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.     

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 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.     

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.     

Establishment and growth of coniferous seedlings in an altered forest floor after long-term exclusion of deer

We evaluated differences between the forest floors and the establishment and growth of coniferous seedlings in fenced (13 years) and unfenced plots on Mt Ohdaigahara where the sika deer (Cervus nippon) population density is high. Large coniferous seedlings (height > 0.05 m) were less abundant in the unfenced plot, as a result of deer browsing. Small coniferous seedlings (height < 0.05 m), however, were more abundant in the unfenced plot, where most seedlings of Abies homolepis were found on bare ground and those of Picea jezoensis var. hondoensis were found on buttresses and fallen logs. The large area of bare ground in the unfenced plot was caused by deer browsing. Deer therefore have an indirect effect on the emergence and growth of small coniferous seedlings by modifying the forest floor.     

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.     

Effects of deer and mice on seedling survival in a temperate mixed forest of Japan

To detect the magnitude of indirect positive effects of deer and mice on seedling survival of some woody species in a Japanese temperate forest, we analyzed the data from an earlier field experiment using a hierarchical Bayesian approach. The forest studied was inhabited by sika deer (Cervus nippon) and mice (Apodemus spp.), and the floor was covered with dwarf bamboo, Sasa nipponica, which negatively affected tree seedlings and was in turn negatively affected by deer and mice. The field experiment was designed as the combination of exclusion or removal of these factors: deer, mice and dwarf bamboo. A hierarchical Bayesian model was constructed and the parameters were estimated by the Markov chain Monte Carlo method. The model successfully showed the indirect positive effects of deer and mice, which improved the survival rate of five cohorts of the seedlings of three tree species by ameliorating the direct negative effect of dwarf bamboo. The cohorts studied were formed by the seedlings of Abies homolepis that emerged in 1997 and 2002, those of Fraxinus lanuginosa f. serrata that emerged in 1998 and 2002, and those of Fagus crenata that emerged in 1999. The positive indirect effect was especially large in F. crenata, which is known to be severely affected by dwarf bamboo. The total effect of deer was shown to be positive for all cohorts except the A. homolepis cohort that emerged in 1997, at which time the dwarf bamboo had not yet fully recovered from the browsing pressure of deer. The total effect of mice was shown to be positive for all of the cohorts. We conclude that these positive effects were due to the large negative effect of dwarf bamboo on the seedlings.     

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.     

Predicting the population dynamics of three understory broad-leaved evergreen species under the influence of Sika deer in primary and secondary forests of mid-western Japan

We examined the population dynamics of three broad-leaved tree species with different susceptibilities to deer predation. Simulation analysis was conducted using a size-structured matrix model for a primary forest plot (PP) and a secondary forest plot (SP) with 56% and 12% evergreen conifer composition in the canopy, respectively. In both plots, populations of Neolitsea sericea, a species that is susceptible to deer predation, initially declined significantly but eventually leveled off. The number of small stems decreased, while that of larger stems increased, indicating that the population dynamics of N. sericea are strongly affected by browsing pressure and that the number of large trees is important for population maintenance. When we examined two deer-resistant species, Pieris japonica and Illicium anisatum, the population of P. japonica increased in the SP and decreased in the PP, whereas that of I. anisatum increased in both plots, likely because mortality tends to increase in persistently dark environments. No significant difference was observed between the present and predicted size distributions of resistant species in the PP. Competition for resources is expected to intensify in the SP as a result of the predicted increase in large stems of the resistant species I. anisatum. Therefore, a specific conservation and management strategy for tree species should be considered for each forest type under the influence of Sika deer.     

Deer Browsing and Impact on Forest Development

Abstract

Browsing by large herbivores can cause serious costs in forest production. The most important costs related to browsing damage are insufficient stand regeneration, volume losses and timber quality impairment. In Sweden, moose (Alces alces)and roe deer (Capreolus capreolus) are the most abundant deer species and therefore cause the majority of browsing damage. Balancing costs and benefits related to deer density is difficult since few data about the final losses in timber volume or quality impairment is available. Simulations of browsing and the impact on stand growth suggest that a minor or moderate browsing intensity has little effect on final harvest volume. The reason is that trees seem to recover some of the lost growth as they grow out of reach of the animals. Balancing between different uses of the forestry resource is complicated by our limited capability to predict the impact of different management regimes.     

运用Cytb 和12s rRNA 基因鉴别梅花鹿(Cervus nippon)和马鹿(Cervus elaphus)的研究

本研究介绍了运用细胞色素b基因和12s核糖体RNA基因部分序列的系统学和核甘酸距离分析来鉴别降解的梅花鹿和马鹿样品。采用PCR和直接测序技术获得了8份嫌疑样品402bp细胞色素b基因序列,并与来自GenBank数据库27份同源的细胞色素b基因序列进行比对。3份嫌疑样品与梅花鹿的核甘酸距离值相同(0.026±0.006),小于梅花鹿与东部马鹿间最小的核甘酸距离值(0.036)。并且梅花鹿和马鹿的系统学分析表明这些样品与梅花鹿聚为一枝,因此可以推测它们来源于梅花鹿。同样的方法得出另3份嫌疑样品来源于马鹿。该结果被387bp12s核糖体RNA基因序列的系统学和核甘酸距离分析进一步证实。该方法是有效的,花费的时间少,能帮助减少同类野生动物案件的发生。图2表1参13。     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Browsing of sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) in the context of life history strategies: a literature review

Sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) constitute small proportions of the deciduous tree volume in Scandinavia, but are highly preferred winter forage for moose and red deer, which occur at historically high densities. Thus, a possible decline of these tree species has been indicated. Against this background, we have reviewed the life histories of relevance for browsing, as well as the basic biology and genetics of sallow and rowan. The species show similarities with respect to short lifespan, small size and sympodial growth pattern, which are risk factors in a browsing context. They also have high juvenile growth rate, important for growing quickly out of reach of browsers. Sallow depends strongly on disturbance for establishment and is more demanding with respect to soil and light conditions than rowan, possibly important for the substantially lower abundance of sallow on the Norwegian Forest Inventory plots. Similarly, the relative recruitment of small size classes of sallow is less than for rowan. Although recruitment is reported to be hampered in wintering areas with high moose or red deer densities, the inventory data, however, dating only back to 1994, do not suggest a general decrease in any of the species. Sallow and rowan saplings show low mortality in moose and deer dominated areas and the species can be characterised as rather resilient to browsing. Of more concern is that browsing can constrain the development of mature rowan and sallow trees locally, with possible consequences for associated epiphytic biodiversity.     

Effects of 13-year fencing on browsing by sika deer on seedlings on Mt. Ohdaigahara, central Japan

We examined the effects of fencing on deer browsing on seedlings 13 years after the building the experimental fence on Mt. Ohdaigahara, central Japan, where the sika deer (Cervus nippon) population is high. There was no difference in the species number of seedlings between 1991 and 2004. The density of seedlings in the fenced plot in 2004 was three times higher than in 1991. Although the unfenced plots had no seedlings higher than 20 cm, there were seedlings up to 120 cm in the fenced plot. These results suggest that 13-year fencing promoted regeneration of seedlings in this area.     

Reduction of herbivore density as a tool for reduction of herbivore browsing on palatable tree species

Browsing of overabundant free-living herbivores often limits the natural as well as artificial regeneration of forest in Central-European Mountains. The aim of the study was to assess the efficiency of the extensive reductions of herbivore populations for protection of forest regeneration in preferred areas. We analysed the relationship between the intensity of shoot browsing and relative density of herbivores in viewpoint of natural regeneration of secondary mountain spruce stands with low proportion of broadleaved trees in Králicky Sněžník Reserve (Czech Republic). The distribution of seasonally migrating herbivores was established by counting faecal pellet groups (standing crop method) in growing season. The intensity of browsing of spruce, beech and rowan was assessed using the proportion of browsed shoots on individual tree sapling. Density of red deer was re-counted from the number of pellet groups and it was 15–56 individual/km². The intensity of spruce browsing was low in the whole area in all seasons. In general, the intensity of rowan and beech shoot browsing was high in the whole area. There were no correlations between deer density and browsing intensity of rowan (browsing was severe in all plots). We conclude that the natural regeneration of attractive trees and shrubs is nearly impossible in habitats where proportion of these food items is too small, even when the density of herbivores is low. Reduction of herbivores density is useful for protecting spruce and other tree species with low preference by herbivores. Forest managers should take into account the high palatability of deciduous tree species and distribution of herbivores on localities. Preference of localities is often based on grasses in the food supply. Regeneration of highly palatable tree species requires both low density of herbivores and sufficient protection of saplings.     

Soil Amendment Effects on Growth and Sprouting Success in a Pennsylvania Northern Red Oak Stand on Extremely Acidic Soils

Abstract

On the Laurel Hill, anecdotal accounts reported coppice regeneration of oaks to be limited. Possible causes of poor sprouting success were large stump size, deer browsing, season of cutting and poor tree vigor. Our study site had been partially cut three years before this study to salvage some northern red oak (Quercus rubra L.) which had died. We attempted to increase Q. rubra vigor, as measured by increasing basal area growth, with soil amendments of 6,600 kg/h dolo-mitic lime, 110 kg/h K2O equivalents and 220 kg/h P2O5 equivalents while accounting for other growth variables. The objective was to improve overall Q. rubra sprouting success by increasing vigor prior to harvest. The soil amendments increased soil available calcium (Ca), magnesium (Mg), potassium (K) and phosphorus (P) and reduced available aluminum (Al). Foliar concentrations of Ca, Mg and K were increased. Basal area increment of treated Q. rubra trees increased by 10 percent and terminal elongation increased by nearly 100 percent. Two years after treatment, the trees were cut during the dormant season and all stumps were protected from deer browsing. After two years, 80 percent of the stumps had sprouted. No effect of lime and fertilizer on the sprouts was found.     

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.     

Winter Browsing by Large Herbivores on Planted Deciduous Seedlings in Southern Sweden

Deciduous seedlings are valuable for the forest industry and, at the same time, often palatable for large herbivores. Therefore, browsing on seedlings of seven deciduous tree species and on Picea abies(L.) Karst., as a reference species, was tested in a field experiment with the species presented at feeding stations. Twenty stations, each with one seedling per species, were established in each of 15 clear-cuts in southern Sweden. The planting took place in October. Depending on species, between 16 and 85% of the seedlings were browsed after one winter. The descending order, based on number of browsed seedlings, was Quercus robur L . > Alnus glutinosa (L.) Gaertn.> Fagus silvatica L.> Tilia cordata Mill.> Prunus avium L.> Betula pendula Roth> Picea abies (L.) Karst. > Fraxinus excelsior L. On 83% (range 45-100%) of the browsed seedlings the leader was damaged. On average, browsing reduced seedling hight by 32% (range 12-47%). Within species, initially taller seedlings were more often selected than shorter ones.     

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.     

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.