Linking sheep density and grazing frequency to persistence of herb species in an alpine environment

Large herbivores are important drivers in ecosystems worldwide. Changes in herbivore densities are predicted to especially affect herbs that are strongly preferred by herbivores. The persistence of herbs could be challenged by enhanced grazing, but also grazing cessation may affect persistence, especially for prostrate herbs, which might be out-competed. To test how different herbivore densities (high, low, and no sheep) affect grazing frequency and plant responses (plant height, flowering frequency, and plant density) at the herb community and species level, we conducted a fully replicated, landscape-scale experiment in an alpine environment. We found that none of the herb species changed their densities after 5 years with experimental changes in grazing pressure. Sheep density affected grazing and flowering frequency at the herb community level. Eight herb species were more grazed at high sheep density as compared to enclosures with no sheep. Herb height decreased at high sheep density as compared to no sheep for the two species with the highest grazing frequency. Increased height and flowering frequency were found for small herbs at high sheep density. Our experiment clearly shows that herbs do not constitute a homogeneous functional group and that, in particular, tall and small herbs are affected in contrasting ways but all species (n = 15) tolerated changes in grazing regimes as densities were maintained at both enhanced grazing and grazing cessation.     

Yaks and sheep trigger different changes in the grasshopper assemblages of the Qilian Mountains via differentially altering plant assemblages

1. The Qilian Mountains represent one of the key livestock‐raising grasslands in China. The two main herbivore species raised in this area – yaks and sheep – are of critical economical value. Grasshoppers compete with these animals for available nutrients, creating multifaceted relationships between livestock, grasshoppers and plants. A clear understanding of such relationships is lacking and is urgently needed to guide conservation efforts. 2. This study aims to document the effects of yak and sheep grazing on grasshopper assemblages and to elucidate the underlying mechanisms of such effects. 3. It is shown here that yaks and sheep impact grasshopper assemblages differently. Grasshopper assemblages exhibited lower density, biodiversity, richness, and evenness of distribution in yak‐grazed pastures than in grazing‐free grasslands. Sheep‐grazed pastures exhibited a dramatically divergent picture, with elevated density, biodiversity and richness, and a slightly decreased evenness of distribution. Grasshoppers were generally larger in grazed pastures than in grazing‐free grasslands, especially in yak‐grazed plots. 4. The present study suggests that differences between yak and sheep pastures in plant assemblage structure and plant traits are probably the underlying forces driving the differences in grasshopper assemblage structure and grasshopper traits, respectively. 5. The study shows that the grasshopper habitat indicator species differ between yak and sheep pastures, raising the possibility that such indicators can be used to monitor grassland usage and degradation in the Qilian Mountains. 6. These results provide novel insights into the dynamic interactions of common domesticated herbivore species, grasshoppers and plants in Qilian Mountains, which augment current knowledge and may ultimately lead to better conservation practices.     

Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Vegetation changes following sheep grazing in abandoned mountain meadows

Abstract. Sheep grazing was investigated as an alternative to traditional management of meadows in the Krkono?e Mts. Until the second World War these meadows were mown in mid‐summer and grazed by cattle for the rest of the season. Subsequent abandonment of the meadows has resulted in decreasing species richness. Degradation phases of the former communities have been replacing the original species‐rich vegetation. Significant changes were apparent six years after the introduction of sheep grazing. In grazed plots the proportion of dominant herbs (Polygonum bistorta and Hypericum maculatum) decreased and grasses (Deschampsia cespitosa, Festuca rubra, Agrostis capillaris, Anthoxanthum alpinum) increased. The increase in grasses was positively correlated with an increase in several herbs. The proportion of some herbs increased despite being selectively grazed (Adenostyles alliariae, Melandrium rubrum, Veratrum lobelianum). Any losses caused by grazing of mature plants were probably compensated by successful seedling establishment. Cessation of grazing resulted in significant changes in vegetation within three years. The cover of nitrophilous tall herbs and grasses (e.g. Rumex alpestris, Holcus mollis, Deschampsia cespitosa, Geranium sylvaticum) increased in the abandoned plots. In the plots grazed for nine years cover of species‐rich mountain meadow species increased (e.g. fine‐leaved grasses, Campanula bohemica, Potentilla aurea, Viola lutea, Silene vulgaris). The main conservation risk is the expansion of a competitive species with low palatability, Deschampsia cespitosa. This species can be suppressed by a combination of grazing and mowing. In order for grazing to be effective, the number of sheep should be proportional to meadow production. This may be difficult to maintain as production is variable and is impossible to predict at the beginning of a growing season. A large part of the biomass may thus remain intact in some years. Negative effects of grazing may be, at least partly, eliminated by a combination of cutting and grazing.     

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.     

Low intensity, mixed livestock grazing improves the breeding abundance of a common insectivorous passerine

Livestock grazing is a major driver of ecosystem change and has been associated with significant declines in various bird species in Britain and worldwide. However, there is little experimental evidence to show how grazing affects bird populations. We manipulated livestock densities in a replicated field experiment and found that mixed sheep and cattle grazing, at low intensity, improved the breeding abundance of a common upland passerine, the meadow pipit Anthus pratensis, after two years. Plots stocked with sheep alone (at high or low density) or not stocked at all held fewer pipit territories. Despite a year-on-year decline in pairs of meadow pipits in intensively grazed plots, we found no effect of sheep number on breeding abundance. Our results support the hypothesis that mixed species of herbivores generate greater heterogeneity in vegetation structure, which modifies prey availability, resulting in a greater abundance of birds. The results of our study should inform the management of grassland areas and enhance the abundance of some bird species, particularly in areas that have seen significant shifts from mixed livestock grazing to grazing dominated by single species of animals.     

Spatial distributions of multiple plant species affect herbivore foraging selectivity

Spatial distribution of food resources is an important factor determining herbivore foraging. Previous studies have demonstrated that clumped distribution of preferred species increases its consumption by herbivores in single‐ or two‐species systems. However, the potential impact of distribution pattern of less preferred species on foraging was ignored. In natural grasslands with high species diversity and complexity, the spatial distribution of preferred species impacts on herbivore foraging may be strongly correlated with the distribution of less preferred species. Our aims were to determine the effect of distribution of both preferred and other plant species on herbivore foraging under conditions close to a native, multi‐species foraging environment, and conceptualize the relationships between spatial distribution of food resources and herbivore consumption. We hypothesized that random distribution of non‐preferred species reduces herbivore consumption of preferred species because the dispersion of less preferred species likely disturbs herbivore foraging. We conducted an experiment using three species with five combinations of clumped and random distribution patterns. Three species Lathyrus quinquenervius, Phragmites australis and Leymus chinensis, were of high, intermediate and low preferences by sheep, respectively. Results showed that distribution of low preferred species, but not that of high preferred one, affected the consumption of preferred species. Sheep obtained higher consumption of high preferred species when low preferred species followed a clumped distribution than a random distribution. Distance between aggregations of high and low preferred species did not affect sheep foraging. It was concluded that the effects of spatial distribution of preferred species on its consumption are dependent on herbivore foraging strategy, and sheep can consume more preferred species when there is a consistent spatial pattern between preferred species and the entire food resource, and that the random dispersion of low preferred species in grassland may reduce herbivore consumption of high preferred species, thus minimizing selective grazing.     

Effects of large herbivores on grassland arthropod diversity

Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio‐temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top‐down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.     

Relationships between the effects of insect herbivory and sheep grazing on seasonal changes in an early successional plant community

Summary The effects of spring grazing by sheep and of natural levels of insect herbivory were studied in 1985 on a limestone field abandoned from arable land for four years. A split-plot design was adopted in which paddocks, arranged in Latin squares, were either left ungrazed or heavily grazed by sheep for ten days in April. Within each paddock plots were either sprayed regularly with Malathion-60 or untreated.Natural levels of insect herbivory, compared to the reduced levels in insecticide-treated plots, had effects of similar magnitude to those from the short burst of spring grazing. Many attributes of the grazed/insecticide-treated sward were either increased or decreased by a factor of two within a season. Both types of herbivore caused changes in the direction of plant succession as well as in its rate. Effects on early successional species were large and similar when caused by either type of herbivore. Effects on later successional species were often smaller, but also showed differences in the action of the two herbivore types, as did effects on sward height, species richness and total cover. The effects of sheep and insect herbivory were not always additive or in the same direction.The results suggest that manipulations of both mammal and insect herbivores may be powerful tools for directing changes in plant community composition.     

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

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

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

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

Stronger effect of gastropods than rodents on seedling establishment,irrespective of exotic or native plant species origin

Experimental evidence about how generalist consumers affect exotic plant invasions is equivocal, but most tests have been limited to few plant species, single herbivore guilds, and single locations. Using a seed‐addition experiment, we studied effects of gastropods and rodents on recruitment success of 37 exotic and 37 native plant species affiliated to three different functional groups (i.e. grasses, legumes and non‐legume herbs). We replicated our seed addition x herbivore exclusion experiment at multiple grassland sites, located within a few km of each other in two regions, coastal central California (USA) and southern Saxony–Anhalt (Germany). The two study regions differed in climate, land‐use, invasion history and species pools which allowed us to disentangle general from context‐specific effects. In both regions, herbivory by gastropods had a stronger impact on the proportion of recruited seedlings and the proportion of recruited species than rodent herbivory, but this effect was much more pronounced in California than in Germany. Especially, seedling recruitment of non‐legume herbs and legumes suffered from gastropod herbivory. Contrastingly, the effect of rodents was negative at the German sites and positive at the Californian sites, likely driven by context‐specific differences in the rodent assemblages. Across both study regions, exotics had higher seedling recruitment than natives, indicating that higher recruitment success constitutes an inherent feature of exotic species. After two years, more exotic than native species established at grassland sites in California while the opposite was true for the German grassland sites. Consistently across regions, native and exotic species did, however, not differ in their response to herbivory, suggesting that generalist consumers suppress recruitment and colonization of plant species irrespective of their origin. Our results demonstrate the importance of a multi‐species, multi‐site approach to separate general responses of exotic and native plants to generalist herbivory from local, regional or species‐specific peculiarities.     

Grazing responses in herbs in relation to herbivore selectivity and plant traits in an alpine ecosystem

Herbivores shape plant communities through selective foraging. However, both herbivore selectivity and the plant’s ability to tolerate or resist herbivory may depend on the density of herbivores. In an alpine ecosystem with a long history of grazing, plants are expected to respond to both enhanced and reduced grazing pressures, and the interaction between plant traits and changes in species abundance are expected to differ between the two types of alteration of grazing regime. To understand the mechanisms behind species response, we investigated the relationship between sheep selectivity (measured in situ), plant traits and experimentally derived measures of change in species abundance as a response to the enhancement (from low to high density) or cessation (from low to zero density) of sheep grazing pressure over a six-year time period for 22 abundant herb species in an alpine habitat in south Norway. Sheep selected large, late-flowering herbs with a low leaf C/N ratio. Species that increased in abundance in response to enhanced grazing pressure were generally small and had high root/shoot ratios, thus exhibiting traits that reflect both resistance (through avoidance) and tolerance (through regrowth capacity) strategies. The abundance of selected species remained stable during the study period, and also under the enhanced grazing pressure treatment. There was, however, a tendency for selected species to respond positively to cessation of grazing, although overall responses to cessation of grazing were much less pronounced than responses to enhanced grazing. Avoidance through short stature (probably associated with increased light availability through the removal of tall competitors) as well as a certain amount of regrowth capacity appear to be the main mechanisms behind a positive response to enhanced grazing pressure in this study. The plant trait perspective clearly improves our insight into the mechanisms behind observed changes in species abundance when the disturbance regime is altered. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Population‐level manipulations of field vole densities induce subsequent changes in plant quality but no impacts on vole demography

Grazing‐induced changes in plant quality have been suggested to drive the negative delayed density dependence exhibited by many herbivore species, but little field evidence exists to support this hypothesis. We tested a key premise of the hypothesis that reciprocal feedback between vole grazing pressure and the induction of anti‐herbivore silicon defenses in grasses drives observed population cycles in a large‐scale field experiment in northern England. We repeatedly reduced population densities of field voles (Microtus agrestis) on replicated 1‐ha grassland plots at Kielder Forest, northern England, over a period of 1 year. Subsequently, we tested for the impact of past density on vole life history traits in spring, and whether these effects were driven by induced silicon defenses in the voles’ major over‐winter food, the grass Deschampsia caespitosa. After several months of density manipulation, leaf silicon concentrations diverged and averaged 22% lower on sites where vole density had been reduced, but this difference did not persist beyond the period of the density manipulations. There were no significant effects of our density manipulations on vole body mass, spring population growth rate, or mean date for the onset of spring reproduction the following year. These findings show that grazing by field voles does induce increased silicon defenses in grasses at a landscape scale. However, at the vole densities encountered, levels of plant damage appear to be below those needed to induce changes in silicon levels large and persistent enough to affect vole performance, confirming the threshold effects we have previously observed in laboratory‐based studies. Our findings do not support the plant quality hypothesis for observed vole population cycles in northern England, at least over the range of vole densities that now prevail here.     

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

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

Temporal variation in density dependent body growth of a large herbivore

Temporal variation both due to density dependent and density independent processes affect performance and vital rates in large herbivores. Annual fluctuations in climate affect foraging conditions and thus body growth of large herbivores during the short growing season in alpine habitats. Also, high animal densities on summer ranges may increase competition for food and reduce body mass gain. Yet, little is known about interactive effects of density and climate on alpine summer ranges, and the time scales these processes operate on. In this fully replicated landscape‐scale experiment, we kept domestic sheep at high and low densities over nine grazing seasons in an alpine habitat, and tested the relative role of density and annual variation for lamb body mass gain during summer and whether effects of density and annual variation interacted. We found that lambs at high density gained less mass over the summer season than lambs at low density. At short time scales the density effect interacted with annual fluctuations in body growth. We documented a long‐term temporal trend in body mass, consistent with the hypothesis that grazing effects affect habitat differentially at high and low density over years. At high density lamb autumn body mass declined during the first three grazing seasons and then stabilized, whereas body mass slightly increased over years at low density. This long‐term trend suggests accumulative density dependent effects from a biomass or quality reduction, and hence delayed food competition at high density and possibly facilitation at low density. Our experiment provides new insight into how density dependent effects on performance of a large herbivore depend on temporal scale of observation.     

Herbivore-induced coexistence of competing plant species

We study a series of spatially implicit lottery models in which two competing plant species, with and without defensive traits, are grazed by a herbivore in a homogeneous habitat. One species (palatable) has no defensive traits, while the other (defended) has defensive traits but suffers reduced reproduction as the result of an assumed trade-off. Not surprisingly, coexistence of these plants cannot occur when the herbivore density is very low (the palatable plant always wins) or very high (the defended plant wins). At intermediate densities, however, herbivory can mediate plant coexistence, even in a homogeneous environment. If the herbivore eats several plants per bite, and its forage-selection depends on the average palatability of the plants it eats, then palatable species in the immediate neighbourhood of defended plants may be more likely to persist (associational resistance) even at higher grazing pressure. If the herbivore shows a positive numerical response to the average palatability of the habitat as a whole, then both plant populations are stabilized and coexistence is promoted, because both species obtain a minority advantage through the negative feedback caused by herbivory. If the herbivore exhibits both of these traits, the system may have at most two non-trivial equilibria, one of which is stable and the other unstable. This means that coexistence in such a system is vulnerable to large fluctuations in herbivore density and identity, and this has implications for conservation in systems where large herbivores are managed to promote plant diversity.     

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.