Forage quality in relation to long-term grazing history,current-year defoliation,and water resource

Forage nitrogen concentrations, nitrogen yields, and in vitro digestibilities were assessed in shortgrass steppe that had been ungrazed, lightly, or heavily grazed for 50 years. Caged plots were defoliated in amounts based upon removals observed in naturallygrazed reference plots or not defoliated. This was done in a year of average precipitation and with a supplemental water treatment to simulate a wet year. In general, current-year defoliation had positive effects, and longterm grazing and supplemental water had negative effects, on forage nitrogen concentrations and digestibilities. However, defoliation interacted with long-term grazing in determning forage nitrogen concentrations, and with grazing and with watering in determining digestibilities. Nitrogen concentration and digestibility increased with defoliation in lightly, but not in heavily, grazed treatments. The dilution effect of supplemental water an digestibilities through increased plant growth was offset by defoliation. The negative effects of long-term grazing on forage quality were small, equally or more than compensated for by defoliation in a year of average precipitation, but more pronounced in the simulated wet year. Nitrogen yields and digestible forage production were usually increased by defoliation, but this depended upon grazing and watering treatments. Increased nitrogen and digestible forage yields and concentrations in response to defoliation were greater than the biomass response in lightly grazed grassland. For both nitrogen and digestibility, yields were greater in grazed than ungrazed treatments in the year of average precipitation, but less in the simulated wet year. Optimizing quantity and year-to-year stability of nitrogen and digestible forage yield may best be achieved with light grazing rather than no or heavy grazing. Clipping was conducted in a manner closely resembling the natural pattern and intensity of defoliation by the cattle, and confirms the potential for a positive feedback of increased forage quality with defoliation observed in pot experiments. Long-term heavy grazing can diminish this response. Quantily (aboveground primary production, ANPP), quantity of quality (digestible and N yields), and quality (concentrations) do not necessarily respond similarly in interactions between current-year defoliation, long-term grazing history, and level of water resource.     

Does selective defoliation mediate competitive interactions in a semiarid savanna? A demographic evaluation

Abstract. Three patterns of target-neighbor plant defoliation were imposed on a late-seral, perennial, C₄-grass, Bouteloua curtipendula, in three long-term grazing regimes to determine the influence of selective defoliation on competitive interactions and species replacement in a semiarid savanna on the Edwards Plateau, Texas, USA. Short-term (3-yr) target plant defoliation did not significantly affect either tiller or plant responses in any of the three grazing regimes. Neighbor plant defoliation, either alone or in combination with target plants, produced a significant defoliation interaction with time for tiller number and basal area per plant, but not for tiller recruitment or mortality. The minimal effect of selective defoliation on the intensity of competitive interactions in this semiarid community indicates that selective grazing has a less definitive role in mediating herbivore-induced species replacement than it does in mesic grasslands and savannas. This interpretation is discussed within the context of long-term (45-yr) change in herbaceous vegetation associated with grazing in this community. Cumulative tiller recruitment in the intensively grazed regime was only 44% of that in the ungrazed regime because of greater plant mortality and fewer surviving plants that recruited tillers. Target plant mortality (50%) only occurred in the intensively grazed regime and the proportion of target plants that initiated tillers decreased by 70, 48 and 32% in the ungrazed, moderately and intensively grazed regimes, respectively, during the final two years of the investigation. The decrease in cumulative tiller recruitment in all grazing regimes was probably mediated by a drought-induced increase in median tiller age the second year of the study. However, tiller per tiller recruitment rate among plants that recruited at least one tiller remained relatively constant among grazing regimes and years. Intensive, long-term grazing has modified the population structure of this late-seral perennial grass to the extent that population responses to both herbivory and periodic drought have been altered in comparison with those of ungrazed and moderately grazed populations. Ecological consequences of a herbivore-induced transition in population structure may be to minimize the effect of selective herbivory on competitive interactions and to function as an avoidance mechanism to reduce the probability of localized population extinction in response to intensive long-term herbivory.     

模拟放牧斑块与氮素添加对半干旱草原群落植物生长的影响

放牧时,动物采食及其排泄物会影响植物的生长,但动物彩食及其排泄物的空间异质性可能会影响这咱效应.在位于我国北方典型农牧交错区的内蒙古多伦县,我们研究了模拟入牧斑块和施氧肥对植物生长的影响,实验采用模拟放牧采食斑块(观割半径分别0、10、20、40和8cm)和土壤施氮(分别为0、5、20Gn\m2)两种处理,植物地上部收获后分为绿体和立柯两部分,并分析其含氮量.结果表明,刈割降低了植物的生物量(40.5%),而施氮可增加生物量(57.8%)刈割交通规则植物生长的抑制作用在面积最水又施肥的斑块上表现更明显.土壤施氮可以促进杜物生长并且影响刈割效应.同时植物的绿-枯比阻碍施氮水平的增回而增回,因此氮会延迟植物的衰老.以上结果表明,刈割(模拟动物采食)斑块的大小会影响草原植物的生长,土壤施氮(模拟动物尿氮)可以提高草原生态系统的初级生产力,并影响刈割效应.     

模拟放牧斑块与氮素添加对半干旱草原群落植物生长的影响(英文)

放牧时,动物采食及其排泄物会影响植物的生长,但动物采食及其排泄物的空间异质性可能会影响这种效应。在位于我国北方典型农牧交错区的内蒙古多伦县,我们研究了模拟放牧斑块和施氮肥对植物生长的影响,实验采用模拟放牧采食斑块(刈割半径分别为0、10、20、40和80 cm)和土壤施氮(分别为0、5、10、20 g N/m2)两种处理,植物地上部收获后分为绿体和立枯两部分,并分析其含氮量。结果表明,刈割降低了植物的生物量(41.5％),而施氮可增加生物量(57.8％)。刈割对植物生长的抑制作用在面积最小又施肥的斑块上表现更明显。土壤施氮可以促进植物生长并且影响刈割效应。同时植物的绿-枯比随施氮水平的增加而增加,因此氮会延迟植物的衰老。以上结果表明,刈割(模拟动物采食)斑块的大小会影响草原植物的生长,土壤施氮(模拟动物尿氮)可以提高草原生态系统的初级生产力,并且影响刈割效应。     

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.     

A hierarchical model for analysing the stability of vegetation patterns created by grazing in temperate pastures

Questions: Does vegetation structure display any stability over the grazing season and in two successive years, and is there any correlation between the stability of these spatial patterns and local sward composition? Location: An upland grassland in the French Massif Central. Method: The mosaic of short and tall vegetation stands considered as grazed and ungrazed patches respectively is modeled as the realization of a Boolean process. This method does not require any arbitrarily set sward‐height thresholds to discriminate between grazed and ungrazed areas, or the use of additional variables such as defoliation indexes. The model was validated by comparing empirical and simulated sward‐height distributions and semi‐variograms. Results: The model discriminated between grazed and ungrazed patches at both a fine (1 m²) and a larger (500 m²) scale. Selective grazing on legumes and forbs and avoidance of reproductive grass could partly explain the stability of fine‐scale grazing patterns in lightly grazed plots. In these plots, the model revealed an inter‐annual stability of large‐scale grazing patterns at the time peak biomass occurred. At the end of the grazing season, lightly grazed plots showed fluctuating patch boundaries while heavily grazed plots showed a certain degree of patch stability. Conclusion: The model presented here reveals that selective grazing at the bite scale could lead to the creation of relatively stable patches within the pasture. Locally maintaining short cover heights would result in divergent within‐plot vegetation dynamics, and thus favor the functional diversity of vegetation.     

Cattle foraging habits shape vegetation patterns of alluvial year-round grazing systems

Year-round grazing with robust cattle is increasingly used as a near-natural tool for the restoration of structurally diverse grassland ecosystems in Western and Central Europe. The aim of this study was to evaluate the general success of year-round grazing and to analyze the interplay between emerging vegetation structures, grazing patterns and abiotic environmental conditions. In summer 2010 vegetation composition, aboveground biomass and soil properties were sampled at 44 quadrats of 4 × 4 m² within two year-round grazed floodplain sites in Northwestern Germany. For plot selection, we predefined structural vegetation types and later statistically determined indicative plant species for each structural type. Our results showed that year-round grazing resulted in the successful creation of eutrophic grassland communities on former agricultural land after 15 years. Soil parameters like phosphorous and potassium concentration and the flooding duration did not or only slightly differ between different structure types. In summer, cattle preferably fed at short-growing patches which were of better digestible biomass than taller patches. Hence, our data clearly demonstrate a positive feedback between grazing intensity and fodder quality leading to a patchy vegetation structure of intensively grazed swards and less frequented areas dominated by high-growing grasses and tall forbs, almost independently from primary differences in soil parameters and other site factors such as flooding duration. The remarkable structural and floristic diversity of year-round grazing systems clearly is a result of these spatially contrasting feeding patterns.     

Production and rain use efficiency in short-grass steppe: grazing history,defoliation and water resource

Abstract. Grassland in the semiarid shortgrass steppe, subjected to 50 years of heavy, light, and no grazing intensity, was clipped to simulate the natural pattern and intensities of defoliation by cattle or not clipped. A level of water resource treatment was superimposed upon the grazing and clipping treatments. Half of the plots were supplemented with additional water to simulate a wet year and half were not supplemented in a year of average precipitation. All three treatments interactively determined above-ground production. Water treatment had the largest overall effect on above-ground production. Current-year defoliation had no direct significant effect on production, but mediated differences between both longterm grazing and watering treatments. Long-term ungrazed compared to grazed grassland was capable of responding to high amounts of precipitation, but was also most affected by low amounts of precipitation and, therefore, displayed greater variability in above-ground production and rain use efficiency. Only in the year of average precipitation, defoliation increased rain use efficiency in long-term lightly, but not heavily, grazed treatment. This suggests a water conservation mechanism of defoliation that is reduced with heavy grazing.     

Effects of grazing pattern and nitrogen availability on primary productivity

A major part of the impact of grazing on primary productivity results from the joint action of tissue removal and nutrient return to the soil via dung and urine. Grazing, however, is not uniformly distributed in space: grazed grasslands show a matrix of grazed and ungrazed patches, which in turn, may or may not be affected by faecal or urine deposition. This paper investigates the effects of grazing spatial pattern and nitrogen availability on primary productivity. We propose that grazed plants located at the edge of a grazed patch are more shaded by their taller ungrazed neighbours than plants at the center. Since the border effect is less important as patch size increases, the effects of grazing will be more positive, or less negative, when grazing pattern is coarse-grained than when it is fine-grained. We also propose that nitrogen availability will affect this response to grazing through its effects on the intensity of competition for light and on the amount of compensatory growth. We performed a field experiment in a grassland community of the Flooding Pampa, Argentina, in which we compared the productivity of undefoliated controls and defoliated patches of different size, with and without nitrogen application. Defoliation reduced primary productivity and this effect was greater in the smallest, fertilized patches. Productivity was highest at patches of intermediate and large sizes. Nitrogen addition increased productivity by two-fold. The integrated photon flux density reaching the base of the canopy was affected by defoliation and by patch size: it was lower in controls than in defoliated patches and increased with patch size. Our results showed that (a) the size of the defoliated patch modified the response of this grassland to defoliation, (b) this response was correlated with light availability, and (c) nitrogen addition, simulating urine depositions, increased primary productivity and affected the response to defoliation of the smallest patches.     

The relation of biomass production with leaf traits varied under different land-use and precipitation conditions in an Inner Mongolia steppe

The analyses of plant leaf traits that strongly influence aboveground net primary production (ANPP) are indispensable for understanding the process of plant biomass formation. However, there are few studies that have attempted to relate patterns of ANPP under contrasting management practices to plant leaf-level traits. To assess how leaf traits affect plant biomass accumulation under different land-use practices, we examined leaf traits and biomass production in three differently managed sites in the Inner Mongolia steppe: a site fenced since 1979 (UG), a winter grazing site (WG), and a heavily grazed site (HG). Low soil water content, leaf area index, and potential growth ability of species at site HG led to low crop growth rate (CGR), net assimilation rate (NAR), and relative growth rate (RGR); resulting in lower ANPP as compared to sites WG and UG. Irrespective of land-use management, prolonged drought significantly decreases ANPP even though it systematically increases mean CGR and RGR. However, leaf N content and leaf weight ratio are the crucial components necessary to determine the RGR at site WG. This suggests that low leaf N and availability of soil N due to haymaking may be responsible for neither over-compensatory nor compensatory growth in this site. The low ANPP in dry years is not due to the low mean CGR and RGR but rather to the short effective growing days (referring to the days the vegetation actually grows), suggesting that production-adjusted grazing regimes may be the most suitable measures for precision land management and avoiding grassland degradation.     

Effects of reindeer on boreal forest floor vegetation: Does grazing cause vegetation state transitions?

Intensive reindeer grazing has been hypothesized to drive vegetation shifts in the arctic tundra from a low-productive lichen dominated state to a more productive moss dominated state. Although the more productive state can potentially host more herbivores, it may still be less suitable as winter grazing grounds for reindeer, if lichens, the most preferred winter forage, are less abundant. Therefore, such a shift towards mosses may have severe consequences for reindeer husbandry if ground-growing lichens have difficulties to recover. We tested if reindeer cause this type of vegetation state shifts in boreal forest floor vegetation, by comparing plant species composition and major soil processes inside and outside of more than 40-year-old exclosures. Lichen biomass was more than twice as high inside exclosures than in grazed controls and almost 5 times higher than in heavily grazed patches. Contrary to our predictions, net N mineralization and plant production were higher in the exclosures than in the grazed controls. The lack of response of phytometer plants in a common garden bioassay indicated that changed soil moisture may drive effects of reindeer on plant productivity in these dry Pine forest ecosystems.     

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.     

Vegetation and soil responses to livestock grazing in Central Asian grasslands: a review of Chinese literature

Grasslands in northern China and the Qinghai-Tibetan plateau are particularly important to both ecosystem functioning and pastoral livelihoods. Although there are numerous degradation studies on the effect of livestock grazing across the region, they are largely only published in Chinese, and most focus on single sites. Based on case studies from 100 sites, covering a mean annual precipitation gradient of 95–744 mm, we present a comprehensive, internationally accessible review on the impact of livestock grazing on vegetation and soils. We compared ungrazed or slightly grazed sites with moderately and heavily grazed sites by evaluating changes in two indicator groups: vegetation (plant species richness, vegetation cover, aboveground biomass, belowground biomass and root/shoot ratio) and soil (pH, bulk density, organic C, total N, total P and available P). Most indicators declined with intensified grazing, while soil pH, bulk density and belowground biomass increased. Available P showed no clear response. Variables within indicator groups were mostly linearly correlated at a given grazing intensity. Relative grazing effects on different indicators varied along specific abiotic gradients. Grazing responses of plant species richness, aboveground biomass, soil bulk density, total N and available P interacted with precipitation patterns, while grazing effects on belowground biomass were influenced by temperature. Elevation had impact on grazing responses of aboveground biomass and soil organic carbon. Complex grazing effects reflect both methodological inconsistency and ecological complexity. Further assessments should consider specific characteristics of different indicators in the context of the local environment.     

Intraspecific adoption and foster feeding of fledglings in the North Island robin

We sampled soils and vegetation within and outside two sheep and rabbit exclosures, fenced in 1979, on steep sunny and shady slopes at 770 m altitude on seasonally-dry pastoral steeplands. The vegetation of sunny aspects was characterised by higher floristic diversity, annual species, and low plant cover. Here the exotic grass Anthoxanthum odoratum dominated on grazed treatments, and the exotic forb Hieracium pilosella on ungrazed. Shady aspects supported fewer, and almost entirely perennial, species. Here Hieracium pilosella dominated grazed treatments, but co-dominated with the exotic forb H. praealtum and the native grass Festuca novae-zelandiae on ungrazed treatments. There was 43% more biomass in exclosures (P < 0.01). Most of the biomass difference (4285 kg/ha) was from greater root mass (2400 kg/ha). 1385 kg/ha of the difference was from herbage and the remainder (500 kg/ha) from litter. Exclosures had 50 to 100% more Ca, Mg, K and P in the biomass (P < 0.05), but the effect on soils was limited to significantly higher concentrations of total N (P < 0.05) and exchangeable Mg (P < 0.01) in 0-7.5 cm soils. We conclude that stopping grazing for 16 years on seasonally-dry steeplands results in greater plant cover, approximately double the biomass of standing vegetation, greater biomass in roots, and more biomass nutrients relative to grazed areas. However, it does not favour native species and has little effect on soil nutrients or soil carbon. Stopping grazing alone therefore cannot be regarded as a comprehensive short- or medium-term vegetation or soil rehabilitation option.     

Plant composition and diversity in a semi-natural Mediterranean island landscape: the importance of environmental factors

Abstract

Islands crucially contribute to the Mediterranean Basin’s high floristic diversity, which, however, is at risk facing climate and land-use changes. Besides the identification of highly diverse areas, the knowledge about factors favouring diversity is of great importance. We analysed plant species diversity and composition related to environmental factors over varied vegetation units on a former Italian prison island in the northwest of Sardinia. Due to a long history of land use with grazing and later abandonment the nowadays protected island features a semi-natural landscape and can serve as an example for strongly anthropogenic altered insular ecosystems. Floristic composition, soil properties, microclimate and ungulate abundance were assessed. Relationships of vegetation composition and diversity with abiotic variables were examined by Canonical Correspondence Analysis, which indicated the importance of air temperature, soil moisture, slope gradient and C/N ratio for floristic differentiation. Most important abiotic factors for plant species richness were relative air humidity and soil moisture, while floristic diversity was mainly determined by air temperature and pH. Furthermore, observation data pointed to an adverse influence of ungulate abundance for plant species diversity. Regarding nature conservation, grazing intensity thus must be critically taken into account, especially for sensitive vegetation units like the coastal garrigue.     

Effects of differential grazing on decomposition rate and nitrogen availability in a productive mountain grassland

Background and aims

Grazing may influence nutrient cycling in several ways. In productive mountain grasslands of central Argentina cattle grazing maintain a mosaic of different vegetation patches: lawns, grazed intensively and dominated by high quality palatable plants, and open and closed tussock grasslands dominated by less palatable species. We investigated if differences in the resources deposited on soil (litter and faeces) were associated with litter decomposition rates and soil nitrogen (N) availability across these vegetation patches.

Methods

We compared the three vegetation patches in terms of litter and faeces quality and decomposability, annual litterfall and faeces deposition rate. We determined decomposition rates of litter and faces in situ and decomposability of the same substrates in a common garden using “litter bags”. We determined soil N availability (with resin bags) in the vegetation patches. Also, we performed a common plant substrates decomposition experiment to assess the effect of soil environment on decomposition process. This technique provides important insights about the soil environmental controls of decomposition (i.e. the sum of soil physicochemical and biological properties, and microclimate), excluding the substrate quality.

Results

The litter quality and faeces deposition rate were higher in grazing lawns, but the total amounts of carbon (C) and nitrogen (N) deposited on soil were higher in tussock grasslands, due to higher litterfall in these patches. The in situ decomposition rates of litter and faeces, and of the two common plant substrates were not clearly related to either grazing pressure, litterfall or litter quality (C, N, P, lignin, cellulose or hemicellulose content). In situ litter decomposition rate and soil ammonium availability were correlated with the decomposition rates of both common plant substrates. This may suggest that difference in local soil environment among patch types is a stronger driver of decomposition rate than quality or quantity of the resource that enter the soil.

Conclusions

Our results show that, although high grazing pressure improves litter quality and increases faeces input, the reduction in biomass caused by herbivores greatly reduces C and N input for the litter decomposition pathway. We did not find an accelerated decomposition rate in grazing lawns as proposed by general models. Our results point to soil environment as a potential important control that could mask the effect of litter quality on field decomposition rates at local scale.     

Combined effects of environment and grazing on vegetation structure in Argentine granite grasslands

Abstract. Effects of grazing and environment on vegetation structure have been widely acknowledged, but few studies have related both factors. We made 57 floristic samples in a highly variable landscape of mountain grasslands in central Argentina; 26 sample were in fence‐lines with contrasting vegetation. For each sample, we recorded topographic and edaphic parameters, as well as grazing intensity indicators. Floristic gradients were analysed with DCA and relations with abiotic and grazing‐related variables were detected with DCCA. Floristic axis 1 was explained by edaphic parameters associated to topography, ranging from communities in well drained soils on upper topographic positions to hydromorphic vegetation in poorly drained soils on lower topographic positions. Species richness decreased as soil moisture increased. Floristic axis 2 was associated with present and long‐term grazing indicators, and reflected shifts in vegetation physiognomy and species evenness. Tall tussock grasslands, with low species evenness and evidences of low or null grazing intensity were located at one extreme. Tussocks were gradually replaced by short graminoids and forbs towards the centre of the gradient, as grazing increased, and evenness reached a maximum. In degraded sites with heavy long‐term grazing intensities, short perennial species were replaced by an annual species, and evenness decreased. The magnitude of changes in floristic composition produced by grazing decreased with increasing soil moisture, and vegetation‐environment relationships were stronger in moderate to highly grazed situations than in lightly or non grazed situations.     

Long-term vegetation dynamics mediated by herbivores,weather and fire in a Juniperus-Quercus savanna

Abstract. Long-term (45-yr) basal area dynamics of dominant graminoid species were analyzed across three grazing intensity treatments (heavily grazed, moderately grazed and ungrazed) at the Texas A&M University Agricultural Research Station on the Edwards Plateau, Texas. Grazing intensity was identified as the primary influence on long-term variations in species composition. Periodic weather events, including a severe drought (1951–1956), had little direct influence on composition dynamics. However, the drought interacted with grazing intensity in the heavily grazed treatment to exacerbate directional changes caused by grazing intensity. Species response to grazing was individualistic and noisy. Three response groups were identified. Taller, more productive mid-grasses were most abundant under moderate or no grazing. Short grasses were most abundant under heavy grazing. Intermediate species were most abundant under moderate grazing and opportunistic to weather patterns. Graminoid diversity increased with the removal or reduction of grazing intensity. The moderately and ungrazed treatments appeared most resistant to short-term weather fluctuations, while the heavily grazed treatment demonstrated significant resilience when grazing intensity was reduced after over 110 yr of overgrazing. Identification of a ‘climax’ state is difficult. Significant directional change, which took nearly 20 yr, appears to continue in the ungrazed treatment after 45 yr of succession. The observed, relatively linear patterns of perennial grass composition within the herbaceous patches of this savanna were generally explained by traditional Clementsian succession. However, when dynamics of the herbaceous community are combined with the woody component of this savanna, the frequency and intensity of fire becomes more important. Across the landscape, successional changes follow several pathways. When vegetation change is influenced by several factors, a multi-scale model is necessary to demonstrate interactions and feedbacks and accurately describe successional patterns. Absence of fires, with or without grazing, leads ultimately to a Juniperus/Quercus woodland with grazing intensity primarily influencing the fuel load and hence fire intensity.     

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.