Arid vegetation in disequilibrium with livestock grazing: Evidence from long‐term exclosures

In recent decades, the conventional equilibrium paradigm for explaining rangeland vegetation dynamics has been challenged. Proponents of an alternative non‐equilibrium paradigm argue that in variable rangeland environments, external climatic events are critical to vegetation dynamics and there is little opportunity for plant–herbivore interactions to reach equilibrium. Understanding which paradigm more effectively describes an ecosystem has important consequences for management. In particular, some authors have argued that a focus on reducing stocking rates in non‐equilibrium systems may be futile, and management should be opportunistic in response to unpredictable rainfall events. We measured herbaceous biomass and plant species richness and abundance at five 14‐year exclosures on Innamincka Regional Reserve. Four were situated in the dunefields land system, and one on the Cooper Creek floodplain. We did not detect any significant differences between grazed and ungrazed treatments in total species richness or abundance, life form richness or abundance, or herbaceous biomass. Only one species, Portulaca oleracea, showed differences in abundance between treatments at more than one site, but the direction of these differences was not consistent. These results suggest that the non‐equilibrium paradigm more accurately describes vegetation dynamics in the dunefields and floodplains of north‐eastern South Australia. It is possible that some species had been lost from the study area prior to the establishment of the exclosures, thereby precluding recovery with protection from grazing; however, a regional analysis of the flora reveals little evidence of this. We argue that the dominance of ephemeral species confers resilience by limiting the development of strong feedbacks between grazing intensity and vegetation dynamics. Current grazing practices seem consistent with the conservation of plant species diversity across the dunefields and floodplains. Future studies should focus on the impacts of cattle grazing on areas of the landscape dominated by palatable perennials, as well as the small number of rare and potentially grazing‐sensitive species identified.     

Restoration of native vegetation following exclosure establishment on communal grazing lands in Tigray,Ethiopia

Questions: Is plant species richness, diversity and above‐ground standing biomass enhanced after establishing exclosures on communal grazing lands? What factors influence the effectiveness of exclosures to restore degraded native vegetation in Tigray, Ethiopia? Location: Northern Ethiopia. Methods: We used a space‐for‐time substitution approach to detect changes in plant species richness, diversity and above‐ground standing biomass after conversion of communal grazing lands to exclosures. We selected replicated (n=3) 5‐, 10‐, 15‐ and 20‐year‐old exclosures and paired each exclosure with an adjacent communal grazing land to ensure that soil and terrain conditions were as similar as possible among each pair. Results: All exclosures displayed higher plant species richness, diversity and biomass than the communal grazing lands. Differences in plant species richness and biomass between an exclosure age and adjacent communal grazing land were higher in oldest than in youngest exclosures. In exclosures, much of the variability in plant species composition and biomass was explained by a combination of edaphic (total nitrogen, phosphorus, texture and soil pH) and site (precipitation and altitude) variables (R²=0.72–0.82). Edaphic and site variables also explained much of the variability in plant species composition in communal grazing lands (R²=0.76–0.82). Our study shows that all exclosures are at an early stage of succession. The increase in economically important indigenous shrub and tree species with exclosure age suggests that, with time, a valuable afromontane forest may develop. Conclusions: Establishment of exclosures on communal grazing lands is a viable option to restore degraded native vegetation. However, before expanding exclosures, the ecological consequences of additional exclosures should be investigated as further expansion of exclosures could increase grazing pressure on remaining grazing areas. Furthermore, consideration of edaphic and site variables will help optimize selection of areas for establishment of exclosures and enhance natural regeneration in exclosures in the future.     

Land rehabilitation and the conservation of birds in a degraded Afromontane landscape in northern Ethiopia

The few remaining Afromontane forest fragments in northern Ethiopia and the surrounding degraded, semiarid matrix form a habitat mosaic of varying suitability for forest birds. To evaluate the effect of recent land rehabilitation efforts on bird community composition and diversity, we studied bird species distributions in ten small forest fragments (0.40–20.95 ha), five grazing exclosures (10-year-old forest restoration areas without wood extraction and grazing livestock) and three grazed matrix sites during the rainy season (July–October 2004) using 277 one-hour species counts. Based on the distribution pattern of 146 bird species, sites were assigned to one of three bird communities (birds of moist forest, dry forest or degraded savanna), each occupying a well-defined position along an environmental gradient reflecting decreasing vegetation structure and density. All three communities were representative of the avifauna of Afrotropical Highland open forest and woodland with a high proportion of invasive and competitive generalist species (31%). Apart from these, exclosures shared more species with forest fragments (20%) than did the grazed matrix (5%), indicating local ecosystem recovery. By increasing habitat heterogeneity, exclosures have the potential to enhance landscape connectivity for forest birds and are, therefore, an effective instrument for conserving species in a fragmented landscape. However, 52 bird species (36%) occurred exclusively within forest patches and many forest birds that use exclosures are unlikely to maintain viable populations when forest fragments disappear, particularly as forest fragments may be a critical resource during the hot dry season. This highlights the high conservation value of small isolated forest fragments for less tolerant, forest-limited and/or biome-restricted species.     

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.     

Response of vertebrates to fenceline contrasts in grazing intensity in semi‐arid woodlands of eastern Australia

Abstract Changes in the abundance, species richness and assemblage composition of vertebrates due to grazing by domestic stock were investigated in the semi‐arid woodlands of eastern Australia. Analyses were based on the differences found at 10 fenceline contrast sites. Two species of amphibians, 22 species of reptiles and two species of small mammal were captured in pit traps during the surveys. Kangaroos (red and eastern grey), sheep, goats and 66 species of birds were recorded along line transects. Analyses revealed that abundance of diurnal reptiles and species richness of diurnal reptiles and birds were significantly lower on heavily grazed sites than they were on lightly grazed sites. At a local scale, the gecko, Gehyra variegata, was more abundant where grazing was heavier, while Diplodactylus conspicillatus, Diplodactylus steindachneri and Rhynchoedura ornata responded to variables indirectly related to grazing intensity (kangaroo density, sheep and goat dung mass and sheep density, respectively). Birds more commonly sighted on lightly grazed areas than heavily grazed areas were the apostlebird, brown treecreeper, crested bellbird, grey butcherbird, hooded robin, jacky winter, little woodswallow, Australian magpie‐lark, mulga parrot, splendid wren, white‐browed treecreeper and yellow‐rumped thornbill. Birds more commonly sighted on heavily grazed areas than on lightly grazed areas were the Australian raven and chestnut‐crowned babbler. Most variation in species composition between sites was due to spatial separation and no regional‐level indicator species of grazing were evident. A combination of direct grazing‐related changes (e.g. loss of ground cover) and indirect effects of the pastoral industry (e.g. introduction of artificial sources of water) lead to changes in fauna at different scales of analysis across regions.     

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.     

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.     

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.     

Land use history and site location are more important for grassland species richness than local soil properties

Lately there has been a shift in Sweden from grazing species‐rich semi‐natural grasslands towards grazing ex‐arable fields in the modern agricultural landscape. Grazing ex‐arable fields contain a fraction of the plant species richness confined to semi‐natural grasslands. Still, they have been suggested as potential target sites for re‐creation of semi‐natural grasslands. We asked to what extent does fine‐scale variation in soil conditions, management history and site location effect local plant diversity in grazed ex‐arable fields. We examined local soil conditions such as texture, pH, organic carbon, nitrogen (N) and extractable phosphate (P) and effects on plant richness in ten pairs of grazed ex‐fields and neighbouring semi‐natural grasslands in different rural landscapes. Each grassland pair where in the same paddock. A multivariate test showed that site location and land use history explained more of differences in species richness than local soil property variables. Plant species richness was positively associated to grazed ex‐fields with low pH, low N and P levels. Sites with high plant richness in semi‐natural grasslands also had more species in the adjacent grazed ex‐fields, compared to sites neighbouring less species‐rich semi‐natural grasslands. Although both soil properties and species richness were different in grazed ex‐fields compared to semi‐natural grassland, the site location within a landscape, and vicinity to species‐rich grasslands, can override effects of soil properties. In conclusion, if properly located, ex‐arable fields may be an important habitat to maintain plant diversity at larger spatio‐temporal scales and should considered as potential sites for grassland restoration.     

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.     

Top‐down control of rare species abundances by native ungulates in a grassland restoration

Ecological restorations are predicted to increase in species diversity over time until they reach reference levels. However, chronosequence studies in grasslands often show that diversity peaks after the first few years and then declines over time as grasses become more dominant. We addressed whether bison grazing and seed additions could prevent this decline in diversity. Exclosures that prevented bison grazing were compared with grazed plots over 4 years, and seed additions were conducted inside and outside exclosures to test for seed and microsite limitations. A previous study conducted 4‐months post seeding found that local species richness was primarily seed limited, but that grazing could sometimes increase seedling emergence. Here, we tested whether increased seedling emergence led to longer‐term increases in the species diversity of the plant community. We found that the seed addition effect grew smaller and the grazing effect grew stronger over time, and that seed additions affected the abundance of added species only when plots were grazed. Grazed plots had higher species diversity and lower biomass and litter buildup compared to non‐grazed plots. Our results suggest that moderate grazing by bison or management that mimics grazing can maintain diversity in grass‐dominated situations. Our results also emphasize the need to follow seed additions over several years to assess correctly whether seed limitation exists.     

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.     

Passive recovery of small vertebrates following livestock removal in the Australian rangelands

In arid environments, grazing by exotic herbivores, including domestic livestock, can greatly influence native, small vertebrate assemblages. Whether the removal of livestock facilitates passive recovery of these assemblages depends on habitat condition and the species present. We explore changes in small mammal and reptile species richness, abundance, and composition in a degraded chenopod shrubland dominated by Acacia victoriae ssp. and open Acacia aneura (mulga) woodland destocked in 1976 and 1984, respectively. Data were obtained between 1997 and 2007, from two grazed and two ungrazed sites in each community. Species richness increased at a faster rate in ungrazed open A. aneura woodland, but did not differ significantly between ungrazed and grazed degraded chenopod shrubland. Subsequent analyses at a finer‐scale detected disparate responses in richness and abundance for microhabitat. At this scale, a greater number of species‐specific responses were also detected, including increased abundance of generalist species and decreased abundance of species requiring low cover. These results reiterate the potential for species‐specific responses to livestock that are more apparent in particular microhabitats. Furthermore, this investigation provides evidence for the gradual passive recovery of small mammal and reptile assemblages in both communities, which is facilitated by the removal of livestock in open A. aneura woodland in fair condition, but not degraded chenopod shrubland in poor condition.     

The effect of clearing on plant composition in mulga (Acacia aneura) dry forest,Australia

Despite high deforestation rates in dry forests there are few assessments of the impacts of this transformation on biodiversity. This study addresses the impact of clearing on plant species composition and diversity in mulga (Acacia aneura) forests of Queensland, Australia using data collected from either side of 58 clearing boundaries, incorporating data on site and management conditions. The physical and climatic conditions of the study sites were narrowly constrained but had greater influence on plant composition and richness than clearing per se. In general, the abundance of perennial forbs and shrubs was higher in cleared forest compared with adjacent intact forest and there was no significant difference in abundance of other plant life forms. Perennial forb richness diminished with the age of clearings, particularly where acid sub‐soils had been exposed and shrubs proliferated. Where deforestation in the mulga dry forests is not accompanied by the establishment of exotic pasture it has a relatively benign effect on plant diversity, mulga itself remains in high density as small trees and the integrity of the forest could be maintained with restoration.     

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

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

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

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

Foraging ecology of roan antelope: key resources during critical periods

The use of landscape zones and grass species by roan antelope, a species threatened with local extirpation within South Africa's Kruger National Park, were investigated. Plant‐based observations of grazing were made within a 300 ha enclosure in the roan range, where 40 roan antelope were confined at high density in the absence of other grazers. The study spanned the dry seasons of two years, one with average rainfall and one with low rainfall. We recorded changes in the extent of grazing of different grass species, height differences between grazed and ungrazed tillers and intensity of cropping per tuft. In the average year, the grazing pressure in the bottomland grassland was twice that in the upland savanna, with two tall grass species bearing the brunt of the grazing through the dry season. Two highly palatable upland grasses were also extensively grazed by the mid dry season. In the dry year, the extent of grazing in the upland exceeded that in the bottomland, and several upland grass species little used the previous year became heavily grazed. Roan antelope appeared to be separated ecologically from more common grazers by their selective use of tall grasses growing in the drainage line grassland during the critical dry season months. However, their grazing expansion into the upland savanna during the dry year potentially brought them into competition with these grazers. Nevertheless, their population performance did not suffer despite the high‐density conditions. Heightened predation pressure following an influx of these grazers, rather than resource limitation, appeared to be primarily responsible for the drastic decline of this species in the park.     

Effects of grazing exclusion on species composition in high-altitude grasslands of the Central Alps

Our study examines the effects of grazing exclusion on low-productive subalpine and alpine grasslands of the Central Alps (UNESCO Biosphere Park Gurgler Kamm, Obergurgl, Austria). A long-term exclusion experiment was established in 2000 in the subalpine, the lower, and the upper alpine zone. With exception of the subalpine zone, domestic herbivores have been grazing during the whole growing season. In grazed and exclosure plots species frequencies were recorded for 7 years. We analysed exclosure effects on species number, community composition, life forms, and functional groups.Species richness did not decrease significantly within the exclosures, but changes in species composition occurred in each zone, although some were transitory in nature. The dynamic trends of the plots were significantly explained by the ‘treatment×year’ effect along the whole altitudinal gradient, but the effects decreased considerably with altitude. In the subalpine and upper alpine exclosures, stress-tolerators, species of low or no nutritive value, and mosses showed a decreasing trend, whereas tall grasses (subalpine exclosures), competitors, and species with high or medium nutritive values (lower alpine exclosures) tended to increase.Overall, our 7-year study revealed that several functional groups reacted to grazing, according to our main expectations. We suggest that these effects will intensify in the long term.     

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

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

Influences of chronic and current season grazing by collared pikas on above-ground biomass and species richness in subarctic alpine meadows

We studied an alpine herbivory gradient established by collared pikas, a small central place foraging lagomorph, to examine the effects of multiple grazing levels on above-ground live biomass (AGB) and species richness (SR) in alpine tundra. The effects of within-season (four sampling periods), multi-season (across three summers) and longer-term dynamics (inferred from spatial location of vegetation with respect to pika haypiles) were examined. Along the grazing gradient, we found support for and against hypotheses that propose biphasic, increasing, or decreasing responses to herbivory, both in terms of AGB and SR. Our results suggest that plant–herbivore predictability is still weak. To further examine the impact of herbivory, we experimentally removed pikas using mesh exclosures placed at increasing distance from the edge of talus occupied by pikas. AGB after the second consecutive year of herbivore exclusion increased by 125% compared to control plots in highly grazed areas adjacent to talus (<1 m). In more lightly grazed sites at distances 1–6 m from talus, AGB increased by more than 40% after pikas were removed. No differences were observed in the ungrazed sites >6 m from talus. AGB was highest in meadow patches previously grazed by pikas compared to those with little grazing history, but this response was only observed after two seasons following release from herbivory. Grazed sites at distances of 1–6 m had the highest SR. These results indicate that multi-year measurements of growth are particularly relevant in ecosystems dominated by long-lived perennials in regions where productivity is low. Infrequent herbivore vacancies may provide local short-term release from pika grazing, thereby contributing to the persistence of productive, highly palatable vegetation.