Response of Polygonum viviparum species and community level to long-term livestock grazing in alpine shrub meadow in Qinghai-Tibet Plateau

Grazing by domestic herbivores is generally recognized as a major ecological factor and an important evolutionary force in grasslands. Grazing has both extensive and profound effects on individual plants and communities. We investigated the response patterns of Polygonum viviparum species and the species diversity of an alpine shrub meadow in response to long-term livestock grazing by a field manipulative experiment controlling livestock numbers on the Qinghai-Tibet Plateau in China. Here, we hypothesize that within a range of grazing pressure, grazing can alter relative allocation to different plant parts without changing total biomass for some plant species if there is life history trade-offs between plant traits. The same type of communities exposed to different grazing pressures may only alter relative species' abundances or species composition and not vary species diversity because plant species differ in resistant capability to herbivory. The results show that plant height and biomass of different organs differed among grazing treatments but total biomass remained constant. Biomass allocation and absolute investments to both reproduction and growth decreased and to belowground storage increased with increased grazing pressure, indicating the increasing in storage function was attained at a cost of reducing reproduction of bulbils and represented an optimal allocation and an adaptive response of the species to long-term aboveground damage. Moreover, our results showed multiform response types for either species groups or single species along the gradient of grazing Intensity. Heavy grazing caused a 13.2％ increase in species richness. There was difference in species composition of about 18％--20％ among grazing treatment. Shannon-Wiener (H') diversity index and species evenness (E) index did not differ among grazing treatments. These results support our hypothesis.     

Interactive influence of rainfall manipulation and livestock grazing on species diversity of the herbaceous layer community in a humid savannah in Kenya

Changes in rainfall regime and grazing pressure affect vegetation composition and diversity with ecological implications for savannahs. The savannah in East Africa has experienced increased livestock grazing and rainfall variability but the impacts associated with those changes on the herbaceous layer have rarely been documented. We investigated the effect of livestock grazing, rainfall manipulation and their interaction on the composition and diversity of the herbaceous community in the savannah for two years in Lambwe, Kenya. Rainfall manipulation plots were set up for vegetation sampling;these plots received either 50% more or 50% less rainfall than control plots. Simpson’s diversity and Bergere Parker indices were used to determine diversity changes and dominance respectively. The frequency of species was used to compute their abundance and their life forms as determined from the literature. Grazing significantly increased species diversity through suppression of dominant species. Rainfall manipulation had no significant impact on plant diversity in fenced plots, but rainfall reduction significantly reduced diversity in grazed plots. In contrast, rainfall manipulation had no impact on dominance in either fenced or grazed plots. The interaction of grazing and rainfall manipulation is complex and will require additional survey campaigns to create a complete picture of the implications for savannah structure and composition.     

Elevational changes in vascular plants richness,diversity, and distribution pattern in Abune Yosef mountain range,Northern Ethiopia

The aim of this research is to investigate the patterns of vascular plant species richness,diversity,and distribution along an elevation gradient in the Abune Yosef mountain range,Ethiopia.Preferential systematic sampling was employed to collect vegetation and environmental data along the elevation gradient.We found that plant species richness declines monotonically from low to high elevations.Specifically,vascular plant species richness and diversity were lower in the Afroalpine grassland(high elevation)than in the Dry evergreen Afromontane forest and Ericaceous forest(low elevations).In contrast,endemic vascular plant richness was significantly higher in the Afroalpine grassland than in the Dry evergreen Afromontane forest and Ericaceous forest.Elevation showed a significant impact on the richness,diversity,and endemism of vascular plants.According to Sorensen's coefficient,the similarity between Dry evergreen Afromontane forest and Ericaceous forest vegetation types is higher(32%)than the similarity between Ericaceous forest and Afroalpine grassland(18%).Only 5%similarity was recorded between the Dry evergreen Afromontane forest and Afroalpine grassland.Growth forms showed different elevationai richness patterns.Trees and liana increased monotonically up to 3300 m.Shrub and herb richness patterns followed a hump-shaped and inverted hump-shaped pattern along the elevation gradient.The elevation patterns of vascular plant species richness,diversity,and growth form in the present study may be attributed to differences in management intensity,spatial heterogeneity,microclimatic variations,and anthropogenic disturbances.     

Disentangling the role of management,vegetation structure,and plant quality for Orthoptera in lowland meadows

Seminatural grasslands provide habitats for various species and are important for biodiversity conservation. The understanding of the diverse responses of species and traits to different grassland managenient methods is therefore urgently needed. We disentangled the role of grassland management (fertilization and irrigation), vegetation structure (biomass, sward height) and plant quality (protein and fiber content) for Orthoptera communities in lowland hay meadows in Germany. We found vegetation structure to be the most important environmental category in explaining community structure of Orthoptera (species richness, total individuals, fiinctional diversity and species composition). Intensively used meadows (fertilized, irrigated, high plant biomass) were characterized by assemblages with few species, low functional diversity, and low conservation value. Thereby, the relatively moderate fertilizer inputs in our study system of up to -75 kg N/ha/year reduced functional diversity of Orthoptera, while this negative effect of fertilization was not detectable when solely considering taxonomic aspects. We found strong support for a prominent role of plant quality in shaping Orthoptera communities and especially the trait composition. Our findings demonstrate the usefulness of considering both taxonomic and functional comp on ents (functio nal diversity) in biodiversity research and we suggest a stronger involvement of plant quality measures in Orthoptera studies.     

封育对科尔沁沙地小叶锦鸡儿群落植被特征及空间异质性的影响

以科尔沁沙地小叶锦鸡儿群落为对象,分析了放牧和不同封育年限下小叶锦鸡儿群落的植被特征及植被分布的小尺度空间异质性.结果表明:放牧和封育样地内植被均以1年生草本植物为主,物种数没有明显差异;封育6年、封育12年样地的植株密度分别为(124.46±5.22)株·m~(-2)和(203.05±10.38)株·m~2,显著高于放牧样地(P<0.05);封育样地的Shannon-Wiener多样性指数、Simpson多样性指数、Pielou均匀度指数均低于放牧样地,并随着封育年限的增加而减小;封育样地植被分布的小尺度空间异质性小于放牧样地,并且封育年限越长,空间异质性越小.

Abstract：

This paper studied the vegetation characteristics and small-scale spatial heterogeneity of Caragana mirophylla community in Horqin Sandy Land in northeast Inner Mongolia of China under grazing and under 6-and 12 years enclosure, aimed to assess the effects of grazing and enclosure on vegetation restoration. In the sampling plots of grazing and different years enclosure, the species composition of C. mirophylla community all dominated by annual herbaceous plants. The species richness in grazed plot and in the plots enclosed for 6 and 12 years was 22, 19, and 20, respectively, with no significant difference. In the plots enclosed for 6 and 12 years, the plant density was (124.46±5.22) plants·m~(-2) and (203. 05±10. 38) plants·m~(-2), respec- tively, being significantly higher than that in grazed plot, which suggested that enclosure was an effective method to accelerate the vegetation restoration in Sandy Land. The Shannon-Wiener index, Simpson species diversity, and Pielou evenness in enclosed plots were lower than those in grazed plot, and decreased with increasing enclosure duration. The small-scale spatial heteroge-neity of vegetation in enclosed plots was smaller than that in grazed plot. The longer the enclosure duration, the smaller the spatial heterogeneity was.     

Spatial patterns of a savanna palm tree Borassus aethiopum and its temporal variability

The spatial distributions and associations of plant species in a stand can provide essential information about their dynamics. However, since tree spatial distribution and association depend on factors that operate at different scales, disentangling the effects of environmental heterogeneity and plant–plant interactions requires the choice of a suitable null model for spatial analysis. We analyzed the spatial distributions of the same savanna palm tree populations that were studied, the underlying hypothesis being that these distributions have changed in 20 years. To remove the effect of large-scale environmental heterogeneity, the inhomogeneous L-function under the heterogeneous Poisson null model was used. We showed: (i) unlike 20 years ago, adults had a regular distribution instead of an aggregated distribution in the grass savanna; (ii) although the spatial distribution of seedlings and juveniles was always aggregated, we observed a decrease in the size of the aggregates (intensities); (iii) except for juveniles, no other stages were associated with nutrient-rich patches, which was also different 20 years ago; (iv) we did not observe any particular difference in structure between two study sites, only that female palm trees were spatially associated with entire-leafed seedlings in the reserve while they were spatially independent in the rural area. Our study supports the hypothesis that the spatial distributions have partially changed, and that the management of spatial heterogeneity has improved and given more precision in the results.     

Variations in the Volatile Organic Compound Emission Potential of Plant Functional Groups in the Temperate Grassland Vegetation of Inner Mongolia, China

The biogenic volatile organic compounds (VOC) emitted by the vegetation of a terrestrial ecosystem play a key role in both regional air quality and tropospheric chemistry. To describe the general emission properties of VOC of different plant functional groups (PFG) in a typical temperate grassland in Inner Mongolia, China, we randomly selected 175 plant species and measured the quantities of isoprene and monoterpene in situ. Results showed that most plants had low VOC emission potential at the species level,especially for some dominant plants, such as Leymus chinensis Tzvel., Stipa grandis Smirn., and Agropyron cristatum Gaertn. At the PFG level, the lowest VOC emission potential was found for perennial rhizome grasses, a major PFG in a typical temperate grassland ecosystem. The effects of overgrazing and subsequent vegetation succession on the emission of VOC by different plant life form functional groups (PLFG)were also discussed.     

Plant biodiversity and its conservation strategy in the in-undation and resettlement districts of the Yangtze Three Gorges, China

The investigation on and conservation project of plant biodiversity in the inundation and resettlement districts of the Three Gorges region were carried out with the ongoing construction of the Three Gorges Dam. Based on the first-hand specimens collected for many years and the historical records, we found that there were about 1784 higher plant species, belonging to 762 genera of 170 families, which accounted for 59.19%, 75.30% and 85.85% of the total amount of species, genera and families in the Three Gorges region, including 27 species specific in these two districts. The shrub and grass communities were mostly distributed along the low-altitude river bank zone of the Yangtze River and were largely influenced by the dynamic water level of the Three Gorges reservoir area. The measurement of Shannon-Wiener and Pielou evenness indexes on 21 Pinus massoniana (Masson pine) communities distributed in the Three Gorges reservoir area was carried out, and results showed that the biodiversity of shrub layers was the highest, followed by grass layers, the middle, and tree layers, the lowest. Among the 11 typical shrub communities in this region, the abundance of shrub layers of Form. Rhus chinensis, Form. Cotinus coggygria var. cinerea, Form. Vitex negundo var. cannabifoliaand and Form. Coriaria sinica was relatively higher, which were 16, 26, 29 and 15, respectively. These four shrubs also showed high biodiversity index (H′), which were 1.791, 3.427, 2.949 and 1.718, respectively. Among the 9 grasslands mainly distributed along the river bank zone, 4 grassland communities with higher altitude (Form. Dicranopteris dichotoma, Form. Miscanthus floridutus, Form. Imperata cylindrica var. major and Form. Arthraxon hispidus) showed higher abundance and biodiversity, which were 1.697, 1.354, 1.144 and 1.018, respectively. Considering the inundation and resettlement districts, there were 22 vegetation types threatened by submersion, including 4 woody plant communities, 9 shrub communities and 9 grassland communities. The 6 shrub communities of Form. Securinega wuxiensis, Form. Bauhinia brachycarpa, Form. Geum aleppicum, Form. Distylium chinense, Form. Buxus henryi and Form. Myrsine africana would be fully submerged and part of Adianthum reniforme var. sinensis and Buxus ichangensis would be also submerged. At present, plant species conservation and monitoring stations in the Three Gorges region have been constructed, and more than 200 plant species have already been transplanted and conserved, including 37 threatened species listed in the China Plant Red Data Book and 11 local constructive species.     

Analysis of phytoplankton-zooplankton relationships in an oligotrophic lake under natural and manipulated conditions

During August, 1987, we performed a series of Limnocorral experiments in lake La Caldera, a small winter-kill lake in which phytoplankton is strikingly nutrient-limited. The effects of biomanipulation on zooplankton-phytoplankton relationships were assessed by monitoring both individual species and whole-assemblage responses. Two sizes of enclosures were used (15 and 350 litres) and two treatments were assayed: 1) removal of zooplankton by 45 μm filter net and 2) doubling the natural grazing pressure by increasing the zooplankton concentration. Results show the two enclosure types to differ strikingly: flagellates disappeared from the small enclosures, resulting in four- to six-fold changes in chlorophylla concentration and three- to four-fold changes in number of individuals. Most species were grazed (a prey selectivity based on criteria other than size was observed) and their net growth rate increased with zooplankton concentration, causing a net increase in the phytoplankton growth, a stimulatory effect probably through nutrient regeneration that overrides the losses due to grazing.     

Analysis of Grassland Vegetation of the Southwest Heilongjiang Steppe (China) Using the Power Law

In 1997, we conducted a vegetation survey in three semi-arid natural grasslands (steppes) withdifferent livestock grazing intensities in Southwest Heilongjiang Province, China. The dominant grasslandspecies was the grass Stipa baicalensis Roshev. Grasslands with light, intermediate, and heavy grazingintensities were located 10, 5, and 2 km from a village, respectively. Villagers use the steppe to raise cattle,horses, sheep, and goats. Each of the three grasslands was surveyed by placing 100 quadrats (50 cm×50 cm)along a 50 m line transect. Each quadrat was divided into four equal areas (25 cm×25 cm; S-quadrats) and allplant species occurring in each of these smaller areas were identified and recorded. These data were summa-rized into frequency distributions and the percentage of S-quadrats containing a given species and thevariance of each species were estimated. The power law was applied to these estimates. The power law wasused to evaluate the spatial heterogeneity and frequency of occurrence for each species in the grasslandcommunity. The lightly grazed grassland exhibited high spatial heterogeneity (caused by large plant size),the highest species diversity, and a high occurrence of S. baicalensis. In contrast, the heavily grazedgrassland exhibited high spatial heterogeneity (caused by patchy populations of small plant size), lowspecies diversity, and a low occurrence of S. baicalensis. We judged that the heavily grazed grassland wasovergrazed and exclusion of livestock from the degraded areas is necessary for recovery.     

放牧强度对短花针茅荒漠草原甲虫群落分布格局的影响

放牧对脆弱的荒漠草原生态系统有着重要影响,且随放牧强度及持续时间不同而变化。鞘翅目昆虫是环境监测与生物多样性变化的指示生物。利用巴氏罐诱法对短花针茅荒漠草原不同放牧强度草地的甲虫群落组成和多样性进行调查,探究放牧对荒漠草原甲虫群落的影响。结果表明:(1)步甲科、金龟科为短花针茅荒漠草原甲虫群落优势类群,埋葬甲科、芫菁科、拟步甲科和花金龟科为常见类群。(2)放牧强度增加不利于维持更多的捕食性甲虫;对照和轻度放牧样地可维持更多的腐食性甲虫。(3)甲虫数量随放牧强度增加而递减;群落多样性以重度放牧草地最大,轻度放牧草地最小;群落优势度为对照、中度、重度显著高于轻度放牧草地。各甲虫类群在不同放牧强度草地出现时间、高峰期均不同。(4)对照、轻度、重度放牧样地的甲虫优势类群群落结构不同于其他生境,但均与中度放牧样地存在相似性。轻度、中度、重度放牧样地的甲虫稀有类群群落结构不同于其他生境,但均与对照样地存在相似性。(5)甲虫群落个体数与植物群落物种丰富度、盖度、植物平均高度、生物量呈显著正相关。Shannon-Wiener多样性指数、Margalef丰富度指数均与植物群落物种丰富度、生物量显著负相关。研究结果为荒漠草原甲虫多样性保护提供参考依据。     

Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Effects of livestock grazing intensity on soil arbuscular mycorrhizal fungi and glomalin-related soil protein in a mountain forest steppe and a desert steppe of Mongolia

Arbuscular mycorrhizal fungi (AMF) are important components of the grassland ecosystems in terms of plant phosphorus uptake and accumulation of glomalin-related soil protein (GRSP). Though Mongolian grasslands are seriously degraded by livestock grazing, the effects of grazing on soil AMF and GRSP remain unclear. Here, we examined community composition and diversity of AMF as well as amount of GRSP at three different grazing intensities: lightly grazed (LG), moderately grazed (MG), and heavily grazed (HG) under two different types of grassland, mountain forest steppe at Hustai and desert steppe at Mandalgobi. The diversity and biomass of AMF-host and non-AMF plants strongly affected the overall AMF community composition and its diversity. When we separately analyzed the factors affecting soil AMF diversity at Hustai and Mandalgobi, decrease in the shoot biomass of Poaceae plants at Hustai and decreases in the species number and shoot biomass of AMF-host plants at Mandalgobi were significantly correlated with AMF diversity. GRSP decreased with increasing grazing intensity, which was significantly correlated with soil pH and total root biomass at Hustai. The decrease in plant biomass caused by grazing thus led to GRSP reduction. Our results showed that change in soil AMF community caused by livestock grazing were associated with change in the biomass and diversity of functional vegetation groups such as Poaeceae, AMF-host and non-AMF plants, indicating the importance to focus on such functional vegetation groups to evaluate the effect of grazing on AMF.

     

Response of Vegetation and Soil Carbon and Nitrogen Storage to Grazing Intensity in Semi-Arid Grasslands in the Agro-Pastoral Zone of Northern China

Overgrazing has been the primary cause of grassland degradation in the semi-arid grasslands of the agro-pastoral transition zone in northern China. However, there has been little evidence regarding grazing intensity impacts on vegetation change and soil C and N dynamics in this region. This paper reports the effects of four grazing intensities namely un-grazed (UG), lightly grazed (LG), moderately grazed (MG) and heavily grazed (HG) on vegetation characteristics and soil properties of grasslands in the Guyuan county in the agro-pastoral transition region, Hebei province, northern China. Our study showed that the vegetation height, canopy cover, plant species abundance and aboveground biomass decreased significantly with increased grazing intensity. Similarly, soil organic carbon (SOC) and total nitrogen (STN) in the 0–50 cm were highest under UG (13.3 kg C m⁻² and 1.69 kg N m⁻²) and lowest under HG (9.8 kg C m⁻² and 1.22 kg N m⁻²). Soil available nitrogen (SAN) was significantly lower under HG (644 kg N hm⁻²) than under other treatments (725–731 kg N hm⁻²) in the 0–50 cm. Our results indicate that the pasture management of “take half-leave half” has potential benefits for primary production and livestock grazing in this region. However, grazing exclusion was perhaps the most effective choice for restoring degraded grasslands in this region. Therefore, flexible rangeland management should be adopted in this region.     

Differential Indirect Effects of Excluding Livestock and Rabbits from Chalk Heath on the Associated Leafhopper (Hemiptera: Auchenorrhyncha) Fauna

Preliminary results are presented of sampling the leafhopper assemblages on a field experiment designed to examine the differential effects of rabbits and livestock (mainly sheep) on the vegetation of chalk heath in southern England. Experimental plots that excluded livestock either allowed entry by rabbits or excluded them. Results were compared with those from plots grazed by both livestock and rabbits. After 7 years, exclusion of grazing herbivores had resulted in predictable increases in vegetation height, but no major changes were detected in the species composition of the vegetation. As expected, ungrazed plots had higher species richness and greater abundances of several individual leafhopper species. However, plots grazed only by rabbits had a leafhopper assemblage that was distinct from either ungrazed or mixed grazing plots. It is suggested that rabbit grazing may have subtle effects on grassland invertebrate assemblages that are not necessarily predictable from an examination of the species composition of the vegetation. Chalk heath vegetation contains an unusual mixture of calcicole and calcifuge plant species, but the leafhopper assemblage included a restricted number of calcareous grassland specialist species and only one species strongly associated with acidic grasslands; most leafhoppers recorded were generalist grassland species.     

Effects of livestock grazing on aboveground insect communities in semi-arid grasslands of southeastern Arizona

Despite the importance of invertebrates in grassland ecosystems, few studies have examined how grassland invertebrates have been impacted by disturbances in the southwestern United States. These grasslands may be particularly sensitive to one common disturbance, livestock grazing, because they have not recently evolved in the presence of large herds of bison, an important mammalian herbivore. This study examined how livestock grazing influenced vegetation-associated insect communities in southeastern Arizona. Insect abundance, richness, diversity, community composition, and key environmental variables were compared between sites on active cattle ranches and sites on a 3160 ha sanctuary that has not been grazed by cattle for over 25 years. Vegetation-associated insect communities were found to be sensitive to livestock grazing. Overall abundance of these insects was lower on grazed grasslands, and certain insect orders appeared to be negatively affected by livestock grazing; beetles were less rich, flies were less diverse, and Hymenoptera were less rich and diverse on grazed sites. Conversely, Hemiptera were more diverse on grazed sites. Species composition of vegetation-associated insect communities also differed and was significantly correlated with percent vegetation cover and number of shrubs. Insect species responsible for these differences were taxonomically diverse, and included herbivores and predators/parasites. When compared to other studies conducted in areas of the United States that fall within the historic range of bison, this study suggests that invertebrates in areas outside this range may be more sensitive to grazing pressure.     

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.