Effects of large herbivores on wood pasture dynamics in a European wetland system

Whether self-regulating large herbivores play a key role in the development of wood-pasture landscapes remains a crucial unanswered question for both ecological theory and nature conservation. We describe and analyse how a ‘partly self-regulating’ population of cattle, horses and red deer affected the development of the woody vegetation in the Oostvaardersplassen nature reserve (Netherlands). Using aerial photographs from 1980 to 2011, we analysed the development of shrubs and trees. Before the large herbivores were introduced in the Oostvaardersplassen in 1983, the woody vegetation increased and vegetation type significantly affected the number of establishments. Cover of woody species increased further from 1983 to 1996, not only by canopy expansion but also by new establishments. After 1996, cover of the woody vegetation decreased from 30% to <1% in 2011 and no new establishments were seen on the photographs. Survival of Sambucus nigra and Salix spp. increased with increasing distance to grassland, which is the preferred foraging habitat of the herbivores. These results support the hypothesis of Associational Palatability. In addition, our results show that the relative decline in cover of S. nigra and Salix spp. over a certain period was negatively correlated with the cover of S. nigra in the beginning of this period, presenting some evidence for the Associational Resistance and Aggregational Resistance hypothesis. Our research shows aspects necessary for the woodland–grassland cycle, such as a strong decline of woody vegetation at high numbers of large herbivores and regeneration of shrubs and trees at low densities. Thorny shrubs, which are important for the cycle, have not yet established in the grasslands. It seems that a temporary decline in herbivore numbers is necessary to create a window of opportunity for the establishment of these woody species.     

Native herbivore exerts contrasting effects on fire regime and vegetation structure

Although native herbivores can alter fire regimes by consuming herbaceous vegetation that serves as fine fuel and, less commonly, accumulating fuel as nest material and other structures, simultaneous considerations of contrasting effects of herbivores on fire have scarcely been addressed. We proposed that a colonial rodent, vizcacha (Lagostomus maximus), reduces and increases fire intensity at different stages in its population cycle in the semiarid scrub of Argentina. Specifically, we hypothesized that, when colonies are active, vizcachas create natural fire-breaks through intense grazing, generating over time patches of large unburned shrubs in grazed zones. In contrast, when colonies are abandoned, recovery of fine fuels and previous accumulation of coarse wood on colonies during territorial displays increases fire intensity, creating patches of high shrub mortality. To test these hypotheses, we estimated stem age of the dominant shrub (Larrea divaricata) and measured aboveground biomass in zones actively grazed by vizcachas and in ungrazed zones, and compared densities of live and dead shrubs on abandoned colonies and adjacent zones following fire. In active colonies, age and biomass of shrubs were much greater in grazed than ungrazed zones. In abandoned colonies that had been burnt, density of dead, burned shrubs was higher and density of live shrubs was lower than in adjacent zones. These results support our hypotheses and reveal a new interaction between native herbivores and fire, in which herbivores augment fire intensity by gathering fuel. Our findings indicate that, through opposing effects on fire, native herbivores enhance the heterogeneity of vegetation in woody-dominated ecosystems.     

Cattle and Weedy Shrubs as Restoration Tools of Tropical Montane Rainforest

Over the last 150 years, a large proportion of forests in Latin America have been converted to pastures. When these pastures are abandoned, grasses may slow re‐establishment of woody species and limit forest regeneration. In this study, we explored the use of cattle in facilitating the establishment of woody vegetation in Colombian montane pastures, dominated by the African grasses Pennisetum clandestinum (Kikuyo) and Melinis minutiflora (Yaraguá). First, we described woody and herbaceous vegetation in grazed and non‐grazed pastures. Second, we tested the effect of grazing and seed addition on the establishment and growth of woody species. We also determined if the effect of grazing was different in P. clandestinum and M. minutiflora pastures. We found that low stocking density of cattle greatly increased density, number of branches per individual (a measure of “shrubiness”), and basal area of woody species, but also reduced woody plant species richness and diversity. In the grazed area, the shrubs Baccharis latifolia (Chilca) and Salvia sp. (Salvia) were the most abundant. The combined effect of grazing and shading from the shrubs reduced herbaceous vegetation by 52 to 92%. In the grazing/seed addition experiment, grazing increased establishment of woody seedlings, particularly of the shrub Verbesina arborea (camargo), but the largest effect was seed addition. Where grasses are an important barrier to regeneration, grazing can facilitate the establishment of shrubs that create a microhabitat more suitable for the establishment of montane forest tree species.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Response of galling invertebrates on Salix lanata to reindeer herbivory

Browsing and defoliation often increase the densities of insect herbivores on woody plants. Densities of herbivorous invertebrates were estimated in a long-term grazing manipulation experiment. More then 30-yr-old fences allow us to compare densities of invertebrate herbivores on Salix lanata in areas heavily grazed and areas lightly grazed by reindeer. The number of gall-forming insects ( Pontania glabrifons) and gall-forming mites were higher on the heavily grazed shrubs than on lightly grazed shrubs. In contrast to most short-term studies, the heavily grazed S. lanata had shorter current annual shoots. No difference in leaf size, leaf nitrogen content, or C:N ratio between grazing intensities were detected. However, the enhanced natural δ15N value indicates that heavily grazed shrubs get a higher proportion of their N directly from reindeer faeces. Leaf weight per unit area and relative fluctuating asymmetry of leaf shape increased in heavily grazed S. lanata . Enhanced relative fluctuating asymmetry might indicate higher susceptibility to herbivores. Long-term grazing seems to increase the density of invertebrate herbivory in the same way as short-term grazing, even if the plant responses differ substantially.     

Effects of browsing in relation to vegetation cover on common yew (<Emphasis Type="Italic">Taxus baccata</Emphasis> L.) recruitment in Mediterranean environments

Common yew (Taxus baccata L.) stands are recognized as prioritary habitats for biodiversity conservation within the European Union. The effects of browsing on the regeneration capacity and spatial dispersal of T. baccata recruits at the European southern limit of the species in the Mediterranean Basin have been herein studied. The efficacy of T. baccata recruitment has been evaluated at six localities in the Northern Sardinia mountains, which have similar altitude, climate, soil, and vegetation but have different types of uses (three were grazed by livestock and three were not). At each site, five habitats have been identified for T. baccata seed dispersal: reproductive female T. baccata canopy, reproductive female Ilex aquifolium canopy, non-fleshy-fruited tree canopy, fleshy-fruited shrubs, and open areas. The density of seedlings was found to be greater under fleshy-fruited trees (reproductive female T. baccata and I. aquifolium) than under shrubs, whereas the sapling density was higher in shrubby habitats, especially at grazed sites due to the mechanical protection afforded by the spiny shrubs against herbivores. Land use (LU) has been found to be the most important factor in determining the spatial distribution of seedlings and saplings in relation to forest habitats. Although browsers had an ephemeral but positive effect on seed germination through their trampling and the resultant scarification, this process eventually became ineffective as was shown by the occurrence of the lowest density of saplings in those habitats where the density of seedlings was the highest. The ultimate and most important effect of browsing was the sharp decrease in the density of saplings, and their almost complete extinction, in non-shrubby habitats. This study highlights the result that, in Mediterranean ecosystems, browsing constitutes the main negative factor on T. baccata seedling-sapling transition and furthermore confirms the necessity to preserve shrubby patches in the vicinity of reproductive female T. baccata and I. aquifolium to permit the regeneration of T. baccata in the presence of livestock. Moreover, at ungrazed sites, T. baccata is able to colonize non-shrubby shady habitats. The application of different management strategies to ungrazed and grazed sites should therefore be the main direction in the management and preservation of T. baccata stands in the Mediterranean region.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Variation in utilization of young leaves by a swallowtail butterfly across a deer density gradient

Phytophagous insects can be affected by plant trait-mediated indirect effects of large herbivores, but little is known regarding how these effects change in response to different densities of large herbivores. To assess the response of an insect to plant qualitative change, the response of a woody vine (Aristolochia kaempferi) to browsing by sika deer (Cervus nippon) and utilization of young leaves by a swallowtail butterfly (Byasa alcinous) were investigated across a deer density gradient. Natural and simulated deer browsing stimulated the regrowth of A. kaempferi and improved nutritional and physical quality of leaves. Young leaves were frequently observed in areas with high deer densities. The proportion of young leaves among the leaves selected for oviposition was higher than their proportion of the total number of leaves. In areas with low deer densities, the utilization of young leaves by B. alcinous increased linearly with deer density, whereas in areas with high deer densities, the utilization of young leaves was around 90%.     

Year-round cattle and horse grazing supports the restoration of abandoned,dry sandy grassland and heathland communities by supressing Calamagrostis epigejos and enhancing species richness

The spread of competitive grasses, changes in species composition and vegetation structure are direct consequences of grassland and heathland abandonment. As an alternative to more costly management measures such as traditional pastoralism, year-round low-intensity grazing with large herbivores is increasingly used to restore and maintain semi-open habitats. However, the suitability of this grazing regime has not yet been investigated for long-abandoned, highly degraded but nutrient-poor sandy grassland and heathland communities. In particular, it is unclear if year-round grazing is suitable for preventing the further spread of highly competitive grasses such as Calamagrostis epigejos while simultaneously maintaining or improving characteristic species richness and vegetation structure. Hence, we conducted a comprehensive field study on two spatial scales (plot-level: 25 m², macroplot-level: 1 ha) to analyse the impacts of year-round low-intensity cattle and horse grazing on the development of the highly competitive grass Calamagrostis epigejos, as well as the vegetation structure and plant species richness of long-abandoned but nutrient-poor dry sandy grassland and heathland communities, their mosaics and Calamagrostis stands within an 800 ha heathland between 2008 and 2015. Finally, we assessed the local conservation status of the habitat types after seven years of grazing in comparison to long-abandoned sites.Grazing successfully reduced the coverage of Calamagrostis epigejos, whereby Calamagrostis stands developed towards species-rich sandy grasslands after seven years of grazing. In addition, the quality of the vegetation structure was improved by enhancing the proportion of bare soil, while litter and grass cover, litter thickness and height of the field layer as well as the coverage of ruderal indicators were significantly reduced on grazed sites in comparison to ungrazed sites in 2015. Moreover, we found an overall positive grazing effect on species richness: Total species number, number of target species as well as subordinated target species significantly increased within the vegetation types over time.Thus, year-round low-intensity cattle and horse grazing is a suitable management tool for restoring, maintaining and even improving long-abandoned, nutrient-poor sandy grassland and heathland communities, and thus to enhance the local conservation status of the habitat types. However, if there is a high initial cover of woody species (e.g. shrubs, tree rejuvenation), then an extensive shrub and tree clearance will be necessary, with manual shrub cutting being crucial to reduce the heavy regrowth of the woody species. In addition, a one-time mowing should be implemented in highly degraded heaths to facilitate the vegetative rejuvenation of degenerate stands of Calluna vulgaris, thus improving its attractiveness for the grazing animals.     

Exotic deer diminish post-fire resilience of native shrub communities on Santa Catalina Island, southern California

Browsing by exotic mule deer on Santa Catalina Island (SCI) off the coast of southern California may diminish the post-fire resilience of native shrublands. To assess this, deer exclosures were established following a wildfire to monitor post-fire recovery of three dominant, native shrub species (Heteromeles arbutifolia, Rhus integrifolia, and Rhamnus pirifolia). Post-fire resprout growth, mortality, and tissue water status as well as pre- and post-fire shrub density and cover were measured inside and outside of deer exclosures. We found that deer browsing significantly limited post-fire resprout growth and led to increased mortality of resprouting H. arbutifolia shrubs (88 % mortality outside compared to 11 % inside exclosures). Post-fire resprouts maintained favorable water status during the study despite drought conditions, indicating that water stress was not a proximate cause of resprout mortality. Deer browsing resulted in a >93 % reduction in canopy coverage of dominant shrub species. The dramatic reduction of native shrubs at this site may create opportunities for displacement by exotic species, resulting in eventual vegetation-type conversion. The observed link between intense browsing and post-fire shrub mortality provides much needed information concerning the environmental impact of exotic deer on SCI and illustrates the interaction between exotic herbivores and fire on an island system.     

Dormant Season Prescribed Fire as a Management Tool for the Control of <Emphasis Type="Italic">Salix caroliniana</Emphasis> Michx. in a Floodplain Marsh

Expansion of woody species into herbaceous wetlands is a serious concern in wetland management. Prescribed fire is often used as a tool to manage woody species, although many species resprout after fire making control problematic. In this study, we assessed the usefulness of repeated dormant season fires for controlling Salix caroliniana (Michx.) in a floodplain marsh in Florida. Salix is a common shrub in southeastern marshes that resprouts prolifically after fire. We compared stem basal area, stem density, and cover of Salix in three adjacent sites in a floodplain marsh in east central Florida. One site was burned once in February 1997, another site was burned in February 1997 and then again in March 1999 and one site was left unburned. At the unburned site, Salix stem basal area, stem density, and cover increased over the course of the study. In the two burned sites, the first fire destroyed large diameter stems and stimulated production of sprouts. As a result, stem basal area and cover decreased but stem density remained unchanged. The second fire caused a decline in stem density and a further decline in cover. Changes in understory species composition and cover could not be attributed to the fires. Our results suggest that dormant season fires are effective in reducing Salix cover and basal area, and that repeated fires have greater effects than a single fire.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.     

Seed rain into upland plant communities in Hong Kong, China

Hong Kong is an extreme example of tropical landscape degradation, with no substantial remnants of the original forest cover and a highly impoverished disperser fauna. Seed availability is a potential limiting factor in vegetation recovery in such landscapes. To assess the quantity and quality of the seed rain of woody taxa, seed traps were placed in the major upland vegetation types: fire-maintained grassland, shrubland, and secondary forest. Within the grassland site, traps were placed under isolated trees, isolated male and female shrubs of Eurya chinensis, and in the open. Seeds were collected every 2 weeks for 2 years. The seed rain was highest under female shrubs in grassland (6455 seeds m⁻² year⁻¹), where it was almost entirely confined to their fruiting period. Next highest were isolated trees (890 seeds), followed by male isolated shrubs (611 seeds), shrubland (558 seeds), forest (129 seeds) and open grassland (47 seeds). The number of seed taxa was highest in shrubland (59), followed by isolated trees (42), forest (42), female isolated shrubs (28), male isolated shrubs (15), and open grassland (9). The seed rain differed in species composition between the forest, shrubland, and grassland sites, while the differences within the grassland site were largely in terms of quantity. Birds (particularly bulbuls, Pycnonotus spp.) are known or inferred to be the major dispersal agents for 85% of the seed taxa trapped, 99% of the total number of seeds trapped, and 99.8% of the seeds trapped in the grassland site. Few taxa and of the total seeds were dispersed by wind and no seed taxa were definitely dispersed by fruit bats. The results suggest that even in the most degraded landscape the seed rain is adequate for the development of woody vegetation cover, but that human intervention will be needed for the restoration of plant diversity.     

Impacts of Tree Rows on Grassland Birds and Potential Nest Predators: A Removal Experiment

Globally, grasslands and the wildlife that inhabit them are widely imperiled. Encroachment by shrubs and trees has widely impacted grasslands in the past 150 years. In North America, most grassland birds avoid nesting near woody vegetation. Because woody vegetation fragments grasslands and potential nest predator diversity and abundance is often greater along wooded edge and grassland transitions, we measured the impacts of removing rows of trees and shrubs that intersected grasslands on potential nest predators and the three most abundant grassland bird species (Henslow’s sparrow [Ammodramus henslowii], Eastern meadowlark [Sturnella magna], and bobolink [Dolichonyx oryzivorus]) at sites in Wisconsin, U.S.A. We monitored 3 control and 3 treatment sites, for 1 yr prior to and 3 yr after tree row removal at the treatment sites. Grassland bird densities increased (2–4 times for bobolink and Henslow’s sparrow) and nesting densities increased (all 3 species) in the removal areas compared to control areas. After removals, Henslow’s sparrows nested within ≤50 m of the treatment area, where they did not occur when tree rows were present. Most dramatically, activity by woodland-associated predators nearly ceased (nine-fold decrease for raccoon [Procyon lotor]) at the removals and grassland predators increased (up to 27 times activity for thirteen-lined ground squirrel [Ictidomys tridecemlineatus]). Nest success did not increase, likely reflecting the increase in grassland predators. However, more nests were attempted by all 3 species (175 versus 116) and the number of successful nests for bobolinks and Henslow’s sparrows increased. Because of gains in habitat, increased use by birds, greater production of young, and the effective removal of woodland-associated predators, tree row removal, where appropriate based on the predator community, can be a beneficial management action for conserving grassland birds and improving fragmented and degraded grassland ecosystems.     

Reintroduction of large herbivores restored plant species richness in abandoned dry temperate grassland

Naturalistic grazing by large herbivores is an increasingly practiced way of managing habitats with conservational value. It has the potential to restore and enhance biodiversity, creating self-sustainable environments vital for organisms requiring regular disturbances to moderate and/or reverse successional changes. European bison, Exmoor pony, and Tauros cattle were introduced in 2015 to a former military training area in Milovice, Czech Republic. The prevailing vegetation type is a forest-steppe savanna with Bromus erectus-dominated xeric grasslands mixed with deciduous shrubs and trees. After the cessation of military use, the area was abandoned which led to successional changes, including the dominance of tall grasses, litter accumulation, and bush encroachment. In 2017–2021, we monitored grassland vegetation in 30 grazed permanent plots (2?×?2 m) and 5 control plots representative of ungrazed, abandoned vegetation adjacent to the grazed areas. Naturalistic grazing increased species richness and the cover of forbs, while the cover of grasses and legumes was minimally affected. Grazing increased functional diversity of plant community, promoted a compositional change to small statured species and an increased incidence of red-list species. Seven years of continuous grazing increased the conservation value of this forest-steppe vegetation, a habitat type rapidly declining in Europe.

     

Herbaceous Forage and Selection Patterns by Ungulates across Varying Herbivore Assemblages in a South African Savanna

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.     

Effects of cattle and rabbit grazing on clonal expansion of spiny shrubs in wood-pastures

Spiny shrubs protect non-defended plants against herbivores. Therefore, they play a role for the diversity in grazed ecosystems. While the importance of these keystone nurse shrubs is presently recognized, little is known about the factors controlling them. This knowledge is required to understand the functioning of grazed ecosystems and for sustainable management.We studied effects of cattle and rabbits on clonal expansion of Prunus spinosa in two ancient wood-pastures in the Netherlands. At each site we set up five blocks in grassland perpendicular to the edges of mature Prunus thickets, each block containing three herbivore treatments: (1) open-to-cattle-and-rabbits, (2) open to rabbits, cattle excluded, (3) cattle and rabbits excluded. We monitored the number and volume of Prunus ramets from 1998 to 2000 and again in 2003, 3 years after exclosure-removal to restore grazing.For 1998–2000 ramet volume, but not ramet number, differed between treatments. Ramet volume was highest when both cattle and rabbits were excluded. Ramet volume did not differ between grazing by rabbits or cattle and rabbits combined, indicating that rabbits alone may be as effective in inhibiting clonal expansion as cattle and rabbits combined. Three years after exclosure-removal ramet number and volume had increased in all treatments. Number of ramets remained unaffected by (former) treatments. Ramet volume remained highest in the former cattle-plus-rabbits exclusion treatment, differing significantly from the ‘open-to-cattle-and-rabbits’ treatment. So, once successfully established during herbivore absence, further expansion is not prevented by cattle and rabbit grazing.This study shows that vertebrate herbivory controls the keystone nurse-shrub in wood-pastures: combined cattle and rabbit grazing, and notably rabbits alone, inhibit expansion. Temporary herbivore absence allows expansion of ramets, which persists after herbivore-reappearance. Sustainable management of wood-pastures should allow spatial-temporal fluctuations of herbivore densities, leading to increased vegetation structure and associated biodiversity.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Fine root dynamic characteristics and effect on plantation’s carbon sequestration of three Salix shrub plantations in Tibetan Plateau alpine sandy land

Desertification land in Gonghe Basin of Tibetan Plateau, China accounts for 91.9% of the total land area. Vegetation restoration and reconstruction with desert shrubs in degraded ecosystem are effective ways to prevent and control desertification. However, the evaluation studies of fine root dynamic characteristics of desert shrubs and their contribution to carbon sequestration of plantation are limited. To gain a better understanding of vegetation restoration, the vertical distribution of fine root biomass, fine root decomposition, fine root turnover was investigated, as well as their coupling effect on carbon sequestration of plantation in three desert vegetation. The results estimated that the total decomposition time of fine roots of Salix cheilophila (S. cheilophila), Salix psammophila (S. psammophila), and Salix microstachya (S. microstachya) are 39.00, 27.99 and 35.95 years. Biomass carbon density for three Salix plantations ranged from 1.42 to 2.39 t/hm², which showed that three Salix plantations in alpine sandy land are an important carbon pool. In addition, fine root biomass carbon density for the three shrub plantations varied significantly. Fine root biomass carbon density for S. psammophila reached the largest among the three plantations, which was 1.48 t/hm², accounting for the ratio of 62% of the plantation total biomass carbon density. The results indicated that the root system of S. psammophila, especially the fine roots, was very developed, which was conducive to soil water transportation and carbon sequestration. Therefore, S. psammophila might be a better species for carbon sequestration of plantation in alpine sandy areas. The carbon input from the fine roots of the three shrub plantations through decomposition and turnover into the plantations accounts for 11.5% to 15.5% of total carbon sequestration of plantations. Therefore, the fine roots dynamics must be considered for long‐term carbon pool estimations in three Salix plantations, otherwise the total carbon sequestration of plantations would be underestimated.     

Fine‐scale habitat heterogeneity influences browsing damage by elephant and giraffe

Effects of large mammalian herbivores on woody vegetation tend to be heterogeneous in space and time, but the factors that drive such heterogeneity are poorly understood. We examined the influence of fine‐scale habitat heterogeneity on the distribution and browsing effects of two of the largest African terrestrial mammals, the elephant and giraffe. We conducted this study within a 120‐ha (500 x 2,400 m) ForestGEO long‐term vegetation monitoring plot located at Mpala Research Center, Kenya. The plot traverses three distinct topographic habitats (“plateau,” “steep slopes,” and “valley”) with contrasting elevation, slope, soil properties, and vegetation composition. To quantify browsing damage, we focused on Acacia mellifera, a palatable tree species that occurs across the three habitat categories. Overall tree density, species richness, and diversity were highest on the steep slopes and lowest on the plateau. Acacia mellifera trees were tallest and had the lowest number of stems per tree on the steep slopes. Both elephant and giraffe avoided the steep slopes, and their activity was higher during the wet season than during the dry season. Browsing damage on Acacia mellifera was lowest on the steep slopes. Elephant browsing damage was highest in the valley, whereas giraffe browsing damage was highest on the plateau. Our findings suggest that fine‐scale habitat heterogeneity is an important factor in predicting the distribution of large herbivores and their effects on vegetation and may interact with other drivers such as edaphic variations to influence local variation in vegetation structure and composition.