The interrelationship between productivity,plant species richness and livestock diet: a question of scale?

Question: What relationship exists between productivity, plant species richness and livestock diet? Are the results dependent on scale? Location: A sheep‐grazed Koelerio‐Corynephoretea sandy habitat of the northern upper Rhine (Germany) as a low productivity model system. Methods: The investigation was carried out for three years at a fine scale (2 m²) and for two years at a broad scale (79 m²). Productivity was measured by means of weighed above‐ground phytomass for fine scale and colour‐infrared (CIR) aerial photographs of the same system for fine and broad scales. For both scales, total numbers of vascular plant species and numbers of endangered vascular plant species were extracted from current vegetation relevés. Additionally, we obtained data on livestock diet (grazed phytomass, crude protein content). Results: Statistical analyses show an influence of the year on all variables; relationships between variables are not significant in every year. Species richness and number of endangered species are negatively related to productivity at fine scale while crude protein content and grazed phytomass are positively related to productivity. At the broad scale the diversity‐productivity relationship shows a ‘hump’ with highest species numbers in middle pioneer stages; numbers of endangered species are highest in all pioneer stages. Conclusions: We found a strong impact of scale and year on the diversity‐productivity relationship. It is inappropriate to analyse only small plots (2 m²), and it is necessary to study different years. This vegetation complex is dependent on grazing impact; thus there is an inversely proportional relationship between nature conservation value (high diversity) and livestock nutrition.     

Long‐term effects of exclusion of grazing stock on degraded herbaceous plant communities in a riparian Eucalyptus camaldulensis forest in south‐eastern Australia

Abstract Stock grazing has degraded many riparian ecosystems around the world. However, the potential for ecosystem recovery following the removal of grazing stock is poorly known. We developed a conceptual model to predict the responses of native and exotic herbaceous plants to grazing exclusion, based on site productivity and the degree of initial vegetation degradation. The effects of excluding grazing stock on richness, cover and composition of herbaceous plants were examined over 12 years in the degraded understorey of a riparian forest in Gulpa Island State Forest in south‐eastern Australia. We predicted that grazing exclusion would lead to limited changes in vegetation cover, richness and composition, owing to presumed low site productivity and the high degree of understorey degradation. Results showed that the cover, richness and composition of native and exotic species varied significantly among years. Over all plots, regions and years, total cover was slightly but significantly lower in grazed than in ungrazed plots (43.4% vs. 50.8%). While the cover of native plants increased over time in both treatments, the rate of increase was slightly greater in ungrazed plots. Grazing exclusion had no effect on the richness of native or exotic species, but had a significant but minor impact on plant composition, with different common species (mostly exotics) being promoted or diminished in ungrazed plots. The composition of grazed and ungrazed areas did not become more different over time. Overall, the effects that could be attributed to grazing exclusion were relatively minor and transient. Results are consistent with predictions based on site productivity and initial degradation, and should not be extrapolated to other more productive, or less degraded, riparian systems.     

Impacts of heavy grazing on plant species richness: A comparison across rangeland biomes of South Africa

The net effect of heavy grazing and land degradation on plant diversity and richness is insufficiently understood for incorporation in national biodiversity assessments. A study was undertaken to determine the effects of heavy grazing primarily on richness of vascular plant species across the arid and semi-arid rangeland biomes of South Africa. Major grazing contrasts were systematically identified for sampling in rangelands of Succulent Karoo, Nama-Karoo, Thicket, Grassland, Kalahari dune savanna and Mopane savanna. The related parameters of species diversity, evenness and turnover were also examined and analysed at the whole site level. The study represents a new site-level comparison of earlier individual studies that also necessitated recalculation and standardization of original data, where appropriate. Impacts of heavy grazing on plant species richness were found to vary from negligible or slightly positive to distinctly negative, depending on site. The sharp reductions in richness may have been associated with special secondary conditions that can occur in arid areas. Species diversity did not track species richness well and was often dominated by species evenness patterns. Moderate to substantial turnover of species occurred, even with negligible change in species richness. Species turnover was largely associated with replacement of species, except on one site where turnover was more evenly split between its replacement and nestedness components. Heavy grazing altered species composition on all study sites, usually with reduced grazing quality and favouring annual plants. Surprisingly few of the replacement species on most of the study areas were alien or exotic. Remarkably, the magnitude of change in species richness across the limited rainfall gradient of the study often greatly exceeded changes associated with the heavy grazing levels at each site. Use of the significant non-linear relationship found between loss of plant canopy cover through grazing and the relative decline in species richness needs further exploration.     

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.     

Biodiversity of semi‐arid Mediterranean grasslands: Impact of grazing and afforestation

Question: What is the impact of grazing and/or afforestation on grassland diversity, species composition and cover parameters? Location: Semi‐arid Mediterranean grasslands of Jordan. Methods: Vegetation, litter, bare soil and rock cover were compared among four management types – free grazing and protected from grazing with three levels of tree cover. Species composition, plant cover, species richness and evenness were used to evaluate differences in vegetation among management types. Species composition differences among management types were also investigated. Results: Semi‐arid Mediterranean grasslands harbour appreciable levels of plant biodiversity. Grazing did not affect plant diversity, indicating the high resilience against and adaptation to grazing; however,grazing affected species composition and cover parameters. Afforestation seems to protect soil through higher litter cover but its impact on plant biodiversity was negative and markedly affected species composition. Conclusions: Neither protection from grazing or massive afforestation alone are sufficient for conserving biodiversity in this system. A management model is suggested where the landscape should be maintained as a mosaic of four management types: complete protection from grazing, grazing rotation, planting sparse trees in eroded areas and revegetating degraded areas using native, herbaceous and grazing tolerant species.     

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.     

The Influence of Historical Land Use and Water Availability on Grassland Restoration

The ecological role of historical land use has rarely been explored in the context of grassland restoration. We conducted a 4‐year field experiment in a steppe and an old field in Inner Mongolia in northern China to examine the influence of historical land use and water availability on ecosystem restoration. Species richness, evenness, and plant cover were higher in the steppe than in the old field. The steppe was more temporally stable compared with the old field in terms of species richness, evenness, plant density, and cover. Water addition increased peak aboveground biomass, belowground net primary productivity, species richness, plant density, and cover in both the steppe and the old field. Water addition also enhanced the stability of ecosystems and the restoration of grassland. Our findings suggested that historical land use determines community structure and influences the process of grassland restoration. Converting grasslands to farmland in semiarid areas can cause the long‐term loss of biodiversity and instability of ecosystem with consequent impacts on ecosystem services. The amendment of limited resources is an effective practice to increase the success of ecosystem restoration.     

Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

Testing the limits of resistance: a 19‐year study of Mediterranean grassland response to grazing regimes

A synthesis of a long‐term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north‐eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land‐use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long‐term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.     

Recovery of understorey vegetation after release from a long history of sheep grazing in a herb‐rich woodland

The effects of stock grazing on native grassy ecosystems in temperate southern Australia are well documented. However, less is known about the potential of ecosystems to recover after a long history of stock grazing and, in particular, whether the removal of stock will have positive, negative or neutral impacts on biodiversity. We examined the response of understorey vegetation to the removal of sheep grazing in a herb‐rich Eucalyptus camaldulensis (red gum) woodland in western Victoria. Using a space‐for‐time chronosequence, woodlands were stratified into groups based on their time‐since‐grazing removal; these were long‐ungrazed (>20 years), intermediate‐time‐since‐grazing (9–14 years), recently ungrazed (5 years) and continuously grazed. We found significantly higher species density in long‐ungrazed sites relative to sites with a more recent grazing history. No differences were found in species density between continuously grazed sites and those ungrazed in the previous 14 years. Species composition differed with time‐since‐grazing removal and indicator species analysis detected several native species (including tall native geophytes and herbs) associated with long‐ungrazed sites that were absent or in low abundance in the more recently grazed sites. Seven of the eight species significantly associated with continuously grazed sites were exotic. Removal of sheep grazing in red gum woodlands can have positive benefits for understorey diversity but it is likely that recovery of key indicators such as native species will be slow.     

Effects of livestock exclusion on in‐stream habitat and benthic invertebrate assemblages in montane streams

1. Stream and riparian ecosystems in arid montane areas, like the interior western United States, are often just narrow mesic strands, but support diverse and productive habitats. Meadows along many such streams have long been used for rangeland grazing, and, while impacts to riparian areas are relatively well known, the effect of livestock grazing on aquatic life in streams has received less attention. 2. Attempts to link grazing impacts to disturbance have been hindered by the lack of spatial and temporal replication. In this study, we compared channel features and benthic macroinvertebrate communities (i) between 16 stream reaches on two grazed allotments and between 22 reaches on two allotments where livestock had been completely removed for 4 years, (ii) before and after the 4‐year grazing respite at a subset of eight sites and (iii) inside and outside of small‐scale fenced grazing exclosures (eight pairings; 10+ year exclosures) in the meadows of the Golden Trout Wilderness, California (U.S.A.). 3. We evaluated grazing disturbance at the reach scale in terms of the effects of livestock trampling on per cent bank erosion and found that macroinvertebrate richness metrics were negatively correlated with bank erosion, while the percentage of tolerant taxa increased. 4. All macroinvertebrate richness metrics were significantly lower in grazed areas. Bank angle, temperature, fine sediment cover and erosion were higher in grazed areas, while riparian cover was lower. Regression models identified riparian cover, in‐stream substratum, bank conditions and bankfull width‐to‐depth ratios as the most important for explaining variability in macroinvertebrate richness metrics. 5. Small‐scale grazing exclosures showed no improvements for in‐stream communities and only moderate positive effects on riparian vegetation. In contrast, metrics of macroinvertebrate richness increased significantly after a 4‐year period of no grazing. 6. The success of grazing removal reported here suggests that short‐term removal of livestock at the larger, allotment meadow spatial scale is more effective than long‐term, but small‐scale, local riparian area fencing, and yields promising results in achieving stream channel, riparian and aquatic biological recovery.     

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.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

Intraspecific variation overrides origin effects in impacts of litter-derived secondary compounds on larval amphibians

Vertebrate herbivores can be key determinants of grassland plant species richness, although the magnitude of their effects can largely depend on ecosystem and herbivore characteristics. It has been demonstrated that the combined effect of primary productivity and body size is critical when assessing the impact of herbivores on plant richness of perennial-dominated grasslands; however, the interaction of site productivity and herbivore size as determinants of plant richness in annual-dominated pastures remains unknown. We experimentally partitioned primary productivity and herbivore body size (sheep and wild rabbits) to study the effect of herbivores on the plant species richness of a Mediterranean semiarid annual plant community in central Spain over six years. We also analyzed the effect of grazing and productivity on the evenness and species composition of the plant community, and green cover, litter, and plant height. We found that plant richness was higher where the large herbivore was present at high-productivity sites but barely changed at low productivity. The small herbivore did not affect species richness at either productivity site despite its large effects on species composition. We propose that adaptations to resource scarcity and herbivory prevented plant richness changes at low-productivity sites, whereas litter accumulation in the absence of herbivores decreased plant richness at high productivity. Our results are consistent with predictions arising from a long history of grazing and highlight the importance of both large and small herbivores to the maintenance of plant diversity of Mediterranean annual-dominated pastures.     

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.     

Interactive effects of fire and grazing on structure and diversity of Mediterranean grasslands

Abstract. The separate and combined effects of fire and cattle grazing on structure and diversity of productive Mediterranean grasslands in northern Israel were examined within a set of climatically and edaphically similar sites. Cover and height of green and dry plants in winter, and species richness and diversity in spring, were measured in paired transects on both sides of cattle fences, and on both sides of boundaries of both incidental and experimentally lit fires. Early in the first growing season after a fire, plant cover as well as height of green plants were reduced, compared to unburnt grassland. These structural effects of fire were similar to the effects of grazing, but they were greater in ungrazed than in grazed grasslands, indicating a fire-grazing interaction. The effects of fire were considerably attenuated in the second growing season after the fire. Species richness and diversity tended to be higher in grazed than in adjacent ungrazed grasslands. Richness consistently increased after a fire only in grazed grasslands with a strong perennial component. In ungrazed grasslands, and in predominantly annual grasslands, fires reduced species richness and diversity at least as often as they increased it. Fire and grazing should be regarded as two agents with distinct and interactive effects on the community, rather than as two alternative mechanisms of a general disturbance factor.     

Effects of grazing on patch structure in a semi‐arid two‐phase vegetation mosaic

Abstract. Question: What are the grazing effects in the spatial organization and the internal structure of high and low cover patches from a two‐phase vegetation mosaic? Location: Patagonian steppe, Argentina. Methods: We mapped vegetation under three different grazing conditions: ungrazed, lightly grazed and heavily grazed. We analysed the spatial patterns of the dominant life forms. Also, in each patch type, we determined density, species composition, richness, diversity, size structure and dead biomass of grasses under different grazing conditions. Results: The vegetation was spatially organized in a two‐phase mosaic. High cover patches resulted from the association of grasses and shrubs and low cover patches were represented by scattered tussock grasses on bare ground. This spatial organization was not affected by grazing, but heavy grazing changed the grass species involved in high cover patches and reduced the density and cover of grasses in both patch types. Species richness and diversity in high cover patches decreased under grazing conditions, whereas in low cover patches it remained unchanged. Also, the decrease of palatable grasses was steeper in high cover patches than in low cover patches under grazing conditions. Conclusions: We suggest that although grazing promotes or inhibits particular species, it does not modify the mosaic structure of Patagonian steppe. The fact that the mosaic remained unchanged after 100 years of grazing suggests that grazing does not compromize population processes involved in maintaining patch structure, including seed dispersal, establishment or biotic interactions among life forms.     

Rapid short-tussock grassland decline with and without grazing, Marlborough, New Zealand

Species abundance, species richness, and ground cover were measured over 10 years on nine paired grazed and exclosure plots in short-tussock grassland in the early stages of invasion by Hieracium species. With and without grazing, H. pilosella and H. caespitosum increased markedly and H. lepidulum increased locally. In contrast, 50% of all other common species and species groups, and total, native, and exotic species richness declined significantly. Exclusion increased or had no effect on rates of increase in Hieracium species and rates of decline in short tussocks, and did not reduce rates of decline in other species. Exclusion had no effect on decline in native species richness, but mainly accelerated declines in total and exotic richness. Declines in 13 key vegetation variables were significantly predicted by increase in Hieracium abundance, suggesting competitive exclusion. With or without grazing, Hieracium species will become more dominant and other species will continue to decline. The effects of large herbivores on plant species diversity can often be predicted from site productivity. Our results indicate the need also to account for species origin, spatial scale, time, and exotic invasion.