Effect of livestock grazing in the partitions of a semiarid plant–plant spatial signed network

In recent times, network theory has become a useful tool to study the structure of the interactions in ecological communities. However, typically, these approaches focus on a particular kind of interaction while neglecting other possible interactions present in the ecosystem. Here, we present an ecological network for plant communities that consider simultaneously positive and negative interactions, which were derived from the spatial association and segregation between plant species. We employed this network to study the structure and the association strategies in a semiarid plant community of Cabo de Gata-Níjar Natural Park, SE Spain, and how they changed in 4 sites that differed in stocking rate. Association strategies were obtained from the partitions of the network, built based on a relaxed structural balance criterion. We found that grazing simplified the structure of the plant community. With increasing stocking rate species with no significant associations became dominant and the number of partitions decreased in the plant community. Independently of stocking rate, many species presented an associative strategy in the plant community because they benefit from the association to certain ‘nurse’ plants. These ‘nurses’ together with species that developed a segregating strategy, intervened in most of the interactions in the community. Ecological networks that combine links with different signs provide a new insight to analyze the structure of natural communities and identify the species which play a central role in them.     

Restoring biopedturbation in grasslands with anthropogenic focal disturbance

Grassland ecosystems evolved with natural disturbance events on multiple spatial scales in which focal, fine-scale soil disturbance by animals often was imbedded within large-scale grazing disturbance. The resulting plant communities adapted to both broad-scale and fine-scale disturbance that resulted in species-rich plant communities. These natural disturbance regimes have been largely replaced by anthropogenic disturbance. While we generally understand grassland response to modern grazing practices, we know much less about plant community response to soil disturbance imbedded within non-focal grazing. Therefore, we used a tracked vehicle to focally disturb soil in a North American mesic mixed prairie that was either undisturbed prairie or prairie with a recent history of disturbance from either grazing or haying. Successional trajectory and recovery time following focal soil disturbance was similar between grazed and hayed plant communities. Species composition did not differ (P < 0.05) between grazed or hayed prairie and the respective undisturbed prairie. Plant species richness and bare ground increased (P < 0.05) following focal soil disturbance in both grazed and hayed communities, but focal soil disturbance combined with either grazing or haying did not change either plant species richness or bare ground more than (P > 0.05) focal soil disturbance alone. Also, the effect of focal soil disturbance was shortlived with recovery in two growing seasons. Our results suggest that anthropogenic focal soil disturbance is a reasonable mechanism to restore soil disturbance to the grassland ecosystem.     

Differences in species richness and life-history traits between grazed and abandoned grasslands in southern Sweden

Disturbance has profound effects on plant community composition. This paper deals with the influence of grazing on species richness and proportions of life‐history attributes of grassland vegetation at six spatial scales (0.001–1000 m²) in two provinces of southern Sweden. The study comprised 33 dry grassland sites, including 22 grazed and 11 abandoned localities, and 28 sites of coastal brackish meadows, divided into five management types (from “heavily grazed” to “abandoned since long time”).
In general grazed sites were species‐richer than abandoned sites, especially at small plot sizes. However, there was a steeper increase in species number towards larger plot sizes in the abandoned sites. Heavy grazing in the coastal meadows resulted in a comparatively low number of species, corroborating the intermediate disturbance hypothesis.
The analysis of life‐history traits indicated the importance of taxonomic group, canopy structure, height, regenerative strategy and, in particular, life form. Leaf anatomy and seed dispersal seemed to be less important. The responses to grazing as regards species traits differed somewhat between grassland types. Grazed sites generally had high proportions of legumes, therophytes, species with basal position of leaves and with regeneration by means of a persistent seed bank. Abandonment of grazing favoured monocots, geophytes, species with vegetative regeneration and (partly) leafy canopy structure. Some differences between grazed and abandoned sites were confined to either the smallest or largest plot sizes, indicating different responses of matrix and interstitial species. Various positive associations (attribute syndromes) or negative associations between individual traits were identified. There was, for example, a positive link between the attributes “geophytes” and “ability of vegetative regeneration”. The recognition of such links is important to avoid misinterpreting certain attributes as functional adaptations to grazing while they are only positively correlated to other attributes of larger significance.     

Effects of grazing,topography, and precipitation on the structure of a semiarid grassland

Structural aspects of the shortgrass steppe plant community, functional groups, and species populations were examined in response to long-term heavy grazing and exclosure from grazing, contiguous wet or dry years, and an environmental gradient of topography. Of the three factors, relatively greater differences in community similarity were observed between catena positions, particularly on the ungrazed treatments. Grazing was intermediate between catena position and short-term weather in shaping plant community structure. Grazed treatments and ridgetops had a less variable species composition through fluctuations in weather.An increase with grazing of the dominant, heavily grazed species was observed. Basal cover and density of total species was also greater on grazed sites. The more uniform grazing lawn structure of the grazed plant communities had an influence on segregation of plant populations along topographical gradients. Segregation was less on grazed catenas, but diversity and the abundance of introduced and opportunistic-colonizer species was also less.Although the shortgrass steppe community was relatively invariant, less abundant species were dynamic and interactions occurred with respect to grazing, weather, and catena position. The effects of grazing may be mitigated by favorable growing seasons but magnified in unfavorable years in populations that are adapted to favorable sites. Grazing can be considered a disturbance at the level of the individual but it may or may not be a disturbance at the level of the population, and it is not a disturbance at the level of the community in this particular grassland.     

Changes in semi-arid plant species associations along a livestock grazing gradient

In semi-arid ecosystems, vegetation is heterogeneously distributed, with plant species often associating in patches. These associations between species are not constant, but depend on the particular response of each species to environmental factors. Here, we investigated how plant species associations change in response to livestock grazing in a semi-arid ecosystem, Cabo de Gata-Níjar Natural Park in South East Spain. We established linear point-intercept transects at four sites with different grazing intensity, and recorded all species at each point. We investigated plant associations by comparing the number of times that each pair of species occurred at the same spatial point (co-occurrences), with the expected number of times based on species abundances. We also assessed associations for each shrub and grass species by considering all their pairs of associations and for the whole plant community by considering all pairs of associations on each site. At all sites, the plant community had a negative pattern of association, with fewer co-occurrences than expected. Negative association in the plant community increased at maximum grazing intensity. Most species associated as expected, particularly grass species, and positive associations were most important at intermediate grazing intensities. No species changed its type of association along the grazing gradient. We conclude that in the present plant community, grazing-resistant species compete among themselves and segregate in space. Some shrub species act as refuges for grazing-sensitive species that benefit from being spatially associated with shrub species, particularly at intermediate grazing intensities where positive associations were highest. At high grazing intensity, these shrubs can no longer persist and positive associations decrease due to the disappearance of refuges. Spatial associations between plant species and their response to grazing help identify the factors that organize plant communities, and may contribute to improving management of semi-arid ecosystems.     

Grazer exclusion alters plant spatial organization at multiple scales,increasing diversity

Grazing is one of the most important factors influencing community structure and productivity in natural grasslands. Understanding why and how grazing pressure changes species diversity is essential for the preservation and restoration of biodiversity in grasslands. We use heavily grazed subalpine meadows in the Qinghai‐Tibetan Plateau to test the hypothesis that grazer exclusion alters plant diversity by changing inter‐ and intraspecific species distributions. Using recently developed spatial analyses combined with detailed ramet mapping of entire plant communities (91 species), we show striking differences between grazed and fenced areas that emerged at scales of just one meter. Species richness was similar at very small scales (0.0625 m²), but at larger scales diversity in grazed areas fell below 75% of corresponding fenced areas. These differences were explained by differences in spatial distributions; intra‐ and interspecific associations changed from aggregated at small scales to overdispersed in the fenced plots, but were consistently aggregated in the grazed ones. We conclude that grazing enhanced inter‐ and intraspecific aggregations and maintained high diversity at small scales, but caused decreased turnover in species at larger scales, resulting in lower species richness. Our study provides strong support to the theoretical prediction that inter‐ and intraspecific aggregation produces local spatial patterns that scale‐up to affect species diversity in a community. It also demonstrates that the impacts of grazing can manifest through this mechanism, lowering diversity by reducing spatial turnover in species. Finally, it highlights the ecological and physiological plant processes that are likely responding to grazing and thereby altering aggregation patterns, providing new insights for monitoring, and mediating the impacts of grazing.     

Vegetation and soil seed bank in a 23-year grazing exclusion chronosequence in a Mediterranean dry grassland

Long-term grazing shaped plant diversity in dry Mediterranean grasslands. Abandonment of grazing affects plant diversity especially in the northern Mediterranean. Considerable efforts are, therefore, under way for grassland conservation and restoration. Yet, we do not know at which temporal scales impacts of grazing abandonment appear and in particular how soil seed banks evolve after longer grazing abandonment. Here, we provide detailed data from one of the very few long-term experiments available. These experiments provide data for up to 23 years (1982–2005) of grazing exclusion built in 1982, 1989, 2000 and 2001. Grazing exclusion decreased species richness, modified vegetation structure and changed soil parameters. Decline in species richness appears in communities that experienced 16 and 23 years of grazing exclusion. Only four to nine plant species of this Mediterranean grassland built persistent soil seed banks appearing after grazing exclusion, compared to 40–50 species in the established vegetation of grazed plots. Hence, similarity between vegetation and soil seed bank decreased with time of grazing exclusion. Even 23 years after abandonment, no woody plants colonised the experiments. We conclude that vegetation will recover fast from grazing abandonment in the short-term. Nevertheless, longer abandonment will impact diversity due to reduced soil seed banks.     

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.     

Effect of spatial and temporal patterns of stress and disturbance intensities in a sub-Mediterranean grassland

Abstract

The present study examined a sub-Mediterranean pastoral system in the central Apennines (Italy) with a long history of grazing, where winter cold stress is alternated with summer drought stress. The research goals were to ascertain whether different floristic structures correspond to different stress conditions (xeric and semimesic), and whether peculiar functional plant traits (such as avoidance and tolerance mechanisms) respond to stress/disturbance intensities, and understand how vegetation reacts to changeable livestock pressure (through floristic and plant trait variations). Cluster analysis indicated that separate communities develop under different stress intensities. Other analyses highlighted how avoidance strategies predominate within the pastoral system. Observations of grazed and ungrazed patches conducted in 10-m transects revealed spiny cushion formation in semimesic grassland, where a brief period of overgrazing occurs in late summer, causing variations in plant community structure. All these results confirm the importance of historical grazing and current land use, showing how small disturbances and stress variations cause ecosystem responses. Best practices for management were identified. In xeric conditions, it is advisable that the intensity of disturbance be lessened, while in semimesic grassland overgrazing should be forbidden during the dry period, because it could facilitate the development of spiny patches, and subsequent spread of Brachypodium rupestre.     

Context‐dependency and anthropogenic effects on individual plant–frugivore networks

Anthropogenic activities, such as grazing by domestic animals, are considered drivers of environmental changes that may influence the structure of interaction networks. The study of individual‐based networks allows testing how species‐level interaction patterns emerge from the pooled interaction modes of individuals within populations. Exponential random graph models (ERGMs) examine the global structure of networks by allowing the inclusion of specific node (i.e. interacting partners) properties as explanatory covariates. Here we assessed the structure of individual plant–frugivore interaction networks and the ecological variables that influence the mode of interactions under different land‐use (grazed versus ungrazed protected areas). We quantified the number of visits, the number of fruits removed per visit and the interaction strength of mammal frugivore species at each individual tree. Additionally we quantified ecological variables at the individual, microhabitat, neighborhood and habitat scales that generated interaction network structure under the different land uses. Individual plant–frugivore networks were significantly modular in both land uses but the number of modules was higher in the grazed areas. We found interaction networks for grazed and ungrazed lands were structured by phenotypic traits of individual trees, by the microhabitat beneath the tree canopy and were affected by habitat modifications of anthropogenic origin. The neighborhood surrounding each individual plant influenced plant–frugivore interactions only at the grazed‐land trees. We conclude that anthropogenic land uses influence the topological patterns of plant–frugivore networks and the frugivore visitation to trees through modification of both habitat complexity and the ecological traits underlying interactions between individual plants and frugivore species.     

Livestock grazing affects vernal pool specialists more than habitat generalists in montane vernal pools

Questions

Do livestock grazing and seasonal precipitation structure species composition in montane vernal pools? Which grazing and precipitation variables best predict cover of vernal pool specialists and species with broader habitat requirements? Is vernal pool species diversity correlated with livestock exclosure, and at what spatial scales?

Location

Montane vernal pools, northeast California, USA.

Methods

Vegetation was sampled in 20 vernal pools, including pools where livestock had been excluded for up to 20 years We compared plant species composition, functional group composition and species diversity among sites that varied in grazing history and seasonal precipitation using CCA and LMM.

Results

Although vernal pool specialists were dominant in montane vernal pools, over a third of plant cover was comprised of species that occur over a broad range of wetland or upland environments. The species composition of vernal pool plant communities was influenced by both livestock grazing and precipitation patterns, however the relative effects of these environmental variables differed by functional group. Livestock exclosures favoured perennial vernal pool specialists over annual vernal pool specialists. In contrast, the cover of habitat generalists was more strongly influenced by seasonal precipitation than livestock grazing. At small spatial scales, species richness and diversity decreased as the number of years a pool had been fenced increased, but this relationship was not significant at a larger spatial scale.

Conclusions

Both livestock grazing and seasonal precipitation structure the montane vernal pool plant community. We found that livestock grazing promotes the cover of annual vernal pool specialists, but at the expense of perennial vernal pool specialists. Wetter vernal pools, however, support higher cover of wetland generalist species regardless of whether pools are grazed.     

Plant life history stage and nurse age change the development of ecological networks in an arid ecosystem

Understanding how ecological networks are organised over the course of an organism's lifetime is crucial for predicting the dynamics of interacting populations and communities across temporal scales. However, most studies so far considered only one life history stage at a time, such as adult, when studying networks of interacting species. Therefore, knowledge about how multiple life history stages affect the development and stability of plant–plant association networks is lacking. We measured the understory adult plant community and the soil seed bank across a plant age gradient of the nurse shrub Retama sphaerocarpa in an arid ecosystem in Spain. Using a multilayer network approach, we built adult understory–nurse and seed bank–nurse networks and analysed how network nestedness, species’ role, and species specificity varied between them and with nurse plant age. We found that seed bank and adult understory networks changed depending on nurse plant age in two different ways. With increasing nurse plant age, adult understory networks became significantly more nested than seed bank networks. The nested architecture of seed bank networks was therefore a poor predictor of adult understory network nestedness. The contribution and specificity of species to network nestedness increased with increasing nurse plant age more in the adult understory than in seed bank networks, despite high species turnover. Our data show that life history and ontogeny affect the development of plant–plant association networks. Niche construction and environmental filtering along nurse ontogeny seem pivotal mechanisms structuring adult understory networks while the assembly of seed bank networks seems rather stochastic. We highlight the importance of mature plant communities for maintaining rare species populations and supporting the stability of ecological communities through time.     

Regional productivity mediates the effects of grazing disturbance on plant cover and patch‐size distribution in arid and semi‐arid communities

Patch‐size distribution and plant cover are strongly associated to arid ecosystem functioning and may be a warning signal for the onset of desertification under changes in disturbance regimes. However, the interaction between regional productivity level and human‐induced disturbance regime as drivers for vegetation structure and dynamics remain poorly studied. We studied grazing disturbance effects on plant cover and patchiness in three plant communities located along a regional productivity gradient in Patagonia (Argentina): a semi‐desert (low‐productivity community), a shrub‐grass steppe (intermediate‐productivity community) and a grass steppe (high‐productivity community). We sampled paddocks with different sheep grazing pressure (continuous disturbance gradients) in all three communities. In each paddock, the presence or absence of perennial vegetation was recorded every 10 cm along a 50 m transect. Grazing effects on vegetation structure depended on the community and its association to the regional productivity. Grazing decreased total plant cover while increasing both the frequency of small patches and the inter‐patch distance in all communities. However, the size of these effects was the greatest in the high‐productivity community. Dominant species responses to grazing explained vegetation patch‐ and inter‐patch‐size distribution patterns. As productivity decreases, dominant species showed a higher degree of grazing resistance, probably because traits of species adapted to high aridity allow them to resist herbivore disturbance. In conclusion, our findings suggest that regional productivity mediates grazing disturbance impacts on vegetation mosaic. The changes within the same range of grazing pressure have higher effects on communities found in environments with higher productivity, markedly promoting their desertification. Understanding the complex interactions between environmental aridity and human‐induced disturbances is a key aspect for maintaining patchiness structure and dynamics, which has important implications for drylands management.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.     

放牧对呼伦贝尔草地植物和土壤生态化学计量学特征的影响

放牧通过畜体采食、践踏和排泄物归还影响草地群落组成、植物形态和土壤养分,植物通过改变养分利用策略适应环境变化。通过分析呼伦贝尔草原放牧和围封样地中的群落植物和土壤的碳氮磷养分及化学计量比,探讨放牧对生态系统化学计量学特征和养分循环速率的影响机制。结果如下:(1)群落尺度上,放牧和围封草地植物叶片C、N和P的含量没有显著差异;但是在种群尺度上,放牧草地植物叶片N含量显著高于围封草地;(2)放牧草地土壤全C、全N、有机C、速效P含量,低于围封草地,硝态N含量高于围封草地;土壤全P和铵态N指标没有显著差异;(3)放牧草地植物C∶N比显著低于围封草地,植物残体分解速率较快,提高了生态系统养分循环速率。     

Grazing-induced effects on soil properties modify plant competitive interactions in semi-natural mountain grasslands

Plant-soil feedbacks are widely recognized as playing a significant role in structuring plant communities through their effects on plant-plant interactions. However, the question of whether plant-soil feedbacks can be indirectly driven by other ecological agents, such as large herbivores, which are known to strongly modify plant community structure and soil properties, remains poorly explored. We tested in a glasshouse experiment how changes in soil properties resulting from long-term sheep grazing affect competitive interactions (intra- and inter-specific) of two graminoid species: Nardus stricta, which is typically abundant under high sheep grazing pressure in British mountain grasslands; and Eriophorum vaginatum, whose abundance is typically diminished under grazing. Both species were grown in monocultures and mixtures at different densities in soils taken from adjacent grazed and ungrazed mountain grassland in the Yorkshire Dales, northern England. Nardus stricta performed better (shoot and root biomass) when grown in grazing-conditioned soil, independent of whether or not it grew under inter-specific competition. Eriophorum vaginatum also grew better when planted in soil from the grazed site, but this occurred only when it did not experience inter-specific competition with N. stricta. This indicates that plant-soil feedback for E. vaginatum is dependent on the presence of an inter-specific competitor. A yield density model showed that indirect effects of grazing increased the intensity of intra-specific competition in both species in comparison with ungrazed-conditioned soil. However, indirect effects of grazing on the intensity of inter-specific competition were species-specific favouring N. stricta. We explain these asymmetric grazing-induced effects on competition on the basis of traits of the superior competitor and grazing effects on soil nutrients. Finally, we discuss the relevance of our findings for plant community dynamics in grazed, semi-natural grasslands.     

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。     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

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） with different livestock grazing intensities in Southwest Heilongjiang Province, China, The dominant grassland species was the grass Stipa baicalensis Roshev. Grasslands with light, intermediate, and heavy grazing intensities 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 all plant species occurring in each of these smaller areas were identified and recorded. These data were summarized into frequency distributions and the percentage of S-quadrats containing a given species and the variance of each species were estimated. The power law was applied to these estimates. The power law was used to evaluate the spatial heterogeneity and frequency of occurrence for each species in the grassland community. 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 grazed grassland exhibited high spatial heterogeneity （caused by patchy populations of small plant size）, low species diversity, and a low occurrence of S. baicalensis. We judged that the heavily grazed grassland was overgrazed and exclusion of livestock from the degraded areas is necessary for recovery.     

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.