivity of degraded alpine grassland at Maqu, in South of Gansu Province

施肥对甘南高寒草甸退化草地植物群落多样性和生产力的影响

Chen W Y , Qi D C , Li G Y

陈文业, 戚登臣, 李广宇

对甘肃玛曲县中度退化草地进行了氮、磷、钾、有机肥施肥处理,研究了其对草地植物群落多样性和生产力的影响.结果表明:施肥各种处理均可不同程度增加退化草地的植被高度和地上生物量,其中施有机肥O和N-P-K使植被高度分别增加2.208倍、2.274倍,地上生物量分别增加了3.315倍、3.161倍;在施P处理中,杂类草减少32.90%,莎草科植物增加 73.43%,有机肥处理中,禾本科植物增加19.69%,不同施肥处理后,莎草科、禾本科和豆科各物种重要值都在增加,而杂类草各物种重要值都在减少; 各种施肥处理(施O除外)均可使退化草地植物群落的物种丰富度减少;在N-P-K处理中物种Simpson多样性指数、Shannon-Wiener指数和Pielou均匀度指数均增加,有机肥处理的丰富度和Simpson优势度指数均增加,N处理中Pielou均匀度和Simpson优势度指数均增加, 在单施P或K处理中,上述4指数均减少了.从生物多样性特征和生产力增加综合考虑,施有机肥和N-P-K是加速高寒草甸退化草地恢复重建的重要措施.

2001

0.0

... 同时,由于粪斑具有特殊气味及寄生虫,粪沉积初期-中期,家畜弃食粪斑及其毗邻区域植物^[26] ...

2014

0.0

. 2014, 31(3):488-498 DOI:doi:10.11829\j.issn.1004-5759.2013-0130

Effects animal dung deposition on grassland nutrients and botanical composition in mixed Lolium perenne and Trifolium repens pasture in Guizhou plateau

畜粪沉积对贵州高原黑麦草+白三叶草地养分和植被构成的影响

Sun H , Yu Y W , Ma X L

孙红, 于应文, 马向丽

The mineral elements of soils and herbages, nutritional values, botanical composition and herbage defoliation in dung deposition and control plots in mixed Lolium perenne and Trifolium repens pasture in Guizhou plateau were quantitatively monitored and systemically analyzed in August 2011. The results indicated that the difference of mineral elements contents between soils and herbages was great and varied with different elements and excretion types. The dung deposition of grazing animals decreased or eliminated the positive correlation between soils and herbage in Na, K, and Mn contents, whereas the dung deposition promoted positive correlation between soils and herbage in Cu and Zn contents. The effects of grazing animal dung deposition on botanical composition of grassland in plant population and community levels varied with patch types and different decomposing stages. Grazing animals dung deposition could promote tiller and growth of L.perenne of D.glomerata , inhibit growth of T.repens , increase height and biomass of herbage and reproductive shoots of L.perenne , and decrease the nutritional values of herbages. The effect of cow dung on botanical composition was more obvious than that of sheep dung. The defoliation of grazing animals for L. perenne and D.glomerata in dung deposition plots was lower than that in controls. The rejected defoliation level of cow in dung deposition plots was higher than that of sheep. The results suggested that the mixed grazing with sheep and cattle could reduce the herbage waste in dung deposition sites and improve the utilization rate of grassland in grassland practical management.

2011年8月, 对贵州高原黑麦草( Lolium perenne)+白三叶(Trifolium repens )草地的牛粪和羊粪沉积处和对照处土壤和牧草矿质成分、牧草营养价值、植被构成和牧草被家畜采食的程度进行定量分析, 为草地管理提供实践依据。结果表明, 主要矿质元素含量在土-草之间差异很大, 因矿质种类和家畜种类不同而异;畜粪沉积降低或消除Na、K和Mn在土-草之间的正相关性, 促进Cu和Zn在土-草之间正相关的形成。畜粪沉积对植被在植物种群和群落水平的影响随畜粪类型和分解阶段不同而异, 粪沉积促进鸭茅( Dactylis glomerata )和黑麦草生长, 抑制白三叶生长, 使牧草高度、生物量和黑麦草生殖枝率提高, 整体营养价值下降。牛粪沉积对植被的影响比羊粪更明显。家畜对粪沉积处黑麦草和鸭茅的采食均低于对照, 牛对粪沉积处植物的弃食程度比羊的高。草地管理中, 实行牛、羊混牧, 可提高草地利用率。

1996

0.0

... 另一方面,家畜具有在食源值与食源报酬之间的空间判断和权衡能力,其也频繁采食草地上数量多、面积大的中、低营养植被斑块^[28] ...

2004

0.0

. 2004, 13(5):33-38 DOI:doi:10.3321/j.issn:1004-5759.2004.05.006

Interspecific associations in Stipa breviflora desert steppe community

短花针茅草原群落种间联结研究

Zhu G L , Yang H Q , Wei Z J

朱桂林, 杨洪琴, 卫智军

种间联结能够反映群落中物种之间的交互作用,在群落结构功能研究中占有重要位置.在1 m×1 m样方尺度上研究了短花针茅荒漠草原群落中出现频率较高的13个物种之间的交互联结性.结果表明,短花针茅群落多种间负相关,多物种之间存在对资源的竞争;11个种对联结显著,其中5个种对联结极显著;显著联结的11个种对中,4对为正联结,7对为负联结;其余种之间相对独立,互为中性联结或联结性不显著.关键种与其它物种之间呈中性联结或联结性不显著,在群落中表现出很强的独立性,分布不受其它物种的影响或所受影响很小,在群落中能够稳定存在并占据优势生态位.伴生种或杂类草与其它物种之间联结显著,正向或负向,在群落中表现出很强的依赖性,分布易受其它物种的影响,在群落中不能稳定存在且只占据劣势生态位.

... 植物种间关联或物种联结性是指不同物种在空间上的相互联结性,体现物种间的依赖性与排斥性,正联结种对对环境或资源有相互依存的相似需求,负联结种对生境或资源具有相互排斥的不同需求^[29] ...

2009

0.0

... 植物群落中优势种和亚优势种对群落的结构和稳定性起重要作用^[30] ...