Vegetation change following removal of keystone herbivores from desert grasslands in New Mexico

Abstract. Responses of plant communities to mammalian herbivores vary widely, due to variation in plant species composition, herbivore densities, forage preferences, soils, and climate. In this study, we evaluated vegetation changes on 30 sites within and adjacent to the Sevilleta National Wildlife Refuge (SNWR) in central New Mexico, USA, over a 20‐yr period following removal of the major herbivores (livestock and prairie dogs) in 1972–1975. The study sites were established in 1976, and were resampled in 1986 and 1996 using line transect methods. At the landscape scale, repeated measures ANOVA of percentage cover measurements showed no significant overall net changes in total perennial plant basal cover, either with or without herbivores present; however, there was an overall increase in annual forbs and plant litter from 1976 to 1996. At the site scale, significant changes in species composition and dominance were observed both through time and across the SNWR boundary. Site histories varied widely, with sites dominated by Bouteloua eriopoda being the most dynamic and sites dominated by Scleropogon brevifolius being the most persistent. Species‐specific changes also were observed across multiple sites: B. eriopoda cover increased while Gutierrezia sarothrae greatly decreased. The non‐uniform, multi‐directional changes of the sites' vegetation acted to prevent detection of overall changes in perennial vegetation at the landscape level. Some sites displayed significant changes after removal of herbivores, while others appeared to respond primarily to climate dynamics. Certain species that were not preferred by livestock or prairie dogs, showed overall declines during drought periods, while other preferred species exhibited widespread increases during wetter periods regardless of herbivore presence. Therefore, the vegetation dynamics cannot be attributed solely to removal of herbivores, and in some cases can be explained by short‐ and long‐term fluctuations in climate. These results emphasize the variety of responses of sites with differences in vegetation to mammalian herbivores under otherwise similar climatic conditions, and illustrate the value of site‐ and landscape‐scale approaches to understanding the impacts of plant‐herbivore interactions.     

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.     

Large and small herbivores have strong effects on tundra vegetation in Scandinavia and Alaska

1. Large and small mammalian herbivores are present in most vegetated areas in the Arctic and often have large impacts on plant community composition and ecosystem functioning. The relative importance of different herbivores and especially how their specific impact on the vegetation varies across the Arctic is however poorly understood.
2. Here, we investigate how large and small herbivores influence vegetation density and plant community composition in four arctic vegetation types in Scandinavia and Alaska. We used a unique set of exclosures, excluding only large (reindeer and muskoxen) or all mammalian herbivores (also voles and lemmings) for at least 20 years.
3. We found that mammalian herbivores in general decreased leaf area index, NDVI, and abundance of vascular plants in all four locations, even though the strength of the effect and which herbivore type caused these effects differed across locations. In three locations, herbivore presence caused contrasting plant communities, but not in the location with lowest productivity. Large herbivores had a negative effect on plant height, whereas small mammalian herbivores increased species diversity by decreasing dominance of the initially dominating plant species. Above‐ or belowground disturbances caused by herbivores were found to play an important role in shaping the vegetation in all locations.
4. Synthesis: Based on these results, we conclude that both small and large mammalian herbivores influence vegetation in Scandinavia and Alaska in a similar way, some of which can mitigate effects of climate change. We also see important differences across locations, but these depend rather on local herbivore and plant community composition than large biogeographical differences among continents.

     

Interactions between spatially separated herbivores indirectly alter plant diversity

Above‐ and belowground herbivores promote plant diversity when selectively feeding on dominant plant species, but little is known about their combined effects. Using a model system, we show that neutral effects of an aboveground herbivore and positive effects of a belowground herbivore on plant diversity became profoundly negative when adding these herbivores in combination. The non‐additive effects were explained by differences in plant preference between the aboveground‐ and the belowground herbivores and their consequences for indirect interactions among plant species. Simultaneous exposure to aboveground‐ and belowground herbivores led to plant communities being dominated by a few highly abundant species. As above‐ and belowground invertebrate herbivores generally differ in their mobility and local distribution patterns, our results strongly suggest that aboveground–belowground interactions contribute to local spatial heterogeneity of diversity patterns within plant communities.     

Impact of herbivores on nitrogen cycling: contrasting effects of small and large species

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.     

Impact of different‐sized herbivores on recruitment opportunities for subordinate herbs in grasslands

Abstract. Potential effects of herbivores on plant species diversity depend on herbivore size, species and density. In this study we examine the effect of different‐sized herbivores (cattle and rabbits) on recruitment of subordinate herbs in grasslands. We show that in a grazed floodplain, grassland plant species richness is mainly determined by the presence of many species of subordinate herbs. These herbs experience high colonization and extinction rates. We conclude that the creation of colonization opportunities for subordinate herbs plays a crucial role in maintaining plant species richness in productive grasslands. We found that cattle disperse large amounts of seeds via their dung, over ten times more than rabbits. Rabbits create more and on average larger bare soil patches than cattle. In a field experiment artificial disturbances improved germination success tremendously for four tested herb species. We found that bare soil is the best regeneration site, while cattle dung gave a too strong nutrient stimulus, resulting in tall vegetation and therefore light limitation. These results can be confirmed with results from field monitoring plots where plant species richness was positively related to the occurrence of bare soil patches. Therefore both large and small herbivores have a major impact on dispersal and colonization, but for different reasons. Cattle are identified as most important for seed dispersal whereas rabbits have a main effect as creators of disturbances. These results emphasize the importance of distinguishing between herbivore species in assessing their (potential) effects.     

Post‐fire pickings: Large herbivores alter understory vegetation communities in a coastal eucalypt forest

Fire and herbivores alter vegetation structure and function. Future fire activity is predicted to increase, and quantifying changes in vegetation communities arising from post‐fire herbivory is needed to better manage natural environments. We investigated the effects of post‐fire herbivory on understory plant communities in a coastal eucalypt forest in southeastern Australia. We quantified herbivore activity, understory plant diversity, and dominant plant morphology following a wildfire in 2017 using two sizes of exclosures. Statistical analysis incorporated the effect of exclusion treatments, time since fire, and the effect of a previous prescribed burn. Exclusion treatments altered herbivore activity, but time since fire did not. Herbivory reduced plant species richness, diversity, and evenness and promoted the dominance of the most abundant plants within the understory. Increasing time since fire reduced community diversity and evenness and influenced morphological changes to the dominant understory plant species, increasing size and dead material while decreasing abundance. We found the legacy effects of a previous prescribed burn had no effect on herbivores or vegetation within our study. Foraging by large herbivores resulted in a depauperate vegetation community. As post‐fire herbivory can alter vegetation communities, we postulate that management burning practices may exacerbate herbivore impacts. Future fire management strategies to minimize herbivore‐mediated alterations to understory vegetation could include aggregating management burns into larger fire sizes or linking fire management with herbivore management. Restricting herbivore access following fire (planned or otherwise) can encourage a more diverse and species‐rich understory plant community. Future research should aim to determine how vegetation change from post‐fire herbivory contributes to future fire risk.     

Community- Weighted Mean Plant Traits Predict Small Scale Distribution of Insect Root Herbivore Abundance

Small scale distribution of insect root herbivores may promote plant species diversity by creating patches of different herbivore pressure. However, determinants of small scale distribution of insect root herbivores, and impact of land use intensity on their small scale distribution are largely unknown. We sampled insect root herbivores and measured vegetation parameters and soil water content along transects in grasslands of different management intensity in three regions in Germany. We calculated community-weighted mean plant traits to test whether the functional plant community composition determines the small scale distribution of insect root herbivores. To analyze spatial patterns in plant species and trait composition and insect root herbivore abundance we computed Mantel correlograms. Insect root herbivores mainly comprised click beetle (Coleoptera, Elateridae) larvae (43%) in the investigated grasslands. Total insect root herbivore numbers were positively related to community-weighted mean traits indicating high plant growth rates and biomass (specific leaf area, reproductive- and vegetative plant height), and negatively related to plant traits indicating poor tissue quality (leaf C/N ratio). Generalist Elaterid larvae, when analyzed independently, were also positively related to high plant growth rates and furthermore to root dry mass, but were not related to tissue quality. Insect root herbivore numbers were not related to plant cover, plant species richness and soil water content. Plant species composition and to a lesser extent plant trait composition displayed spatial autocorrelation, which was not influenced by land use intensity. Insect root herbivore abundance was not spatially autocorrelated. We conclude that in semi-natural grasslands with a high share of generalist insect root herbivores, insect root herbivores affiliate with large, fast growing plants, presumably because of availability of high quantities of food. Affiliation of insect root herbivores with large, fast growing plants may counteract dominance of those species, thus promoting plant diversity.     

Effects of spontaneous field margin vegetation on the regulation of herbivores in two winter crops

Non-crop vegetation of field margins provides resources for natural enemies of crop herbivores. However, it is still not well known whether this resource provisioning effect is strong enough to improve herbivore regulation within crop fields and which plant species and functional groups favour this ecosystem service. A better understanding of the interactions between field margin vegetation and herbivore regulation is crucial to evaluate management strategies and to design suppressive plant mixtures. We surveyed 64 wheat and oilseed rape fields of Western France for two years (16 fields per year and crop) in order (1) to identify plant diversity or group effects on herbivore regulation within crop fields and (2) to identify species within plant groups that improve regulation. Herbivores, herbivore damage and natural enemies were monitored on crop plants at a distance of 5 and 50 m from the field margin. At the same time, the cover and phenological stage of all vascular plants were estimated in the adjacent field margin. The study demonstrated a positive relationship between the cover of entomophilous plant species that were flowering at the survey date and response variables related to herbivore regulation. Plant species richness and the cover of plant species taxonomically close to crop plants had a small influence on herbivores and natural enemies in wheat whereas related wild Brassicaceae increased herbivory and decreased herbivore regulation in oilseed rape. Within the entomophilous flowering plants, several species were significantly related to a better herbivore regulation in univariate analyses. Multivariate ordination techniques allowed the identification of plant species influencing several response variables of herbivore regulation at the same time. Our study demonstrated the importance of entomophilous species that flowered at peak infestation of crop herbivores. Spontaneous field margins rich in flowering entomophilous species provide an important ecosystem service without expensive sowing of seed mixtures.     

The more the merrier? Multi-species grazing of small herbivores mediates plant community impacts

Small herbivores play keystone functional roles in grassland ecosystems. Recognising the combined effects where herbivores co-exist is important for guiding grassland restoration and biodiversity conservation. On the Tibetan Plateau, both plateau pikas and Himalayan marmots are regarded as pests by Tibetan people and local government, but little is known about their combined effects. We conducted a field study to determine the combined effects of pikas and marmots on plant biodiversity and vegetation structure. Plateau pikas alone consistently reduced the plant height and diversity and increased the vegetation cover of physically unpalatable plants. However, the co-existence of marmots with pikas decreased the vegetation cover of physically unpalatable plants, while increasing the cover of palatable plants and plant diversity, ultimately changing the impact of pikas alone and modulating aspects of the plant community. These results illustrate that increasing the abundance and richness of small herbivores in grassland ecosystems may promote plant diversity and benefit vegetation restoration rather than aggravate the degradation of grasslands.     

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.     

放牧对草地昆虫多样性的影响研究进展

在草地生态系统中,大型草食动物放牧是重要的管理方式之一,对草地生物多样性起着关键的驱动作用。昆虫是草地生态系统中生物多样性的重要组成成分,对生态系统的食物网结构以及其功能与稳定性起着关键作用。已有研究结果表明,大型草食动物与昆虫存在密切联系,放牧对草地昆虫多样性或有正向、或负向、或无明显作用,这依赖于放牧管理方式、昆虫类群以及草地类型。放牧必然通过直接(采食、践踏或粪尿)或间接(植物群落组成或植被结构)作用对昆虫多样性产生显著的影响。当前,关于大型草食动物放牧对草地昆虫多样性影响研究较多,但是,从研究系统性、深入性和延续性来说还存在一定问题。本文在综述国内外对放牧对草地昆虫多样性的影响研究基础上,提出了今后的研究方向,对于理解放牧管理的草地昆虫多样性变化规律,以及为积极探索维持草地昆虫多样性的长期有效的科学管理措施提供理论指导。     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Native herbivores and environmental heterogeneity as mediators of an exotic grass invasion

Given that many exotic plant species throughout the world are having large ecological and economic effects, it is vital to understand the forces that mediate their success in novel landscapes. Both native herbivores and recipient ecosystems can have substantial effects on the performance of exotic plant species, and may interact with each other or vary in their effects over time. Unfortunately, few studies have evaluated the importance of these kinds of context‐dependent effects. Here, we use a 17‐year‐old exclosure experiment stratified across a coastal grassland in northern California to address the relative importance of a reintroduced mammalian herbivore, tule elk (Cervus canadensis nannodes), and environmental heterogeneity in mediating the growth, abundance, and recruitment of a problematic grass invader, Holcus lanatus. We found that elk reduced Holcus abundance, aboveground biomass, percent cover, frequency, and seedling recruitment, but that these effects often varied among habitat types, with effects being greater in open grasslands than shrub‐dominated grasslands. The performance of Holcus populations also varied significantly among habitat types, with the invader usually having the greatest success in Baccharis‐dominated grasslands. Our results suggest that environmental heterogeneity had much greater influence on Holcus success than elk, and that these effects were due largely to soil pH and moisture. The negative effects of elk on Holcus appeared after 4 years and did not intensify after an additional 13 years. Furthermore, despite their negative effects, these prominent herbivores did not prevent the spread of Holcus into previously uninvaded areas. Our research highlights the importance of assessing the individual and interactive effects of native herbivores and environmental heterogeneity on the success of invasive, exotic plant species. It emphasizes the reality that the negative effects of herbivores on exotic plant species will often vary across heterogeneous landscapes and may be insufficient to prevent the expansion of these invaders.     

Contrasting effects of large herbivore grazing on smaller herbivores

Assemblages of large herbivores may compete for food or facilitate one another. However, small vertebrate herbivore species co-occurring with large herbivores may be affected by large herbivore grazing through changes in plant species composition, nutrient content and vegetation structure. These changes can be either positive or negative for the smaller herbivores, but this may depend on the species of small herbivores. We experimentally tested the impact of cattle grazing on habitat choice of European rabbits (Oryctolagus cuniculus) and common voles (Microtus arvalis). We excluded cattle for 7 years and measured changes in vegetation parameters, and the response of rabbits and voles. Rabbits were facilitated by cattle, whereas voles strongly preferred vegetation without cattle. The facilitation effect was stronger at low rabbit densities. Vegetation biomass and nitrogen concentration were not affected by cattle grazing, but vegetation height increased significantly where cattle were excluded. Plant species composition also changed following cattle exclusion; however, the main food plants of rabbits and voles remained abundant in each grazing treatment. We conclude that the response of both rabbits and voles predominantly reflect the differences in vegetation height in the presence and absence of cattle, but in a contrasting fashion. The difference in response between rabbits and voles may result from reduced perceived predation risk, which is lowest in high vegetation for voles, but in short vegetation for rabbits, which depend on their burrows for safety. The use of large herbivores in grassland conservation management can thus have a contrasting effect on different species of small herbivores.     

Plant community response to loss of large herbivores: comparing consequences in a South African and a North American grassland

Loss of biodiversity poses one of the greatest threats to natural ecosystems throughout the world. However, a comprehensive understanding of the impacts of species losses from upper trophic levels is still emerging. Here we compare the impacts of large mammalian herbivore species loss on grassland plant community structure and composition in a South African and North American grassland. Herbaceous plant communities were surveyed at sites without large mammalian herbivores present and at sites with a single species of herbivore present in both locations, and additionally at one site in South Africa with multiple herbivore species. At both the North American and South African locations, plant communities on sites with a single herbivore species were more diverse and species rich than on sites with no herbivores. At the multi-herbivore site in South Africa, plant diversity and richness were comparable to that of the single herbivore site early in the growing season and to the no herbivore site late in the growing season. Analyses of plant community composition, however, indicated strong differences between the multi-herbivore site and the single and no herbivore sites, which were more similar to each other. In moderate to high-productivity ecosystems with one or a few species of large herbivores, loss of herbivores can cause a significant decrease in plant diversity and richness, and can have pronounced impacts on grassland plant community composition. In ecosystems with higher herbivore richness, species loss may also significantly alter plant community structure and composition, although standard metrics of community structure may obscure these differences.     

Responses of insect herbivores and herbivory to habitat fragmentation: a hierarchical meta‐analysis

Loss and fragmentation of natural habitats can lead to alterations of plant–animal interactions and ecosystems functioning. Insect herbivory, an important antagonistic interaction is expected to be influenced by habitat fragmentation through direct negative effects on herbivore community richness and indirect positive effects due to losses of natural enemies. Plant community changes with habitat fragmentation added to the indirect effects but with little predictable impact. Here, we evaluated habitat fragmentation effects on both herbivory and herbivore diversity, using novel hierarchical meta‐analyses. Across 89 studies, we found a negative effect of habitat fragmentation on abundance and species richness of herbivores, but only a non‐significant trend on herbivory. Reduced area and increased isolation of remaining fragments yielded the strongest effect on abundance and species richness, while specialist herbivores were the most vulnerable to habitat fragmentation. These fragmentation effects were more pronounced in studies with large spatial extent. The strong reduction in herbivore diversity, but not herbivory, indicates how important common generalist species can be in maintaining herbivory as a major ecosystem process.     

Scale as modifier in vegetation diversity experiments: effects on herbivores and predators

We conducted a meta-analysis to elucidate the role of experimental scale in field trials exploring the role of habitat diversifications on herbivore and predator abundance. Literature from a period of 18 years from 6 journals yielded a total of 25 predator and 41 herbivore abundance observations in experimental treatments with higher plant diversity compared to a control. These were divided into three size bins based on reported plot size. A clear pattern emerged for herbivore abundance whereby diversification experiments performed in small plots yielded a large negative effect on herbivores, intermediate sized plots showed an intermediate effect, and the largest plots exhibited a negligible effect. Predators were more abundant in medium sized diversified plots than in small ones, but, again, no effect could be discerned in the largest plots. These results suggest that past literature reviews and meta-analyses illustrating general trends of herbivore declines associated with increased vegetation diversity may be misleading. In particular, the effects of diversification appear to vary widely as a function of the spatial scale of vegetation plots. We speculate that effects of vegetation diversification may be enhanced when insects can move uninhibited among control and treatment plots in small scale experiments. In smaller plots herbivores may aggregate in the control plots that present a more concentrated resource, and generalist predators move to the more diverse habitat. At larger scales the ability of insects to "choose" between simple and diverse plots is diminished .     

Browsing herbivores improve the state and functioning of savannas: A model assessment of alternative land‐use strategies

Changing climatic conditions and unsustainable land use are major threats to savannas worldwide. Historically, many African savannas were used intensively for livestock grazing, which contributed to widespread patterns of bush encroachment across savanna systems. To reverse bush encroachment, it has been proposed to change the cattle‐dominated land use to one dominated by comparatively specialized browsers and usually native herbivores. However, the consequences for ecosystem properties and processes remain largely unclear. We used the ecohydrological, spatially explicit model EcoHyD to assess the impacts of two contrasting, herbivore land‐use strategies on a Namibian savanna: grazer‐ versus browser‐dominated herbivore communities. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture, and water use by plants. Our results showed that plant types that are less palatable to herbivores were best adapted to grazing or browsing animals in all simulated densities. Also, plant types that had a competitive advantage under limited water availability were among the dominant ones irrespective of land‐use scenario. Overall, the results were in line with our expectations: under high grazer densities, we found heavy bush encroachment and the loss of the perennial grass matrix. Importantly, regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in browsing scenarios. Browsing herbivores increased grass cover, and the higher total cover in turn improved water uptake by plants overall. We concluded that, in contrast to grazing‐dominated land‐use strategies, land‐use strategies dominated by browsing herbivores, even at high herbivore densities, sustain diverse vegetation communities with high cover of perennial grasses, resulting in lower erosion risk and bolstering ecosystem services.     

The potential role of large herbivores in nature conservation and extensive land use in Europe

The concept of ecosystem restoration is gaining momentum in western Europe. This is necessary because in most managed nature reserves one or more of the following processes, which are analogous to those that have led to the dramatic loss of biological diversity in Europe, are still operating: continuing nutrient output, continuing high level of disturbance and fixing the system in some successional stage. This is partly because most management activities have been derived from, or copy, former agricultural practices. The study of natural ecosystems has revealed the key role large herbivores have in maintaining structural diversity in the vegetation and so biological diversity. Because of this they have been used as tools in achieving a variety of conservation goals. Here, various effects large herbivores can have on plant species composition, structural diversity of the vegetation and fauna are briefly reviewed. Attention is given to pasture-woodlands in southern Europe, which often have a relatively high biological diversity and share some key features with natural ecosystems: very low nutrient input, extensive grazing with large herbivores and the presence of natural tree cover. In a number of European countries attempts are being made to restore normal functioning multi-(herbivore) species ecosystems.