Identifying threshold densities for wild deer in the UK above which negative impacts may occur

• 1 At high densities, deer populations may have adverse effects upon and within their environment. In this review we explore published and unpublished information to derive density thresholds for deer species in relation to impacts upon agriculture, forestry, conservation habitats, road traffic, and human and livestock health in the UK. Impact levels are affected by many factors other than absolute density. We therefore seek to establish the range of densities within which negative impacts might start to occur and which should trigger objective monitoring of actual impacts.
• 2 In commercial forestry, a threshold of 4 deer per 100ha has been suggested. Unfenced native woodlands seem to regenerate naturally if there are fewer than 4–5 large deer or fewer than 25 roe deer Capreolus capreolus per 100ha; open habitats may suffer only light or moderate impacts from red deer Cervus elaphus at landscape densities of 7–8 per 100ha.
• 3 Woodland bird species may have declined where deer densities are high but absolute thresholds seem impossible to establish. One study suggests maximum diversity at about 8 white‐tailed deer Odocoileus virginianus per 100ha.
• 4 Deer–vehicle collisions are affected by various factors in addition to deer density, but British and American studies suggest that accident frequencies decline at densities below 7–8 per 100ha.
• 5 Fallow deer Dama dama populations may maintain bovine TB (bTB) infection at much lower densities (25/100ha) than red or roe deer (91/100ha and 200/100ha, respectively) assuming 100% prevalence. Even at 30% prevalence a density of 75 fallow deer per 100ha could maintain bTB within the population.
• 6 We conclude that deer density alone is unlikely to be a good predictor of impact, and suggest that long‐term management should be based on assessment both of actual impacts and apparent density of deer.

     

Appropriate vegetation indices for measuring the impacts of deer on forest ecosystems

Increasing deer density can cause serious degradation of forests in the Americas, Europe, and Asia. To manage deer impacts, evaluating their current impacts on forest ecosystems is necessary, usually via vegetation indices. However, the relationship between vegetation indices and absolute deer density, while taking into account tree size, snow depth, light condition, and the type of understory vegetation, has never been investigated. We examined the relationship between various vegetation indices and absolute deer density in 344 study plots in the deciduous broad-leaved forest of Yamanashi Prefecture, central Japan. In each plot, debarking and browsing, along with the coverage and maximum height of understory vegetation, were surveyed. Estimated deer densities for 82 5 × 5-km mesh units ranged from 0.8 deer/km² to 32.7 deer/km². The percentages of debarked trees within a plot ranged from 0 to 84%. Debarking was promoted by high deer density, small tree size, and thick snow. The effect of tree size on debarking was stronger than that of deer density. Occurrence of browsing on understory vegetation was higher at higher deer densities, and where understory vegetation was dominated by evergreen dwarf bamboo. Coverage and maximum height of understory vegetation were unaffected by deer density but increased with canopy openness and the dominance of dwarf bamboo in the understory. Overall, we predict that debarking of small trees living in heavy snow areas should occur even at low deer densities (<10 deer/km²). Browsing on dwarf bamboo should occur at intermediate deer densities (10–30 deer/km²), while debarking of thick trees living in low snow areas should occur only at high deer densities (≥30 deer/km²). Our study shows that debarking and browsing on understory vegetation are appropriate indices for evaluating deer impacts on forest ecosystems, but that tree size, snow depth, and the type of understory vegetation should also be considered.     

Expanding and invading plant species in sagebrush steppe affect multiple aspects of small-mammal ecology

Invasion and expansion of non-native and native plants have altered vegetation structure in many terrestrial ecosystems. Small mammals influence multiple ecosystem processes through their roles as ecosystem engineers, predators, and prey, and changes to vegetation structure can affect habitat use, community composition, and predator-prey interactions for this assemblage of wildlife. In the sagebrush (Artemisia spp.) shrublands of the western United States, invasion by non-native grasses and expansion of native conifer trees beyond their historical range has altered vegetation structure. These changes may potentially affect distributions and interactions of deer mice (Peromyscus maniculatus), which are generalist omnivores, and Columbia Plateau pocket mice (Perognathus parvus), more specialized granivores. To assess the extent to which altered habitat affects small-mammal density, survival, and home-range size, we examined these aspects of small-mammal ecology along a gradient of cheatgrass (Bromus tectorum) invasion and western juniper (Juniperus occidentalis) establishment in sagebrush shrublands in southwestern Idaho, USA. From 2017–2019, we used a spatially explicit mark-recapture design to examine attributes of small-mammal ecology along an invasion gradient. We did not find support for an effect of cheatgrass cover on density or survival of either species. Home-range size of deer mice was 2.3 times smaller in heavier cheatgrass cover (60%) compared to areas with little or no cheatgrass cover. Density of deer mice was highest (5 individuals/ha) in areas with 10% juniper cover and decreased with increasing juniper cover, whereas density of pocket mice was positively influenced by shrub cover. Survival of deer mice declined as juniper stem density increased. Conversely, survival of pocket mice increased with increasing juniper stem density. We found evidence for interspecific interactions between these 2 species, in the form of a density-dependent effect of deer mice on pocket mouse home-range size. Home-range size for pocket mice was 2 times smaller in areas with the highest estimated density of deer mice compared to areas with low densities of deer mice. Our data provides unique information about how small mammals in the sagebrush steppe are affected by expanding and invasive plant species and potential ways that habitat restoration efforts, in the form of conifer removal, may influence small mammals. Understanding the response of small mammals to conifer expansion or removal may shed light on the demographic and numerical responses of other wildlife associated with the sagebrush biome, including predators.     

Predation has a greater impact in less productive environments: variation in roe deer, Capreolus capreolus, population density across Europe

Aim We aimed to describe the large‐scale patterns in population density of roe deer Caprelous capreolus in Europe and to determine the factors shaping variation in their abundance. Location Europe. Methods We collated data on roe deer population density from 72 localities spanning 25° latitude and 48° longitude and analysed them in relation to a range of environmental factors: vegetation productivity (approximated by the fraction of photosynthetically active radiation) and forest cover as proxies for food supply, winter severity, summer drought and presence or absence of large predators (wolf, Canis lupus, and Eurasian lynx, Lynx lynx), hunter harvest and a competitor (red deer, Cervus elaphus). Results Roe deer abundance increased with the overall productivity of vegetation cover and with lower forest cover (sparser forest cover means that a higher proportion of overall plant productivity is allocated to ground vegetation and thus is available to roe deer). The effect of large predators was relatively weak in highly productive environments and in regions with mild climate, but increased markedly in regions with low vegetation productivity and harsh winters. Other potentially limiting factors (hunting, summer drought and competition with red deer) had no significant impact on roe deer abundance. Main conclusions The analyses revealed the combined effect of bottom‐up and top‐down control on roe deer: on a biogeographical scale, population abundance of roe deer has been shaped by food‐related factors and large predators, with additive effects of the two species of predators. The results have implications for management of roe deer populations in Europe. First, an increase in roe deer abundance can be expected as environmental productivity increases due to climate change. Secondly, recovery plans for large carnivores should take environmental productivity and winter severity into account when predicting their impact on prey.     

Big game or big loss? High deer densities are threatening woody plant diversity and vegetation dynamics

Land-use change and current game management have favored an increasing population of wild ungulates (especially deer) in many regions of the Northern Hemisphere. Here, we assess the impact of high deer densities (>30 ind km^?2) on the highly diverse woody vegetation of Mediterranean ecosystems, where big game have been favored for the last decades. We examine whether prolonged deer browsing (over 30 years) affected plant composition, diversity and dynamics of the original (non-browsed) vegetation. Deer browsing led to an average decrease of 30.4 % in woody plant diversity (species richness), due to a lack of regeneration for the most preferred plant species. Species belonging to early stages of succession (mostly Labiatae and Cistaceae) were non-preferred by deer. Conversely, the most preferred species belonged mainly to late stages of plant succession. Deer impact on Mediterranean shrublands is causing biotic homogenization of plant communities and is threatening vegetation dynamics by forcing it to return to earlier succession stages. Strict deer population control favouring larger trophies but lower offspring numbers together with an adequate habitat management (increasing grass and acorn availability) would be the most efficient measures to reverse this diversity loss. Restoration work seems only appropriate for the most vulnerable species. We highlight the need of sampling deer-free areas with low or null historical browsing to assess the real impact of deer on woody plant diversity and vegetation dynamics.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Germination of native and introduced plants from scats of Fallow Deer (Dama dama) and Eastern Grey Kangaroo (Macropus giganteus) in a south‐eastern Australian woodland landscape

Introduced deer occur in many forests and woodlands in Australia and potentially play an important role in influencing the floristics and structure of these landscapes through eating plants and disseminating seeds. In a glasshouse trial, we tested whether field‐collected scats of Fallow Deer (Dama dama) contained viable plant seeds. Scats of deer obtained from a woodland study area in Kosciuszko National Park, New South Wales, produced seedlings from a range of native and introduced plant species. Forbs and herbs were dominant in these samples, confirming the grazing behaviour of deer at the time scats were collected. Samples of scats from Eastern Grey Kangaroos (Macropus giganteus), collected contemporaneously from the same sites as deer scats, also produced plant germinants. By volume, deer scats produced a greater diversity of plant germinants, including native and weed species, than did kangaroo scats. Although no weed species emanating from deer or kangaroo scats were of national significance, several species were of regional environmental significance, including Common Mullein (Verbascum thapsis), which was only found germinating out of deer scat, Stinkgrass (Eragrostis cilianensis) and Purpletop (Verbena bonariensis). In addition to dispersing viable seeds, Fallow Deer may also influence vegetation structure through their browsing. Further research is necessary to elucidate their respective role in dispersing native and introduced plants as well as any impacts that foraging behaviour might be having on woodland landscapes, to better inform management of the resident deer population.     

Suppression of Regeneration in New Zealand Mountain Beech Forests is Dependent on Species of Introduced Deer

We compared the impacts on forest regeneration of introduced sika (Cervus nippon) and red (Cervus elaphus) deer in New Zealand. Plot data were used to compare mountain beech (Nothofagus solandri var. cliffortioides) regeneration between a region with sika deer, and four regions without sika deer. All regions surveyed had red deer present. In the region where sika deer had been present for more than a decade, there was evidence of poor mountain beech seedling regeneration. In the four regions without sika deer, there was evidence of a strong regenerative response at stands with low occupancy by trees. When compared to larger deer species, sika deer have a digestive morphology allowing greater dietary versatility, which may result in them impeding forest regeneration where red deer do not. In contrast to mountain beech, some small-leaved shrub species may have been competitively advantaged by intensive browsing from sika deer. This is contra to a current view that small-leaved shrub species with interlacing branches were able to tolerate browsing from extinct ratite birds, but not introduced deer. Sika deer have been introduced into countries where other deer species are indigenous, such as Canada, Denmark, Great Britain, Czech Republic, Ireland and the United States of America. Because of their dietary advantage, sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites where impacts on tree regeneration are likely to be greatest.     

Test of Localized Management for Reducing Deer Browsing in Forest Regeneration Areas

ABSTRACT White-tailed deer (Odocoileus virginianus) browsing in forest regeneration sites can affect current and future stand structure and species composition. Removal of deer social units (localized management) has been proposed as a strategy to alleviate deer overbrowsing in forest systems. We conducted an experimental localized removal in a high-density deer population in the central Appalachians of West Virginia, USA, during winter 2002. We removed 51 deer within a 1.1-km² area that encompassed 2 forest regeneration sites (14 ha). During the summer following removal, we detected decreases in distance from the removal area in 8 of 30 (26.7%) adult females having pretreatment mean telemetry locations <2.5 km from the center of the removal area. We measured browsing rates during the summers of 2001–2004 from forest regeneration sites to examine efficacy of localized management. Browsing rates declined annually in both removal and control areas, due in part to increased timber harvesting on the larger study site, suggesting that increasing forage availability may be more effective at reducing impacts on forest regeneration than localized reductions in deer populations. Three years after the initial removal, we removed an additional 31 deer from the original 1.1-km² removal area. Home range shifts of adjacent deer coupled with the large number of animals collected in the second removal suggests that localized management only produces temporary voids within high-density deer herds. Localized management may not effectively reduce negative impacts of deer in areas of high deer density.     

Short-term effects of reduced white-tailed deer density on insect communities in a strongly overbrowsed boreal forest ecosystem

Browsing by overabundant deer modifies plant communities and alters forest regeneration, which can indirectly impact associated insect fauna. We tested the hypothesis that the response of insect communities to changes in deer abundance should depend on the strength of their association with plants, which we considered as a key functional trait. Seven years after a deer density control experiment was established in partly harvested forests on Anticosti Island (Quebec, Canada), we evaluated the effects of reducing white-tailed deer (Odocoileus virginianus) density from >20 down to 15, 7.5 and 0 deer km^?2, on four insect taxa representing different levels of dependence on plants. As predicted by our hypothesis, the sensitivity of insect taxa to deer density decreased along a gradient representing their degree of association with plants. Carabidae remained unaffected, while Apoidea and Syrphidae communities differed between uncontrolled and reduced deer densities, but not as clearly as for Lepidoptera. As expected, insect communities responded faster in harvested than in forested areas because vegetation changes more rapidly in open habitats. For most insect taxa, dominant species were the most strongly affected by deer density reduction, but it was clearly stronger for predator taxa (Syrphidae and Carabidae). A fast recovery of rare species was observed for macro Lepidoptera. Reducing deer density down to 15 deer km^?2 is sufficient to restore insect diversity on Anticosti Island, but it is unlikely to be efficient in all situations, particularly when competing tree regeneration is firmly established.     

Getting the biggest birch for the bang: restoring and expanding upland birchwoods in the Scottish Highlands by managing red deer

High deer populations threaten the conservation value of woodlands and grasslands, but predicting the success of deer culling, in terms of allowing vegetation to recover, is difficult. Numerical simulation modeling is one approach to gain insight into the outcomes of management scenarios. We develop a spatially explicit model to predict the responses of Betula spp. to red deer (Cervus elaphus) and land management in the Scottish Highlands. Our model integrates a Bayesian stochastic stage‐based matrix model within the framework of a widely used individual‐based forest simulation model, using data collected along spatial and temporal gradients in deer browsing. By initializing our model with the historical spatial locations of trees, we find that densities of juvenile trees (<3 m tall) predicted after 9–13 years closely match counts observed in the field. This is among the first tests of the accuracy of a dynamical simulation model for predicting the responses of tree regeneration to herbivores. We then test the relative importance of deer browsing, ground cover vegetation, and seed availability in facilitating landscape‐level birch regeneration using simulations in which we varied these three variables. We find that deer primarily control transitions of birch to taller (>3 m) height tiers over 30 years, but regeneration also requires suitable ground cover for seedling establishment. Densities of adult seed sources did not influence regeneration, nor did an active management scenario where we altered the spatial configuration of adults by creating “woodland islets”. Our results show that managers interested in maximizing tree regeneration cannot simply reduce deer densities but must also improve ground cover for seedling establishment, and the model we develop now enables managers to quantify explicitly how much both these factors need to be altered. More broadly, our findings emphasize the need for land managers to consider the impacts of large herbivores rather than their densities.     

Linking direct and indirect pathways mediating earthworms, deer, and understory composition in Great Lakes forests

Ahistorical drivers such as nonnative invasive earthworms and high deer densities can have substantial impacts on ecosystem processes and plant community composition in temperate and boreal forests of North America. To assess the roles of earthworm disturbance, deer, and environmental factors in the understory, we sampled 125 mixed temperate-boreal forest sites across the western Great Lakes region. We utilized structural equation modeling (SEM) to address the hypothesis that earthworm disturbance to the upper soil horizons and selective herbivory by deer are associated with depauperate understory plant communities dominated by graminoid and nonnative species. Evidence of earthworm activity was found at 93 % of our sites and 49 % had high to very high severity earthworm disturbance. The SEM fit the data well and indicated that widespread nonnative earthworm disturbance and high deer densities had similar magnitudes of impact on understory plant communities and that these impacts were partially mediated by environmental characteristics. One-third of the variation in earthworm disturbance was explained by soil pH, precipitation, and litter quality. Deer density and earthworm disturbance both increased graminoid cover while environmental variables showed direct and indirect relationships. For example, the positive relationship between temperature and graminoids was indirect through a positive temperature effect on deer density. This research characterizes an integrated set of key environmental variables driving earthworm disturbance and deer impacts on the forest understory, facilitating predictions of the locations and severity of future change in northern temperate and boreal forest ecosystems.     

Recovery of a dwarf birch (Betula nana) population following reduction in grazing by red deer (Cervus elaphus)

Summary

In Britain Betula nana is a nationally scarce plant, its distribution being limited by climate, heather burning and grazing by deer and sheep. The recovery of a population of B. nana was monitored for several years inside a Scots pine regeneration exclosure in N.E. Scotland. Browsing pressure was reduced in the exclosure but a few red deer (amounting to 3–4 km^?2) were deliberately retained. B. nana stem density increased after red-deer density was reduced. Over the same period there was a reduction in browsing inside the exclosure. B. nana stem density was negatively related to heather height, deer presence and tree canopy cover. Comparisons of growth were made with plants outside the exclosure between 1998 and 2000, the mean annual increment of stems, allowing for browsing loss, being respectively 1.25 and 0.45 cm for inside and outside the exclosure. Most stem basal diameters inside the exclosure were smaller than outside. Reducing red deer density to 3–4 km^?2 on similar upland sites with peaty soils could lead to regeneration of B. nana.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Influence of Landscape Composition and Structure on Antler Size of White-Tailed Deer

Abstract: Cause for spatial variation in phenotypic quality of white-tailed deer (Odocoileus virginianus) populations is of great interest to wildlife managers. Relating phenotypic variation of populations to large-scale land-use patterns may provide insight into why populations exhibit spatial variation and elucidate how management can influence population phenotype. We used an information-theoretic approach to relate average antler size of 203 deer populations to composition and structure of the habitat occupied by those populations. We used interspersion, edge, and diversity indices to represent habitat structure and percentage of area in vegetation types to represent habitat composition. Landscape composition was a better predictor of deer population antler size than was landscape structure. Percentages of the management unit in agriculture, pasture, and pine forest were variables commonly found in the region-specific set of best models. Model-averaged estimates of agriculture and pasture parameters were always positive and estimates of pine forest parameters were always negative, which suggests that land-use types that promote growth of early successional herbaceous plants will positively influence antler size and, most likely, body growth and reproduction of white-tailed deer populations. Conversely, our findings suggest landscapes dominated by pine forests did not provide optimal amounts of quality forages for white-tailed deer. Pine forest effects should be mitigated using a combination of increased harvest to lower deer density and silvicultural practices like thinning, prescribed burning, and selective herbicide applications that stimulate growth of high-quality forages beneath the forest canopy to improve deer phenotypic quality.     

Deer do not affect short‐term rates of vegetation recovery in overwash fans on Fire Island after Hurricane Sandy

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Additive Partitioning of Diversity Reveals No Scale-dependent Impacts of Large Ungulates on the Structure of Tundra Plant Communities

Large herbivores can change ecosystem functioning by impacting plant diversity. However, although such impacts are expected to be scale-dependent in ecosystems with wide-roaming ungulates, scaling issues rarely enter empirical assessments. We here test the hypothesis that the impact of increased reindeer abundance on plant diversity in alpine tundra is scale-dependent. Based on potentially high productivity of the focal habitat units and hence the possibility of positive grazer impacts on plant diversity we predicted higher α and β diversity at the habitat scale where reindeer densities are high. We also explored whether there were differences in diversity patterns at larger scales, including the scale of reindeer management districts. We estimated grazing disturbance as high versus low reindeer density in selected districts (a total extent of 7421 km²) of Northern Norway where reindeer-induced vegetation shifts are debated. We focus on dominance patterns because they can quantify the vegetation state and thus performed additive partitioning of Simpson diversity on multiple scales assessing also species’ contributions to diversity. Contrary to our predictions, we found only weak scale-dependent effects of reindeer grazing on plant diversity. Under high reindeer densities there was evidence for a landscape-scale homogenization of the vegetation, but the predicted α and β diversity increases at the habitat scale were not found. Consistently through all scales considered, four shrub species contributed the most to plant diversity. These results contradict the idea that reindeer at high stocking densities induce shifts in plant species dominance in productive habitats. We conclude that context-dependencies such as spatial scales of management units and habitat types need to be explicitly considered in evaluations of the impacts of large ungulates on plant diversity.     

Deer exclusion unveils abiotic filtering in forest understorey plant assemblages

Background and AimsThe 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.MethodsWe 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.Key ResultsOur 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.ConclusionsWe 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.     

Windows of opportunity: white-tailed deer and the dynamics of northern hardwood forests of the northeastern US

Herbivory, lighting regimes, and site conditions are among the most important determinants of forest regeneration success, but these are affected by a host of other factors such as weather, predation, human exploitation, pathogens, wind and fire. We draw together >50 years of research on the Huntington Wildlife Forest in the central Adirondack Mountains of New York to explore regeneration of northern hardwoods. A series of studies each of which focused on a single factor failed to identify the cause of regeneration failure. However, integration of these studies led to broader understanding of the process of forest stand development and identified at least three interacting factors: lighting regime, competing vegetation and selective browsing by white-tailed deer (Odocoileus virginianus). The diverse 100–200 year-old hardwood stands present today probably reflect regeneration during periods of low deer density (<2.0 deer/km²) and significant forest disturbance. If this hypothesis is correct, forest managers can mimic these “natural windows of opportunity” through manipulation of a few sensitive variables in the system. Further, these manipulations can be conducted on a relatively small geographic scale. Control of deer densities on a scale of 500 ha and understory American beech (Fagus grandifolia) on a scale of <100 ha in conjunction with an even-aged regeneration system consistently resulted in successful establishment of desirable hardwood regeneration.     

Interactions between fire and introduced deer herbivory on coastal heath vegetation

The coastal heathlands of the Royal National Park are impacted by both fire and herbivory by introduced deer, and to date these two factors have been dealt with independently in the management of natural areas. In recent years, there has been increasing recognition for a more integrated approach to manage these two disturbance agents. Fire and its role in Australian heathlands are well known, while impacts from introduced deer and the combined effects of fire and introduced deer are still poorly understood. In this study, we investigated the effects of fire and Javan rusa deer (Cervus timorensis) on both vegetation cover and floristics. The percentage cover of plants at different height layers and the presence/absence of individual species were recorded at sites representing two different burn histories (1993/1994 and 2000/2001) and deer presence or absence. Fire significantly reduced vegetation cover at low (<50 cm) and intermediate heights (50–100 cm), while deer presence affected grasses and sedges, and low vegetation at more recently burnt sites. Rusa deer also affected composition of the plant species assemblages, but no such effect was found for fire. Understanding the influence of each disturbance factor independently and together in the heathlands will be critical for implementing a more robust framework for future management.