Impact of bark stripping by sika deer, Cervus nippon, on subalpine coniferous forests in central Japan

We investigated the effect of bark stripping by sika deer, Cervus nippon, on forest regeneration in subalpine coniferous forests on Mt. Ohdaigahara and in the Ohmine Mountains of central Japan. Bark stripping by sika deer occurred in five major tree species: Abies homolepis; Abies Veitchii; Tsuga diversifolia; Picea jezoensis var. hondoensis; and Chamaecyparis obtusa. The percentage of damaged trees on Mt. Ohdaigahara was higher than in the Ohmine Mountains, probably because of the higher deer density. On Mt. Ohdaigahara, the DBH distributions of stems for P. jezoensis var. hondoensis, A. homolepis, T. diversifolia andC. obtusa were bell-shaped with fewer smaller and larger trees. On the other hand, in the Ohmine Mountains the distributions for P. jezoensis var. hondoensis and A. Veitchii showed a reverse-J shaped with more smaller trees. Larger overstory conifers on Mt. Ohdaigahara were killed by bark stripping when 100% barked, although in the Ohmine Mountains ca. 50% of the trees survived even when 100% barked. After the disappearance of the overstory conifers on Mt. Ohdaigahara, the dwarf bamboo, Sasa nipponica, expanded into the forest floor due to changes in light reaching the forest floor. Since S. nipponica is the main forage of deer in this area, this increase caused a corresponding increase in the deer population, which in turn, could cause a further decline in the coniferous forests.     

Bark stripping by sika deer on veitch fir related to stand age, bark nutrition, and season in northern Mount Fuji district, central Japan

We carried out this study to clarify the relationships between the seasonal changes of bark stripping and food quality, and between bark-stripping intensity and bark nutrition with age of veitch fir (Abies veitchii) by sika deer (Cervus nippon) in the northern Mount Fuji district, from June 2000 to July 2003. We found that sika deer gnawed and ate all of the bark stripped, and a seasonal change in bark stripping occurred from December to May or June with a peak in March–April. The stripping period largely overlapped with the periods of low food availability and poor food nutrition (indicated by fecal chemical component index) of sika deer, from January to April. Both bark nutritional quality and bark-stripping intensity related to stand age and tree size of veitch fir negatively. Consequently, bark-stripping intensity related to bark nutritional quality positively. February is the worst forage period in terms of both quantity (due to deep snow) and quality (indicated by fecal chemical components). There was a time lag of 1 to 2 months in the peak of bark stripping in March–April when compared with the poor forage period in February. This time lag may suggest that sika deer need more nutritious and easily digested food from March due to increased nutrient demands that result from depleted body condition in both sexes, gestation of pregnant females, and the recovery of active metabolism.     

Effects of bark stripping by sika deer, Cervus nippon, on wind damage to coniferous trees in subalpine forest of central Japan

We evaluated the effect of bark stripping by sika deer (Cervus nippon), and subsequent wood decay, on tree fall in a coniferous forest on Mt Ohdaigahara in central Japan from July to September 2006. This valuable primeval coniferous forest is declining because of bark stripping. Broken trunks of Abies homolepis and Picea jezoensis var. hondoensis had more serious decay and larger bark-stripping wounds on the trunk than standing trees, suggesting that bark stripping causes trunk decay and results in broken trunks and uprooting by typhoons.     

Development of a method for forest cafeteria tests for evaluating chemical deterrents against bark stripping by sika deer (Cervus nippon yesoensis)

Damage caused by sika deer (Cervus nippon yesoensis) is a serious problem in commercial and environmental (non-harvested) forests in Hokkaido, northern Japan. Cafeteria tests in forests may be useful for evaluating the efficacy of chemical deterrents against bark stripping by deer. To develop a method for forest cafeteria tests in the continuous snow cover period, two experiments were carried out. In the experiments, logs were produced from tree trunks, and used as carriers of chemical deterrents. Carriers were installed in forests and fed to deer. The first experiment was to find suitable sites and installation methods for carriers. Criteria for the local suitability and the installation methods were as follows: a) Sites where deer are active should be selected; b) Carriers should be installed along actively used deer trails; c) Installation sites of carriers should be changed in response to deer movement; d) Carriers should be produced from tree species that deer naturally prefer; and e) Each carrier should be partially buried in the snow. The second experiment evaluated the feasibility of a cafeteria test method based on the results of the first experiment. The method was used for 13 sets of the cafeteria test, in which the deterrent effectiveness of 5 chemicals (wood tar, rosin, wood vinegar, and 2 pyroligneous liquors) was examined. We obtained results from all the sets. The chemicals tested did not deter bark stripping by deer. Nevertheless, the method used in the present study was practical for the cafeteria tests.     

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.     

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.     

Expansion of dwarf bamboo, Sasa nipponica, grassland under feeding pressure of sika deer, Cervus nippon, on subalpine coniferous forest in central Japan

The long-term changes of the area of Sasa nipponica grassland, bark stripping damage to trees by sika deer, Cervus nippon, and sika deer densities on Mt. Ohdaigahara were assessed with aerial photographs, information from previous studies, and field experiments. The grasslands expanded year after year. The expansion rate was highest from 1982 to 1992, and lowest from 1992 to 1997. Bark-stripping damage on coniferous trees was heavier than that on broadleaf trees in both the early 1980s and 1998. The deer density in the eastern part of the area had been high (14.4–64.3 km⁻²) from 1984 until 1998. We presume that rich growth of S. nipponica and heavy feeding pressure by sika deer over the long term caused expansion of S. nipponica grasslands by killing canopy trees by bark stripping and preventing forest regeneration.     

Food habits of sika deer as assessed by fecal analysis in Mt. Ohdaigahara,central Japan

Food habits of sika deer,Cervus nippon centralis temminck, in Mt. Ohdaigahara, central Japan, were investigated by the fecal analysis method from September, 1990, to June, 1991. The fecal analysis showed that the percentage ofSasa nipponica Makino et Shibata in the fecal composition was approximately 50% throughout the year, indicating that it was the main food plant. Bark and twigs were also found in the feces in all seasons, suggesting that the deer ate bark throughout the year. Even though the amount of the bark eaten per capita was small, the high density of deer may possibly cause serious bark damage. Barking can be one of the main causes of the mortality ofPicea jezoensis Sieb. et Zucc. var.hondoensis (Mayr) Rehder andAbies homolepis Sieb. et Zucc. in Mt. Ohdaigahara. A part of this study was presented at the 105th Annual Meeting of the Japanese Forestry Society (1994).     

Chemical components of hardwood barks stripped by the alien squirrel Callosciurus erythraeus in Japan

The Pallas squirrel (Callosciurus erythraeus; common name in Japan: Formosan squirrel) was introduced in the 1950s and has established populations throughout southeastern Kanagawa Prefecture, Japan. Some tree species in natural forests, parks, and gardens in residential areas have been intensively debarked by the squirrels, especially in winter and spring. The amounts of chemical components, such as polyphenols, resins, flavanols, and sugars, in the bark were compared among species and individual trees collected in the forest of Yokohama Nature Sanctuary. Interspecific differences in the extent of stripping were not related to the amounts of the four chemical components. For individual trees, the bark with feeding scars tended to contain more sugar than that without scars.     

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.     

Over-winter weight dynamics in supplementally fed free-ranging sika deer (Cervus nippon)

Continuous body weight dynamics in supplementally fed free-ranging sika deer (Cervus nippon) during the three consecutive overwinters (from January to April), 1992–1994 were surveyed. Almost all individuals lost weight continuously throughout each winter (adult deer 5–13%; young deer 3–19%) and several deer lost weight even in April. Weight gain after loss began in the mid-April when ambient temperature increased and when plant growth began. Weight dynamics varied according to sex, age and reproductive status. Adult deer showed significant large weight loss than the young. Young deer lost weight (calves 18±11 g/day, yearling and two-year-olds 27±29 g/day) more slowly than the adults (65±23 g/day). Following this study of over-winter body weight dynamics, it seems that early springs are the most critical periods for the survival of free-ranging sika deer because of their poor nutritional condition. A portion of this report was presented at the 106th Annual Meeting of the Japanese Forestry Society (1994).     

The effects of selective logging on stand structure and the regeneration of subboreal forests in Hokkaido, northern Japan

We investigated the effects of selective logging on stand structure and regeneration in selectively logged subboreal forests in Taisetsuzan National Park in Hokkaido in northern Japan. The basal area decreased and the size structure of trees altered in the stands studied due to repeated, intense selective logging, in which larger trees were cut down as a priority. Sapling density in the stands was much lower than that in primary forests. In the simple and multiple regression analyses that were used to estimate the effects of selective logging on sapling density, sapling density had a significant positive correlation with tree density and had little correlation with the density of logged stumps or the height ofSasa (dwarf bamboo) growing on the forest floor. These results suggest that the establishment sites around canopy trees influenced the establishment of saplings, rather than the gaps caused by selective logging. However, both the coefficient of determination and the standardized partial regression coefficient of multiple regression analysis were higher for the stand with a dense cover ofSasa than for the stand with a sparse cover ofSasa. Thus, the success of regenerating forests with selective logging depends on both the site of advanced regeneration and the light conditions that regulate growth.     

Population density and range use of sika deer,Cervus nippon, on Mt. Ohdaigahara, central Japan

We investigated the population density and range use of sika deer,Cervus nippon, on Mt. Ohdaigahara, central Japan, from July 1996 to July 1997. The deer population density estimated by the block count was high at 17.5–30.9 deer/km². Although the deer remained this area from spring to fall, they moved out in winter, probably because of deep snow. A spotlight count showed that the deer prefer the eastern area as spring-summer range, where the forest consists of coniferous trees over a floor cover ofSasa nipponica, andS. nipponica grassland. The deer also use dense coniferous forest for cover, especially in summer and fall. Part of this study was presented at the 108th Annual Meeting of the Japanese Forestry Society (1997). This study was financially supported by the Environment Agency, Japan.     

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.     

Basal area growth and mortality of Betula maximowicziana affected by crown dieback in a secondary forest in Hokkaido, northern Japan

Since the late 1990s, decline of B. maximowicziana Regel has been observed in mature secondary forests in various parts of Hokkaido, northern Japan. To develop a method of thinning for large-timber production of B. maximowicziana, we measured basal area growth and the mortality of 217 trees during a four-year period (1999–2003) in a 90-year-old secondary forest with serious tree damage. We analyzed growth and mortality in relation to the degree of crown dieback (DC), symmetric and asymmetric competition from neighbors, and initial tree size. Individual basal area growth decreased with increasing DC, with increasing symmetric (two-sided) competition, and with decreasing initial tree size. During the four-year period, 4.1% of the observed trees died. Logistic regression analysis revealed that mortality rate increased with increasing DC and with increasing symmetric competition. These results suggest that both growth and mortality rates were affected by the same factors (i.e., DC and symmetric competition). We concluded that the resource for which individuals were competing at the study site was underground, most likely water. Modeled growth and mortality rates can be used to improve the management of damaged forests. A management plan for the damaged study site is proposed.     

Over-expression of Tryptophan Decarboxylase Gene in Poplar and its Possible Role in Resistance Against Malacosoma disstria

Amines and their derivatives are known to influence insect behavior involved in feeding and reproduction. In order to examine the feasibility of improving the resistance of poplar to insect pests by the introduction of a plant-derived amine-generating transgene, explants from the hybrid poplar clone ‘INRA 717 1B4’ (P. tremula ×P. albo) were transformed with a Camptotheca acuminata tryptophan decarboxylase (TDC, EC 4.1.1.28) cDNA driven by the CaMV 35S promoter. The enzyme TDC catalyzes the decarboxylation of tryptophan to tryptamine, which, in addition to being a bioactive amine itself, is known to act as a precursor of various other indole derivatives. Putative transgenic lines were confirmed by PCR for the TDC1 gene sequence and by the expression analysis of the transgene mRNA and encoded protein. No visible phenotypic changes were associated with ectopic TDC1 expression. Chemical and radiotracer analyses of the transgenic plants revealed tryptamine accumulation as high as 4 mM in leaf tissue, and suggested that the tryptamine produced by ectopically expressed TDC was not further metabolized. Insect bioassays with the TDC transgenic plants showed that the tryptamine accumulation was consistently associated with adverse effects on feeding potential and physiology of Malacosoma disstria (forest tent caterpillar).     

Virulence of ophiostomatoid fungi associated with the spruce bark beetleIps typographus f.japonicus in Yezo spruce

Yezo spruce trees (Picea jezoensis), approximately 40-year-old were inoculated with eight ophiostomatoid fungi associated withIps typographus f.japonicus to compare relative virulence of the fungi. Among them,Ophistoma penicillatum formed the longest necrotic lesion on inner bark around inoculation points, followed byO. aenigmaticum, Ceratocystis polonica, andO. bicolor, whileC. polonica formed a larger dry zone in sapwood than the other fungi. Yezo spruce trees were also mass inoculated withC. polonica, O. penicillatum, O. piceae singly or mixed to demonstrate the ability of the fungi to kill Yezo spruce trees. The trees inoculated withC. polonica, O. penicillatum or the mixed inoculum showed discoloration of needles in the early summer of the next year and died by autumn. However, the trees inoculated withO. piceae or the control inocula did not die, except for one tree. These results indicated thatC. polonica andO. penicillatum were more virulent thanO. piceae and suggested that they might be at least partially responsible for the mortality of the beetle-infested Yezo spruce trees. Part of this study was supported by the Sumitomo Foundation, Japan to Y. Yamaoka and I. Takahashi. Part of this study was presented at the 107th meeting of the Japanese Forestry Society, April 1–4, 1996, Tsukuba, Ibaraki, at the 42nd annual meeting of the Mycological Society of Japan, May 16–17, 1998, Kyoto, and at the 110th meeting of the Japanese Forestry Society, April 2–5, 1999, Matsuyama, Ehime. Contribution No. 143, Laboratories of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Water relations of Yezo spruce and Todo fir in declined stands of boreal Forest in Hokkaido, Japan

Recently, symptoms of decline have been widely observed in Yezo spruce (Picea jezoensis) and Todo fir (Abies sachalinensis) in Hokkaido. In order to clarify the mechanism of decline, the water status of Yezo spruce and Todo fir trees in the Tokyo University Forest in Hokkaido were investigated. The decline is observed mainly in stands damaged by Typhoon XV of 1981 (D-stands), but is not observed in undamaged stands (C-stands). Sampled trees in both types of stands were selected, the severity of their decline estimated, diurnal water potentials measured and water relation parameters such as water potential at turgor-loss point ( ) and osmotic potential at full saturation ( ) were estimated by pressure-volume analysis. The diurnal trends of leaf water potential and midday water potential ( ) of the sample trees were clearly different between the D-stands and the C-stand. Water stress in trees was much more severe in the D-stands than in the C-stand. and of the declining trees were lower in the D-stands than in the C-stand. However, and were not significantly different between declining trees and healthy-looking trees in D-stands. It was concluded that stand-level water stress, induced by the destruction of the canopy by the typhoon, was associated with the decline symptoms. The measurement of water relation parameters proved to be useful for diagnosis and prediction of decline at a stand level in this area.     

Effects of bark harvest and other human activity on populations of the African cherry (Prunus africana) on Mount Oku, Cameroon

Bark extracts from the African cherry, Prunus africana (Rosaceae), are a popular treatment for enlarged prostates. Harvests of the bark began in Cameroon in the 1970s. Because of concerns regarding the sustainability of the trade, the species is included on the IUCN Red List and in CITES Appendix II. This study followed five P. africana populations in the Kilum-Ijim Forest Preserve on Mount Oku, Cameroon, examining growth, mortality and reproductive parameters, as well as response to harvest and other human activities. During the first part of the study (1998–1999), the forest had limited human activity; by the second part (2007–2008), more activity was apparent, including wildfires, grazing and a forest-wide bark harvest in 2005/2006. Over the study period, population structure differed from a typical J-shaped frequency curve for long-lived species, which may reflect past harvesting. After the 2005/2006 harvest, the population structure had shifted slightly toward the smaller size classes. In addition, the number of surviving trees was reduced in all size classes. About half of the reproductive trees died during the study. Size class was not a significant predictor of death, but the location of the harvest (plot) was. All trees affected by wildfires died, suggesting that the species is not adapted to fire. Trees that were harvested without disrupting the vascular cambium survived better and had minimal loss of crown. Thus, the fate of the trees in a given plot may lie in the care taken by an individual harvester. Average growth (0.34 cm per year) was not significantly different among the size classes. Crown die-back significantly reduced fruit production, obscuring the asynchronous alternating fruiting pattern. Seedling numbers followed a similar alternating pattern, but survival was negligible due to grazing. The combined factors of mortalities of a large percentage of reproductive trees (especially the largest ones), highly reduced fruit production and poor seedling survival offer a bleak prognosis for future regeneration and long-term persistence of the species in this forest. Only after decades of harvest are existing standing crop inventories and scientifically based annual quotas now being determined. It is known that the trees are easily domesticated. Efforts have been intensified to train villagers and community forest managers in vegetative propagation techniques and nursery practices, offering some hope that the species can be successfully managed to provide for sustainable harvests and dependable rural livelihoods.