放牧干扰下高寒草甸物种、生活型和功能群多样性与生物量的关系

为明晰放牧强度对高寒草甸物种、生活型以及功能群多样性和生物量的时间累积作用以及对多样性与生物量之间相互关系的影响。选择青藏高原东北缘高寒草甸,在6个放牧强度样地连续4 a(2012—2015)进行物种、生活型、功能群多样性和生物量调查。采用重复测量方差分析和线性回归法分析放牧强度和放牧年份对物种、生活型、功能群多样性以及多样性与生物量之间的相关关系的影响。结果表明:(1)放牧强度和放牧年份均对生物量产生显著影响。随放牧强度增加,生物量显著降低。但随放牧年份延长,不同放牧强度区生物量无一致的变化规律。(2)放牧年份对物种、生活型和功能群丰富度、均匀度和优势度的影响均达到显著水平。但放牧强度和放牧年份只对物种丰富度产生交互作用,放牧强度对物种丰富度的影响具有时间的累积效应。(3)放牧干扰下不同层次多样性,仅物种丰富度与所有多样性指数显著相关,物种丰富度可以作为物种多样性测度的代表性指标。(4)多样性与生物量关系的研究,对丰富度而言,仅低放牧强度样地(Plot2)中物种丰富度随生物量增加而显著降低,其余放牧强度样地的物种、生活型和功能群丰富度均与生物量无关。对均匀度而言,高放牧强度样地(Plot4、Plot5、Plot6)生活型均匀度随生物量的增加而显著降低。对优势度而言,高放牧强度样地生活型优势度随生物量的增加而显著增加。生活型多样性可作为放牧干扰下生物量变化快速预测的有效指标。     

放牧干扰下若尔盖高原沼泽湿地植被种类组成及演替模式

以若尔盖高原退化沼泽植被为研究对象,应用多重比较、双因素方差分析、物种累积曲线、PCA排序、方差分解等方法分析了不同放牧压力、放牧季节下物种丰富度、多度、生活型组成、群落演替的变化。结果表明:不同放牧季节物种丰富度格局不尽相同,其中6、9、10月在各牧压梯度间无显著差异,7、8月均以极重度和极度阶段(中生草甸)的最高,原生沼泽的最低。在物种累积速率上,沿牧压梯度以极重度和极度阶段的最高,沿放牧季节以7、8月最高;双因素方差分析结果进一步表明物种丰富度与放牧季节无显著关系,但与放牧压力关系显著。放牧压力和放牧季节共解释了物种多度总方差的47.6%,其中放牧压力解释了50.1%,放牧季节以及二者方差交集均为负值;沿牧压梯度,沼泽植被逆向演替模式倾向于沼泽→草甸,沼泽化草甸阶段不明显,但是演替方向未发生变化,建群种替代规律为:乌拉草→木里苔草→栗褐苔草,生活型组成中直立型植物比例较少,莲座型和匍匐型植物增加。总之,放牧季节对物种丰富度无显著影响,但在一定程度上改变了其牧压梯度格局,降低了物种累积速率。放牧压力改变了群落物种丰富度、生活型组成和演替模式,但放牧可能仅为沼泽植物群落物种多样性格局和演替的驱动力之一。     

青藏高原高寒草地植物物种丰富度及其与环境因子和生物量的关系

在青藏高原进行了大范围的群落调查 ,研究高原的两种主要草地群落类型———高寒草甸和高寒草原的植物物种丰富度及其变化。结果表明 :(1)在 5 0个样地 2 5 0个 1m× 1m的样方中 ,共出现 2 6 7种植物 ,其中高寒草甸179种 ,高寒草原 135种。在高寒草甸 ,1m2 样方内物种数最多为 32种 ,最少的仅为 3种 ;在高寒草原 ,物种数最多为 18种 /m2 ,最少的仅为 2种 /m2 。 (2 )物种丰富度随经度和纬度的增加呈增加趋势 ;随海拔的上升呈减少趋势。对物种丰富度与环境因子之间进行逐步回归 ,发现物种丰富度与生长季降水和温暖指数呈显著正相关。 (3)物种丰富度与地上生物量呈显著正相关。     

青藏高原高寒灌丛植被对长期放牧强度试验的响应特征

在青藏高原中国科学院海北高寒草甸生态系统定位研究站对金露梅高寒灌丛草场植被开展了长期不同放牧强度试验,分别在短期(4年)、中期(11年)和长期(18年)放牧阶段研究不同放牧干扰强度对草地植物物种多样性、群落结构、地上生物量和草场质量的影响.研究表明,在不同放牧阶段,随着放牧强度增加植物群落的高度和盖度都降低.在中期放牧干扰阶段,物种多样性数和均匀度指数随着放牧强度增加呈现典型的单峰曲线模式;在长期放牧干扰阶段,随着放牧强度增加,占优势地位的灌木和禾草被典型杂类草替代,其中的重度放牧干扰简化了高寒灌丛植被群落结构,减少了地上现存生物量,特别是可食优良牧草生物量.植被对放牧的响应除了与放牧强度和放牧时间阶段密切相关外,还与该地区水热条件的变化有一定的相关性.针对长期放牧干扰的反应特性可将金露梅灌丛草场中植物划分为增加型、敏感型、忍耐型和无反应型4种类型.除了丰富度指数、多样性指数和均匀度指数外,其它一些特征参数并不支持著名的中度干扰假说.本研究发现,长期重度放牧促进了青藏高原高寒草地退化,适度放牧有利于高寒灌丛草场的生物多样性保护和牧草利用;"取半留半"的放牧原则在青藏高原草场放牧管理实践中值得推荐,它将有利于防止草场退化,提高牧草利用率和维持较高的生物多样性.     

大兴安岭森林草原过渡带典型植物群落动态变化特征

大兴安岭森林草原过渡带是大兴安岭森林与呼伦贝尔草原接壤的关键区域。由于过渡带本身的脆弱性,气候变化和过度放牧等人为因素的干扰导致区域内植物群落结构和功能发生剧烈变化。然而,气候变化和放牧双重作用对森林草原过渡带草地植物群落动态影响的研究还鲜有报道。以内蒙古陈巴尔虎旗森林草原过渡带为研究对象,分别于2008、2009、2010和2019年沿着草甸草原-森林草原过渡区域-林间草地-森林的梯度进行野外调查,对比分析11年间受气候变化和过度放牧的双重干扰下,该区域植物物种丰富度、Raunkiear生活型功能群、牧草饲用价值以及群落生物量随时间的变化动态。结果表明：从总体上看,与2008-2010年均值相比,2019年植物物种数量明显下降,其中禾本科牧草减少10余种;植物Raunkiear生活型功能群组成发生显著变化,其中一、二年生植物物种数增加4%,地面芽植物降低8%;牧草饲用价值中优等牧草所占比例下降5%,劣等牧草比例增加7%;总地上生物量显著下降35%。在过渡带各区段中,草甸草原群落结构和功能受气候和过度放牧影响更加明显。与2008-2010年均值相比,2019年草甸草原群落物种丰富度降低12%,优等牧草比例降低4%,劣等牧草比例增加7%,植物地上生物量更是显著降低62%。对区域气候和放牧因素分析可得,年降水量和年均温与过渡带地上生物量均显著相关,其中降水是导致过渡带整体上生产力下降的主要因素;而过度放牧是导致草甸草原群落生产力下降的主要因素。     

藏香猪翻拱型放牧对不同类型高寒草甸植物特征的影响及土壤环境驱动力

生态系统脆弱的高寒草甸是生态学研究的热点。藏香猪翻拱土壤并取食地下草根层,是藏族自治区一种特有的放牧类型,关于该放牧类型对藏区高寒草甸的影响尚缺乏研究。以滇西北高原国际重要湿地——纳帕海流域的典型高寒草甸为对象,研究了不同高寒草甸类型(陆生草甸与沼泽化草甸)中的植物群落结构特征对藏香猪翻拱型放牧的响应。结果表明,翻拱型放牧显著降低了植被盖度(63.5%)、地上生物量(84.6%)与地下生物量(97.4%),并促进植物生物量向地上部分的再分配。翻拱型放牧下随着植被盖度与生物量降低,植物竞争强度减小,植株高度与空间利用率提高。然而,不同草甸类型中的植物群落结构对翻拱型放牧表现出差异响应。陆生草甸上的群落组成基本未变,而沼泽化草甸上的优势植物华扁穗草(Blysmus sinocompyessus)向1年生水生植物水蓼(Polygonum hydropiper)演替。土壤含水率是导致植物响应差异的关键驱动力。综上,藏香猪翻拱型放牧对植物生物量及盖度等特征均产生不利影响,导致高寒草甸生物量积累锐减,而植物群落结构的变化是放牧干扰与土壤水分、空间竞争协同作用的结果,反映出植物在不同环境中对放牧干扰的差异化响应。研究区域环境特征与放牧干扰的耦合关系,可以为藏香猪放牧作用下的高寒草地植物多样性保护及恢复提供理论依据。     

高寒草甸放牧扰动与两种植物的反应研究

对不同放牧强度下矮嵩草草甸植物垂穗披碱草和鹅绒委陵菜的生长和繁殖特性进行了5年研究。结果表明,垂穗披碱草株高、分蘖数、地上物量、地下生物量、有性繁殖效力在不放牧或轻度放牧下最大,营养生长效力在重度放牧中最大,而贮藏效力在重度放牧中最小。鹅绒委陵菜叶数、匍匐茎长度、地上生物量、匍匐茎生物量、开花数、生殖生物量、有性繁殖效力及营养繁殖效力在放牧区显著高于不放牧的对照区。说明适当的放牧干扰有利于垂穗披碱草营养生长,但放牧过重会抑制其有性繁殖;由于放牧采食减少了种间竞争影响,从而提高了鹅绒委陵菜营养,生长效力和有性繁殖效力。两物种地下贮藏物的比例随着放牧强度增加显著减少。放牧强度对高寒草甸的群落演替方向有显著影响。     

青藏高原高寒草甸植物群落物种组成和生物量沿环境梯度的变化

生物多样性与生态系统功能的关系及其机制是生态学领域的重大科学问题. 人们越来越关注环境因子对多样性-生产力关系的影响. 植物群落组成、物种丰富度、物种特征、生物量的分布结构和植物枯枝落叶对高寒草甸物种多样性和生产力有着重要的影响. 因此, 我们利用2001~2004年中国科学院海北生态系统定位站高寒草甸群落的实测资料, 研究了不同环境梯度(土壤含水量和营养)下, 植物群落生物量, 物种丰富度及组成的变化. 结果表明, 植物群落物种组成的不同反应在生物量的分布上, 以藏嵩草为优势种的藏嵩草沼泽化草甸群落总生物量(地上、地下)最高(13196.96±719.69 g/m²), 次之是以杂类草和莎草科为主的小嵩草草甸(2869.58±147.52 g/m₂), 以禾本科和杂类草为主的矮嵩草草甸最低(2153.08±141.95 g/m²). 藏嵩草沼泽化草甸中, 草本植物枯枝落叶显著高于小嵩草、矮嵩草草甸, 土壤含水量对草本植物枯枝落叶有较大的影响. 不同类型草甸群落中, 地上生物量与土壤有机质、全氮和群落盖度之间均呈显著正相关(P < 0.05); 藏嵩草沼泽化草甸中, 总生物量与物种丰富度呈负相关(r_s = -0.907, P < 0.05)、地下生物量与土壤含水量呈正相关(r_s = -0.900, P < 0.05); 而在小嵩草和矮嵩草草甸中它们之间均没有达到显著水平, 说明不同类型高寒草甸群落生产力除受物种多样性、功能群内物种密度和均匀度的影响, 同时也受物种本身特征和外部环境资源的影响. 不同类型草甸群落生物量的分布与土壤含水量和土壤养分的变化相一致.     

青藏高原高寒草甸物种多样性的海拔梯度分布格局及对地上生物量的影响

植物物种多样性在海拔梯度上的变化规律以及物种多样性与生产力的关系是生态学研究的热点, 至今还没有得出一般性规律。本文以青海省海南藏族自治州贵德县的拉脊山(36°21′ N, 101°27′ E, 海拔3,389-3,876 m)和果洛藏族自治州的玛沁县军牧场山体(34°22′ N, 100°30′ E, 海拔4,121-4,268 m)为研究对象, 对植物高度、盖度、地上生物量和物种多样性随海拔高度的变化进行调查和统计分析, 以探讨青藏高原高寒草甸的物种多样性和地上生物量在海拔梯度上的变化规律及两者的关系。结果表明: (1)两条山体样带上地上生物量与物种多样性随海拔的变化规律一致: 随着海拔的升高, 地上生物量线性降低; Shannon-Wiener指数、Simpson指数和物种丰富度都呈单峰曲线, 在中间海拔最大, 而Pielou指数随海拔的升高线性增加。结合目前针对青藏高原高寒草甸的研究数据, 发现物种丰富度随海拔高度的变化均呈单峰曲线, 说明随着海拔的升高物种多样性先升高后降低可能是青藏高原物种多样性分布的普遍规律。(2)地上生物量与物种多样性的关系在两条山体样带上表现一致: 地上生物量随Shannon- Wiener指数、Simpson指数和Pielou指数的升高而线性降低, 但与物种丰富度不相关。综合两条山体样带所有样方数据, 发现地上生物量与Shannon-Wiener指数、Simpson指数不相关, 而随物种丰富度的升高线性增加。结合目前在青藏高原的相关研究数据, 发现地上生物量与物种丰富度呈S型曲线(logistic model)。     

呼伦贝尔草原不同草地利用方式下生物多样性与生物量的关系

生物多样性与生产力的关系是当前生态学中研究的重点之一,以呼伦贝尔草原为研究对象,通过连续两个生长季的野外监测,从草地植物功能型的角度探讨了在不同利用方式下草地物种丰富度与地上生物量的关系,结果表明：（1）不同草地利用方式显著影响草地生物多样性和生产力,在3种不同利用方式中,生物多样性总体的趋势是割草〉围封〉放牧,其中Shannon-Wiener指数、Simpson指数和物种丰富度均差异显著;割草草地地上生物量最高,围封草地次之,放牧草地最少。（2）将草地植物按照植物功能型分类,放牧草地1、2年生植物占优势,随着物种丰富度的增加,1、2年生植物生物量没有明显的变化趋势;割草草地以禾本科植物和非禾本科植物为主,随着物种丰富度的增加,禾本科植物生物量呈下降趋势,而非禾本科植物变化不明显;围封草地中禾本科植物占优势,其他功能型植物分布较均匀,多度、频度和生物量等差异不显著。（3）3种草地利用方式中只有围封草地物种丰富度和地上生物量存在显著的正相关,即随着物种丰富度的增加,生物量也随之升高。其他两种利用方式下,物种丰富度对地上生物量没有显著影响。     

Moderate grazing promotes the root biomass in Kobresia meadow on the northern Qinghai–Tibet Plateau

Grazing is an important modulator of both plant productivity and biodiversity in grassland community, yet how to determine a suitable grazing intensity in alpine grassland is still controversy. Here, we explore the effects of different grazing intensities on plant biomass and species composition, both at community level and functional group level, and examines the productivity–species richness relationship under four grazing patterns: no grazing (CK), light grazing (LG), moderate grazing, (MG) and heavy grazing (HG), attempt to determine a suitable grazing intensity in alpine grassland. The results were as follows. The total aboveground biomass (AGB) reduced with increasing grazing intensity, and the response of plant functional groups was different. AGB of both sedges and legumes increased from MG to HG, while the AGB of forbs reduced sharply and the grass AGB remained steady. There was a significant positive relationship between productivity and species richness both at community level and functional group level. In contrast, the belowground biomass (BGB) showed a unimodal relationship from CK to HG, peaking in MG (8,297.72 ± 621.29 g/m²). Interestingly, the grassland community tends to allocate more root biomass to the upper soil layer under increasing grazing intensities. Our results suggesting that moderate levels of disturbance may be the optimal grassland management strategy for alpine meadow in terms of root production.     

Changes in plant biomass and species composition of alpine <Emphasis Type="Italic">Kobresia</Emphasis> meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the importance of species composition, species richness, the type of different growth forms, and plant biomass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, herbaceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a decrease from 13196.96±719.69 g/m2 in the sedge-dominated K. tibetica swamp to 2869.58±147.52 g/m2 in the forb and sedge dominated K. pygmaea meadow, and to 2153.08±141.95 g/m2 in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of belowground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P＜0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P＜0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P＜0.05); belowground biomass was positively correlated with soil moisture (P＜0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was positively correlated to soil organic matter and soil total nitrogen (P＜0.05). This suggests that the distribution of biomass coincided with soil moisture and edaphic gradient in alpine meadows.     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

Changes in plant biomass and species composition of alpine Kobresia meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the im-portance of species composition, species richness, the type of different growth forms, and plant bio-mass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, her-baceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a de-crease from 13196.96±719.69 g/m2 in the sedge-dominated K. tibetica swamp to 2869.58±147.52 g/m2 in the forb and sedge dominated K. pygmaea meadow, and to 2153.08±141.95 g/m2 in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of below-ground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P<0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P<0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P<0.05); belowground biomass was positively correlated with soil moisture (P<0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was posi-tively correlated to soil organic matter and soil total nitrogen (P<0.05). This suggests that the distribu-tion of biomass coincided with soil moisture and edaphic gradient in alpine meadows.     

高寒小嵩草草甸牦牛优化放牧强度的研究

高寒小嵩草草甸牦牛放牧强度试验表明:(1)不同放牧强度下各植物类群的地上生物量和总的地上生物量之间差异极显著,莎草科植物地上生物量的百分比组成之间差异极显著,禾本科和杂类草地上生物量的百分比组成之间差异显著,而且禾本科和莎草科(除对照外)植物的地上生物量及其百分比组成随放牧强度的增加而减小,杂类草的变化与之相反;(2)优良牧草比例和草地质量指数与放牧强度之间均呈负相关,而优良牧草比例的年度变化和牦牛个体增重的年度变化之间呈正相关;(3)群落的相似性系数随放牧强度的增加而减小.通过建立植被变化度量指标,认为优良牧草比例的年度变化是评价高寒小嵩草草甸放牧价值的直接度量指标,而相似性系数的变化和草地质量指数的变化与牦牛生产力没有明显的联系,不能反映草场植被放牧价值的变化,只能指示植物群落整体的相对变化程度;牦牛的放牧强度约为1.86头/hm2是小嵩草高寒草甸暖季草场可持续生产而不退化的最大放牧强度.     

Effects of resource additions on species richness and ANPP in an alpine meadow community

Aims Theories based on resource additions indicate that plant species richness is mainly determined by the number of limiting resources. However, the individual effects of various limiting resources on species richness and aboveground net primary productivity (ANPP) are less well understood. Here, we analyzed potential linkages between additions of limiting resources, species loss and ANPP increase and further explored the underlying mechanisms.Methods Resources (N, P, K and water) were added in a completely randomized block design to alpine meadow plots in the Qinghai-Tibetan Plateau. Plant aboveground biomass, species composition, mean plant height and light availability were measured in each plot. Regression and analysis of variance were used to analyze the responses of these measures to the different resource-addition treatments.Important findings Species richness decreased with increasing number of added limiting resources, suggesting that plant diversity was apparently determined by the number of limiting resources. Nitrogen was the most important limiting resource affecting species richness, whereas P and K alone had negligible effects. The largest reduction in species richness occurred when all three elements were added in combination. Water played a different role compared with the other limiting resources. Species richness increased when water was added to the treatments with N and P or with N, P and K. The decreases in species richness after resource additions were paralleled by increases in ANPP and decreases in light penetration into the plant canopy, suggesting that increased light competition was responsible for the negative effects of resource additions on plant species richness.     

The relationship between species richness and productivity in four typical grasslands of northern China

The relationship between plant species richness and primary productivity has long been acentral topic in biodiversity research.In this paper,we examine the relationship between species richness and productivity in four typical grasslands of Northern China at different spatial scales.At the community scale,a positive correlation was found for six of seven communities.A unimodal pattern was found only for one community (Stipa glareosa community),while at a large scale (vegetation type or landscape/region),the relationship was also found significantly positive.Species richness ranged from 4 to 35 species,and community aboveground productiand aboveground productivity were found in alpine meadow,followed by meadow steppe,typical steppe and desert steppe.     

Spatial heterogeneity and plant species richness at different spatial scales under rabbit grazing

Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.