Introduced deer reduce native plant cover and facilitate invasion of non-native tree species: evidence for invasional meltdown

Invasive species are a major threat to native communities and ecosystems worldwide. One factor frequently invoked to explain the invasiveness of exotic species is their release in the new habitat from control by natural enemies (enemy-release hypothesis). More recently, interactions between exotic species have been proposed as a potential mechanism to facilitate invasions (invasional meltdown hypothesis). We studied the effects of introduced deer on native plant communities and exotic plant species on an island in Patagonia, Argentina using five 400 m² exclosures paired with control areas in an Austrocedrus chilensis native forest stand. We hypothesized that introduced deer modify native understory composition and abundance and facilitate invasion of introduced tree species that have been widely planted in the region. After 4 years of deer exclusion, native Austrocedrus and exotic Pseudotsuga menziesii tree sapling abundances are not different inside and outside exclosures. However, deer browsing has strongly inhibited growth of native tree saplings (relative height growth is 77% lower with deer present), while exotic tree sapling growth is less affected (relative height growth is 3.3% lower). Deer significantly change abundance and composition of native understory plants. Cover of native plants in exclosures increased while cover in controls remained constant. Understory composition in exclosures after only 4 years differs greatly from that in controls, mainly owing to the abundance of highly-browsed native species. This study shows that introduced deer can aid the invasion of non-native tree species through negatively affecting native plant species.     

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.     

Herbivore-induced expansion of generalist species as a driver of homogenization in post-disturbance plant communities

We used a controlled experiment to investigate how disturbance scale (canopy gap area) and herbivory influence post-disturbance plant community dynamics. Twenty canopy gaps were installed in a temperate hemlock-hardwood forest during the winter of 2002–2003: seven small gaps (50–150 m²), seven medium gaps (151–250 m²), and six large gaps (251–450 m²). Within each gap, we established 4–12 sample plots (depending on gap size); 1–3 of which were enclosed with wire mesh white-tailed deer (Odocoileus virginianus) exclosures in 2005. Gaps were revisited and intensively sampled in 2007. After five growing seasons, ground-layer plant communities in non-exclosed plots were more similar compositionally than exclosure plots. Non-exclosed plots in small and medium gaps were more similar to non-exclosed plots in large gaps than they were to exclosed plots in their respective gap size class. Shade-tolerant forbs and trees were less common outside exclosures, while generalist species associated with higher understory light levels and exotics were more prevalent outside exclosures. Our results suggest that even in forests with relatively low deer densities (6.5–9.3 deer km⁻²), white-tailed deer herbivory may influence the developmental trajectory of post-disturbance plant communities and be a mechanism for decreasing β-diversity along environmental gradients.     

Long-Term Effects of White-Tailed Deer Exclusion on the Invasion of Exotic Plants: A Case Study in a Mid-Atlantic Temperate Forest

Exotic plant invasions and chronic high levels of herbivory are two of the major biotic stressors impacting temperate forest ecosystems in eastern North America, and the two problems are often linked. We used a 4-ha deer exclosure maintained since 1991 to examine the influence of a generalist herbivore, white-tailed deer (Odocoileus virginianus), on the abundance of four exotic invasive (Rosa multiflora, Berberis thunbergii, Rubus phoenicolasius and Microstegium vimineum) and one native (Cynoglossum virginianum) plant species, within a 25.6-ha mature temperate forest dynamics plot in Virginia, USA. We identified significant predictors of the abundance of each focal species using generalized linear models incorporating 10 environmental and landscape variables. After controlling for those predictors, we applied our models to a 4-ha deer exclusion site and a 4-ha reference site, both embedded within the larger plot, to test the role of deer on the abundance of the focal species. Slope, edge effects and soil pH were the most frequent predictors of the abundance of the focal species on the larger plot. The abundance of C. virginianum, known to be deer-dispersed, was significantly lower in the exclosure. Similar patterns were detected for B. thunbergii, R. phoenicolasius and M. vimineum, whereas R. multiflora was more abundant within the exclosure. Our results indicate that chronic high deer density facilitates increased abundances of several exotic invasive plant species, with the notable exception of R. multiflora. We infer that the invasion of many exotic plant species that are browse-tolerant to white-tailed deer could be limited by reducing deer populations.     

Impacts of feral horses and deer on an endangered woodland of Kosciuszko National Park

In recent years, the impacts of rapidly increasing populations of feral horses and deer on the vegetation and stability of soils have become highly visible and widespread in Kosciuszko National Park. We investigated these impacts in the White Cypress Pine (Callitris glaucophylla Joy Thomps. & L.A.S. Johnson) – White Box (Eucalyptus albens Benth) woodlands of the lower Snowy River valley. This woodland is a component of the White Box‐Yellow Box‐Blakely's Red Gum Grassy Woodland and Derived Native Grasslands complex that is nationally listed as a critically endangered ecological community. To investigate the severity of the impacts of feral horse (Equus caballus) and deer (Dama dama and Rusa unicolor) in the valley in 2013 and 2017/18, we surveyed fenced exclosures and paired grazed plots that were first established and surveyed in 1984 and re‐surveyed in 1987. Using LFA and VAST methodologies (not used in 1987), we compared the relative response of environmental variables in plots inside and outside the exclosures in an attempt to ascertain recent herbivore impacts. While there was no evidence of horses or deer from dung surveys in 1987, in 2018, 84% of the dung was from horses, 13% from deer, 1% from rabbits and 2% from macropods. Total herbivore dung density increased fourfold since the 1987 survey. On the understanding that all plots had the same starting condition in 1984 with respect to prior herbivory, we deduce that horses and deer are having significant ecological impacts. There was a far greater cover of understorey plants and the midstorey was denser and taller inside the exclosures. Outside the exclosures, the vegetation cover was far more sparse and soil erosion was active and extensive. The total number of invertebrates captured in small pitfall traps was nearly twice as many within the exclosures compared to the grazed plots. The dense even‐aged regrowth overstorey stands of White Cypress Pine, inside and outside the exclosures, have changed little in 34 years.     

Resource availability and plant diversity explain patterns of invasion of an exotic grass

Aims In this study, we examine two common invasion biology hypotheses—biotic resistance and fluctuating resource availability—to explain the patterns of invasion of an invasive grass, Microstegium vimineum.Methods We used 13-year-old deer exclosures in Great Smoky Mountains National Park, USA, to examine how chronic disturbance by deer browsing affects available resources, plant diversity, and invasion in an understory plant community. Using two replicate 1 m 2 plots in each deer browsed and unbrowsed area, we recorded each plant species present, the abundance per species, and the fractional percent cover of vegetation by the cover classes: herbaceous, woody, and graminoid. For each sample plot, we also estimated overstory canopy cover, soil moisture, total soil carbon and nitrogen, and soil pH as a measure of abiotic differences between plots.Important findings We found that plant community composition between chronically browsed and unbrowsed plots differed markedly. Plant diversity was 40% lower in browsed than in unbrowsed plots. At our sites, diversity explained 48% and woody plant cover 35% of the variation in M. vimineum abundance. In addition, we found 3.3 times less M. vimineum in the unbrowsed plots due to higher woody plant cover and plant diversity than in the browsed plots. A parsimonious explanation of these results indicate that disturbances such as herbivory may elicit multiple conditions, namely releasing available resources such as open space, light, and decreasing plant diversity, which may facilitate the proliferation of an invasive species. Finally, by testing two different hypotheses, this study addresses more recent calls to incorporate multiple hypotheses into research attempting to explain plant invasion.     

Impact of Habitat-Specific GPS Positional Error on Detection of Movement Scales by First-Passage Time Analysis

Advances in animal tracking technologies have reduced but not eliminated positional error. While aware of such inherent error, scientists often proceed with analyses that assume exact locations. The results of such analyses then represent one realization in a distribution of possible outcomes. Evaluating results within the context of that distribution can strengthen or weaken our confidence in conclusions drawn from the analysis in question. We evaluated the habitat-specific positional error of stationary GPS collars placed under a range of vegetation conditions that produced a gradient of canopy cover. We explored how variation of positional error in different vegetation cover types affects a researcher''s ability to discern scales of movement in analyses of first-passage time for white-tailed deer (Odocoileus virginianus). We placed 11 GPS collars in 4 different vegetative canopy cover types classified as the proportion of cover above the collar (0–25%, 26–50%, 51–75%, and 76–100%). We simulated the effect of positional error on individual movement paths using cover-specific error distributions at each location. The different cover classes did not introduce any directional bias in positional observations (1 m≤mean≤6.51 m, 0.24≤p≤0.47), but the standard deviation of positional error of fixes increased significantly with increasing canopy cover class for the 0–25%, 26–50%, 51–75% classes (SD = 2.18 m, 3.07 m, and 4.61 m, respectively) and then leveled off in the 76–100% cover class (SD = 4.43 m). We then added cover-specific positional errors to individual deer movement paths and conducted first-passage time analyses on the noisy and original paths. First-passage time analyses were robust to habitat-specific error in a forest-agriculture landscape. For deer in a fragmented forest-agriculture environment, and species that move across similar geographic extents, we suggest that first-passage time analysis is robust with regard to positional errors.     

Deer Herbivory as an Ecological Constraint to Restoration of Degraded Riparian Corridors

Ungulate herbivory can impact riparian vegetation in several ways, such as by reducing vigor or reproductive output of mature plants, and through increased mortality of seedlings and saplings. Much work has focused on the effects of livestock grazing within riparian corridors, while few studies have addressed the influence of native ungulate herbivory on riparian vegetation. This study investigated the effect of deer herbivory on riparian regeneration along three streams with degraded riparian corridors in Mendocino County, California. We utilized existing stream restoration efforts by private landowners and natural resource agencies to compare six deer exclosures with six upstream control plots. Livestock were excluded from both exclosure and control plots. Three of the deer exclosures had been in place for 15 years, one for 6 years, and two for 4 years. The abundance and size distribution of woody riparian plant species such as Salix exigua, S. laevigata, S. lasiolepis, Alnus rhombifolia, and Fraxinus latifolia were quantified for each exclosure and control plot. The mean density of saplings in deer exclosures was 0.49 ± 0.15/m², while the mean density of saplings in control plots was 0.05 ± 0.02/m². Within exclosures, 35% of saplings were less than 1 m and 65% were greater than 1 m; within control plots, 97% of saplings were less than 1 m in height. The fact that little regeneration had occurred in control plots suggests that deer herbivory can substantially reduce the rate of recovery of woody riparian species within degraded riparian corridors. Exclusionary fencing has demonstrated promising results for riparian restoration in a region with intense deer herbivory.     

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.     

Clonal growth and plant species abundance

Background and Aims

Both regional and local plant abundances are driven by species'' dispersal capacities and their abilities to exploit new habitats and persist there. These processes are affected by clonal growth, which is difficult to evaluate and compare across large numbers of species. This study assessed the influence of clonal reproduction on local and regional abundances of a large set of species and compared the predictive power of morphologically defined traits of clonal growth with data on actual clonal growth from a botanical garden. The role of clonal growth was compared with the effects of seed reproduction, habitat requirements and growth, proxied both by LHS (leaf–height–seed) traits and by actual performance in the botanical garden.

Methods

Morphological parameters of clonal growth, actual clonal reproduction in the garden and LHS traits (leaf-specific area – height – seed mass) were used as predictors of species abundance, both regional (number of species records in the Czech Republic) and local (mean species cover in vegetation records) for 836 perennial herbaceous species. Species differences in habitat requirements were accounted for by classifying the dataset by habitat type and also by using Ellenberg indicator values as covariates.

Key Results

After habitat differences were accounted for, clonal growth parameters explained an important part of variation in species abundance, both at regional and at local levels. At both levels, both greater vegetative growth in cultivation and greater lateral expansion trait values were correlated with higher abundance. Seed reproduction had weaker effects, being positive at the regional level and negative at the local level.

Conclusions

Morphologically defined traits are predictive of species abundance, and it is concluded that simultaneous investigation of several such traits can help develop hypotheses on specific processes (e.g. avoidance of self-competition, support of offspring) potentially underlying clonal growth effects on abundance. Garden performance parameters provide a practical approach to assessing the roles of clonal growth morphological traits (and LHS traits) for large sets of species.     

Consumers and establishment limitations contribute more than competitive interactions in sustaining dominance of the exotic herb garlic mustard in a Wisconsin, USA forest

The understory is a diverse component of temperate forest ecosystems, contributing significantly to forest ecosystem services. Despite their importance, many native understories face stresses from current and past land use, habitat fragmentation, invasive species, and overabundant herbivores. We established a four block, three factor experiment to evaluate the relative contribution of native plant establishment, competitive effects from the invasive herb garlic mustard (Alliaria petiolata), and herbivory from white-tailed deer (Odocoileus virginianus) to better understand the mechanisms promoting low native plant richness and cover and understory dominance by the biennial exotic herb garlic mustard in a NE Wisconsin, USA forest. Four years of garlic mustard removal failed to increase native plant richness or cover in non-restored plots. However, deer access and the introduction of native plants (restoration treatment) both significantly enhanced native plant cover and richness, with restored species cover in fenced plots approximately 216 % that of open-access plots, and the majority of these species flowered at significantly higher proportions inside of fenced areas. In contrast, deer access did not significantly alter the cover, or seed production of garlic mustard. We also found no significant effect of garlic mustard presence on the cover or flowering of restored native species. We conclude that multiple factors, including limited natural establishment by native species and selective herbivory drove low native, high exotic dominance at our site, suggesting that a shift in focus from invasive plant removal to combined native plant restoration and herbivore control is needed to maximize the recovery of this degraded forest understory.     

Deer density affects ground-layer vegetation differently in conifer plantations and hardwood forests on the Boso Peninsula, Japan

Deer overabundance reduces forest ground-layer vegetation and can cause cascading impacts on a forest ecosystem. To predict these effects, we must elucidate the relationship between deer density and the status of ground-layer vegetation. This relationship was studied in the Boso Peninsula, where the deer population density exhibits a clear geographical gradient. We examined species richness and cover of ground-layer vegetation at several cedar plantations and hardwood forests in the Boso Peninsula. We also examined whether deer impacts were altered by light condition, soil water content and forest type (cedar and hardwood). Species richness of ground-layer vegetation was maximized at an intermediate level of deer density, suggesting an intermediate disturbance effect. This phenomenon was compatible with the observation that evergreen species, which were competitive dominants, decreased with increasing deer density, whereas less competitive deciduous species increased until herbivory was intermediate. As deer density increased, cover of ground-layer vegetation gradually decreased, but species unpalatable to deer increased in abundance, suggesting indirect positive effects of deer for unpalatable species. Cedar plantations tended to have greater species richness and ground-layer cover than hardwood forests with similar deer levels. Canopy openness, an indicator of light conditions, increased species richness of hardwood forests and ground-layer cover of cedar plantations, even under deer herbivory. Topographic wetness index, an indicator of soil water content, significantly increased the ground-layer cover of cedar plantations under deer herbivory. These results emphasize the importance of environmental productivity and forest type in the management of ground-layer vegetation experiencing deer overabundance.     

Equivalence in the strength of deer herbivory on above and below ground communities

Herbivores exert a strong influence on the species composition and richness of plant communities, but the magnitude of their effect on belowground communities remains poorly understood. While an increasing number of studies acknowledge the importance of documenting belowground effects of herbivores, very few of these evaluate variation in the strength of the response from aboveground to belowground communities. Our study documents the long-term consequences of sustained deer herbivory for plant and arthropod communities adjacent to 15 exclosures that have been in place since 1996. We hypothesized that herbivory would alter the composition and diversity of communities, but the strength of the effects of herbivory would weaken from plants, to leaf-litter invertebrates, and to belowground microarthropod communities. First, we found that herbivory negatively impacted plant seedling and sapling abundance and performance, reduced the abundance of ants and the taxonomic richness of arthropods in the litter layer and reduced the richness of soil microarthropod communities. Second, in contrast to our hypothesis, the magnitude of effect size did not vary among trophic levels, indicating that effects of deer herbivory cascade from plants to the leaf-litter and soil arthropod communities with equal strength. While much recent research has focused on how specific traits of plants may mediate the effects of herbivory on associated species, our results suggest that indirect effects of herbivory might influence many components of belowground communities.     

Regional-Scale Assessment of Deer Impacts on Vegetation Within Western Connecticut,USA

Abstract: High densities of white-tailed deer (Odocoileus virginianus) are believed to cause broad-scale forest regeneration failure and loss of plant diversity. But, the empirical basis for such presumptions is limited. We, therefore, conducted a survey in western Connecticut, USA, woodlots to examine how spatial variation in deer densities influences variation in impacts on plant species abundance, identity and diversity, and tree regeneration. We also used a Geographic Information System to quantify trends between land-cover type and deer density. Deer density was not correlated with any vegetation or land use variable. This suggests that deer density is not a leading factor determining variation in vegetation impacts across western Connecticut.     

The herb community of a tropical forest in central Panamá: dynamics and impact of mammalian herbivores

Mammals are hypothesized to either promote plant diversity by preventing competitive exclusion or limit diversity by reducing the abundance of sensitive plant species through their activities as browsers or disturbance agents. Previous studies of herbivore impacts in plant communities have focused on tree species and ignored the herbaceous community. In an experiment in mature-phase, tropical moist forest sites in central Panamá, we studied the impact of excluding ground-dwelling mammals on the richness and abundance of herbs in 16, 30×45-m plots. Within each plot, we censused the herbaceous community in 28, 2×2-m subplots (1,792 m² total area sampled). We identified over 54 species of herbs averaging 1.21 ramets m⁻² and covering approximately 4.25% of the forest floor. Excluding mammals for 5 years had no impact on overall species richness. Within exclosures, however, there was a significant two-fold increase in the density of rare species. Overall herbaceous density and percent cover did not differ between exclosures and adjacent control plots, although cover did increase over time. Mammalian exclusion significantly increased the total cover of three-dominant herb species, Pharus latifolius, Calathea inocephala, and Adiantum lucidum, but did not affect their density. This study represents one of the most extensive herbaceous community censuses conducted in tropical forests and is among a few that quantify herbaceous distribution and abundance in terms of both density and cover. Additionally, this work represents the first community level test of mammalian impacts on the herbaceous community in a tropical forest to date. Our results suggest that ground dwelling mammals do not play a key role in altering the relative abundance patterns of tropical herbs in the short term. Furthermore, our results contrast sharply with prior studies on similar temporal and spatial scales that demonstrate mammals strongly alter tree seedling composition and reduce seedling density. Thus, we question the pervasiveness of top–down control on tropical plant communities and the paradigm that defaunation will inexorably lead to widespread, catastrophic shifts in plant communities.     

Impact on plant communities by white-tailed deer in Mississippi,USA

Background: The abundance of white-tailed deer (Odocoileus virginianus) in the eastern United States has escalated during the twentieth century, potentially impacting plant communities.

Methods: We measured understorey plant cover and biomass five years after excluding deer from mature forests of three ecological regions in Mississippi, USA. We extended the significance of P values to 0.10 to detect developing impacts.

Results: Deer impacts were limited and varied by ecological region. We recorded 151 species in cover transects. Consistent exclosure treatment effects were detected in two regions where there was greater cover of two deer forages and less cover of three non-forages. Species richness was greater in exclosures in one region, but otherwise species richness and diversity indices did not differ. We recorded 127 species in biomass quadrats. Exclosure treatment effects on biomass were inconsistent. Out of five species with significant differences, three had more biomass in controls, including two deer forages. Except for greater total biomass in controls of one region, there were no differences by growth form or total vegetation for canopy coverage or biomass. Ordination of community canopy cover demonstrated similarity of paired exclosure and controls.

Conclusions: Exclosure treatment effects on canopy cover and species richness in two regions indicated limited negative impacts from deer foraging. A time frame of more than five years may be required for exclusion to allow recovery of vegetation, even with relatively open canopies and a long growing season.     

Bird predation on herbivorous insects: indirect effects on sugar maple saplings

Insectivorous birds have been shown to have direct effects on abundances of herbivorous arthropods, but few studies have tested the indirect effects of birds on plant performance through consumption of herbivorous insects. In a 3-year study at the Hubbard Brook Experimental Forest, New Hampshire, we tested whether bird predation indirectly affects leaf herbivory levels and leaf and shoot biomass production of understory sugar maple (Acer saccharum) saplings. Trees were randomly assigned to one of four treatments: an insecticide application to reduce herbivory levels, exclosures that prevented bird access, addition of Lepidoptera larvae, and controls. Trees sprayed with an insecticide supported significantly fewer Lepidoptera larvae than other treatments throughout the study. Also, trees in exclosures supported more Lepidoptera larvae than controls during one count each year, and pooled across all counts during the second year. As predicted, the mean proportion of leaf area consumed varied significantly among treatments and was least in the insecticide treatment, followed by controls, exclosures, and Lepidoptera additions. Significant differences among treatments in herbivory levels, however, did not lead to differences in leaf or shoot biomass production. Thus, bird predation decreased Lepidoptera abundances and decreased herbivory levels, but did not increase biomass production during the following year. Over 85% of the herbivores in our study were Homoptera nymphs that were not folivorous and are not important bird prey items, potentially dampening the indirect effects of bird predation on biomass production. A comparison of these results with previous studies suggests that the indirect effects of bird predation on plant biomass production may depend on the plant species, abundance and composition of the herbivore community, and primary productivity of the ecosystem.     

Herbivory on planted oak seedlings across a habitat edge created by timber harvest

Edge habitats create environmental gradients that affect plant community composition and herbivore behavior. Silvicultural disturbance creates edge habitat with direct (via changes in light) and indirect (via changes in herbivore behavior) consequences for the growth and survival of tree seedlings, and thus, the composition of the future forest stands. Herbivores, particularly ungulates, can be a major limiting factor in oak regeneration, and silvicultural disturbance may alter the abundance or behavior of herbivores following harvest. We measured the severity of herbivory on experimentally planted white (Quercus alba) and black oak (Quercus velutina) seedlings by white-tailed deer (Odocoileus virginianus) and eastern cottontail rabbits (Sylvilagus floridanus), as well as foliar damage from insects, across gradients created by clearcuts in a deciduous forest in Indiana, USA. Overall browse pressure on oaks was low in our study. Nonetheless, spatial variation in herbivory depended on herbivore taxa; herbivory by rabbits was highest inside harvest openings, whereas foliar damage by insects peaked in the forest. Intensity of deer herbivory was constant across the edge. In addition, we observed indirect interactions among herbivore species mediated by a seedling’s browsing history. Herbivore damage by deer was positively related to past browsing by rabbits, and foliar damage from insects was positively related to past browsing by both deer and rabbits. Increasing woody plant competition reduced herbivory on seedlings by both deer and rabbits. Given the lack of spatial variability in deer herbivory and low overall herbivory by rabbits, we suspect that interactions between timber harvesting and herbivory did not have a strong impact on oak seedlings at our study sites.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

Oral Vaccination of White-Tailed Deer (Odocoileus virginianus) with Mycobacterium bovis Bacillus Calmette-Guerin (BCG)

Wildlife reservoirs of Mycobacterium bovis represent serious obstacles to the eradication of tuberculosis from livestock, particularly cattle. In Michigan, USA tuberculous white-tailed deer transmit M. bovis to other deer and cattle. One approach in dealing with this wildlife reservoir is to vaccinate deer, thus interfering with the intraspecies and interspecies transmission cycles. Thirty-three white-tailed deer were assigned to one of two groups; oral vaccination with 1×10⁸ colony-forming units of M. bovis BCG Danish (n = 17); and non-vaccinated (n = 16). One hundred eleven days after vaccination deer were infected intratonsilarly with 300 colony-forming units of virulent M. bovis. At examination, 150 days after challenge, BCG vaccinated deer had fewer gross and microscopic lesions, fewer tissues from which M. bovis could be isolated, and fewer late stage granulomas with extensive liquefactive necrosis. Fewer lesions, especially those of a highly necrotic nature should decrease the potential for dissemination of M. bovis within the host and transmission to other susceptible hosts.