Impact of grazing regime on a Mongolian forest steppe

Question: What is the impact of grazing regime on plant species abundance, plant growth form, plant productivity and plant nutrient concentrations in a forest steppe? Location: Hustai National Park in the forest steppe region of Mongolia. Methods: On the Stipa steppe we applied three different grazing regimes by using; (1) one type of exclosure which excluded grazing by large mammalian herbivores, mainly takh (Przewalski horse), (2) another type of exclosure that excluded both large and small (Siberian marmots) mammalian herbivores, and (3) control plots which were freely grazed. We measured species frequencies, tiller densities, plant biomass and nitrogen concentrations of the vegetation. Results: Exclusion from grazing by takh and marmots significantly increased plant standing crop, but marmot grazing and full grazing did not show significant differences. Protection from grazing decreased forage quality, shown by a lower N-concentration of the standing crop. However, this was solely the result of the lower live-dead ratio of the vegetation. The frequency of the rhizomatous Leymus chinensis decreased under reduced grazing, as did the frequency of the total of rhizomatous species. The frequency of Stipa krylovii increased under reduced grazing, as did its basal areas, tiller density and tussock height. Conclusion: Reduced grazing leads to a lower abundance of rhizomatous species and an increase in tussock species.     

Effect of cattle grazing on soil salinity and vegetation composition along an elevation gradient in a temperate coastal salt marsh of Samborombón Bay (Argentina)

Salt marshes of Samborombón Bay (Argentina) have been grazed sporadically at very low stocking rates, but in the last decade, grazing intensity increased due to agriculture expansion. We investigated the effect of cattle grazing on vegetation and soil salinity on the most extended Spartina densiflora community. This community develops along an elevation gradient where the frequency and duration of tidal flooding and soil salinity increased as elevation decreased. Vegetation and soil data were collected from a national park excluded to cattle grazing for 30 years and from an adjacent commercial livestock farm continuously grazed by cattle. As elevation level decreased, plant cover, richness and diversity of functional groups and species decreased. As we expected, grazing altered soil salinity and vegetation composition in different extent along the elevation gradient. Grazing changed vegetation structure more intensively in the high elevation level because it reduced the competitive exclusion exerted by S. densiflora, allowing the increase in floristic richness. Grazing increased soil salinity and the contribution of salt-tolerant species only in the medium but not in the low elevation level probably because the higher frequency and duration of tidal flooding counterbalanced the increase in evaporation promoted by biomass removal in the low respect to the medium elevation level. While grazing may cause positive impacts for plant conservation in the high elevation level, it may cause negative consequence for livestock production because of the reduction in forage quality along the entire elevation gradient.     

Effect of fencing and grazing on a Kobresia-dominated meadow in the Qinghai-Tibetan Plateau

Grazing is one of the most important factors influencing community structure and productivity in natural grasslands. Fencing to exclude grazers is one of the main management practices used to protect grasslands. Can fencing improve grassland community status by restraining grazing? We conducted a field community study and indoor soil analyses to determine the long-term effects of fencing and grazing on the above-ground community and soil in a Kobresia-dominated meadow in the Qinghai-Tibetan Plateau, NW China. Our results showed that fencing significantly improved above-ground vegetation productivity but reduced plant density and species diversity. Long-term fencing favored the improvement of forage grass functional groups and restrained the development noxious weed functional groups. There were significant positive effects of fencing on below-ground organic matter, total nitrogen, available nitrogen, total phosphorus and available phosphorus. The productivity of grazed meadow showed a weak decrease over time. There were long-term decreasing trends for plant density both in fenced and grazed meadows. Our study suggests that grazing can be considered as a useful management practice to improve species diversity and plant density in long-term fenced grasslands and that periodic grazing and fencing is beneficial in grassland management.     

Impact of selective grazing on plant production and quality through floristic contrasts and current-year defoliation in a wet grassland

Grazing impacts the structure and functional properties of vegetation through floristic changes (i.e., long-term effect) and current defoliation (i.e., short-term effect). The aim of this study was to assess the relative importance of these two grazing effects on productivity (ANPP) and plant quality (C/N ratio) among plant patches submitted to a variety of grazing intensity for several years. Long-term grazing effect was measured by comparing ANPP and C/N ratio among plant patches with contrasting floristic composition. Short-term impact of grazing was measured by comparing ANPP and C/N in plant patches, with and without defoliation. Floristic contrasts led to a lower ANPP in highly grazed patches than in lightly grazed ones. This result may be related to the increasing proportion of grazing-tolerant and grazing-avoiding species with increasing grazing intensity. Vegetation C/N contrasts were recorded among grazed patches but did not linearly relate to grazing intensity. Short-term effect of current-year defoliation on ANPP was limited as vegetation compensated for biomass removal. No evidence for grazing-enhancement of ANPP was found even at moderate grazing intensity. Long-term floristic changes with grazing thus appeared to be the main driving factor of variations in ANPP. In contrast, C/N ratio showed no general and consistent variation along the grazing gradient but varied consistently depending on the community investigated, thus suggesting an effect of the species pool available.     

Land use and drought interactively affect interspecific competition and species diversity at the local scale in a semiarid steppe ecosystem

Few studies have considered interactive effects of grazing and drought on species composition and the relative contribution of species to total biomass, although it is important to understand the short-term dynamics and community succession in grazed ecosystems. We monitored species diversity and relative biomass contribution at one site protected from grazing since 1979 (UG79), and at winter grazing (WG) and heavily grazed (HG) sites. Continuous heavy grazing resulted in lower plant height and more but small individuals (tillers or stolons). Drought significantly reduced total plant density on all sites. Grazing affected species diversity more than drought. Species richness at site UG79 was significantly higher than at sites WG and HG, while drought only tended to reduce species diversity. Drought stress and grazing disturbance interactively controlled species competition and functional groups. Both perennial grasses and forbs had greater contribution to total biomass at site UG79, and perennial grasses contributed more than 97% of total biomass at site WG. The contribution to total biomass of annual forbs and semi-shrubs significantly increased at site HG after two dry years. The significant decrease in Potentilla acaulis and a substantial increase in annual species at this site indicate that the perennial vegetation of this ecosystem is in great danger of extinction under conditions of prolonged drought.     

Plant species richness responses to grazing protection and degradation history in a low productivity landscape

Questions: Does species richness and abundance accumulate with grazing protection in low productivity ecosystems with a short evolutionary history of grazing, as predicted by emerging theory? How do responses to grazing protection inform degradation history? Location: Mulga (Acacia aneura) dry forest, eastern Australia, generally considered chronically degraded by livestock grazing. Methods: Three paired exclosures (ungrazed, and macropod‐grazed) were compared with open‐grazed areas after 25 years using quadrats located on either side of the fences. Additionally, the regional flora for mulga dry forest was assessed to identify species that may have declined and could be threatened by grazing. Results: Low herbaceous biomass accumulation (<1.3 t ha^?1) with full grazing protection confirmed a low productivity environment. For most plant life forms the highest species richness was in macropod‐grazed exclosures, an intermediate grazing disturbance that best approximates the evolutionary history of the environment. This was the net outcome of species that both declined and increased in response to grazing. Regeneration and subsequent self‐thinning of mulga was promoted with grazing protection, but did not confound interpretation of species richness and abundance responses. At the regional scale only 11 native species out of 407 comprising the mulga dry forest flora were identified as rare and potentially threatened by grazing. Conclusions: Significant increases in richness or abundance of native plants with grazing protection, persistence of perennial grasses, regeneration of mulga and scant evidence of a major decline in the regional flora are not consistent with established assertions that long‐grazed mulga dry forest has crossed functional thresholds that limit recovery. Further, a peak in species richness under intermediate (macropod) grazing is counter to the shape of the response predicted by emerging theory for recovery of species richness in a low productivity environment. The finding prompts a more thorough understanding of the distinction between environments with inherently low productivity and those degraded by grazing.     

Importance of grazing and soil acidity for plant community composition and trait characterisation in coastal dune grasslands

Question: Does grazing by large herbivores affect species composition or community‐wide variation in plant functional traits? Location: Dune grasslands at the Belgian coast. Methods: Plant cover and soil data were collected in 146 plots that were randomly selected at 26 grazed and ungrazed grassland sites. Plant community composition was assessed by Detrended Correspondence Analysis and mean values of plant trait categories were calculated across the plots. Results: Differentiation of plant composition and community‐wide plant trait characteristics was largely determined by grazing, soil acidity and their interaction. In ungrazed situations, a clear floristic distinction appears between acidic (non‐calcareous) and alkaline (calcareous) grasslands. In grazed situations, these floristic differences largely disappeared, indicating that grazing results in a decrease of natural variation in species composition. At higher soil pH, a larger difference in plant community composition and community‐wide plant traits was observed between grazed and ungrazed plots. In ungrazed situations, shifts in plant functional traits along the acidity gradient were observed. Conclusions: Grazing is responsible for shifts in plant community composition, and hence a decrease in plant diversity among grasslands at opposing acidity conditions in coastal dune grasslands. Therefore, care should be taken when introducing grazing as a system approach for nature conservation in dune grasslands as it may eliminate part of the natural variation in plant diversity along existing abiotic gradients.     

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.

     

Effects of grazing exclusion on plant species richness and phytomass accumulation vary across a regional productivity gradient

Question: Does long‐term grazing exclusion affect plant species diversity? And does this effect vary with long‐term phytomass accumulation across a regional productivity gradient? Location: Lowland grassy ecosystems across the state of Victoria, southeast Australia. Methods: Floristic surveys and phytomass sampling were conducted across a broad‐scale productivity gradient in grazing exclusion plots and adjacent grazed areas. Differences in species richness, evenness and life‐form evenness between grazed and ungrazed areas were analysed. The environmental drivers of long‐term phytomass accumulation were assessed using multiple linear regression analysis. Results: Species richness declined in the absence of grazing only at the high productivity sites (i.e. when phytomass accumulation was >500 g m^?2). Species evenness and life‐form evenness also showed a negative relationship with increasing phytomass accumulation. Phytomass accumulation was positively associated with both soil nitrogen and rainfall, and negatively associated with tree cover. Conclusions: Competitive dominance is a key factor regulating plant diversity in productive grassy ecosystems, but canopy disturbance is not likely to be necessary to maintain diversity in less productive systems. The results support the predictions of models of the effects of grazing on plant diversity, such as the dynamic equilibrium model, whereby the effects of herbivory are context‐dependent and vary according to gradients of rainfall, soil fertility and tree cover.     

Linking grazing response of species abundance to functional traits in the Tibetan alpine meadow

Whether grazing response of species abundance can be predicted by plant functional trait remains a challenge untested in a specific ecosystem. We measured species abundance and relevant functional traits for 30 common component species in a moderately grazed and a control community over 3 years (2005, 2006 and 2007) in a Kobresia capillifolia dominated alpine meadow on the Tibetan Plateau. Our objective was to examine species response to grazing and to test the relationship between the response of species abundance and functional traits in grazed and control habitats. Our results showed: i) in terms of species relative abundance and saturated height (the maximum height of an adult individual), most component species significantly decrease in response to moderate grazing and the effect differed among species and between functional groups. ii) The response of species abundance was significantly negatively correlated with saturated height, but not correlated with specific leaf area or seed size. We concluded that the response direction of species abundance to grazing can be predicted by functional traits, whereas it is a weak predictor of the extent of grazing response in species abundance.     

Productivity: key factor affecting grazing exclusion effects on vegetation and soil

In this study, we inquire into the effects of short-term goat grazing abandonment on plant species and functional composition, bare ground and net primary productivity (NPP) in two traditionally grazed pastures located in the Canarian Network of Natural Protected Areas and the Natura 2000 Network. In addition, we analyse soil chemical properties, biomass tannin content and energetic value to find out how grazing abandonment affects soil fertility and forage quality of these agroecosystems. Grazing exclusion effects on plant species and functional composition, as well as on soil fertility depended on the productivity of the studied pasture. Erect forbs and shrubs (endemic to Macaronesian region and native) were favoured by grazing removal in the most productive pasture, while soil fertility decreased in the driest and least productive site. An increase in NPP after exclusion was consistent among study sites. Although we consider goat grazing as necessary for maintaining traditional agroecosystems, we also suggest controlling it over time, allowing some periods of rest to give endemic shrub species time to recover from near propagule sources.     

Grazing as a mediator for maintenance of offspring diversity: Sexual and clonal recruitment in alpine grassland communities

To understand the effects of grazing on grassland plants sexual and clonal recruitment, we conducted a demographic field investigation of species recruitment along a grazing gradient in the Tibetan alpine grassland. Grazing intensity had significant effects on quantity and diversity of sexual and clonal recruitment. Sexual recruitment increased significantly, but clonal offspring production decreased significantly with increased grazing intensity. Grazing intensity had different, significant effects on offspring recruitment of the various functional groups in the community, grasses (GG), sedges (SG), legumes (LG) and forbs (FG). Higher grazing intensity reduced offspring recruitment of GG and SG; it increased offspring recruitment of LG and FG. Seedlings were significantly more abundant in lightly grazed, moderately grazed and heavily grazed meadows than in non-grazed grasslands. Offspring diversity from sexual recruitment was significantly higher than that from clonal recruitment in grazed than in non-grazed grasslands. Our studies indicate that moderate grazing had positive effects on seedling recruitment and offspring diversity, but heavy gazing may alter community succession by affecting recruitment patterns among the four plant functional groups.     

Grazing without grasses: Effects of introduced livestock on plant community composition in an arid environment in northern Patagonia

Question: How does grazing intensity affect plant density, cover and species richness in an Patagonian arid ecosystem? Location: Monte steppe ecoregion, SW Argentina. Methods: I analysed the effect of grazing on plant density, cover and species richness using a stocking rate gradient within the same habitat. Six paddocks were used with stocking rates ranging between 0.002 – 0.038 livestock/ha. Plant density, species richness, plant cover and percentage of grazed branches were determined by sampling plots within each paddock. The percentage of grazed branches was used as an independent measurement of grazing intensity. Results: Higher stocking rates were related to lower plant density, species richness and plant cover. The paddock with the lowest grazing intensity had 86% more plants per unit area, 63% more plant cover and 48% higher species richness. The percentage of grazed branches and the quantity of dung increased with stocking rate. Conclusions: Introduced livestock seriously affect native vegetation in the Patagonian Monte. The damage observed in this xerophytic plant community suggests that plant adaptations to aridity do not provide an advantage to tolerate or avoid grazing by vertebrate herbivores in this region. Plant degradation in this arid environment is comparable to the degradation found in more humid ecosystems.     

Comparison of biodiversity and ground cover between a commercial rotationally grazed property and an adjacent nature reserve in semi‐arid rangeland

Continuous livestock grazing can have negative effects on biodiversity and landscape function in arid and semi‐arid rangelands. Alternative grazing management practices, such as rotational grazing, may be a viable option for broad‐scale biodiversity conservation and sustainable pastoral management. This study compared ground cover, plant species composition and floristic and functional diversity along gradients of grazing intensity between a pastoral property rotationally grazed by goats and an adjacent nature reserve (ungrazed by commercial livestock) in semi‐arid south‐eastern Australia. Understorey plant species composition differed significantly between the rotationally grazed property and the nature reserve, with a greater proportion and frequency of palatable species recorded in the nature reserve. Understorey plant species richness, diversity, functional biodiversity measures and ground cover declined with increasing grazing pressure close to water points under commercial rotational grazing management. However, at a whole‐paddock scale, there were few differences in plant biodiversity and ground cover between the rotationally grazed property and the nature reserve, despite differences in overall plant species composition. Flexible, adaptive, rotational grazing should be investigated further for its potential to achieve both socio‐economic and biodiversity conservation outcomes in semi‐arid rangelands to complement existing conservation reserves.     

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。     

Effects of grazing on soil seed bank dynamics: An approach with functional groups

Abstract. The relationship between intensity and timing of cattle grazing on changes in the size and composition of the soil seed bank were investigated in a 3‐yr study in a Mediterranean grassland in northeastern Israel. Treatments included manipulations of stocking rates and of grazing regimes, in a factorial design. The retrieved soil seed bank community was rich in species, with 133 species accounting for 80% of the 166 species recorded at the site. Within the seed bank, 89% of the species were annuals. Seed bank dynamics was analysed in terms of plant functional groups and germination strategies. In terms of total seed bank density and including all functional groups, 42% of the seeds present in the soil did not germinate under watering conditions. The dormancy level differed greatly among functional groups. The seed bank of annual legumes, crucifers, annual thistles and annual forbs had a large fraction of non‐germinated seeds and characterized areas grazed early in the growing season under high and very high grazing intensity. These functional groups were considered to have a higher potential for persistent seed banks production. In contrast, short and tall annual grasses and tall perennial grasses, that were dominant in ungrazed or moderately grazed paddocks, generally had seed banks with a very small fraction of non‐germinated seeds. Seed bank densities varied widely between grazing treatments and years. Under continuous grazing, heavy grazing pressure reduced seed bank densities of grasses and crucifers in comparison to moderate grazing. The greatest reduction on the seed bank densities resulted from heavy grazing concentrated during the seed‐set stages.     

Moderate patchiness optimizes heterogeneity,stability, and beta diversity in mesic grassland

Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire–grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%–33% of area burned) and three‐ to four‐year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.     

·荒漠草地植物多样性对草食动物采食的响应机制

草食动物采食对草地植物多样性和生态系统功能的影响机制是放牧生态学研究的核心问题。该研究以内蒙古锡林郭勒盟苏尼特右旗荒漠草原的长期放牧控制实验为平台, 从既有草地植物多样性和动物偏食性两个层面系统地研究了荒漠草地植物多样性对草食动物采食的响应机制。结果显示: 1)荒漠草地植物对草食动物采食呈现4种响应模式: 放牧“隐没种”、放牧“敏感种”、放牧“无感种”、“绝对优势种”; 2)在群落尺度上, 物种多样性指数随放牧强度增加而减少, 与不放牧小区相比, 重度放牧(HG)与适度放牧(MG)小区植物多样性均下降, 且这一规律同样适用于功能群多样性, 灌木半灌木这一功能群内物种多样性对放牧干扰较敏感; 3)在草地既有植物的基础上, 以不放牧小区为参考系, 草食动物对植物功能群偏食性的排序为: 一二年生草本(AB) >多年生杂类草(PF) >灌木半灌木(SS) >多年生禾草(PG), 且偏食性物种主要分布于AB和PF中; 4)植物多样性与动物偏食性基本呈显著负相关关系(p < 0.05)。     

Can heavy grazing on communal land elevate plant species richness levels in the Grassland Biome of South Africa?

A common perception, particularly in South Africa, is that heavily and continuously grazed communal land leads to degradation and loss of plant diversity when compared to commercial rangeland farming or conservation areas. We focus on whether this applies to the Grassland Biome of South Africa and whether the opposite can occur, namely, an increase in plant species richness under heavy grazing. A study of a contrast between a communal area of the former Ciskei and a neighbouring nature reserve showed that intense utilization under communal use led to a significant increase in plant species richness. However, this increase was scale-dependent with the greatest significant difference occurring at sample plot scale (50 m²) but converging at the broader scale of the whole study site. Species that increased with heavy grazing included those from arid Karroid areas as well as some from more mesic grassland and savanna areas. The contribution of beta diversity to gamma diversity across the grazing contrast was relatively low which reflects the relatively high proportion of species shared between treatments. Total plant canopy cover declined sharply with heavy grazing. In terms of plant canopy cover, grazing favoured annual over perennial plants, prostrate over erect plants, and stoloniferous over tussock plant architecture. This pattern was not supported when expressed in terms of number of species belonging to these grazing response groups or traits.     

Response of galling invertebrates on Salix lanata to reindeer herbivory

Browsing and defoliation often increase the densities of insect herbivores on woody plants. Densities of herbivorous invertebrates were estimated in a long-term grazing manipulation experiment. More then 30-yr-old fences allow us to compare densities of invertebrate herbivores on Salix lanata in areas heavily grazed and areas lightly grazed by reindeer. The number of gall-forming insects ( Pontania glabrifons) and gall-forming mites were higher on the heavily grazed shrubs than on lightly grazed shrubs. In contrast to most short-term studies, the heavily grazed S. lanata had shorter current annual shoots. No difference in leaf size, leaf nitrogen content, or C:N ratio between grazing intensities were detected. However, the enhanced natural δ15N value indicates that heavily grazed shrubs get a higher proportion of their N directly from reindeer faeces. Leaf weight per unit area and relative fluctuating asymmetry of leaf shape increased in heavily grazed S. lanata . Enhanced relative fluctuating asymmetry might indicate higher susceptibility to herbivores. Long-term grazing seems to increase the density of invertebrate herbivory in the same way as short-term grazing, even if the plant responses differ substantially.