Exclusion Of Large Herbivores Research Articles

Large herbivore diversity slows sea ice-associated decline in arctic tundra diversity.

Biodiversity is declining globally in response to multiple human stressors, including climate forcing. Nonetheless, local diversity trends are inconsistent in some taxa, obscuring contributions of local processes to global patterns. Arctic tundra diversity, including plants, fungi, and lichens, declined during a 15-year experiment that combined warming with exclusion of large herbivores known to influence tundra vegetation composition. Tundra diversity declined regardless of experimental treatment, as background growing season temperatures rose with sea ice loss. However, diversity declined slower with large herbivores than without them. This difference was associated with an increase in effective diversity of large herbivores as formerly abundant caribou declined and muskoxen increased. Efforts that promote herbivore diversity, such as rewilding, may help mitigate impacts of warming on tundra diversity.

Large herbivores facilitate the persistence of rare taxa under tundra warming

Ecological rarity, characterized by low abundance or limited distribution, is typical of most species, yet our understanding of what factors contribute to the persistence of rare species remains limited. Consequently, little is also known about whether rare species might respond differently than common species to direct (e.g., abiotic) and indirect (e.g., biotic) effects of climate change. We investigated the effects of warming and exclusion of large herbivores on 14 tundra taxa, three of which were common and 11 of which were rare, at an inland, low-arctic study site near Kangerlussuaq, Greenland. Across all taxa, pooled commonness was reduced by experimental warming, and more strongly under herbivore exclusion than under herbivory. However, taxon-specific analyses revealed that although warming elicited variable effects on commonness, herbivore exclusion disproportionately reduced the commonness of rare taxa. Over the 15-year duration of the experiment, we also observed trends in commonness and rarity under all treatments through time. Sitewide commonness increased for two common taxa, the deciduous shrubs Betula nana and Salix glauca, and declined in six other taxa, all of which were rare. Rates of increase or decline in commonness (i.e., temporal trends over the duration of the experiment) were strongly related to baseline commonness of taxa early in the experiment under all treatments except warming with grazing. Hence, commonness itself may be a strong predictor of species’ responses to climate change in the arctic tundra biome, but large herbivores may mediate such responses in rare taxa, perhaps facilitating their persistence.

Root, not aboveground litter, controls soil carbon storage under grazing exclusion across grasslands worldwide

AbstractGrazing exclusion is a common management strategy for improving grasslands degraded due to overgrazing. Although previous meta‐analyses proved that grazing exclusion improved soil carbon (C) storage of grasslands in China, we know little about its effect on grassland ecosystems elsewhere and whether litter accumulation induced by grazing exclusion regulates soil C storage. Using meta‐analysis, we integrated 91 publications to examine the effects of grazing exclusion on community litter and soil C storage, accompanied by analysis of the regulating roles of body size of herbivore, climatic conditions, and grazing exclusion duration. We found that grazing exclusion enhanced litter biomass and decomposition rate but did not change litter chemical quality. Generally, grazing exclusion enhanced plant production and soil C storage across grassland worldwide. Exclusion of medium and small herbivores had stronger effects on aboveground litter production and soil C storage than exclusion of large herbivores, and grazing exclusion's enhancement on litter production and soil C storage tended to increase as grazing exclusion proceeded. Notably, grazing exclusion‐induced increase in aboveground litter biomass was attributed to the increase in biomass of graminoids, while root biomass, not aboveground litter biomass, primarily regulated the increase in soil C storage. The stimulating effect of grazing exclusion on soil C storage was enhanced possibly due to enhanced root biomass response along a precipitation gradient. Generally, our study provided new insights that management practice under grazing exclusion should take plant community dynamics, grazing exclusion duration, preceding grazer type, and climate conditions into account simultaneously.

Large herbivores transform plant-pollinator networks in an African savanna

Pollination by animals is a key ecosystem service1,2 and interactions between plants and their pollinators are a model system for studying ecological networks,3,4 yet plant-pollinator networks are typically studied in isolation from the broader ecosystems in which they are embedded. The plants visited by pollinators also interact with other consumer guilds that eat stems, leaves, fruits, or seeds. One such guild, large mammalian herbivores, are well-known ecosystem engineers5-7 and may have substantial impacts on plant-pollinator networks. Although moderate herbivory can sometimes promote plant diversity,8 potentially benefiting pollinators, large herbivores might alternatively reduce resource availability for pollinators by consuming flowers,9 reducing plant density,10 and promoting somatic regrowth over reproduction.11 The direction and magnitude of such effects may hinge on abiotic context-in particular, rainfall, which modulates the effects of ungulates on vegetation.12 Using a long-term, large-scale experiment replicated across a rainfall gradient in central Kenya, we show that a diverse assemblage of native large herbivores, ranging from 5-kg antelopes to 4,000-kg African elephants, limited resource availability for pollinators by reducing flower abundance and diversity; this in turn resulted in fewer pollinator visits and lower pollinator diversity. Exclusion of large herbivores increased floral-resource abundance and pollinator-assemblage diversity, rendering plant-pollinator networks larger, more functionally redundant, and less vulnerable to pollinator extinction. Our results show that species extrinsic to plant-pollinator interactions can indirectly and strongly alter network structure. Forecasting the effects of environmental change on pollination services and interaction webs more broadly will require accounting for the effects of extrinsic keystone species.

Exclusion of large herbivores affects understorey shrub vegetation more than herb vegetation across 147 forest sites in three German regions.

BackgroundMany studies have analysed the effect of browsing by large herbivores on tree species but far fewer studies have studied their effect on understorey shrubs and herbs. Moreover, while many studies have shown that forest features and management intensity strongly influence understorey vegetation, the influence of such variation on the effect of large-herbivore exclusion is not known.This studyIn this study, we analysed changes of species richness, Shannon diversity, evenness and cover of understorey herbs and shrubs after excluding large herbivores for seven years on 147 forest sites, differing in management intensity and forest features, in three regions of Germany (Schwäbische Alb, Hainich-Dün, Schorfheide-Chorin). Further, we studied how the effect of large-herbivore exclusion on understorey vegetation was influenced by forest management intensity and several forest features.ResultsAs expected, exclusion of large herbivores resulted in highly variable results. Nevertheless, we found that large-herbivore exclusion significantly increased cover and Shannon diversity of shrub communities, while it did not affect herb communities. Forest management intensity did not influence the effect of large-herbivore exclusion while some forest features, most often relative conifer cover, did. In forests with high relative conifer cover, large-herbivore exclusion decreased species richness and cover of herbs and increased Shannon diversity of herbs and shrubs, while in forests with low relative conifer cover large-herbivore exclusion increased species richness and cover of herbs, and decreased Shannon diversity of herbs and shrubs.ConclusionWe suggest that browsing by large herbivores should be included when studying understorey shrub communities, however when studying understorey herb communities the effects of browsing are less general and depend on forest features.

Structural anti-herbivore defense reduction of two Patagonian spiny shrubs in response to long time exclusion of large herbivores

This work focuses on the structural defenses of the two dominant spiny shrub species of the Patagonian shrub-grass steppe. We compared the amount of structural defenses (spines and thorns) of Mulinum spinosum (Cav.) Pers. (Apiaceae) and Adesmia volckmannii Phil. (Fabaceae) between grazed and long term non-grazed plots. M. spinosum showed a higher spinescence than A. volckmannii. Moreover, both species plants located in grazed plots showed higher spinescence than those located in non-grazed ones. Therefore, we can conclude that (1) M. spinosum has higher spinescence than A. volckmannii, probably because sheep prefer M. spinosum flowers than A. volckmannii leaves; and (2) both M. spinosum and A. volckmannii showed an increase of spinescence induced by herbivory. These responses may be evidences of co-evolution between these shrub species and large wild browsers that guarantee an adequate capacity to tolerate herbivory disturbance. Such capacity would be important to maintain community functioning, because shrubs provide secure sites for seed germination and, in some cases, are able to set symbiotic interactions with N-fixing bacteria.

Spatial vegetation patterns and neighborhood competition among woody plants in an East African savanna.

The majority of research on savanna vegetation dynamics has focused on the coexistence of woody and herbaceous vegetation. Interactions among woody plants in savannas are relatively poorly understood. We present data from a 10-yr longitudinal study of spatially explicit growth patterns of woody vegetation in an East African savanna following exclusion of large herbivores and in the absence of fire. We examined plant spatial patterns and quantified the degree of competition among woody individuals. Woody plants in this semiarid savanna exhibit strongly clumped spatial distributions at scales of 1-5m. However, analysis of woody plant growth rates relative to their conspecific and heterospecific neighbors revealed evidence for strong competitive interactions at neighborhood scales of up to 5m for most woody plant species. Thus, woody plants were aggregated in clumps despite significantly decreased growth rates in close proximity to neighbors, indicating that the spatial distribution of woody plants in this region depends on dispersal and establishment processes rather than on competitive, density-dependent mortality. However, our documentation of suppressive effects of woody plants on neighbors also suggests a potentially important role for tree-tree competition in controlling vegetation structure and indicates that the balanced-competition hypothesis may contribute to well-known patterns in maximum tree cover across rainfall gradients in Africa.

The effect of structurally complex corals and herbivory on the dynamics of Halimeda

The calcareous green alga Halimeda is a key contributor to carbonate sediment production on coral reefs. As herbivores have a direct negative effect on the abundance of Halimeda, protection from herbivory is critical for Halimeda growth. Branching corals such as Acropora are likely to provide refugia for Halimeda from grazers, yet studies are scarce. Here, we investigated the vulnerability of two Halimeda species to herbivory using fish exclusion cages and assessed the contribution of coral structural complexity to seasonal changes in Halimeda biomass and morphometrics. While up to 50 % Halimeda abundance was depleted outside cages due to herbivory and the exclusion of large herbivores resulted in an increase in net growth up to threefold, Halimeda recruitment was positively affected by herbivory, more than two times greater outside cages. However, these responses differed between species and seasons; only one species was affected in winter but not summer. Coral structural complexity facilitated an increase of total algal biomass particularly in summer. At the individual level, thalli growing inside the Acropora canopy were always significantly larger (thallus biomass, volume and height) than those growing in exposed areas. We estimated that the carbonate production of Halimeda was nearly three times greater inside refuges provided by Acropora. Because Halimeda species differ in growth rates and susceptibility to grazing, we predict that the ongoing degradation of the habitat complexity provided by branching corals will alter Halimeda community structure and its contribution to local sediment budgets.

Small herbivores suppress algal accumulation on Agatti atoll, Indian Ocean

Despite large herbivorous fish being generally accepted as the main group responsible for preventing algal accumulation on coral reefs, few studies have experimentally examined the relative importance of herbivore size on algal communities. This study used exclusion cages with two different mesh sizes (1 × 1 cm and 6 × 6 cm) to investigate the impact of different-sized herbivores on algal accumulation rates on the shallow ( 8 cm body depth) while allowing smaller fishes to access the plots. In contrast to the conclusions of most previous studies, the exclusion of large herbivores had no significant effect on the accumulation of benthic algae and the amount of algae present within the coarse-mesh cages was relatively consistent throughout the experimental period (around 50 % coverage and 1–2 mm height). The difference in algal accumulation between the fine-mesh and coarse-mesh cages appears to be related to the actions of small individuals from 12 herbivorous fish species (0.17 ind. m−2 and 7.7 g m−2) that were able to enter through the coarse mesh. Although restricted to a single habitat, these results suggest that when present in sufficient densities and diversity, small herbivorous fishes can prevent the accumulation of algal biomass on coral reefs.

Competition on the range: science vs. perception in a bison-cattle conflict in the western USA.

Summary Competition between livestock and wild ungulates is commonly perceived to occur on shared rangelands. In the Henry Mountains (HM) of Utah, a free‐ranging population of bison Bison bison has raised concerns among ranchers holding grazing permits on these public lands. Bison are the most conspicuous potential competitors with cattle, but lagomorphs (mainly jackrabbits Lepus californicus) are also abundant in this area. The local ranching community is applying political pressure on state and federal agencies to resolve ‘the bison problem’, but the relative grazing impacts of bison, cattle and lagomorphs have not previously been quantified.We constructed 40 grazing exclosures (each 5·95 m2) in the conflict area: 20 excluded bison + cattle (‘partial’) and 20 excluded bison + cattle + lagomorphs (‘full’). All exclosures, each with a paired open reference plot, were monitored for 1 year, and above‐ground plant production was measured. GPS telemetry (bison) and scheduled grazing (cattle) allowed visitation to be quantified for each ungulate species based on the number of ‘animal days’ in the area. Rancher perceptions of wildlife–cattle interactions were recorded in a questionnaire survey.Ranchers perceived bison as a high‐level competitor with cattle, whereas lagomorphs were perceived as low‐level competitors.Grazed reference plots yielded an average (±SE) of 22·7 g m−2 (±5·16) of grass, compared to 36·5 g m−2 (±7·33) in the partial exclosures and 43·7 g m−2 (±7·61) in the full exclosures. Exclusion of large herbivores thus resulted in a 13·8 g m−2 increase in grass biomass relative to the reference plots (P = 0·005), with the additional exclusion of lagomorphs resulting in a further 7·18 g m−2 increase (P = 0·048).Overall, lagomorphs accounted for 34·1%, bison 13·7% and cattle 52·3% of the total grass biomass removed by all herbivores on the shared range. Synthesis and applications. Cattle face a greater competitive challenge from lagomorphs than from bison in the study area. This case study illustrates the need for science‐based management of social–ecological systems in which even long‐term resource users might underestimate the complexities of trophic interactions. Attention should be redirected at the lagomorphs and their main predators, coyotes Canis latrans, which are currently subject to population control. To reduce negative perceptions among local ranchers, options should be explored to incorporate benefit‐sharing into the management of the bison population.

Large herbivores and season independently affect woody stem circumference increment in a semi-arid African savanna

Seasonality is a characteristic feature of savannas. Assuming plants are dormant in the dry season, long-term browsing by large herbivores (or their exclusion) in savannas should affect stem growth in the wet season alone. In addition, plant survival should improve with exclusion. A total of 236 dendrometers on six woody species in three large (>25 ha) plots (all large herbivores excluded, elephants excluded, control—no large herbivores excluded) were observed from 2006 to 2013 in Kruger National Park, South Africa. Average monthly increment (AMI) per dendrometer was calculated and the interaction between exclusion and season (dry or wet) was tested species-wise. The AMIs of Acacia exuvialis, Combretum apiculatum, Dichrostachys cinerea and Grewia flavescens were generally reduced by herbivore exclusion, while the unpalatable Euclea divinorum was unaffected. Survival rates of C. apiculatum and E. divinorum were not significantly associated with exclusion, while other species’ survival rates benefitted from exclusion. The dry-season AMIs of all species were significantly lower than wet-season AMIs, but dry-season AMIs were positive for two species common on footslopes (A. grandicornuta and E. divinorum), while being almost zero for species common on midslopes and crests, possibly because of hydrological variation on the catena. The results suggested that stem circumference growth of prominent woody species in semi-arid savannas is prone to reduction by exclusion of large herbivores, except for generally unpalatable species. Large herbivores may affect woody stem growth in semi-arid savannas by altering competition among plants.

Exclusion of large herbivores: Long-term changes within the plant community

Semi-natural woodlands are a globally important relict ecosystem for biodiversity, which are threatened through a range of anthropogenically induced changes, including overgrazing by large herbivores. Fencing to conserve biodiversity is increasingly used as a management tool, so the long-term impacts of large herbivore removal requires investigation. The objective of this research is to investigate the effect of large herbivore exclusion on vegetation, through time, using empirical long-term vegetation data collected over ∼40years. The responses of ground flora communities to this removal will be assessed, and it is predicted that a change in woodland vegetation will occur through time. Plant community composition was surveyed in a network of 7 large-herbivore exclosures, within protected temperate oak woodland in Ireland, at intervals up to 41years. Ground flora species abundance was quantified and time since fencing was used to standardise the survey data, with three time groups being derived from this to assess beta diversity changes through time. With total removal of large herbivores from the oak woodland ecosystem, this study has identified significant changes in ground flora composition and abundance, and a general homogenisation of the vegetation community with increasing time since large herbivore removal. Large-scale long-term fencing of oak woodlands should be replaced by large herbivore management programmes, in order to ensure the conservation of diverse woodland ecosystems.

Rapid structural and compositional change in an old-growth subtropical forest: using plant traits to identify probable drivers.

Recent studies have shown directional changes in old-growth tropical forests, but changes are complex and diverse, and their drivers unclear. Here, we report rapid net structural and compositional changes in an old-growth subtropical forest and we assess the functional nature of these changes to test hypothetical drivers including recovery from past disturbances, reduction in ungulate browsing, CO2 fertilization, and increases in rainfall and temperature. The study relies on 15 years of demographic monitoring within 8 ha of subtropical montane forest in Argentina. Between 1992 and 2007, stem density markedly increased by 50% (12 stems ha−1 y−1) and basal area by 6% (0.13 m2 ha−1 y−1). Increased stem density resulted from enhanced recruitment of understory treelets (Piper tucumanum, Eugenia uniflora, Allophylus edulis) into small size classes. Among 27 common tree species, net population growth was negatively correlated with maximum tree size and longevity, and positively correlated with leaf size and leaf nutrient content, especially so when initial population size was controlled for. Changes were inconsistent with predictions derived from past disturbances (no increase in shade-tolerant or long-lived late-succesional species), rainfall or temperature increase (no increase in evergreen or deciduous species, respectively). However, the increase in nutrient-rich soft-leaved species was consistent with exclusion of large herbivores two decades before monitoring started; and CO2 fertilization could help explain the disproportionate increase in small stems. Reductions in populations of large vertebrates have been observed in many otherwise undisturbed tropical forests, and our results suggest they can have important structural and functional repercussions in these forests.

Herbivores strongly influence algal recruitment in both coral- and algal-dominated coral reef habitats

Coral reefs can exist as coral- and macroalgae-dominated habitats often separated by only a few hundred metres. While herbivorous fish are known to depress the abundance of algae and help maintain the function of coral-dominated habitats, less is known about their influence in algae-dominated habitats. Here, we quantified herbivorous fish and benthic algal communities over a 6 mo period in coral-dominated (back-reef) and algal-dominated (lagoon) habitats in a relatively undisturbed fringing coral reef (Ningaloo, Western Australia). Simulta - neously, we tested the effects of herbivorous fish on algal recruitment in both habitats using recruitment tiles and fish exclusion cages. The composition of established algal communities differed consistently between habitats, with the back-reef hosting a more diverse community than the Sargassum-dominated lagoon. However, total algal biomass and cover only differed between habitats in autumn, coinciding with maximum Sargassum biomass. The back-reef hosted high coral cover and a diverse herbivorous fish community, with herbivore biomass an order of magni- tude greater than the lagoon. Despite these differences in herbivore composition, exclusion of large herbivores had a similar positive effect to foliose macroalgae recruitment on experimental tiles in both back-reef and lagoon habitats. Additionally, territorial damselfish found in the back- reef increased turf algae cover and decreased crustose coralline algae cover on recruitment tiles. Collectively, our results show that disparate herbivorous fish communities in coral- and algae- dominated habitats are similarly able to limit the recruitment of foliose macroalgae, but suggest that when herbivorous fish biomass and diversity are relatively low, macroalgal communities are able to escape herbivore control through increased growth.

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.

Large herbivores favour species diversity but have mixed impacts on phylogenetic community structure in an African savanna ecosystem

Summary There has been much debate on the impact of large herbivores on biodiversity, especially given that large mammals are becoming locally extinct in many places. The use of evolutionary information on community structure has typically been limited to evaluating assembly processes, for example, competition or habitat filtering, whereas a lack of long‐term experiments has precluded the test of predictions considering more complex biotic interactions. Reconstructing the complete phylogeny of the trees and shrubs of the Kruger National Park from DNA data, we tested for phylogenetic signal in antiherbivory traits and compared the phylogenetic structure of communities under various degrees of herbivore pressure using experimental plots spanning several decades. We show that all antiherbivory traits examined demonstrated weak but significant phylogenetic signal, and that exclusion of large herbivores results in impoverished species diversity in restructured communities. Surprisingly, we also show that reduction in species diversity coupled with community reorganization does not necessarily result in a decrease in phylogenetic diversity, and that community responses to herbivore exclusion depend on initial structure. Synthesis. Extinction of large mammal herbivores will have cascading effects on plant diversity; however, impacts on plant community structure are contingent on initial conditions. This research has implications for best practice when managing large herbivores and natural habitats.

Elk herbivory alters small mammal assemblages in high‐elevation drainages

Heavy herbivory by ungulates can substantially alter habitat, but the indirect consequences of habitat modification for animal assemblages that rely on that habitat are not well studied. This is a particularly important topic given that climate change can alter plant-herbivore interactions. We explored short-term responses of small mammal communities to recent exclusion of Rocky Mountain elk (Cervus elaphus) in high-elevation riparian drainages in northern Arizona, where elk impacts on vegetation have increased over the past quarter century associated with climate change. We used 10-ha elk exclosures paired with unfenced control drainages to examine how browsing influenced the habitat use, relative abundance, richness and diversity of a small mammal assemblage. We found that the small mammal assemblage changed significantly after 5years of elk exclusion. Relative abundance of voles (Microtus mexicanus) increased in exclosure drainages, likely due to an increase in habitat quality. The relative abundances of woodrats (Neotoma neomexicana) and two species of mice (Peromyscus maniculatus and P.boylii) decreased in the controls, while remaining stable in exclosures. The decline of mice in control drainages was likely due to the decline in shrub cover that they use. Thus, elk exclusion may have maintained or improved habitat for mice inside the exclosures while habitat quality and mouse abundance both declined outside the fences. Finally, small mammal species richness increased in the exclosures relative to the controls while species diversity showed no significant trends. Together, our results show that relaxation of heavy herbivore pressure by a widespread native ungulate can lead to rapid changes in small mammal assemblages. Moreover, exclusion of large herbivores can yield rapid responses by vegetation that may enhance or maintain habitat quality for small mammal populations.

Post‐dispersal seed removal in four Mediterranean oaks: species and microhabitat selection differ depending on large herbivore activity

AbstractQuercus forest regeneration is limited by different factors, post‐dispersal predation being one of the most critical stages. We analysed seed removal of four coexisting Quercus species (Q. ilex, Q. suber, Q. faginea and Q. pyrenaica) in a Mediterranean forest located in Southern Spain. Marked and weighed acorns from each of the species were placed in experimental units with or without exclusion of large herbivores and in two microhabitat types (Q. ilex shade or open). Acorn removal was monitored for 120 days in order to test the effect of exclusion of large herbivore and microhabitat type on seed removal rates and species selection. Interestingly, the results of microhabitat and species selection differed depending on the presence of large herbivores. Removal was faster in sites without exclusion, where most seeds (≈85%) disappeared during the first 9 days. In these sites, no differences in seed removal were found between different microhabitats, but seeds of two species, those with higher seed mass (Q. suber and Q. pyrenaica) were most preferred. However, under exclusion of large herbivores, seed removal was affected by the microhabitat, this being greater in Q. ilex shaded microhabitats, which showed a higher structural diversity. Also, species selection was completely different under exclusion of large herbivores, and seeds of Q. ilex and Q. faginea were removed faster. These results highlight the importance of large herbivore activity on seed removal and its effect on microhabitat and species selection. Therefore, specific selection by seed consumers may modify seedling recruitment and may have an important influence on the relative abundance of coexisting Quercus species.

The fencing paradigm in woodland conservation: consequences for recruitment of a semi‐arid tree

AbstractQuestionHow does long‐term fencing against large domestic herbivores affect regeneration of the dominant tree, Prosopis flexuosa, and hence the structure of semi‐arid woodlands?LocationWoodlands in the Central Monte Desert biome of Argentina, Ñacuñán Man and the Biosphere Reserve area (34° 20′ S, 67° 58′ W) and surrounding cattle‐grazed ranches.MethodsWe compared seedling emergence and survival, the spatial distribution of seedlings and saplings, and the population stage‐based structure of P. flexuosa between paired sites inside and outside the Reserve of Ñacuñán (Argentina), which has been fenced to exclude domestic cattle for ca. 40 yr.ResultsReserve sites had lower tree recruitment and seedling emergence, in spite of having greater seed production and seedling survival. Outside the reserve, survival was higher for seedlings in high‐density clumps than for isolated seedlings. Seedling clumps occurred mostly near adult individuals, where cattle dung was abundant, suggesting an effect of cattle on seed dispersal. The balance between the effects of cattle exclusion on seedling emergence and on seedling survival was reflected in the stage structure of P. flexuosa woodlands, as populations at fenced sites were dominated by adult individuals, whereas those at cattle‐grazed areas presented greater proportions of seedlings and saplings.ConclusionFencing is a common practice used worldwide to exclude conservation areas from human disturbance. For example, it is assumed that disturbed woodlands may recover after fencing, thanks to increased tree recruitment after the exclusion of large herbivores. However, the actual effectiveness of fencing as a tool for forest conservation in overgrazed environments could be controversial because sequential effects of herbivores during the tree regeneration cycle may hamper predictions of the overall balance of the recruitment process. Our results suggest that dispersal by cattle influences the regeneration of P. flexuosa woodlands, and that the positive effects of cattle on seedling emergence compensate for the negative effects on seed production and seedling and sapling survival. Future management of P. flexuosa woodlands in the Central Monte Desert should consider that tree recruitment is closely related to land use, and that cattle exclusion does not necessarily guarantee woodland recovery in the long term.

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

AbstractThe 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.