Impact of deer on temperate forest vegetation and woody debris as protection of forest regeneration against browsing

A 3-year field experiment with paired exclosure (fenced areas, excluding deer) and control plots (unfenced areas, free access to deer), with two treatments with and without woody debris, was carried out at two sites in a temperate forest in eastern France. The aim of the experiment was to assess the effect of browsing by roe deer (Capreolus capreolus) and red deer (Cervus elaphus) on the diversity and richness of plant species and to test the effectiveness of using woody debris to protect seedlings and saplings from deer browsing. In presence of deer, both plant species richness and diversity were reduced the first year of the study, but this negative impact of deer then disappeared after 3 years. Deer browsing mostly affected species composition of plant communities. We observed a decrease in the abundance of preferred species such as Carpinus betulus, Rubus fructicosus, Rubus idaeus, Anemone nemorosa and Epilobium angustifolium, and palatable species such as Acer spp., Carex spp., Festuca spp. and Mycelis muralis, whereas unpalatable species such as Lamium spp., or species particularly resilient to browsing such as grasses (Brachypodium spp. and Luzula spp.) increased in abundance. The use of woody debris as protection against browsing by deer did not limit damage to seedlings and saplings of the main commercially valuable species, Abies alba and Quercus spp. Instead of limiting deer impact, use of woody debris seemed to increase the negative effect of deer browsing on regeneration in control plots relatively to those without protection.     

Quantitative estimates of tree species selectivity by moose (Alces alces) in a forest landscape

Abstract

An extensive literature is available on browsing preference for certain tree species. However, useful predictive tools for estimating the impact of deer on forests production and biodiversity can still be improved. A step in that direction is not only to rank preference among tree species but also to quantify the relative risk of being browsed. The foraging selectivity of moose was evaluated using three different statistical methods developed to study habitat utilization. The general pattern for the three methods was consistent. From the results, groups of forage species were clustered and a quantitative index of selectivity was calculated for the groups. The selectivity index showed that rowan (Sorbus aucuparia), willow (Salix ssp.) and aspen (Populus tremula) had a 14 times higher probability of being browsed than a group consisting of Scots pine (Pinus sylvestris) and downy birch (Betula pubescens), while juniper (Juniperus communis) and silver birch (Betula pendula) had a 3.5 times higher probability than Scots pine and downy birch. Since the most preferred species were the least abundant, one should be cautious about the generality of the index between areas, as it may indicate that preference depends on plant species composition. The method used can easily be applied in forest management. Information on quantitative selectivity indices may improve the possibility of managing moose in accordance with acceptable browsing damage.     

Deer prevent western redcedar (Thuya plicata) regeneration in old-growth forests of Haida Gwaii: Is there a potential for recovery?

The current increase in deer populations in many forests has fostered a growing concern about their impact on forest ecology. Sitka black-tailed deer (Odocoileus hemionus sitchensis) were introduced to Haida Gwaii (British Columbia, Canada) in the late 19th century, and they have dramatically affected the regeneration of woody species in both old- and second-growth forests since then. The lack of recruitment in western redcedar (Thuja plicata) in old-growth forests has been attributed to deer. The objectives of this study were to (1) experimentally confirm that deer browsing causes a lack of western redcedar recruitment and (2) assess the potential for and speed of recovery after a prolonged exclusion of deer. We installed a set of 20 enclosures and monitored them over a period of 8 years from 1997 to 2005. We compared temporal changes in redcedar cover and in the survival and growth of marked seedlings in plots that were or were not accessible to deer. Redcedar cover in the vegetation layer accessible to deer was generally low (from 3 to 5%) but higher inside the enclosures (an average difference of 2.3%). Protected seedlings survived better, were higher, presented more leafed shoots, and had less stems than unprotected individuals, features that suggest that deer were key to the lack of regeneration. However, growth was very slow (protected marked seedlings grew 2.5 cm on average in 8 years) and, under the current conditions, the time required for a protected seedling to escape deer would probably take over two decades. This very slow growth rate under closed canopy conditions probably reflects a gap-phase regeneration strategy and/or sensitivity to competition with other woody species. The combination of a very slow growth with a high palatability and a lack of physical defences, in contrast to the other dominant conifers in this ecosystem, probably explain why redcedar regeneration can be eliminated from old-growth forest by abundant deer populations.     

Intensity of browsing on trees and shrubs under experimental variation of cattle stocking densities in southern Bolivia

Mountain forests are traditionally used for cattle grazing during the dry season in southern Bolivia. To evaluate browsing intensity and damage to young trees and shrubs, a forest grazing area was monitored for about 3 months in both 2006 and 2007. Three similar paddocks of about 3 ha each, consisting of grassland and forest in a ratio of about 1:3 were stocked with 0.9, 1.6 and 2.6 tropical livestock units per ha (equivalents of 250 kg of body weight), reflecting low, medium and high stocking densities. Six 5 m × 5 m plots were mapped out within each paddock in 2006. Three additional plots per paddock were added in 2007. Within plots, the individuals of 18 woody species were coded and evaluated weekly for intensity of browsing. Intensity was quantified using five-scale categories reflecting the proportions of plant tissue removed (0 = no browsing, 1 = 1–25%, 2 = 26–50%, 3 = 51–75% and 4 = 76–100%). Across all plant species, the average category of browsing intensity, as determined in the post-grazing evaluation, was affected (P < 0.001) by stocking density. Browsing intensity increased in an approximately linear manner from 1.58 to 2.18 and 2.77 with low, medium and high stocking density, respectively. There was no significant difference between years. Most woody species followed the same general response pattern to stocking density, although some species were only noticeably browsed at high stocking density while a few others were intensively browsed at all stocking densities. The onset of browsing on woody species was observed only after some weeks had passed, indicating that herbaceous plants were preferred as a forage resource. The proportion of fatally damaged individuals, as assessed about 8 months after grazing in 2006, were 10.6%, 8.6% and 11.4% for low, medium and high stocking density, respectively. The percentage of completely browsed individuals increased with stocking density, but 80% of those plant individuals recovered following a resting period of 8 months. However, although most woody species recovered from browsing and fatal damage was infrequent, long-term changes in woody plant species composition and structure caused by grazing cannot be excluded. High stocking density in particular may reduce the vigour of heavily browsed species.     

Effects of white-tailed deer on vegetation structure and woody seedling composition in three forest types on the Piedmont Plateau

White-tailed deer (Ododcoileus virginiana) can substantially affect the structure and species composition of a forest. The tolerance of a forest community to browsing may vary by type as a result of varying biotic and abiotic factors of the environment. To date, no studies have compared the effects of browsing among forest communities within a physiographic region. We investigated the effects of browsing on vegetation structure and woody seedling composition in three forest types (oak–hickory, Virginia pine–eastern red cedar, bottomland hardwood) in Manassas National Battlefield Park (MNBP), Virginia, USA. We compared forb cover, vertical plant cover (0–1.5 m tall), and survival of tagged seedlings in 10 exclosed (2 m × 6 m) and 10 unexclosed plots in each forest type during a 5-year period. No differential effects of browsing were found among forest types. In all forest types, deer (67 deer/km²) suppressed forb and vertical plant cover to levels less than would be expected in the absence of deer. Seedling survival rates of most species were significantly reduced by browsing. By the 4th year of the study, box elder (Acer negundo), hickory (Carya spp.), and red maple (Acer rubrum) had been eliminated from unexclosed plots, and red and white oaks (Quercus spp.) dramatically reduced. Ash (Fraxinus spp.), black cherry (Prunus serotina), and hackberry (Celtis occidentalis), although significantly impacted, remained the most abundant species throughout the study. These findings suggest that white-tailed deer may be modifying the structure of the forest interior to the extent that it adversely affects wildlife species dependent on a dense understory to thrive. We predict that the future composition of forests in MNBP will shift towards stands with fewer species and a greater dominance of ash, black cherry, and hackberry, particularly in the oak–hickory and bottomland hardwood forests, where the majority of current dominants are most affected.     

Fertilization—Effects on deciduous tree growth and browsing by moose

Knowledge of plant–herbivore interactions is fundamental for understanding foraging patterns of herbivores and their effects on the ecosystem as well as the human use of natural resources. The nutrient resources available for plants may determine not only growth rate, but also the amounts of secondary compounds in the plant tissues. As a consequence, browsing pressure on plant communities by large herbivores can be affected by fertilization and nitrogen deposition. We conducted a field fertilization experiment to test the hypothesis that fertilization leads to increased sapling growth and increased browsing by a large herbivore, the moose (Alces alces). The study was conducted on fertilized and unfertilized plots (50 m × 50 m) in northern Sweden. The plots were located in Scots pine (Pinus sylvestris) stands of young succession stages with an intermixture of deciduous species. Fertilization with 600 kg calcium–ammonium–nitrate per ha (=200 kg N ha⁻¹), was conducted in spring on one of the plots in each of 25 paired plots. The other plot in each pair was kept as control. The effect of fertilization on plants and browsing pressure was evaluated on five deciduous tree species in June and August the year after fertilization. Both browsing from preceding winter (bites) and current summer (leaf strips and bites) was recorded. Shoot size and other shoot variables increased with fertilization. Fertilization increased winter browsing for most species, but few differences were statistically significant. The late summer recording (August) showed pronounced differences in browsing pressure (leaf strips and bites) between control and treatment. In June, only a trend of increased browsing (leaf strips) was found for all species on fertilized plots. Generally, the pressure of leaf stripping was low in June, probably due to the short time the annual shoots had been exposed to moose browsing. Number of summer bites (recorded in August) was significantly higher on fertilized plots for the Betula spp., and the trend was similar for the other tree species. Within species, there was no difference in strip length between the treatments. The result indicates that fertilization can be used as a habitat improvement for ungulates within wildlife management. But, fertilization is also likely to lead to an increased risk of browsing damage, which is undesirable if forests are being fertilized for increased forest production.     

Effects of deer abundance on broad-leaf tree seedling establishment in the understory of <Emphasis Type="Italic">Abies sachalinensis</Emphasis> plantations

Browsing by sika deer (Cervus nippon) has significant negative effects on regeneration in many forests in Japan; however, the effects of browsing on regeneration processes have not been determined quantitatively. Our study was conducted in Abies sachalinensis plantations in seven tracts with differences in deer abundance on Hokkaido, northern Japan, to identify indicators for the effects of deer on the regeneration of broad-leaf species from observing seedlings. Five 5 × 20-m plots were located within each tract, and the densities of seedlings 30–200 cm tall and percentages of browsed seedlings were determined. We used sightings per unit effort (SPUE) by hunters and spotlight survey counts (SLCs) as indices of deer abundance for each tract. Seedling density was negatively correlated with deer abundance and coverage of dwarf bamboo, and basal areas of overstory trees also affected number of seedlings. Percentage of browsed seedlings was positively correlated with deer abundance, and it was affected by deer preferences among seedling species. However, the percentage of browsed seedlings was more clearly related to deer abundance than seedling density. There were few seedlings of tree species ≥100 cm tall in tracts with the highest deer abundance. Based on these results, deer abundances of SPUE >6 sightings per hunter-day or SLC >15 animals per 10 km are likely to prevent regeneration of broad-leaf species. The percentage of browsed seedlings and density and browsing damage on tree seedlings ≥100 cm tall are useful indicators of the effects of deer.     

Slow responses of understory plants of maple-dominated forests to white-tailed deer experimental exclusion

We examined the response of understory plants in mature maple-dominated forests of southern Québec, Canada, following about 30 years of high deer densities, using a deer exclosure experiment. An exclosure and a paired control of 625 m² each were established on six sites in 1998. An exclosure and a paired control of 16 m² were added at each of the same sites in 2003 but under a recent canopy gap to determine if light could enhance plant responses. We measured plant richness and abundance, and aboveground biomass of different plant groups for 8 years in the understory plots and for 3 years in the canopy gaps. Four herbaceous species were also monitored individually in the same plots. No significant differences between treatments were found in plots under forest cover, except for lateral obstruction at 0–50 cm height which was higher in the exclosures. Under canopy gaps, however, tree seedling and total plant abundance were higher in deer exclosures than in control plots. Trillium erectum recovered partially as individuals were taller, had larger leaves and more frequently produced a flower or a fruit in the absence of deer browsing under forest cover. To a lesser extent, Erythronium americanum and Maianthemum canadense also exhibited signs of recovery but were still at the single-leaf stage after 8 years of recovery. In general, the different plant groups exhibited little recovery following deer exclusion, possibly because of the low light levels that prevailed in the understory of undisturbed maple-dominated forests. The higher latitude of the present study could also contribute to the slow recovery rates of the different groups of plants compared to studies conducted in northeastern USA. Variability among sites and years had an effect on detection of statistically significant differences. Trends are however appearing over time, suggesting that many understory plants are recovering very slowly following deer exclusion. Our results emphasize the importance of studying large herbivore–forest interactions on different groups of plants, but also on specific species, and under different latitudes to be fully understood.     

The tree species matrix,influence on the level of herbivore browsing in mixed forest stands in southwest Sweden†

Herbivory pressure on a forest stand and each plant individual may be affected by the spatial distribution of conspecific and allospecific plants on the site; whether the plants are standing in solitary or groupwise settings; or by the differences in the preference of plants in relation to each other. The study was conducted in southwest Sweden, where 120 plots (1508?m²) were visited and 1280 individual woody plants evaluated and measured. We tested the hypothesis that preferred woody plants can protect unpreferred conspecific and allospecific woody species against herbivory in a system with one dominant, generalist herbivore, the fallow deer (Dama L.), and that the effectiveness of this protection varies depending on the relative preference of the neighboring conspecific and allospecific woody species placed in conspecific groups or standing solitary. Our results support the hypothesis that preferred woody plants can protect unpreferred conspecific and allospecific woody species against herbivory in the case of Picea abies dominating stands.     

Effects of deer on the post-fire recovery of a Mediterranean plant community in Central Portugal

We studied the effect of dense populations of red and fallow deer on a burned Mediterranean plant community in Portugal. We sampled a set of 12 open and 12 fenced plots 2, 3, 4, and 8 years after fire occurrence. Each plot was sub-sampled using a quadrat grid to estimate the relative plant frequency (proportion of quadrats with plants). The effect of deer on temporal trends of overall plant frequency and diversity, and on the abundance of the four most common woody species, was modeled using Generalized Linear Mixed Models. The effect of deer on plant assemblage composition was assessed by use of Redundancy Analysis. In open plots the relative plant frequency increased along the study period from 0.59 to 0.85, whereas in fenced plots there was a significantly faster recovery from 0.35 to 0.96. Cistus salvifolius and Rubus ulmifolius were significantly favored by fencing but a similar effect was not observed for Erica scoparia and Ulex jussiaei. The Redundancy Analysis showed a divergent floristic composition of the two sets of plots. The effect of deer was significant for the Evenness index but not for the Shannon and the Richness indices. Our results show that a high deer density may significantly delay the post-fire recovery of a Mediterranean community of woody plants and cause shifts in the direction of plant succession.     

Interactions among forestry regeneration treatments,plant vigour and browsing damage by deer.

In a large field experiment we studied the influence of regenerationmethods on the extent to which roe deer (Capreoluscapreolus L.) browse on spruce (Picea abies L.Karst) seedlings. Our objective was to evaluate if treatments that are good atstimulating regeneration may increase browsing damage by deer. Data werecollected between 1993 and 1995, and in the winters of 1993–94 and1994–95 the frequency of browsed seedlings was 13.2% and 13.4%,respectively. The frequency of browsing damage varied considerably betweensitesand clearcuts. Browsing damage was more frequent when the regeneration methodsincluded insecticide treatments. Among insecticide-free treatments, browsingwasheavier on scarified plots than on herbicide-treated or control plots (in whichseedlings were planted on untreated ground and given no post-plantingtreatment). Containerised seedlings were browsed more than bare-rooted ones.Neither the age of clearcut when planting, nor removal of slash, had any effecton browsing. It was demonstrated that regeneration methods that increased plantvigour (as assessed by leader growth and needle colour index) led to morebrowsing damage. Thus, our results support the plant vigour hypothesis, whichstates that many herbivores prefer to feed on vigorous plants because they aremore nutritious. However, the difference in browsing damage betweencontainerised and bare-rooted seedlings could not be explained solely by theplant vigour hypothesis.     

Effects of elevation and postharvest disturbance on the composition of vegetation established after the clear-cut harvest of conifer plantations in southern Shikoku, Japan

Large areas of previously clear-cut conifer plantations have been recently abandoned in Japan. We investigated the vegetation in the clear-cut sites and examined the environmental factors affecting species composition of the vegetation. We set up 32 study sites, each composed of several study plots (5 × 5 m), ranging from 220 m to 1060 m a.s.l. Elevation and warmth index (cumulated thermal quantity) were the primary factors affecting the species composition, with clear-cut areas showing a smaller effect in the nonmetric multidimensional scaling (NMS) ordination. Two-way indicator species analysis (TWINSPAN) divided the 32 study sites into ten vegetation groups, clustering the sites by elevation or by postharvest disturbances (i.e., replanting or browsing of Sika deer). Deciduous trees and shrubs were significant in the vegetation cover at higher elevations, while they were less so in areas of high Sika deer populations. We also investigated the abundance of old-growth species, which are expected to regenerate where the clear-cut site is abandoned. Evergreen Quercus and Castanopsis saplings were abundant at low elevations (<600 m), suggesting that they will successfully regenerate. The sapling densities of Abies firma and Betula grossa were significantly large where a clear-cut site was adjacent to natural forest, which is expected to act as a seed source. This implies that degraded deciduous forests may establish after clear-cutting at intermediate and high elevations (>600 m) if the clear-cut site is distinct from seed sources. It is argued that the preservation of natural forests is critical for the regeneration of old-growth species.     

Forest structure and woody plant species composition along a fire chronosequence in mixed pine–oak forest in the Sierra Madre Oriental, Northeast Mexico

Although wildfires are occurring frequently in the pine–oak forests in the Sierra Madre Oriental (northeastern Mexico), data on post-fire succession and forest structure are still rare. Our objectives were to (1) assess the changes in woody plant species composition after fire and to (2) to relate successional patterns to environmental variability. Based on their fire history 23 plots were selected in the Parque Ecológico Chipinque (PECH). Changes in forest structure across the chronosequence of burned stands were deduced from density, height and diameter measurements of trees and shrubs (>5 cm in diameter) in all plots of 1000 m². Differences in woody plant species composition among the plots were evaluated using Shannon evenness measure and the Whittaker's measure and by Hierarchical cluster analysis and Detrended Correspondence Analysis. Hierarchical cluster analysis showed a high similarity among all recently burned plots, independed of the aspect. Multivariate analysis showed that local environmental factors, including time since fire, continue to structure species composition. Oak species (mainly Quercus rysophylla) resprouted successfully after fire and dominated young post-fire stands. Pine species (Pinus pseudotrobus and Pinus teocote) only appeared 18 years after fire and were the dominating species in mature stands (62 years after fire). In contrast, woody plant species composition in older stands tended also to be influenced by factors such as aspect and by the potential solar radiation (PSR) during the growing season. The results demonstrate that in the PECH, natural regeneration is sufficient and woody plant species composition will be similar to pre-fire conditions after 60 years of succession. We conclude that the park managers should consider incorporating natural disturbance regimes into their management practices.     

Dispersal of vascular plants by game in northern Germany. Part I: Roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) and wild boar (<Emphasis Type="Italic">Sus scrofa</Emphasis>)

The endozoochorous and epizoochorous dispersal of vascular plant species by roe deer (Capreolus capreolus) and wild boar (Sus scrofa) was investigated in forest areas of Schleswig-Holstein and Lower Saxony, Germany. The primary aim of this study was to learn about which plant species and in what numbers diaspores are dispersed by the two wild ungulate species. In particular, the significance of zoochory for species composition and biodiversity of forests was evaluated. Fresh faecal pellets were collected from April–November 2001 in the forests of the two study areas. In addition, the coats and hooves of shot roe deer and wild boar were brushed out. The number of viable seeds was determined by greenhouse germination (seedling emergence method). The samples were spread in trays over sterilized soil from the study forests and were kept under controlled conditions in the greenhouse for 12 months. A total of 2,473 individuals from 77 vascular plant species were recorded. While roe deer exceeded wild boar concerning seed contents in the faeces, the significance of roe deer for epizoochorous dispersal was relatively low compared with wild boar. An analysis of the habitat preference of the vascular plant species dispersed by the two ungulate species revealed a high proportion of species growing in forests as well as in the open landscape, and also of non-forest species, while woody plants and herbaceous species closely tied to forest habitats were severely underrepresented. We also discuss consequences for forest ecology and nature conservation.     

Young conifer stands form a deer browsing refuge for an oak admixture: silvicultural implications for forest regeneration under herbivore pressure

There is little knowledge how ungulate pressure on forest regeneration may be mitigated by silvicultural methods. The knowledge is especially needed for artificially regenerated, deciduous tree species. We studied factors affecting browsing incidence by deer in the Pisz Forest District in Poland, an area where 10,000 ha of forest was damaged by a 2002 hurricane. In 2006, we established three experimental plots (in total, 22.6 ha), in which the main species was Scots pine (Pinus sylvestris) admixed with pedunculate oak (Quercus robur). The data on browsing were collected in 2008–2015. In general, oak browsing incidence was unrelated to oak planting density. On a plantation scale, it was significantly affected by the pine age. Although in each variant all the oaks were browsed for four consecutive years (2009–2012), in 2013 browsing incidence began to decrease. When the pines grew higher and formed a physical barrier, it was harder for deer—roe deer (Capreolus capreolus), red deer (Cervus elaphus) and moose (Alces alces)—to move through and locate the oaks. Moreover, within plantations, oak browsing incidence was higher in the patches with shorter pines. Browsing of individual saplings or small groups of saplings was also negatively affected by the height of neighbouring pine saplings. Oak density influenced deer selectivity depending on the tree height. In a low oak tree density, browsing incidence was unrelated to oak height, while in higher tree density, deer selected oaks of the height between 40 and 100 cm. We postulate that deciduous admixture in a coniferous (unattractive) stand can be planted with a few year delay. Older coniferous trees should impede locating of attractive tree species by deer and the browsing incidence.     

Effects of sika deer (Cervus nippon) and dwarf bamboo (Sasamorpha borealis) on seedling emergence and survival in cool-temperate mixed forests in the Kyushu Mountains

Effects of sika deer (Cervus nippon) and dwarf bamboo (“sasa;” Sasamorpha borealis) on seedling emergence and survival were investigated in cool-temperate mixed forests in the Kyushu mountain range, Japan. We compared the effects of sika deer between two sites with different sika deer densities. One site (no-sasa forest) has sparse cover of S. borealis and a high density of sika deer, and the other site (sasa forest) has dense cover of S. borealis with a low density of sika deer. In the no-sasa forest, more seedlings emerged and the survival rate of the seedlings was higher compared with the sasa forest. Compared with the sasa forest, the canopy in the no-sasa forest was more open, the organic layer was shallower and drier, the mean daily soil temperature was higher, and soil temperature fluctuated more widely. Those environmental parameters did not differ between unfenced and fenced plots. The comparison between the sasa and no-sasa forests suggested that removal of S. borealis by sika deer had an indirect positive effect on seedling emergence and survival. However, the fenced-plot experiments indicated that sika deer inhibited seedling emergence and survival. These contradictory conclusions could result from the much larger negative effects of S. borealis on seedling emergence and survival compared with the effects of sika deer browsing.     

Effects of Sika Deer Browsing on the Structure and Regeneration of the Abies firmaForest on Kinkazan Island,Northern Japan

Abstract

Two Abies firmaforests, one on an island (Kinkazan Island) inhabited by Sika deer (Cervus nippori)and another on the opposite peninsula in northern Japan, were compared. Species richness was poorer on the island than on the peninsula, and only low growing plants (rosettes or prostrate forms) and unpalatable plants were found there. The biomass in the forest floor in the island plots was only about 10% ofthat on the peninsula plots. Densities of woody plants in the island plots were much lower (350-600 stems/ha) than those in the peninsula plots (1400-2300 stems/ha). The size class distributions of woody plants were biased toward larger size classes in the island plots, indicating that these stands were older and lacked young trees. In contrast, the tree sizes were variable among the peninsula plots, suggesting that the peninsula forest was logged in different years. Sika deer seemed to have prevented the regeneration of the A. firma forest on the island by removing most of the seedlings and saplings. Proper population control of the deer is necessary for healthy forest regeneration.     

Browsing damage on broadleaved trees in semi-natural temperate forest in Sweden,with a focus on oak regeneration

Absence of, or poor, oak (Quercus spp.) regeneration is a problem in uneven-aged, mixed closed-canopy broadleaved forests. Browsing by ungulates on small trees may contribute to poor oak regeneration in such forests. This possibility was investigated in 25 Swedish stands, and browsing damage was analysed in relation to landscape and stand factors. The proportion of browsed small (<20 cm tall) oak seedlings and other seedlings was low, and apparently a minor mortality factor. For saplings (20–130 cm tall), accumulated browsing damage was generally higher on oak than on five major competing tree species: Fraxinus excelsior, Corylus avellana, Tilia cordata, Acer platanoides and Sorbus aucuparia. Leaf removal was rare in late summer, except for rowan. The amount of cover (shelter) for ungulates near plots was positively correlated with oak browsing intensity; within plots, a high density of ash saplings may reduce browsing on oak saplings. In these forests, browsing probably retards growth of oak saplings relative to competing trees. Oak may persist as a minor stand component, but monitoring is needed to study future changes.     

Trillium grandiflorum height is an indicator of white-tailed deer density at local and regional scales

High densities of white-tailed deer (Odocoileus virginianus) in the U.S.A. and Canada are reducing or preventing forest regeneration, and as a result, deer function as a keystone species in some sites. Management decisions about deer require reliable population density data, which are challenging to obtain at both regional and local scales. We tested the broad-scale applicability of the indicator species approach in which forage plant height was used to estimate deer density. The maximum heights of marked and unmarked plants of the widely distributed, spring-flowering polycarpic herb, white trillium (Trillium grandiflorum) were measured across southern Ontario, Canada, over a 15-year period. A significant negative relationship was found at 10 sites between maximum plant height and estimates of deer population densities, which were derived from counts of live or culled animals and varied from 7 to 40 deers km⁻². Maximum plant height could be reliably measured within a 4-week period. The underlying mechanism driving the negative relationship between plant height and deer density was attributed to deer preferentially selecting taller plants that grew less than ungrazed plants in the subsequent year. In 16 additional sites with locally high deer populations, the mean maximum height of T. grandiflorum appeared to be a more reliable indicator of deer density than estimates based on hunter returns across the broader regional scale of the Wildlife Management Unit. The ability to assess local scale white-tailed deer densities based on measuring heights of an easily identifiable, widespread plant, provides local residents and landowners with a tool for estimating the potential impacts of deer browsing and grazing in local woodlots and forests, improving local knowledge about herbivory pressure.     

Pits and rocky microsites in a subalpine forest stand facilitate regeneration of spruce saplings by suppressing dwarf bamboo growth inside a deer-proof fence

We determined patterns of microsite suppression in dwarf bamboo Sasa nipponica when grazing deer were absent. This bamboo species is able to outcompete Hondo spruce (Picea jezoensis var. hondoensis) saplings under many environmental circumstances. We set up two 10 × 100 m plots inside a deer-proof fence within a subalpine forest on Mt. Ohdaigahara, central Japan, and two similarly sized plots outside the fence. Within the plots, we surveyed microsites where spruce saplings grew. We measured height and shoot elongation of all spruce saplings, and culm height and cover ratios of dwarf bamboo growing around each spruce sapling. Spruce sapling density and average height were higher inside the deer-proof fence than outside, as were bamboo height and cover. Thus, there was a negative effect of deer browsing on vegetation parameters outside the fence and a suppression of the negative effect of bamboo on spruce sapling growth inside the fence. Spruce sapling height was higher in tree-fall pits than in other microsites inside the fence, whereas both dwarf bamboo height and cover were lower in pits and rocky sites than elsewhere. In soil and collar microsites, spruce sapling shoot growth was lower and bamboo height and cover were higher than in pits and rocky sites. Inside the fence, dwarf bamboo cover was high, but pits and rocks suppressed its growth, allowing spruce saplings to flourish. To restore heavily damaged spruce forests with advanced saplings, it will be necessary to construct deer-proof fences and create and maintain microsites with pits and rocks.