Effects Of Large Herbivores Research Articles

Large herbivores suppress liana infestation in an African savanna

African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages-declining wildlife populations and their displacement by livestock-may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana, Cynanchum viminale (Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana-tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock.

Effects of large herbivores on fire regimes and wildfire mitigation

Abstract Abandonment of agricultural land is widespread in many parts of the world, leading to shrub and tree encroachment. The increase of flammable plant biomass, that is, fuel load, increases the risk and intensity of wildfires. Fuel reduction by herbivores is a promising management strategy to avoid fuel build‐up and mitigate wildfires. However, their effectiveness in mitigating wildfire damage may depend on a range of factors, including herbivore type, population density and feeding patterns. Here, we review the evidence on whether management with herbivores can reduce fuel load and mitigate wildfires, and if so, how to identify suitable management that can achieve fire mitigation objectives while providing other ecosystem services. We systematically reviewed studies that investigated links between herbivores, fire hazard, fire frequency and fire damage. We found that, in general, herbivores reduce fuel load most effectively when they are mixed feeders, when grazing and browsing herbivores are combined and when herbivore food preferences match the local vegetation. In some cases, the combination of herbivory with other management strategies, such as mechanical clearing, is necessary to reduce wildfire damage. Synthesis and Applications. We conclude that herbivores have the capacity to mitigate wildfire damage, and we provide guidance for grazing management for wildfire mitigation strategies. As areas undergoing land abandonment are particularly prone to wildfires, the maintenance or promotion of grazing by domestic or wild herbivores is a promising tool to reduce wildfire risk in a cost‐effective way, while also providing other ecosystem services. Relevant land‐use policies, including fire suppression policies, agricultural and forest(ry) policies could incentivise the use of herbivores for better wildfire prevention.

Deer exclusion unveils abiotic filtering in forest understorey plant assemblages.

The role of deer (family Cervidae) in ecosystem functioning has traditionally been neglected by forest ecologists due to the animal's scarcity in most parts of the northern hemisphere. However, the dramatic rebound in deer populations throughout the 20th century has brought deer browsing to the forefront of forest ecological questioning. Today there is ample evidence that deer affect tree regeneration, understorey plant and animal diversity, and even litter decomposition. However, the mechanisms underlying the effects of deer on forest ecosystems remain unclear. Among others, the relative role of abiotic factors versus biotic interactions (e.g. herbivory) in shaping plant assemblages remains largely unknown. We used a large-scale experiment with exclosures distributed along abiotic gradients to understand the role of black-tailed deer (Odocoileus hemionus sitchensis) on the forest understorey on the Haida Gwaii archipelago (western Canada), a unique context where most of the key ecological effects of deer presence have already been intensively studied. Our results demonstrate that 20 years of deer exclusion resulted in a clear increase in vascular plant richness, diversity and cover, and caused a decline in bryophyte cover. Exclusion also unveiled abiotic (i.e. soil water availability and fertility) filtering of plant assemblages that would otherwise have been masked by the impact of abundant deer populations. However, deer exclusion did not lead to an increase in beta diversity, probably because some remnant species had a competitive advantage to regrow after decades of over browsing. We demonstrated that long-term herbivory by deer can be a dominant factor structuring understorey plant communities that overwhelms abiotic factors. However, while exclosures prove useful to assess the overall effects of large herbivores, the results from our studies at broader scales on the Haida Gwaii archipelago suggest that exclosure experiments should be used cautiously when inferring the mechanisms at work.

Ungulate herbivory reduces abundance and fluctuations of herbivorous insects in a boreal old-growth forest

Ecological theory predicts the strongest ecosystem effects of herbivory when dominant and ecologically important species are consumed. Bilberry, Vaccinium myrtillus, is such a key plant species, attractive to many other species in the boreal forests, for example ungulate and invertebrate herbivores. Large herbivores may remove substantial biomass and alter plant quality and therefore affect abundance and populations of invertebrate animals sharing the same food plant. We combined experimental exclusion of ungulates with a browsing intensity gradient to investigate the 15-year effect of ungulate (Cervus elaphus and Ovis aries) browsing on bilberry plant size and on bilberry-feeding herbivorous larvae (Lepidoptera and Symphyta), in a Norwegian old growth boreal forest ecosystem. Bilberry ramets in exclosure plots had nearly nine times higher dry mass and three times higher abundance of invertebrates feeding on them than in ungulate-access plots. Sweep-netting data verified these findings as larval numbers were twice as high in exclosure plots. The pattern in the large herbivore effects on bilberry size and abundance of herbivorous larvae were identical along the browsing gradient. Differences in larval abundance between treatments, as indicated by leaf-chewing, increased during the 15-year study period, and the community fluctuations were larger when ungulate herbivores were excluded. The browsing effect was moderated by plant quality as larval densities were lowest on both heavily-browsed and non-browsed plants, and highest on ramets that had 50–74% of annual shoots browsed. Our study supports previous findings in that bilberry is relatively disturbance tolerant and may recover quickly, but that ungulates may compete with herbivorous larvae for food biomass. Additionally, our results strongly indicates that population insect community peaks and fluctuations are dampened by ungulate consumption. Our findings add to the understanding on how ungulates may structure forest ecosystems directly and indirectly.

Synthesizing the effects of large, wild herbivore exclusion on ecosystem function

AbstractWild large herbivores are declining worldwide. Despite extensive use of exclosure experiments to investigate herbivore impacts, there is little consensus on the effects of wild large herbivores on ecosystem function.Of the ecosystem functions likely impacted, we reviewed the five most‐studied in exclosure experiments: ecosystem resilience/resistance to disturbance, nutrient cycling, carbon cycling, plant regeneration, and primary productivity.Experimental data on large wild herbivores' effects on ecosystem functions were predominately derived from temperate grasslands (50% grasslands, 75% temperate zones). Additionally, data were from experiments that may not be of adequate size (median size 400 m2despite excluding all experiments below 25 m2) or duration (median duration 6 years) to capture ecosystem‐scale responses to these low‐density and wide‐ranging taxa.Wild herbivore removal frequently impacted ecosystem functions; for example, net carbon uptake increased by three times in some instances. However, the magnitude and direction of effects, even within a single function, were highly variable.A focus on carbon cycling highlighted challenges in interpreting effects on a single function. While the effect of large herbivore exclusion on carbon cycling was slightly positive when its components (e.g. pools vs. fluxes of carbon) were aggregated, effects on individual components were variable and sometimes opposed.Given modern declines in large wild herbivores, it is critical to understand their effects on ecosystem function. However, this synthesis highlights strong variability in direction, magnitude, and modifiers of these effects. Some variation is likely due to disparity in what components are used to describe a given function. For example, for the carbon cycle we identified eight distinctly meaningful components, which are not easily combined yet are potentially misrepresentative of the larger cycle when considered alone. However, much of the observed difference in responses likely reflects real ecological variability across complex systems.To move towards a general predictive framework we must identify where variation in effect is due to methodological differences and where due to ecosystem context. Two critical steps forward are (a) additional quantitative synthetic analyses of large herbivores' effects on individual functions, and (b) improved, increased systematic exclosure research focusing on effects of large herbivores' exclusion on functions.A freePlain Language Summarycan be found within the Supporting Information of this article.

Effects of large herbivores on tundra vegetation in a changing climate, and implications for rewilding.

In contrast to that of the Pleistocene epoch, between approximately 2.6 million and 10 000 years before present, the extant community of large herbivores in Arctic tundra is species-poor predominantly due to human extinctions. We here discuss how this species-poor herbivore guild influences tundra ecosystems, especially in relation to the rapidly changing climate. We show that present herbivore assemblages have large effects on tundra ecosystem composition and function and suggest that the effect on thermophilic species expected to invade the tundra in a warmer climate is especially strong, and that herbivores slow ecosystem responses to climate change. We focus on the ability of herbivores to drive transitions between different vegetation states. One such transition is between tundra and forest. A second vegetation transition discussed is between grasslands and moss- and shrub-dominated tundra. Contemporary studies show that herbivores can drive such state shifts and that a more diverse herbivore assemblage would have even higher potential to do so. We conclude that even though many large herbivores, and especially the megaherbivores, are extinct, there is a potential to reintroduce large herbivores in many arctic locations, and that doing so would potentially reduce some of the unwanted effects of a warmer climate.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Identifying foundation species in North American forests using long‐term data on ant assemblage structure

AbstractFoundation species are locally abundant and uniquely control associated biodiversity, whereas dominant species are locally abundant but are thought to be replaceable in ecological systems. It is important to distinguish foundation from dominant species to direct conservation efforts. Long‐term studies that remove abundant species while measuring community dynamics have the potential to (1) aid in the identification of foundation vs. dominant species and, (2) once a foundation species is identified, determine how long its effects persist within a community after its loss. Long‐term data on ant assemblages within two canopy‐manipulation experiments—the Harvard Forest Hemlock Removal Experiment (HF‐HeRE) and the Black Rock Future of Oak Forests Experiment (BRF‐FOFE)—provide insights into how ant assemblages change and reassemble following the loss of Tsuga canadensis or Quercus spp. Previous research documented foundation species effects on ants in the HF‐HeRE for up to four years after T. canadensis loss. Six additional years of data at HF‐HeRE presented for the first time here show that removal of T. canadensis resulted in taxonomic and some measures of functional shifts in ant assemblages that persisted for ten years, further supporting the hypothesis that T. canadensis is a foundation species at Harvard Forest. In contrast, ant assemblages at BRF‐FOFE varied little regardless of whether oaks or other tree species were removed from the canopy, suggesting that Quercus species do not act as foundation species at Black Rock Forest. Deer and moose exclosures within each experiment also allowed for comparisons between effects on ants of foundation or dominant tree species relative to effects of large herbivores. At HF‐HeRE, effects of T. canadensis were stronger than effects of large herbivores on taxonomic and functional diversity of ant assemblages. At BRF‐FOFE, in contrast, effects of Quercus species were weaker than effects of large herbivores on ant taxonomic diversity and some measures of ant functional diversity. These findings illustrate the importance of distinguishing between the roles of irreplaceable foundation species and replaceable dominant ones in forested ecosystems along with other drivers of biodiversity (e.g., herbivory).

Determinants of patchiness of woody vegetation in an African savanna

AbstractQuestionsHow is woody vegetation patchiness affected by rainfall, fire and large herbivore biomass? Can we predict woody patchiness and cover over large‐scale environmental gradients?LocationHluhluwe‐iMfolozi Park, South Africa.MethodsWe quantified variation in local patchiness as the lacunarity of woody cover on satellite‐derived images. Using Random Forest regression we analysed how both average woody cover and its patchiness depend on annual rainfall, fire frequency and grazer and browser metabolic biomass densities.ResultsFire frequency and rainfall were the clearest predictors, whereas effects of large herbivores on woody vegetation were smaller and more complex. Under low rainfall conditions (500 mm·yr−1) trees had less total cover and were more regularly spaced across the landscape (lower patchiness). Woody cover and vegetation patchiness increased with rainfall whereas fire frequency decreased woody cover and further increased patchiness.ConclusionThese results suggest a switch from competition between neighbouring trees under low rainfall conditions towards benefits among neighbours with increasing fire frequencies. Whereas overall woody cover and its patchiness are two independent aspects of savanna woody vegetation patterns, both need to be investigated to obtain a good understanding of the functioning and diversity of savanna ecosystems.

A novel mechanism for grazing lawn formation: large herbivore‐induced modification of the plant–soil water balance

Summary Large herbivores play a key role in creating spatial heterogeneity through the formation of grazing lawns. Recent research suggests that the currently accepted nutrient‐based theory on the formation of these grazing lawns cannot universally explain their formation in all ecosystems where they are found. We developed and investigated an alternative hypothesis on grazing lawn formation and maintenance based on herbivore effects on the plant–soil water balance. We propose that large herbivores change the soil water balance in grazing lawns through defoliation and soil compaction, causing a shift in vegetation composition towards a drought‐tolerant plant community. Investigating this idea in a tropical savanna, we indeed found profound differences in grazing lawn soil properties and water balance. In particular, defoliation increased soil temperatures and potential evaporation rates while soil compaction increased bulk density and decreased water infiltration rates, especially on fine‐textured soils. Soil moisture was therefore generally much lower in grazing lawns than in adjacent bunch grass areas. Furthermore, we found that grazing lawn species show drought‐tolerant traits, with higher leaf sodium levels, suggesting evolutionary adaptation to these herbivore‐induced dry conditions. However, leaf water potentials did not differ between grazing lawn and bunch grass species. Synthesis. This study shows that large herbivores might form grazing lawns through previously underestimated effects on water balance. Thus, future studies on large herbivore effects on vegetation should increasingly focus on additional pathways of soil compaction and defoliation. While nutrient‐based processes driving grazing lawn formation may operate during the wet season in savannas, we suggest that water balance‐based processes are additionally important during the dry season.

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.

Nutrient and secondary metabolite concentrations in a savanna are independently affected by large herbivores and shoot growth rate

Carbon-based secondary metabolites (CBSMs) such as tannins are assumed to function as plant defences against herbivores. CBSMs are thought to be inversely related to growth rate and nutrient concentrations because a physiological trade-off exists between cellular growth and differentiation, but CBSM concentrations can be altered by herbivory-induced changes in the trade-off. We predicted that a significant interaction exists between herbivory and growth phase, such that the effects of large herbivores (or their exclusion) on nutrient or CBSM concentrations are greatest during phases of rapid shoot or leaf growth. Leaf samples were collected during phases of different growth rate from six woody species 4 years after establishment of a large-scale long-term herbivore exclusion experiment in Kruger National Park, South Africa. Samples were analysed for N, P, condensed tannins and total phenolics. Interactions between growth phase and herbivores were rare. However, the assumption that elevated nutrients and reduced CBSMs occurs during fast phases of growth was supported by four species (consistent with the growth-differentiation balance hypothesis), but not the other two. Large herbivores generally did not affect nutrients, but CBSMs in four species were reduced by large herbivores other than elephants, while CBSMs in two species were reduced by elephants. Carbon limitation ultimately prevailed among woody plants taller than 2 m under long-term browsing. Large herbivores and plant growth phase are independent and important determinants of nutrients or CBSMs in African savannas, but the effects depend on the interacting assemblages of species, which poses challenges to the application of current general hypotheses of plant defence.

Herbivore trampling as an alternative pathway for explaining differences in nitrogen mineralization in moist grasslands

Studies addressing the role of large herbivores on nitrogen cycling in grasslands have suggested that the direction of effects depends on soil fertility. Via selection for high quality plant species and input of dung and urine, large herbivores have been shown to speed up nitrogen cycling in fertile grassland soils while slowing down nitrogen cycling in unfertile soils. However, recent studies show that large herbivores can reduce nitrogen mineralization in some temperate fertile soils, but not in others. To explain this, we hypothesize that large herbivores can reduce nitrogen mineralization in loamy or clay soils through soil compaction, but not in sandy soils. Especially under wet conditions, strong compaction in clay soils can lead to periods of soil anoxia, which reduces decomposition of soil organic matter and, hence, N mineralization. In this study, we use a long-term (37-year) field experiment on a salt marsh to investigate the hypothesis that the effect of large herbivores on nitrogen mineralization depends on soil texture. Our results confirm that the presence of large herbivores decreased nitrogen mineralization rate in a clay soil, but not in a sandy soil. By comparing a hand-mown treatment with a herbivore-grazed treatment, we show that these differences can be attributed to herbivore-induced changes in soil physical properties rather than to above-ground biomass removal. On clay soil, we find that large herbivores increase the soil water-filled porosity, induce more negative soil redox potentials, reduce soil macrofauna abundance, and reduce decomposition activity. On sandy soil, we observe no changes in these variables in response to grazing. We conclude that effects of large herbivores on nitrogen mineralization cannot be understood without taking soil texture, soil moisture, and feedbacks through soil macrofauna into account.

The effects of large herbivores on the landscape dynamics of a perennial herb

Models assessing the prospects of plant species at the landscape level often focus primarily on the relationship between species dynamics and landscape structure. However, the short-term prospects of species with slow responses to landscape changes depend on the factors affecting local population dynamics. In this study it is hypothesized that large herbivores may be a major factor affecting the short-term prospects of slow-responding species in the European landscape, because large herbivores have increased in number in this region in recent decades and can strongly influence local population dynamics. The impact of browsing by large herbivores was simulated on the landscape-level dynamics of the dry grassland perennial polycarpic herb Scorzonera hispanica. A dynamic, spatially explicit model was used that incorporated information on the location of patches suitable for S. hispanica, local population dynamics (matrices including the impact of large herbivores), initial population sizes and dispersal rate of the species. Simulations were performed relating to the prospects of S. hispanica over the next 30 years under different rates of herbivory (browsing intensity) and varying frequencies of population destruction (e.g. by human activity). Although a high rate of herbivory was detected in most populations of S. hispanica, current landscape-level dynamics of S. hispanica were approximately in equilibrium. A decline or increase of over 20 % in the herbivory rate promoted rapid expansion or decline of S. hispanica, respectively. This effect was much stronger in the presence of population destruction. Browsing by large herbivores can have a dramatic effect on the landscape dynamics of plant species. Changes in the density of large herbivores and the probability of population destruction should be incorporated into models predicting species abundance and distribution.

Large herbivore effects on songbirds in boreal forests: lessons from deer introduction on Anticosti Island

Over recent decades, deer populations have reached densities exceeding their natural range of variability in several regions worldwide. Abundant deer populations can change the composition and structure of understory plant communities in forests and trigger trophic cascades affecting animal communities. To investigate such indirect effects on boreal forest songbirds we compared the understory vegetation and songbird communities of balsam fir stands on Anticosti Island (Quebec, Canada), where white-tailed deer (Odocoileus virginianus) were introduced, to those of the neighbouring Mingan Archipelago, where deer are absent. Mean cover of trees at 0–2 m height was on average sevenfold higher on Mingan than on Anticosti, and songbirds dependent on the understory were more common on Mingan than on Anticosti. Songbird community composition was more variable on Mingan, suggesting that the opening of the forest understory on Anticosti through intensive browsing homogenized the songbird community by reducing the occurrence of understory-dependent species. These indirect effects on songbirds were similar, although less dramatic, than those described in temperate forests. We interpret this as a consequence of lower productivity and complexity in understory vegetation of boreal forests. Our results emphasize the negative effects of abundant deer populations on animal diversity and underline the role played in the dynamics of communities by top down factors that regulate herbivore populations.

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.

Effects of large herbivores on murid rodents in a South African savanna

Abstract:Our study presents experimentally based results on how large herbivore species affect savanna vegetation and thus murid rodents in the Hluhluwe-iMfolozi Park in KwaZulu-Natal, South Africa. We permanently excluded groups of large herbivore guilds of various body sizes (ranging from white rhino to hares) from sixteen 40 × 40-m plots of vegetation by using different fence types. We determined grass species composition and vegetation height and collected capture–mark–recapture data on murid rodents. Nutrient concentrations of the dominant grass species and rodent diet compositions were analysed. We found that herbivore species of different body sizes had different effects on murid rodents. The exclusion of medium-sized herbivores, such as warthog, impala and nyala increased the abundance of high-quality grass species, especiallyPanicum maximum. However, the dominant rodent speciesLemniscomys rosaliapreferred the most abundant grass species, rather than high-quality grasses. The absence of large bulk feeders, such as zebra, buffalo and white rhino led to an increase in vegetation height. In response, tall vegetation promoted both rodent abundance and species diversity and altered rodent species composition. Ultimately, our results indicate that the greatest effect on murid rodents came from the reduction of vegetation cover by large bulk feeders, which likely increased rodent predation risk.

Positive and negative effects of grass, cattle, and wild herbivores on Acacia saplings in an East African savanna

Plant-plant interactions can be a complex mixture of positive and negative interactions, with the net outcome depending on abiotic and community contexts. In savanna systems, the effects of large herbivores on tree-grass interactions have rarely been studied experimentally, though these herbivores are major players in these systems. In African savannas, trees often become more abundant under heavy cattle grazing but less abundant in wildlife preserves. Woody encroachment where cattle have replaced wild herbivores may be caused by a shift in the competitive balance between trees and grasses. Here we report the results of an experiment designed to quantify the positive, negative, and net effects of grasses, wild herbivores, and cattle on Acacia saplings in a Kenyan savanna. Acacia drepanolobium saplings under four long-term herbivore regimes (wild herbivores, cattle, cattle + wild herbivores, and no large herbivores) were cleared of surrounding grass or left with the surrounding grass intact. After two years, grass-removal saplings exhibited 86% more browse damage than control saplings, suggesting that grass benefited saplings by protecting them from herbivory. However, the negative effect of grass on saplings was far greater; grass-removal trees accrued more than twice the total stem length of control trees. Where wild herbivores were present, saplings were browsed more and produced more new stem growth. Thus, the net effect of wild herbivores was positive, possibly due to the indirect effects of lower competitor tree density in areas accessible to elephants. Additionally, colonization of saplings by symbiotic ants tracked growth patterns, and colonized saplings experienced lower rates of browse damage. These results suggest that savanna tree growth and woody encroachment cannot be predicted by grass cover or herbivore type alone. Rather, tree growth appears to depend on a variety of factors that may be acting together or antagonistically at different stages of the tree's life cycle.

Tree Regeneration and Sapling Damage of Pedunculate Oak Quercus robur in a Grazed Forest in Galicia, NW Spain: A Comparison of Continuous and Rotational Grazing Systems

Forest grazing has been recognised as being a useful tool in fire-risk reduction, in addition to having the potential to preserve or enhance forest biodiversity if managed correctly. Concern for natural regeneration of forest trees in Europe has also prompted interest in the effects of large herbivores on seedling and sapling growth and mortality. An investigation was carried out into sapling damage and density of natural regeneration of oak (Quercus robur) in a mature, pony-grazed, Pinus radiata forest in Galicia, NW Spain under two different grazing regimes (continuous and rotational). In all treatments significantly more oak seedlings and saplings were recorded in areas of grass sward than areas dominated by taller ground flora species. Damage to oak saplings was assessed from the form (height and canopy) relative to stem diameter. The height and average canopy diameter of similar-aged saplings were found to be significantly greater in ungrazed (control) than both continuous and rotationally grazed treatments. Height and canopy diameters of similar-aged oak were not significantly different between the two grazing treatments. Significant differences were observed in tree form, with unbrowsed saplings having the greatest height to canopy width ratio and those in the continuously browsed plots having the smallest. An obvious decrease in the goodness of fit (R2) of regression analyses were found in continuously grazed areas compared to rotational and control plots for both height and canopy data. The differences in damage observed were not significantly different enough to suggest one method of grazing over the other as being better for minimising sapling damage. Management requirements are more likely to dictate grazing regime. Overall, stock density is likely to have a more significant effect on damage than stocking system.

Cumulative effects of wild ungulate and livestock herbivory on riparian willows.

We examined the effects of wild ungulates (deer and elk) and domestic sheep browsing on the growth, structure, and reproductive effort of two common willow species, Salix boothii and S. geyeriana, in a montane northeast Oregon riparian zone. With the use of exclosures, large herbivore effects on willows were studied in an area browsed by native mammals only and an adjacent area in which domestic sheep also lightly grazed during summer months. Growth variables were repeatedly measured on individual plants over a 5-year period to understand physiognomic and flowering responses of native willows to different levels of browsing pressure. At the beginning of the study, all willows were intensely browsed but were significantly taller in the area browsed only by native mammals than in the area also grazed by sheep (69 versus 51cm, respectively). Willows inside exclosures responded with pronounced increases in height, crown area, and basal stem diameters while the stature of browsed plants outside exclosures stayed constant or declined. In the area browsed by both sheep and wild herbivores, the size of browsed plants remained at pre-treatment levels (<60cm in height) for the duration of the study. There was no significant difference in growth rates of enclosed willows, indicating that current herbivory was the primary cause of growth retardation in the study area. Foliar area was strongly correlated with basal stem numbers for enclosed plants but much less so for browsed plants. Willows inside exclosures had more than twice as much foliar area per stem. Stem diameters were a positive function of crown area: stem-number ratios, suggesting lower photosynthetic potential was correlated with diminished radial growth among browsed plants. No flowering was observed until 2years after exclusion when plants inside all exclosures and browsed willows in the wild ungulate area responded with a large pulse in flowering. Browsed plants in the sheep + wild ungulate area did not flower. The number of catkins produced per plant was significantly associated with willow height and plants <70cm in height did not flower, thus suggesting a size threshold for reproduction in these species. Our results suggest that even relatively light levels of domestic livestock grazing, when coupled with intense wild ungulate browsing, can strongly affect plant structure and limit reproduction of riparian willows.

Effects of large herbivores and fire on the regeneration of Acacia erioloba woodlands in Chobe National Park, Botswana

Abstract Acacia erioloba woodlands provide important forage and shade for wildlife in northern Botswana. Mortality of mature trees caused by browsing elephants has been well documented but the lack of regeneration of new trees has received little attention. Annual growth of new shoots and changes in height were measured to determine the influence of elephants and small ungulate browsers, rainfall and fire on the growth and survival of established A. erioloba seedlings from 1995 to 1997 in the Savuti area of Chobe National Park. All above‐ground vegetation was removed from 40% of established seedlings in 1995 and 28% in 1997 by browsing elephants, and the mean height of remaining seedlings decreased from &gt;550 mm to &lt;300 mm. When seedlings browsed by kudu, impala and steenbok but not elephants are considered, mean seedling height increased &lt;50 mm per year, even though mean new shoot growth remaining at the end of the dry season was 100–200 mm. Fires burned portions of the study area in 1993 and 1997, killing above‐ground vegetation, but most established A. erioloba seedlings survived, producing coppice growth from roots. While elephants and fire caused the greatest reduction in established seedling height and number, small browsers suppressed growth, keeping seedlings vulnerable to fire and delaying growth to reproductive maturity.