Habitat use of large ungulates in northeastern Germany in relation to forest management

Estimating large herbivore density has been a major area of research in recent decades. Previous studies monitoring ungulate density, however, focused mostly on determining animal abundance, and did not interpret animal distribution in relation to habitat parameters. We surveyed large ungulates in the Biodiversity Exploratory Schorfheide-Chorin using faecal pellet group counts. This allowed us to explore the link between relative ungulate abundance, habitat use, and browsing damage on trees in a region with several types of forest, including unharvested and age-class beech forests, as well as age-class pine forests. Our results demonstrate that roe deer and fallow deer relative abundance is negatively correlated with large tree cover, and positively correlated with the cover of small shrubs (Rubus spec., Vaccinium spec.), and winter food supply. Habitat use of roe deer and fallow deer, as estimated by counting faecal pellet groups, revealed a preference for mature pine forests, and avoidance of deciduous forests. This differential habitat use is explained by different distributions of high quality food resources during winter. The response of deer to understory cover differed between roe deer and fallow deer at high cover percentages. The amount of browsing damage we observed on coniferous trees was not consistent with the relative deer abundance. Browsing damage was consistently higher on most deciduous trees, except for beech saplings which sustained less damage when roe deer density was low. Because roe deer is a highly selective feeder, it was reported to affect tree diversity by feeding only on trees with high nutritional value. Consequently, we propose that managing the number of all deer species by hunting is necessary to allow successful forest regeneration. Such an adjustment to deer numbers would need to account for both current tree diversity and alternative food resources. Our findings may be applicable to other forest landscapes in northeastern Germany including mature pine stands and differently harvested deciduous forests.     

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.     

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.     

Spatial heterogeneity of winter forest resources used by deer

Monitoring red deer (Cervus elaphus) and roe deer (Capreolus capreolus) populations is a key aspect of wildlife management in Europe, so the utilization of resources must be considered. However, the spatial distribution of resources used has often been overlooked. We studied spatial heterogeneity of winter forest resources used by deer and applied an indirect method to evaluate the browsing pressure of these cervids in a forest of the Cévennes National Park (Lozère, France). We analyzed the pattern of browsing by principal component analysis (PCA) and graphical display. We demonstrated that heather (Calluna vulgaris), bilberry (Vaccinium sp.) and beech (Fagus sylvatica), which figured in more than 50% of the browsing observed, were unevenly consumed in the study area. Moreover, the deer consumed more beech outside high beech forest. Our methodology provides a helpful tool to monitor browse–ungulate interaction by taking into account the availability of key browse resources.     

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.     

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.     

Short-term browsing by roe deer has little effect on survival and growth of sessile oak seedlings

We studied the effect of roe deer (Capreolus capreolus) browsing on the growth and mortality of sessile oak (Quercus petraea) seedlings. First, we measured the height and basal diameter, and counted the number of shoots of the 849 seedlings of an experimental plantation, 60 of which were individually protected by fencing. We then introduced two tame roe deer females every day for three weeks from late May to mid-June. Measurements and counts were repeated after roe deer removal, then once again the following winter. We found no significant effect of browsing on mortality, total height, or basal diameter of the seedlings. However, the browsed seedlings exhibited a lower number of shoots despite regrowth observed after roe deer removal. We propose to use the tolerance of oak to deer browsing to promote natural and inexpensive methods, to protect oak during regeneration, such as the spatial arrangement of plants or the use of slash or surrounding vegetation in favourable environments.     

Habitat selection by ungulates in relation to predation risk by wolves and humans in the Białowieża Forest, Poland

Predation risk is an important factor influencing the distribution of ungulates and their impact on forest structure. However, simultaneous predation risk by wolves and humans is rarely considered in the analyses of habitat selection by ungulates. We counted ungulate pellets on transects to analyse the influence of wolves and humans on ungulate density distribution in the Białowieża Forest, Poland. We assessed whether (1) forest exploitation influenced ungulate habitat selection, (2) ungulate density was higher in areas without human hunting, (3) ungulates avoided the surroundings of a major road, (4) prey density was higher in the strife zone between home ranges of wolf packs both in the presence and absence of human hunting, (5) ungulates avoided areas selected by wolves, and (6) wolf kill sites were in high prey density areas. Red deer (Cervus elaphus) and wild boar (Sus scrofa) selected unexploited over exploited forests and areas without hunting, whereas roe deer (Capreolus capreolus) preferred exploited forests. Wild boar and European bison (Bison bonasus) avoided the area within 300 m of a major road, whereas we could not detect any avoidance by red or roe deer. Prey density was not higher in wolf strife zones, regardless of human hunting. Ungulates did not avoid areas selected by wolves. Wolves killed red deer in areas with prey density of about 4 red deer/km², regardless of whether the average red deer density in those areas was higher or lower. We conclude that habitat alteration by forest exploitation and hunting by humans influenced the density distribution of ungulates more than predation risk by wolves.     

Ranking temperate woody species along a gradient of browsing by deer

As a result of diverse management policies, densities of red and roe deer have varied considerably for the last few decades in Europe. Their selective browsing on woody species can modify the interactions between competing species and thus change overall patterns of plant diversity in forests. However, no classification of woody species according to browsing by deer yet exists in Europe. The selectivity index, which measures the balance between availability and consumption of plant species by herbivores, could be used for such a classification. The aim of this study was, first, to rank woody species according to this selectivity index, then to test whether their rank changed when browsing pressure varied in space or time. We also explored morphological and ecological life traits (Ellenberg indicator values) that might explain differences in selectivity.In 2006, we re-sampled a network of 217 coupled floristic and browsing survey plots, which had been previously described twice (1976, 1981). The plots were located in the forest of Arc-en-Barrois (France), a large woodland area divided into a North and a South Forest which have had different histories of browsing pressure.For the three samples and the two forests, we calculated a selectivity index for 19 woody species. Dogwood (Cornus sp.) and field rose (Rosa arvensis) appeared to be the most selected species whereas beech (Fagus sylvatica), common mezereon (Daphne mezereum) and wild-service tree (Sorbus torminalis) were always avoided. The rank order of species according to their selectivity index remained stable over the three sample years and between the two forests (Spearman's rank correlations: ρ ≈ 0.8 spatially, and range from 0.3 to 0.8 temporally). This indicates that variations in browsing pressure did not strongly modify deer selectivity patterns. Selectivity was positively correlated with wood density and negatively correlated with maximum height of the species, but unrelated to other plant traits. Finally, selectivity can be seen as an intrinsic characteristic of those woody species, and could be used as an additional life trait when analyzing vegetation communities.     

Leader Browsing by Red and Roe Deer on Young Sitka Spruce Trees in Western Scotland I. Damage Rates and the Influence of Habitat Factors

Leader browsing was monitored for several years on some 2000trees at 14 sites in Glenbranter Forest, Argyll. We recordedtrees individually to obtain cumulative totals for browsingand other damage, and to relate incidence to height and state(i.e. whether the trees had single or multiple leaders, oldor new leaders). Rates of leader browsing peaked in late winter and also fora few weeks during the spring flush of growth; they were leastfrom mid June to September. Incidence was closely related toheight, trees from 30 to 50 cm tall suffering the most damage,while trees taller than 80 cm rarely incurred leader browsing. Annual rates of leader browsing were greatest in the secondand third year after planting but varied considerably amongsites. Newly established crops were no more damaged than second-rotationrestocked crops. Multiple regression showed that the incidenceof leader browsing was significantly negatively related to thedensity of trees <60 cm tall and to percentage cover of ericoidplants, and significantly positively related to deer presence,as measured by the accumulation of pellet groups of both deerspecies. Individual trees were browsed two to three times on averagebefore reaching a safe height, but a few trees experienced upto eight leader browsings. In recovery from browsing, treesoften produced several leaders; we found that trees with multipleleaders or new leaders were more susceptible to leader browsingthan trees with single or old leaders. So once a tree has beenbrowsed the likelihood of future browsing is increased. Received 5 December 1989.     

Early-stage impacts of sika deer on structure and function of the soil microbial food webs in a temperate forest: A large-scale experiment

Modification of forest vegetation caused by an overabundance of mammalian herbivores has been reported in temperate and subarctic regions all over the world. However, the indirect effects of these herbivores on the structure and functions of soil decomposer systems are not fully understood, especially in temperate forests. We investigated the early effects of sika deer invasion on soil decomposer systems in a Japanese temperate forest using two large-scale experimental enclosures with low and high densities of deer (LD: 25 ha, 4 deer km⁻²; HD: 6.25 ha, 16 deer km⁻²) including control plots without deer (WD). Three years after deer introduction the understory cover of dwarf bamboo (Sasa nipponica) declined due to deer browsing in both enclosures. At the same time, measurements were made of the soil microbial community, soil nematode community, soil nitrogen (N) mineralization rate, and carbon (C) and N content in dwarf bamboo leaves. In LD, soil microbial biomass was lower from WD, probably due to the decrease of fresh aboveground litter from dwarf bamboo. Surprisingly, there were no cascading effects on total abundance of soil nematodes and soil total N mineralization potential which were unaffected by deer in the LD treatment, while soil NH₄⁺-N content was lower and soil nematode community structure was different (abundance of 4 families was higher and that of 3 families was lower, but the functional structure was not different) from WD. Specifically, the responses to deer introduction varied between microbes and nematodes, and the change of balance in the microbial food webs may have altered N mineralization processes. In contrast, in the HD treatment, all the variables measured were not significantly different from those of WD treatment. Intensive browsing by deer may have cancelled out the effects of the decrease in aboveground litter input on the soil decomposer systems through other pathways, such as a transitory increase in belowground litter input caused by induced changes in allocation patterns of bamboo. No changes in total N mineralization potential, leaf N, and composition of understory vegetation in both enclosures indicated that deer introduction did not facilitate nor retard N cycling regardless of deer density. This study showed that sika deer browsing can affect soil decomposer systems at an early stage of invasion even at low density, which contrasts with previous studies on the subject. Linking our findings of early-stage effects of deer on soil decomposer systems to longer-term dynamics of understory vegetation and tree regeneration will be needed to evaluate the adequacy of deer management practices with respect to the sustainability of soil nutrient supplies.     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Ungulate herbivory of boreal and temperate forest regeneration in relation to seedling mineral nutrition and secondary metabolites

Fertilization at planting may improve reforestation productivity on sites frequented by ungulates by stimulating rapid growth above the browse line, allowing seedlings to escape animal damage. Additionally, stored nutrients may aid seedlings in recovery from browse damage. High plant nutrient content may, however, deem seedlings more susceptible to ungulate browsing because of increased foliar protein content and, thus, nutritional value. Research examining the relationship between fertilization of forest trees and ungulate herbivory has been conducted almost exclusively in boreal and temperate zones of North America and Europe, and has produced two common themes. The first is that nearly all trials used broadcast application of fertilizer. This method supplies nutrients to both the target species and competing vegetation, resulting in significant increases in nutritional value for all potential forage. The second common theme is that there is a higher likelihood of browse for fertilized plants. Exceptions to this thread exist, however, because browse preference of fertilized seedlings is apparently species-specific due to differences among species in resource allocation. Palatability appears to interact with the production of plant chemical defenses (i.e., alkaloids, phenylpropanoids, terpenoids). Variations in plant mineral nutrition and secondary metabolite production can play a significant role determining forage preference; however, ungulates under starvation or stress will browse plants of high toxicity or low nutritional value to maintain fitness.     

Patterns of Stem Damage by Moose (Alces alces) in Young Pinus sylvestris Stands in Sweden

The incidence of recent moose browsing of the main stem on young Scots pine (Pinus sylvestris L.) trees, and correlations with tree or stand characteristics, were examined using data from a large-scale survey in Sweden. On average, 10% of all Scots pine main stems showed one or more types of recent damage, with browsing of the apical leader accounting for about 75% of all damage recorded. Stripping of bark off the main stem occurred most frequently in the southern area, where site productivity is highest. Apical leader browsing was negatively correlated with height of the main stems, length of the apical leader and incidence of pre-commercial thinning, whereas bark stripping and stem breaking were positively correlated with the same variables. All types of recent stem damage correlated positively with the extent of previous stem damage and also with the extent of recent browsing of lateral shoots.     

Deer density effects on vegetation in aspen forest understories over site productivity and stand age gradients

Studies examining the interacting effects of ungulate herbivore pressure and site productivity on vegetation are mostly on grassland-grazing systems and have shown conflicting patterns. Here we examine the effects of deer density (>30 years differences in density between two landowners), site productivity (site index, SI) and stand age on subcanopy vegetation characteristics in 60 closed canopy, clear-cut origin Populus tremuloides dominated stands, Michigan, USA. Stand age effects were included because age varied among stands and can affect subcanopy vegetation patterns. Compared with fewer deer, stands with more deer had greater total forest floor vegetation mass, and its major components bracken fern (Pteridium aquilinum), sedge (mostly Carex pensylvanica) and trees/shrubs <0.25 m tall, but lower forb mass and lower forest floor vegetation species richness and diversity. Deer density and SI had strong interacting effects on total forest floor mass, forb mass, and species richness. Forb mass increased with SI, but only in stands with fewer deer, whilst total vegetation mass was greater in stands with more deer at lower SI and declined with SI more sharply than for stands with fewer deer. Species richness increased with SI but more so at lower than higher deer density. Deer density and age had interacting effects on mass of trees/shrubs <0.25 m tall and sedge. Compared with fewer deer, stands with more deer had greater sedge and tree/shrub mass, and sedge mass decreased and tree/shrub mass increased more sharply with age. In lower deer stands there was a dense subcanopy tree and shrub strata within and beyond the reach of deer 0.9-10 m tall whereas in higher deer stands this vegetation layer was nearly absent. We conclude that higher deer browse pressure in early successional Populus stands (1) strongly limits the recruitment of woody stems to larger (>0.9 m tall) size classes, which could affect long-term successional trajectories, and (2) diminishes forb density and species richness, especially at higher site productivity, but increases total forest floor vegetation mass (mostly bracken fern and sedge), especially at lower site productivity. Given associations of bracken fern and sedge with poorer and/or more open sites and assuming high palatability of forbs, this pattern may result from the combination of selective herbivory and higher light availability caused by limited recruitment of trees and shrubs to taller strata.     

Is deer herbivory directly proportional to deer population density? Comparison of deer feeding frequencies among six forests with different deer density

The intensity of deer herbivory, rather than simply the deer population density, directly affects the forest ecosystem, but a linear relationship between these two factors has generally been assumed. To assess their relationship, we investigated deer population density and tree sapling vegetation in six forests with different deer density on Yakushima Island, Japan. The feeding frequency was used as an index of deer herbivory. Palatable saplings showed high feeding frequency and became rare in deer-abundant areas, while unpalatable saplings showed low feeding frequency and increased with increasing deer density. In addition, feeding frequency on sapling vegetation did not continue to increase with increasing deer population and was limited to only 0.24-0.32 in deer-abundant areas (more than 20 deer/km²). These data suggest that deer shift their main food items from living palatable saplings to other alternatives such as litter fall rather than living unpalatable saplings. Clearly, the nonlinear relationship between deer density and deer herbivory on forest vegetation could result from a change in the food eaten by deer, and it is therefore necessary to assess the impacts of deer based on not only the size of the deer population but also the intensity of direct herbivory.     

Establishment patterns of woody species in low intensity-grazed pastures after the cessation of intensive agricultural use

Low intensity grazing with large herbivores is increasingly used to steer the development of mosaic landscapes (including wood-pastures) and woodland expansion on former agricultural land. We studied the establishment and early growth pattern of woody species in grasslands (formerly used for high intensity summer cattle grazing) in which a low intensity-grazing regime was set up between 4 and 11 years ago. We tested whether the developing vegetation patches, consisting of tall herbs and scrub, influenced this pattern. Further, we investigated if increased susceptibility to trampling disturbance by large herbivores on wet soils in floodplains influenced sapling establishment.Formerly intensively used grasslands were studied at six sites, four in floodplains and two in interfluvial zones on nutrient rich, heavy soils. In total, we sampled 141 plots with grassland, tall herb or scrub vegetation and recorded individuals of established tree and shrub species, their height, diameter, browsing damage and distance to potential seed sources. In the floodplains, we recorded soil disturbance by large herbivores (hoofprints) in wet and moist soils.Sapling frequencies, with Fraxinus excelsior as the most common species (>50% of all individuals), were two to three times higher in tall herb and scrub patches compared to grassland. Only the spiny Crataegus monogyna established in higher frequencies in grassland plots. In floodplains, we found two to three times more established woody saplings in wet tall herb plots, compared to moist and wet grassland plots and moist tall herb plots.Browsing reduced sapling height in grassland and tall herb patches. Only in scrub patches did average height (±2.5 m) reach above the browse line. We found 30–70% less browsing damage to saplings in scrub compared to damage in grassland and tall herb plots. In particular, damage to the apical shoot was strongly reduced.Our findings show that tree establishment occurs in spatial association with non-grassland patches that offer protection against browsing. Fast growing spiny shrubs like Rubus sp. offer protection for palatable saplings, allowing establishment and growing out beyond the browse line. Alternatively, increased establishment occurs in wet tall herb patches, where soil disturbance by moderate trampling favours germination and, subsequently, saplings are protected at least temporarily against browsing by unpalatable tall herbs.     

Changes in spatial patterns of sika deer distribution and herbivory of planted seedlings: a comparison before and after deer population control by culling

Sika deer (Cervus nippon) sometimes cause extensive damage to planted tree seedlings. To evaluate the effects of culling on the spatial distribution patterns of sika deer and browse damage to planted seedlings, we compared the data collected before and after experimental culling in a cool, temperate, mixed forest on Kyushu Island, Japan. Experimental culling, conducted in an area of 1 km², removed five, four and two sika deer in April, June and October 2011, respectively. During the year before culling, the spatial pattern of the number of sika deer caught on camera corresponded to the predicted sika deer density. Sika deer immediately browsed planted seedlings after the initial planting. The cumulative number of browsed seedlings increased over time, especially in winter. The spatial pattern of the cumulative number of sika deer caught on camera corresponded to that of browsed seedlings at the year’s end. During the year when culling was conducted, the number of sika deer caught on camera decreased around the center of the study site where the culling was conducted and the number of browsed seedlings decreased. During the year following culling, the cumulative number of browsed seedlings was very similar to that in the year before the culling, while the same low number of sika deer was caught on camera. These results indicate that the effects of deer culling resulted in decreased levels of sika deer appearance and browse damage for more than 1 year and for several months, respectively.     

Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

Abstract

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km^?2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.     

Altitudinal differences in bark stripping by sika deer in the subalpine coniferous forest of Mt. Fuji

Deer expansion is a growing concern for forest ecosystem management. In Japan, upward expansion to subalpine and alpine areas has reached alarming proportions in recent years. We examined bark stripping by sika deer along an altitudinal gradient in the subalpine coniferous forest at three altitude ranges (1800-2000 m, 2000-2200 m, and 2200-2400 m) on the southern slope of Mt. Fuji. We tested differences in densities and diameter at breast height (DBH) of trees and those with bark-stripped stems of all tree species among the three altitude ranges. Then, we compared the relative densities of deer, based on pellet counts, to determine the impact of deer in relation to deer use and forest stand patterns across the altitudinal range studied. The results of the study show that differences in bark stripping by sika deer depended largely on the elevation and the species. Larger stems were sparsely distributed in the lowest elevation zone between 1800 and 2000 m. The relative density of deer was highest in the areas exhibiting high bark-stripping intensity on small regenerating trees of the dominant coniferous species, Abies veitchii, and on broadleaf species. In the highest elevation zone between 2200 and 2400 m, smaller stems were densely distributed, and the relative deer density was lowest where the bark-stripping intensity on small stems was lower for all three species studied. The damage to subalpine tree stems corresponded to the availability of palatable tree species with a small diameter, as reflected by the successional stage along the elevational gradient of subalpine forest on Mt. Fuji. These results suggest that the continuous impact of bark stripping on the dominant tree species might cause severe changes in forest succession.