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Susceptible conditions for debarking by deer in subalpine coniferous forests in central Japan

Susceptible conditions for debarking by deer in subalpine coniferous forests in central Japan Background: Recently, deer have expanded their distribution to higher altitude ranges including subalpine forests. However, culling deer and construction of deer fence in subalpine forests are difficult because of steep slopes and complex topography. Thus it is necessary to clarify the factors which are associated with debarking by deer for the effective protection of subalpine forests. In this study, we examined which factors are associated with debarking by sika deer (Cervus nippon) in subalpine coniferous forests. Methods: We conducted our survey in Minami-Alps National Park, central Japan. We established 24 10 m × 40 m plots and surveyed the occurrence of debarking on saplings >30 cm in height and <3 cm in diameter at breast height (DBH) and on trees >3 cm in DBH, as well as sapling density within each plot. Minimum distances to nearest grassland of plots were calculated (tentatively assuming grassland would attract deer and would cause high debarking pressure in the surrounding subalpine forests). Results: The mean percentage of debarked live saplings was higher than that of live trees. The mean percentage of debarked saplings which had already died was 81.6 %. Debarking of saplings increased with lower elevation, taller sapling size, and marginally increased near grassland. Sapling density was lower in plots with low basal area of conspecific trees near grassland and differed among species. Sapling density marginally decreased with decreasing elevation and increasing stand tree density. Debarking of trees was positively related to small DBH and low elevation, and marginally increased near grassland and differed among species. Conclusions: Our results suggest that tall saplings in subalpine forests of low elevation or near subalpine grassland were susceptible to debarking by deer and monitoring of these areas may permit the early detection of the impacts of deer in subalpine coniferous forests. Keywords: Abies, Cervus nippon, Debarking, Grassland, Picea, Sapling density, Subalpine region Background et al. 2013) in forests. To deal with the overabundance In recent years, the population densities of large ungu- of deer and its effect on forest ecosystems, control of lates, especially deer species, have increased worldwide nuisance deer and the construction of fences to protect (Stewart and Burrows 1989; Fuller and Gill 2001; Rooney vegetation have been conducted. However, such counter- 2001; Apollonio et al. 2010; Iijima et al. 2013). An in- measures are difficult to perform in subalpine forests be- crease of deer density has been shown to result in more cause of difficult access (e.g., steep slopes, complex prevalent debarking of trees (Akashi and Nakashizuka topography, great distances from roads). Deer seasonally 1999; Nagaike and Hayashi 2003; Iijima and Nagaike migrate to higher elevations, where delayed bud flush 2015) and browsing of saplings and understory vegeta- caused by low temperature provides fresh leaves even in tion (Gill and Beardall 2001; Beguin et al. 2009; Suzuki mid-summer and they can escape from predators and more intense hunting pressure in low-elevation areas (Mysterud et al. 2011). These factors also have caused * Correspondence: hayato.iijima@gmail.com the number of deer in many higher elevation areas to Yamanashi Forest Research Institute, Saisyoji 2290-1, Fujikawa, Yamanashi 400-0502, Japan © 2015 Iijima and Nagaike. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 2 of 7 increase (Takatsuki 2009), and debarking in subalpine attractive habitat (e.g., subalpine grassland) may affect forests has increased (Yokoyama et al. 2001; Takeuchi deer density. Deer generally prefer herbaceous plants to et al. 2011) in recent years. Therefore, factors correlated woody plants (Takatsuki 1986; Winnie 2012), and se- with debarking by deer in subalpine forests should be vere browsing of herbaceous plants in subalpine grass- clarified to help effectively conserve these forests. lands by deer has been reported (Schütz et al. 2003; Many studies have examined the factors influencing Nagaike 2012; Nagaike et al. 2014). Thus, subalpine debarking and browsing in lower elevation forests. Debark- forests which were surrounded by large subalpine grass- ing was shown to depend on the size of trees (e.g., Nagaike lands would be more susceptible for debarking by deer. and Hayashi 2003; Koda and Fujita 2011; Borkowski and Thus far, however, direct evaluation of the relationship Ukalski 2012), their species (e.g., Kay 1993; Akashi and between these factors and the intensity of debarking in Nakashizuka 1999; Moore et al. 1999; Takeuchi et al. subalpine forest is rare. 2011), proportion of coniferous stands (Ligot et al. 2013), The objective of this study was to clarify the conditions the distance from forest road (McLaren et al. 2000), and which are correlated with debarking by sika deer in sub- snow depth (Iijima and Nagaike 2015). In addition to these alpine forests in order to improve the effective manage- factors, spatial variation of deer impact was observed in ment and conservation of these subalpine forests. We lower elevation forests: the higher the deer density, the hypothesized that the occurrence of debarking and sapling higher the proportion of debarked trees (Iijima and density would increase and decrease in low altitude and Nagaike 2015) and browsed saplings (Akashi et al. 2011) abundant grassland. and the lower the sapling density (Beguin et al. 2009). The spatial variation of deer density across low-elevation Methods areas was explained by the presence of attractive habi- Study area tat (e.g., artificial grassland; Kamei et al. 2010; Iijima et The present study was conducted in subalpine forests in al. 2013). However, the attractive habitat in subalpine Minami-Alps National Park (357.5 km )incentral Japan forests, which are located on steeper slopes and have (Fig. 1), where sika deer (Cervus nippon) density has been more complex topography than lower elevation areas, increasing in recent years (Izumiyama and Mochizuki has not been well studied. 2008; Izumiyama et al. 2009). The park is characterized by Takeuchi et al. (2011) reported that elevation was an steep slopes and a concentration of many mountains important factor affecting expansion of deer density into within a small area (Fig. 1). Subalpine coniferous forests subalpine zones. In addition to elevation, the amount of dominate from 2000 to 2500 m elevation, mainly Fig. 1 Map of the study area. Gray polygons indicate the locations of subalpine grasslands (from the Natural Environment Information geographic information system provided by the Biodiversity Center of Japan: http://www.biodic.go.jp/trialSystem/top.html). Solid and dashed lines denote prefectural boundaries and trails, respectively. Solid squares indicate surveyed plots. This map was drawn by QGIS (QGIS Development 2015) Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 3 of 7 composed of Abies mariesii, Abies veitchii, Picea jezoensis along the two long sides of each plot (8 subplots per a var. hondoensis,and Tsuga diversifolia (Fig. 2). Subalpine plot). Height and the occurrence of debarking on grasslands consisting of herbaceous species and dwarf tree saplings >30 cm in height and <3 cm in DBH were community (Pinus pumila) are present at elevations measured and recorded in each subplot. In this study, we higher than 2500 m, and mixed conifer-broadleaf forests could not determine the timing of the occurrence of are present below 2000 m. Sika deer are widely distributed debarking and did not distinguish new and old debarking. in Japan. Their shoulder heights range from 90 to 190 cm and they have a highly varied diet (they eat graminoids in The minimum distance from each plot to subalpine northern Japan and eat leaves and fruits in southern Japan; grassland Ohdachi et al. 2009). They can reach bark and leaves up The location of subalpine grassland in this region was to ca. 2 m in height. obtained from the Natural Environment Information geographic information system (http://www.biodic.go.jp/ Field surveys trialSystem/top.html, accessed 20 April 2015) provided We established 24 rectangular plots (each of 10 m × 40 m) by the Biodiversity Center of Japan (Fig. 1). The mini- in natural coniferous forests at elevations of 2000 m (low- mum distance from each plot to subalpine grassland elevation zone) and 2500 m (high-elevation zone) and sur- was calculated by QGIS ver 2.8.1 (Quantum GIS De- veyed in September 2011 (Fig. 1). For all trees with the velopment 2015). diameter at breast height (DBH) > 3 cm in the plots, we noted the species and measured DBH and the presence of Statistical analysis debarking. Furthermore, for saplings survey, 8 subplots Sapling density and the occurrence of debarking of sap- (each of 1 m × 2 m) were established at 10-m intervals lings and trees were analyzed using a generalized linear Abies mariesii Abies veitchii Picea jezoensis Tsuga diversifolia Others Low High Elevation zones Fig. 2 Species composition of the dominant trees at low- and high-elevation zones. BA is basal area of trees at breast height. “Others” include Larix kaempferi, Betula corylifolia, Betula ermanii, Prunus maximowiczii, and Sorbus commixta The percentage of BA (%) 0 20 40 60 80 100 Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 4 of 7 mixed model (GLMM). For all models, we adopted each variables. A binomial distribution was adopted as the error plot as random effect. distribution with a logit link function. For analysis of sapling density, minimum distance All analyses were conducted using R ver 3.1.3 (R Core from each plot to subalpine grassland, elevation zone, Team 2015) and the glmmML package (Brostrom and tree density in each plot (indicating understory light Holmberg 2011). Model selection was performed using conditions), and species of each sapling were used as the Akaike information criterion (AIC, Akaike 1973) in explanatory variables. We used the number of saplings conjunction with a backward elimination procedure. We as response variable. The means of the number of sap- determined that explanatory variables used in conjunc- lings and sapling density were equal because surveyed tion with all models which had delta AIC compared to areas of the number of saplings were same for all plots. the lowest AIC < 2 are significant and explanatory variable We also added basal area (BA) of conspecific trees (as used in conjunction with any models which had delta AIC an index of seed source) as an explanatory variable compared to the lowest AIC < 2 are marginally significant. because the abundance of the seed source may affect At the same time, we calculated Akaike weight (w) and the sapling density. A Poisson distribution was adopted as relative importance of variable (Burnham and Anderson the error distribution with a log-link function. Picea 2002) for comparing the importance of each variable. jezoensis was omitted from the species data because Akaike weight is defined as none of the surveyed individuals had been debarked, exp − which made it impossible to estimate the coefficient. 2 w ¼ For analysis of the occurrence of debarking of saplings, exp − r¼1 minimum distance from each plot to subalpine grass- land, elevation zone, species, and sapling height were where Δ = AIC − AIC is the difference between an i i min included as explanatory variables. A binomial distribu- AIC of each model and the minimum AIC among all tion was adopted as the error distribution with a logit candidate models (including null model and full model). link function. This value, referred to as the Akaike weight, provides a For analysis of the occurrence of debarking of trees, relative weight of evidence for each model. The relative minimum distance from each plot to subalpine grassland, importance of predictor variable can be calculated as the elevation zone, DBH, and species were used as explanatory sum of the Akaike weights over all of the models in Fig. 3 Stand conditions Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 5 of 7 Table 1 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining the occurrence of debarking on saplings a b,c AIC ΔAIC Elevation zone Dist. grassland Height of saplings Species 2500 m Abies veitchii Tsuga diversifolia 398.028 0.000 −2.634 0.037 −4 399.976 1.948 −2.619 −1.269 × 10 0.037 a b Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0. Picea jezoensis was excluded from the analysis of saplings because no P. jezoensis saplings were debarked, so it was impossible to estimate the coefficient which the parameter of interest appears. The relative dead saplings were much higher than those for live sap- importance range from 0 (least important) to 1 (most lings (Fig. 3). These results suggest that debarking by important). deer first occurs on saplings. Tree and sapling sizes affected the occurrence of Results debarking but the effects of sizes showed opposite pat- The percentages of debarked live trees differed among terns for trees and saplings (Tables 1 and 3). Higher per- plots (Fig. 3). The differences among sapling types was centages of debarked small trees were also reported in large, with dead saplings being debarked at very high many previous studies (Akashi and Nakashizuka 1999; rates (mean = 81.6 %; Fig. 3). Yokoyama et al. 2001; Nagaike and Hayashi 2003; Jiang Debarking of saplings was significantly related to a taller et al. 2005; Kiffner et al. 2008; Takeuchi et al. 2011; height of saplings and lower elevation zone, marginally Borkowski and Ukalski 2012), although the reasons were related to minimum distance to grassland, and was not not clarified. One possibility for the selective debarking affected by species (Table 1). Sapling density significantly of small trees is the relative ease of debarking smaller increased with increasing BA of conspecific trees and in- trees (Ando et al. 2004). In contrast to trees, the rela- creasing minimum distance to grassland and marginally tionship between sapling size and the occurrence of increased with increasing elevation and decreasing tree debarking has rarely been examined. Takatsuki and density (Table 2). Sapling density of A. mariesii was sig- Gorai (1994) compared the size structure of trees be- nificantly higher than that of other species (Table 2). tween two distinct areas in Japan that differed in deer Debarking of trees was significantly related to small density. They showed that forests with high deer density DBH and low elevation zone, and marginally decreased lacked saplings > 90 cm in height, whereas there were with long distance to grassland. Debarking of trees abundant saplings shorter than 90 cm in forests both marginally differed by species (Table 3). Overall, Size of under high and low deer densities. In the present study, saplings and trees had strong effects on the occurrence one reason for the low debarking intensity on small sap- of debarking (Table 4). Elevation zone, minimum dis- lings (Table 1) is that they are more difficult to find than tance to grassland, and species were marginal factors of large saplings. Consequently, saplings >30 cm in height the occurrence of debarking of saplings and trees and and < 3 cm in DBH might be more susceptible to sapling density (Table 4). debarking by deer, suggesting that these might be a pos- sible optimal size for debarking by sika deer. Discussion P. jezoensis saplings appear to be less palatable for sika We showed the relationship between debarking of trees deer because no P. jezoensis saplings was debarked and and saplings in subalpine coniferous forests and various we found marginal effect of species on debarking of trees factors. The percentages of debarked live saplings were (Table 3). Generally, Abies species were preferentially higher than those of live trees, and the percentages for debarked over co-occurring Picea species (Heuze et al. Table 2 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining sapling density a b c AIC ΔAIC Elevation zone BA of conspecific Dist. grassland Species Tree density 2500 m trees Abies veitchii Picea jezoensis Tsuga diversifolia −4 1382.409 0.000 0.048 9.310 × 10 −0.555 −2.675 −1.237 −4 1382.624 0.214 0.745 0.048 9.561 × 10 −0.559 −2.676 −1.236 −0.018 −4 1383.631 1.272 0.210 0.048 9.372 × 10 −0.557 −2.680 −1.239 −4 1384.286 1.877 0.048 9.311 × 10 −0.554 −2.672 −1.239 −0.002 a b c Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. BA, basal area. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0 Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 6 of 7 Table 3 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining the occurrence of debarking on trees a b c AIC Δ AIC Elevation zone DBH of Dist. grassland Species 2500 m trees Abies veitchii Picea jezoensis Tsuga diversifolia −4 526.047 0.000 −0.684 −0.014 −4.146 × 10 −4 526.261 0.214 −0.771 −0.014 −4.196 × 10 0.127 −1.379 −0.947 527.907 1.856 −0.592 −0.014 527.974 1.926 −0.666 −0.014 0.076 −1.453 −0.959 a b Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. DBH, diameter at breast height. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0 2005), and the different palatability among tree species as rich food sources (Takatsuki 1986; Winnie 2012). suggests that deer debarking may alter the species com- Trees and saplings in subalpine forests near subalpine position of forests. However, P. jezoensis has been heavily grassland would be eaten by deer before sprout of debarked in some areas in Japan (Yokoyama et al. 2001; grasses in early spring and after defoliation of grasses in Ando et al. 2003), indicating that palatability of debark- late autumn. Some sika deer migrate to high-elevation ing may be affected by surrounding vegetation (Moore area before sprout of grasses and after defoliation to es- et al. 1999; McLaren et al. 2000). Therefore, P. jezoensis cape the competition with other deer in low-elevation saplings at the sites examined in the present study may areas (Shin-ichiro Hamasaki, personal communication). be debarked in the future if the deer density increases. Minimum distance to subalpine grassland, however, In addition to size and species as factors for debarking, had only a marginal effect on the occurrence of debark- elevation zones and grassland area also were associated ing of saplings and trees (Tables 1 and 3). As the result with the damage to subalpine forests by deer. Low of intensive debarking on saplings (Fig. 3), low sapling sapling density and high risk of debarking of trees and density near subalpine grassland occurred (Table 2) and saplings were found in low elevation plots (Table 2). then it might be obscure the relationship between the Other studies in Japan also found the intensities of distance to grassland and the occurrence of debarking debarking on trees and saplings in subalpine coniferous of saplings. In the future, the intensity of debarking of forests to be greater at lower elevations (Takeuchi et al. tree would increase if deer density remains at this level 2011). In our study area, sika deer migrate to high- or increases. elevation areas in summer and to low-elevation areas from late autumn to late spring (Izumiyama and Mochizuki 2008) although there are some deer in low-elevation area Conclusions even in summer. Then, opportunities of debarking for Our analysis revealed that large saplings within the range saplings and trees were larger in low-elevation zone than of 30 cm in height and 3 cm in DBH in subalpine forests in high-elevation because deer stay longer time in low were most susceptible for debarking by sika deer. In elevation-area than in high elevation-area. In addition to addition to the size, saplings in low elevation zone or near elevation, we found that the large subalpine grassland subalpine grasslands were more susceptible to debarking caused low sapling density (Table 2) and high risk of the by sika deer. This information may be useful to improve occurrence of debarking of saplings and trees (Tables 1 monitoring deer impact, or determining where to con- and 3). Deer would likely be attracted to subalpine grass- struct deer fences in the subalpine region. At the same land because it has numerous herbaceous plants that serve time, deer population control in the subalpine region should be conducted in forests of low elevation zone or Table 4 Relative importance of variables in models near subalpine grasslands. Debarking of Sapling density Debarking of saplings trees Competing interests Elevation zone 1.000 0.506 0.785 This study was funded by the Comprehensive Research Organization for BA of conspecific species Not included 1.000 Not included Science and Technology of Yamanashi Prefectural Government and Mitsui & CO., LTD. The authors declare that they have no competing interests about DBH of trees Not included Not included 1.000 non-financial aspects. Dist. grassland 0.274 1.000 0.674 Height of saplings 1.000 Not included Not included Authors’ contributions Species 0.137 1.000 0.463 HI and TN substantially contributed about setting the experimental design and collecting data. HI and TN deeply discussed and approved the content Tree density Not included 0.457 Not included of manuscript. Both authors read and approved the final manuscript. Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 7 of 7 Acknowledgements Moore NP, Hart JD, Langton SD (1999) Factors influencing browsing by fallow We thank Chiaki Ootsu for field assistance, as well as the staff of the deer Dama dama in young broad-leaved plantations. Biol Cons 87:255–260 “Komorebi-sansou” mountain hut. Mysterud A, Loe LE, Zimmermann B, Bischof R, Veiberg V, Meisingset E (2011) Partial migration in expanding red deer populations at northern latitudes: Received: 23 July 2015 Accepted: 20 December 2015 A role for density dependence? Oikos 120:1817–1825 Nagaike T, Hayashi A (2003) Bark-stripping by sika deer (Cervus nippon)in Larix kaempferi plantations in central Japan. For Ecol Manage 175:563–572 Nagaike T (2012) Effects of browsing by sika deer (Cervus nippon) on subalpine References vegetation at Mt. Kita, central Japan. Ecol Res 27:467–473 Akaike H (1973) Information theory and an extension of the maximum likelihood Nagaike T, Ohkubo E, Hirose K (2014) Vegetation recovery in response to the principle, in 2nd International Symposium on Information Theory. Tsahkadsor, exclusion of grazing by sika deer (Cervus nippon) in seminatural grassland on Armenian SSR, pp 267–281 Mt. Kushigata, Japan. ISRN Biodiversity: Article ID 493495 Akashi N, Nakashizuka T (1999) Effects of bark-stripping by sika deer (Cervus nippon) Ohdachi SD, Ishibashi Y, Iwasa MA, Saitoh T (2009) The Wild Mammals of Japan. on population dynamics of a mixed forest in Japan. For Ecol Manage 113:75–82 Shoukadoh Book Sellers, Kyoto Akashi N, Unno A, Terazawa K (2011) Effects of deer abundance on broad-leaf QGIS Development Team (2015) Quantum GIS Geographic Information System. tree seedling establishment in the understory of Abies sachalinensis Open Source Geospatial Foundation Project. http://qgis.osgeo.org plantations. J For Res 16:500–508 R Core Team (2015) R: A Language and Environment for Statistical Computing. R Ando M, Yokota H, Shibata E (2003) Bark stripping preference of sika deer, Cervus Foundation for Statistical Computing, Vienna, Austria, http://www.R-project.org/ nippon, in terms of bark chemical contents. For Ecol Manage 177:323–331 Rooney TP (2001) Deer impacts on forest ecosystems: a North American Ando M, Yokota H, Shibata E (2004) Why do sika deer, Cervus nippon, debark perspective. Forestry 74:201–208 trees in summer on Mt. Ohdaigahara, central Japan? Mamm Study 29:73–83 Schütz M, Risch AC, Leuzinger E, Krüsi BO, Achermann G (2003) Impact of herbivory Apollonio M, Andersen R, Putman R (2010) European ungulates and their by red deer (Cervus elaphus L.) on patterns and processes in subalpine management in the 21st century. Cambridge University Press, NY. grasslands in the Swiss National Park. For Ecol Manage 181:177–188 Beguin J, Pothier D, Prévost M (2009) Can the impact of deer browsing on tree Stewart GH, Burrows LE (1989) The impact of white-tailed deer Odocoileus regeneration be mitigated by shelterwood cutting and strip clearcutting? For virginianus on regeneration in the coastal forests of Stewart Island, New Ecol Manage 257:38–45 Zealand. Biol Cons 49:275–293 Borkowski J, Ukalski K (2012) Bark stripping by red deer in a post-disturbance Suzuki M, Miyashita T, Kabaya H, Ochiai K, Asada M, Kikvidze Z (2013) Deer area: the importance of security cover. For Ecol Manage 263:17–23 herbivory as an important driver of divergence of ground vegetation Brostrom G, Holmberg H (2011) Generalized linear models with clustered data: communities in temperate forests. Oikos 122:104–110 fixed and random effects models. Comput Stat Data Anal 55:3123–3134 Takatsuki S (1986) Food habits of sika deer on Mt. Goyo, northern Honshu. Ecol Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a Res 1:119–128 practical information-theoretic approach, 2nd edn. Springer, New York Takatsuki S, Gorai T (1994) Effects of sika deer on the regeneration of a Fagus Fuller RJ, Gill RMA (2001) Ecological impacts of increasing numbers of deer in crenata forest on Kinkazan Island, northern Japan. Ecol Res 9:115–120 British woodland. Forestry 74:193–199 Takatsuki S (2009) Effects of sika deer on vegetation in Japan: a review. Biol Cons Gill RMA, Beardall V (2001) The impact of deer on woodlands: the effects of 142:1922–1929 browsing and seed dispersal on vegetation structure and composition. Takeuchi T, Kobayashi T, Nashimoto M (2011) Altitudinal differences in bark Forestry 74:209–218 stripping by sika deer in the subalpine coniferous forest of Mt. Fuji. For Ecol Heuze P, Schnitzlerm A, Klein F (2005) Consequences of increased deer browsing Manage 261:2089–2095 in winter on silver fir and spruce regeneration in the southern Vosges Winnie JAJ (2012) Predation risk, elk, and aspen: tests of a behaviorally mediated Mountains: implication for forest management. Ann For Sci 62:175–181 trophic cascade in the Greater Yellowstone Ecosystems. Ecology 93:2600–2614 Iijima H, Nagaike T, Honda T (2013) Estimation of deer population dynamics by Yokoyama S, Maeji I, Ueda T, Ando M, Shibata E (2001) Impact of bark stripping Bayesian state-space model with multiple abundance indices. J Wildl Manage by sika deer, Cervus nippon, on subalpine coniferous forests in central Japan. 77:1038–1047 For Ecol Manage 140:93–99 Iijima H, Nagaike T (2015) Appropriate vegetation indices for measuring the impacts of deer on forest ecosystems. Ecol Ind 48:458–463 Izumiyama S, Mochizuki T (2008) Seasonal range use of sika deer, which inhabits the subalpine zone in the southern Japan Alps. Bull Shinshu Univ Alps Field Center 6:25–32 (in Japanese with English summary) Izumiyama S, Mochizuki T, Takii A (2009) GPS tracking of Sika deer which inhabits the sub-alpine zone in the Southern Japan Alps. Bull Shinshu Univ Alps Field Center 7:63–71 (in Japanese with English summary) Jiang Z, Ueda H, Kitahara M, Imaki H (2005) Bark stripping by sika deer on Veitch fir related to stand age, bark nutrition, and season in northern Mount Fuji district, central Japan. J For Res 10:359–365 Kamei T, Takeda K, Koh K, Izumiyama S, Watanabe O, Ohshima K (2010) Seasonal pasture utilization by wild sika deer (Cervus nippon) in a sown grassland. Grass Sci 56:65–70 Kay S (1993) Factors affecting severity of deer browsing damage within coppiced woodlands in the south of England. Biol Cons 63:217–222 Submit your manuscript to a Kiffner C, Rößiger E, Trisl O, Schulz R, Rühe F (2008) Probability of recent bark journal and benefi t from: stripping damage by red deer (Cervus elaphus) on Norway spruce (Picea abies) in a low mountain range in Germany: a preliminary analysis. Silva 7 Convenient online submission Fennica 42:125–134 7 Rigorous peer review Koda R, Fujita N (2011) Is deer herbivory directly proportional to deer population density? Comparison of deer feeding frequencies among six forests with 7 Immediate publication on acceptance different deer density. For Ecol Manage 262:432–439 7 Open access: articles freely available online Ligot G, Gheysen T, Lehaire F, Hébert J, Licoppe A, Lejeune P, Brostaux Y (2013) 7 High visibility within the fi eld Modeling recent bark stripping by red deer (Cervus elaphus) in South 7 Retaining the copyright to your article Belgium coniferous stands. Ann For Sci 70:309–318 McLaren BE, Mahoney SP, Porter TS, Oosenbrug SM (2000) Spatial and temporal patterns of use by moose of pre-commercially thinned, naturally-regeneration Submit your next manuscript at 7 springeropen.com stands of balsam fir in central Newfoundland. For Ecol Manage 133:179–196 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png "Forest Ecosystems" Springer Journals

Susceptible conditions for debarking by deer in subalpine coniferous forests in central Japan

"Forest Ecosystems" , Volume 2 (1): 7 – Dec 1, 2015

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Springer Journals
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2015 Iijima and Nagaike.
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2197-5620
DOI
10.1186/s40663-015-0059-y
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Abstract

Background: Recently, deer have expanded their distribution to higher altitude ranges including subalpine forests. However, culling deer and construction of deer fence in subalpine forests are difficult because of steep slopes and complex topography. Thus it is necessary to clarify the factors which are associated with debarking by deer for the effective protection of subalpine forests. In this study, we examined which factors are associated with debarking by sika deer (Cervus nippon) in subalpine coniferous forests. Methods: We conducted our survey in Minami-Alps National Park, central Japan. We established 24 10 m × 40 m plots and surveyed the occurrence of debarking on saplings >30 cm in height and <3 cm in diameter at breast height (DBH) and on trees >3 cm in DBH, as well as sapling density within each plot. Minimum distances to nearest grassland of plots were calculated (tentatively assuming grassland would attract deer and would cause high debarking pressure in the surrounding subalpine forests). Results: The mean percentage of debarked live saplings was higher than that of live trees. The mean percentage of debarked saplings which had already died was 81.6 %. Debarking of saplings increased with lower elevation, taller sapling size, and marginally increased near grassland. Sapling density was lower in plots with low basal area of conspecific trees near grassland and differed among species. Sapling density marginally decreased with decreasing elevation and increasing stand tree density. Debarking of trees was positively related to small DBH and low elevation, and marginally increased near grassland and differed among species. Conclusions: Our results suggest that tall saplings in subalpine forests of low elevation or near subalpine grassland were susceptible to debarking by deer and monitoring of these areas may permit the early detection of the impacts of deer in subalpine coniferous forests. Keywords: Abies, Cervus nippon, Debarking, Grassland, Picea, Sapling density, Subalpine region Background et al. 2013) in forests. To deal with the overabundance In recent years, the population densities of large ungu- of deer and its effect on forest ecosystems, control of lates, especially deer species, have increased worldwide nuisance deer and the construction of fences to protect (Stewart and Burrows 1989; Fuller and Gill 2001; Rooney vegetation have been conducted. However, such counter- 2001; Apollonio et al. 2010; Iijima et al. 2013). An in- measures are difficult to perform in subalpine forests be- crease of deer density has been shown to result in more cause of difficult access (e.g., steep slopes, complex prevalent debarking of trees (Akashi and Nakashizuka topography, great distances from roads). Deer seasonally 1999; Nagaike and Hayashi 2003; Iijima and Nagaike migrate to higher elevations, where delayed bud flush 2015) and browsing of saplings and understory vegeta- caused by low temperature provides fresh leaves even in tion (Gill and Beardall 2001; Beguin et al. 2009; Suzuki mid-summer and they can escape from predators and more intense hunting pressure in low-elevation areas (Mysterud et al. 2011). These factors also have caused * Correspondence: hayato.iijima@gmail.com the number of deer in many higher elevation areas to Yamanashi Forest Research Institute, Saisyoji 2290-1, Fujikawa, Yamanashi 400-0502, Japan © 2015 Iijima and Nagaike. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 2 of 7 increase (Takatsuki 2009), and debarking in subalpine attractive habitat (e.g., subalpine grassland) may affect forests has increased (Yokoyama et al. 2001; Takeuchi deer density. Deer generally prefer herbaceous plants to et al. 2011) in recent years. Therefore, factors correlated woody plants (Takatsuki 1986; Winnie 2012), and se- with debarking by deer in subalpine forests should be vere browsing of herbaceous plants in subalpine grass- clarified to help effectively conserve these forests. lands by deer has been reported (Schütz et al. 2003; Many studies have examined the factors influencing Nagaike 2012; Nagaike et al. 2014). Thus, subalpine debarking and browsing in lower elevation forests. Debark- forests which were surrounded by large subalpine grass- ing was shown to depend on the size of trees (e.g., Nagaike lands would be more susceptible for debarking by deer. and Hayashi 2003; Koda and Fujita 2011; Borkowski and Thus far, however, direct evaluation of the relationship Ukalski 2012), their species (e.g., Kay 1993; Akashi and between these factors and the intensity of debarking in Nakashizuka 1999; Moore et al. 1999; Takeuchi et al. subalpine forest is rare. 2011), proportion of coniferous stands (Ligot et al. 2013), The objective of this study was to clarify the conditions the distance from forest road (McLaren et al. 2000), and which are correlated with debarking by sika deer in sub- snow depth (Iijima and Nagaike 2015). In addition to these alpine forests in order to improve the effective manage- factors, spatial variation of deer impact was observed in ment and conservation of these subalpine forests. We lower elevation forests: the higher the deer density, the hypothesized that the occurrence of debarking and sapling higher the proportion of debarked trees (Iijima and density would increase and decrease in low altitude and Nagaike 2015) and browsed saplings (Akashi et al. 2011) abundant grassland. and the lower the sapling density (Beguin et al. 2009). The spatial variation of deer density across low-elevation Methods areas was explained by the presence of attractive habi- Study area tat (e.g., artificial grassland; Kamei et al. 2010; Iijima et The present study was conducted in subalpine forests in al. 2013). However, the attractive habitat in subalpine Minami-Alps National Park (357.5 km )incentral Japan forests, which are located on steeper slopes and have (Fig. 1), where sika deer (Cervus nippon) density has been more complex topography than lower elevation areas, increasing in recent years (Izumiyama and Mochizuki has not been well studied. 2008; Izumiyama et al. 2009). The park is characterized by Takeuchi et al. (2011) reported that elevation was an steep slopes and a concentration of many mountains important factor affecting expansion of deer density into within a small area (Fig. 1). Subalpine coniferous forests subalpine zones. In addition to elevation, the amount of dominate from 2000 to 2500 m elevation, mainly Fig. 1 Map of the study area. Gray polygons indicate the locations of subalpine grasslands (from the Natural Environment Information geographic information system provided by the Biodiversity Center of Japan: http://www.biodic.go.jp/trialSystem/top.html). Solid and dashed lines denote prefectural boundaries and trails, respectively. Solid squares indicate surveyed plots. This map was drawn by QGIS (QGIS Development 2015) Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 3 of 7 composed of Abies mariesii, Abies veitchii, Picea jezoensis along the two long sides of each plot (8 subplots per a var. hondoensis,and Tsuga diversifolia (Fig. 2). Subalpine plot). Height and the occurrence of debarking on grasslands consisting of herbaceous species and dwarf tree saplings >30 cm in height and <3 cm in DBH were community (Pinus pumila) are present at elevations measured and recorded in each subplot. In this study, we higher than 2500 m, and mixed conifer-broadleaf forests could not determine the timing of the occurrence of are present below 2000 m. Sika deer are widely distributed debarking and did not distinguish new and old debarking. in Japan. Their shoulder heights range from 90 to 190 cm and they have a highly varied diet (they eat graminoids in The minimum distance from each plot to subalpine northern Japan and eat leaves and fruits in southern Japan; grassland Ohdachi et al. 2009). They can reach bark and leaves up The location of subalpine grassland in this region was to ca. 2 m in height. obtained from the Natural Environment Information geographic information system (http://www.biodic.go.jp/ Field surveys trialSystem/top.html, accessed 20 April 2015) provided We established 24 rectangular plots (each of 10 m × 40 m) by the Biodiversity Center of Japan (Fig. 1). The mini- in natural coniferous forests at elevations of 2000 m (low- mum distance from each plot to subalpine grassland elevation zone) and 2500 m (high-elevation zone) and sur- was calculated by QGIS ver 2.8.1 (Quantum GIS De- veyed in September 2011 (Fig. 1). For all trees with the velopment 2015). diameter at breast height (DBH) > 3 cm in the plots, we noted the species and measured DBH and the presence of Statistical analysis debarking. Furthermore, for saplings survey, 8 subplots Sapling density and the occurrence of debarking of sap- (each of 1 m × 2 m) were established at 10-m intervals lings and trees were analyzed using a generalized linear Abies mariesii Abies veitchii Picea jezoensis Tsuga diversifolia Others Low High Elevation zones Fig. 2 Species composition of the dominant trees at low- and high-elevation zones. BA is basal area of trees at breast height. “Others” include Larix kaempferi, Betula corylifolia, Betula ermanii, Prunus maximowiczii, and Sorbus commixta The percentage of BA (%) 0 20 40 60 80 100 Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 4 of 7 mixed model (GLMM). For all models, we adopted each variables. A binomial distribution was adopted as the error plot as random effect. distribution with a logit link function. For analysis of sapling density, minimum distance All analyses were conducted using R ver 3.1.3 (R Core from each plot to subalpine grassland, elevation zone, Team 2015) and the glmmML package (Brostrom and tree density in each plot (indicating understory light Holmberg 2011). Model selection was performed using conditions), and species of each sapling were used as the Akaike information criterion (AIC, Akaike 1973) in explanatory variables. We used the number of saplings conjunction with a backward elimination procedure. We as response variable. The means of the number of sap- determined that explanatory variables used in conjunc- lings and sapling density were equal because surveyed tion with all models which had delta AIC compared to areas of the number of saplings were same for all plots. the lowest AIC < 2 are significant and explanatory variable We also added basal area (BA) of conspecific trees (as used in conjunction with any models which had delta AIC an index of seed source) as an explanatory variable compared to the lowest AIC < 2 are marginally significant. because the abundance of the seed source may affect At the same time, we calculated Akaike weight (w) and the sapling density. A Poisson distribution was adopted as relative importance of variable (Burnham and Anderson the error distribution with a log-link function. Picea 2002) for comparing the importance of each variable. jezoensis was omitted from the species data because Akaike weight is defined as none of the surveyed individuals had been debarked, exp − which made it impossible to estimate the coefficient. 2 w ¼ For analysis of the occurrence of debarking of saplings, exp − r¼1 minimum distance from each plot to subalpine grass- land, elevation zone, species, and sapling height were where Δ = AIC − AIC is the difference between an i i min included as explanatory variables. A binomial distribu- AIC of each model and the minimum AIC among all tion was adopted as the error distribution with a logit candidate models (including null model and full model). link function. This value, referred to as the Akaike weight, provides a For analysis of the occurrence of debarking of trees, relative weight of evidence for each model. The relative minimum distance from each plot to subalpine grassland, importance of predictor variable can be calculated as the elevation zone, DBH, and species were used as explanatory sum of the Akaike weights over all of the models in Fig. 3 Stand conditions Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 5 of 7 Table 1 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining the occurrence of debarking on saplings a b,c AIC ΔAIC Elevation zone Dist. grassland Height of saplings Species 2500 m Abies veitchii Tsuga diversifolia 398.028 0.000 −2.634 0.037 −4 399.976 1.948 −2.619 −1.269 × 10 0.037 a b Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0. Picea jezoensis was excluded from the analysis of saplings because no P. jezoensis saplings were debarked, so it was impossible to estimate the coefficient which the parameter of interest appears. The relative dead saplings were much higher than those for live sap- importance range from 0 (least important) to 1 (most lings (Fig. 3). These results suggest that debarking by important). deer first occurs on saplings. Tree and sapling sizes affected the occurrence of Results debarking but the effects of sizes showed opposite pat- The percentages of debarked live trees differed among terns for trees and saplings (Tables 1 and 3). Higher per- plots (Fig. 3). The differences among sapling types was centages of debarked small trees were also reported in large, with dead saplings being debarked at very high many previous studies (Akashi and Nakashizuka 1999; rates (mean = 81.6 %; Fig. 3). Yokoyama et al. 2001; Nagaike and Hayashi 2003; Jiang Debarking of saplings was significantly related to a taller et al. 2005; Kiffner et al. 2008; Takeuchi et al. 2011; height of saplings and lower elevation zone, marginally Borkowski and Ukalski 2012), although the reasons were related to minimum distance to grassland, and was not not clarified. One possibility for the selective debarking affected by species (Table 1). Sapling density significantly of small trees is the relative ease of debarking smaller increased with increasing BA of conspecific trees and in- trees (Ando et al. 2004). In contrast to trees, the rela- creasing minimum distance to grassland and marginally tionship between sapling size and the occurrence of increased with increasing elevation and decreasing tree debarking has rarely been examined. Takatsuki and density (Table 2). Sapling density of A. mariesii was sig- Gorai (1994) compared the size structure of trees be- nificantly higher than that of other species (Table 2). tween two distinct areas in Japan that differed in deer Debarking of trees was significantly related to small density. They showed that forests with high deer density DBH and low elevation zone, and marginally decreased lacked saplings > 90 cm in height, whereas there were with long distance to grassland. Debarking of trees abundant saplings shorter than 90 cm in forests both marginally differed by species (Table 3). Overall, Size of under high and low deer densities. In the present study, saplings and trees had strong effects on the occurrence one reason for the low debarking intensity on small sap- of debarking (Table 4). Elevation zone, minimum dis- lings (Table 1) is that they are more difficult to find than tance to grassland, and species were marginal factors of large saplings. Consequently, saplings >30 cm in height the occurrence of debarking of saplings and trees and and < 3 cm in DBH might be more susceptible to sapling density (Table 4). debarking by deer, suggesting that these might be a pos- sible optimal size for debarking by sika deer. Discussion P. jezoensis saplings appear to be less palatable for sika We showed the relationship between debarking of trees deer because no P. jezoensis saplings was debarked and and saplings in subalpine coniferous forests and various we found marginal effect of species on debarking of trees factors. The percentages of debarked live saplings were (Table 3). Generally, Abies species were preferentially higher than those of live trees, and the percentages for debarked over co-occurring Picea species (Heuze et al. Table 2 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining sapling density a b c AIC ΔAIC Elevation zone BA of conspecific Dist. grassland Species Tree density 2500 m trees Abies veitchii Picea jezoensis Tsuga diversifolia −4 1382.409 0.000 0.048 9.310 × 10 −0.555 −2.675 −1.237 −4 1382.624 0.214 0.745 0.048 9.561 × 10 −0.559 −2.676 −1.236 −0.018 −4 1383.631 1.272 0.210 0.048 9.372 × 10 −0.557 −2.680 −1.239 −4 1384.286 1.877 0.048 9.311 × 10 −0.554 −2.672 −1.239 −0.002 a b c Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. BA, basal area. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0 Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 6 of 7 Table 3 AIC and coefficients of models which had ΔAIC < 2 compared to the lowest AIC model for explaining the occurrence of debarking on trees a b c AIC Δ AIC Elevation zone DBH of Dist. grassland Species 2500 m trees Abies veitchii Picea jezoensis Tsuga diversifolia −4 526.047 0.000 −0.684 −0.014 −4.146 × 10 −4 526.261 0.214 −0.771 −0.014 −4.196 × 10 0.127 −1.379 −0.947 527.907 1.856 −0.592 −0.014 527.974 1.926 −0.666 −0.014 0.076 −1.453 −0.959 a b Coefficient of elevation zone was calculated as a relative value when the coefficient of the low-elevation zone (2000 m) was set to 0. DBH, diameter at breast height. Coefficient of species was calculated as a relative value when the coefficient of Abies mariesii was set to 0 2005), and the different palatability among tree species as rich food sources (Takatsuki 1986; Winnie 2012). suggests that deer debarking may alter the species com- Trees and saplings in subalpine forests near subalpine position of forests. However, P. jezoensis has been heavily grassland would be eaten by deer before sprout of debarked in some areas in Japan (Yokoyama et al. 2001; grasses in early spring and after defoliation of grasses in Ando et al. 2003), indicating that palatability of debark- late autumn. Some sika deer migrate to high-elevation ing may be affected by surrounding vegetation (Moore area before sprout of grasses and after defoliation to es- et al. 1999; McLaren et al. 2000). Therefore, P. jezoensis cape the competition with other deer in low-elevation saplings at the sites examined in the present study may areas (Shin-ichiro Hamasaki, personal communication). be debarked in the future if the deer density increases. Minimum distance to subalpine grassland, however, In addition to size and species as factors for debarking, had only a marginal effect on the occurrence of debark- elevation zones and grassland area also were associated ing of saplings and trees (Tables 1 and 3). As the result with the damage to subalpine forests by deer. Low of intensive debarking on saplings (Fig. 3), low sapling sapling density and high risk of debarking of trees and density near subalpine grassland occurred (Table 2) and saplings were found in low elevation plots (Table 2). then it might be obscure the relationship between the Other studies in Japan also found the intensities of distance to grassland and the occurrence of debarking debarking on trees and saplings in subalpine coniferous of saplings. In the future, the intensity of debarking of forests to be greater at lower elevations (Takeuchi et al. tree would increase if deer density remains at this level 2011). In our study area, sika deer migrate to high- or increases. elevation areas in summer and to low-elevation areas from late autumn to late spring (Izumiyama and Mochizuki 2008) although there are some deer in low-elevation area Conclusions even in summer. Then, opportunities of debarking for Our analysis revealed that large saplings within the range saplings and trees were larger in low-elevation zone than of 30 cm in height and 3 cm in DBH in subalpine forests in high-elevation because deer stay longer time in low were most susceptible for debarking by sika deer. In elevation-area than in high elevation-area. In addition to addition to the size, saplings in low elevation zone or near elevation, we found that the large subalpine grassland subalpine grasslands were more susceptible to debarking caused low sapling density (Table 2) and high risk of the by sika deer. This information may be useful to improve occurrence of debarking of saplings and trees (Tables 1 monitoring deer impact, or determining where to con- and 3). Deer would likely be attracted to subalpine grass- struct deer fences in the subalpine region. At the same land because it has numerous herbaceous plants that serve time, deer population control in the subalpine region should be conducted in forests of low elevation zone or Table 4 Relative importance of variables in models near subalpine grasslands. Debarking of Sapling density Debarking of saplings trees Competing interests Elevation zone 1.000 0.506 0.785 This study was funded by the Comprehensive Research Organization for BA of conspecific species Not included 1.000 Not included Science and Technology of Yamanashi Prefectural Government and Mitsui & CO., LTD. The authors declare that they have no competing interests about DBH of trees Not included Not included 1.000 non-financial aspects. Dist. grassland 0.274 1.000 0.674 Height of saplings 1.000 Not included Not included Authors’ contributions Species 0.137 1.000 0.463 HI and TN substantially contributed about setting the experimental design and collecting data. HI and TN deeply discussed and approved the content Tree density Not included 0.457 Not included of manuscript. Both authors read and approved the final manuscript. Iijima and Nagaike Forest Ecosystems (2015) 2:33 Page 7 of 7 Acknowledgements Moore NP, Hart JD, Langton SD (1999) Factors influencing browsing by fallow We thank Chiaki Ootsu for field assistance, as well as the staff of the deer Dama dama in young broad-leaved plantations. Biol Cons 87:255–260 “Komorebi-sansou” mountain hut. Mysterud A, Loe LE, Zimmermann B, Bischof R, Veiberg V, Meisingset E (2011) Partial migration in expanding red deer populations at northern latitudes: Received: 23 July 2015 Accepted: 20 December 2015 A role for density dependence? Oikos 120:1817–1825 Nagaike T, Hayashi A (2003) Bark-stripping by sika deer (Cervus nippon)in Larix kaempferi plantations in central Japan. For Ecol Manage 175:563–572 Nagaike T (2012) Effects of browsing by sika deer (Cervus nippon) on subalpine References vegetation at Mt. Kita, central Japan. Ecol Res 27:467–473 Akaike H (1973) Information theory and an extension of the maximum likelihood Nagaike T, Ohkubo E, Hirose K (2014) Vegetation recovery in response to the principle, in 2nd International Symposium on Information Theory. Tsahkadsor, exclusion of grazing by sika deer (Cervus nippon) in seminatural grassland on Armenian SSR, pp 267–281 Mt. Kushigata, Japan. ISRN Biodiversity: Article ID 493495 Akashi N, Nakashizuka T (1999) Effects of bark-stripping by sika deer (Cervus nippon) Ohdachi SD, Ishibashi Y, Iwasa MA, Saitoh T (2009) The Wild Mammals of Japan. on population dynamics of a mixed forest in Japan. For Ecol Manage 113:75–82 Shoukadoh Book Sellers, Kyoto Akashi N, Unno A, Terazawa K (2011) Effects of deer abundance on broad-leaf QGIS Development Team (2015) Quantum GIS Geographic Information System. tree seedling establishment in the understory of Abies sachalinensis Open Source Geospatial Foundation Project. http://qgis.osgeo.org plantations. J For Res 16:500–508 R Core Team (2015) R: A Language and Environment for Statistical Computing. R Ando M, Yokota H, Shibata E (2003) Bark stripping preference of sika deer, Cervus Foundation for Statistical Computing, Vienna, Austria, http://www.R-project.org/ nippon, in terms of bark chemical contents. For Ecol Manage 177:323–331 Rooney TP (2001) Deer impacts on forest ecosystems: a North American Ando M, Yokota H, Shibata E (2004) Why do sika deer, Cervus nippon, debark perspective. Forestry 74:201–208 trees in summer on Mt. Ohdaigahara, central Japan? Mamm Study 29:73–83 Schütz M, Risch AC, Leuzinger E, Krüsi BO, Achermann G (2003) Impact of herbivory Apollonio M, Andersen R, Putman R (2010) European ungulates and their by red deer (Cervus elaphus L.) on patterns and processes in subalpine management in the 21st century. Cambridge University Press, NY. grasslands in the Swiss National Park. For Ecol Manage 181:177–188 Beguin J, Pothier D, Prévost M (2009) Can the impact of deer browsing on tree Stewart GH, Burrows LE (1989) The impact of white-tailed deer Odocoileus regeneration be mitigated by shelterwood cutting and strip clearcutting? For virginianus on regeneration in the coastal forests of Stewart Island, New Ecol Manage 257:38–45 Zealand. Biol Cons 49:275–293 Borkowski J, Ukalski K (2012) Bark stripping by red deer in a post-disturbance Suzuki M, Miyashita T, Kabaya H, Ochiai K, Asada M, Kikvidze Z (2013) Deer area: the importance of security cover. For Ecol Manage 263:17–23 herbivory as an important driver of divergence of ground vegetation Brostrom G, Holmberg H (2011) Generalized linear models with clustered data: communities in temperate forests. Oikos 122:104–110 fixed and random effects models. Comput Stat Data Anal 55:3123–3134 Takatsuki S (1986) Food habits of sika deer on Mt. Goyo, northern Honshu. Ecol Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a Res 1:119–128 practical information-theoretic approach, 2nd edn. Springer, New York Takatsuki S, Gorai T (1994) Effects of sika deer on the regeneration of a Fagus Fuller RJ, Gill RMA (2001) Ecological impacts of increasing numbers of deer in crenata forest on Kinkazan Island, northern Japan. Ecol Res 9:115–120 British woodland. Forestry 74:193–199 Takatsuki S (2009) Effects of sika deer on vegetation in Japan: a review. Biol Cons Gill RMA, Beardall V (2001) The impact of deer on woodlands: the effects of 142:1922–1929 browsing and seed dispersal on vegetation structure and composition. Takeuchi T, Kobayashi T, Nashimoto M (2011) Altitudinal differences in bark Forestry 74:209–218 stripping by sika deer in the subalpine coniferous forest of Mt. Fuji. For Ecol Heuze P, Schnitzlerm A, Klein F (2005) Consequences of increased deer browsing Manage 261:2089–2095 in winter on silver fir and spruce regeneration in the southern Vosges Winnie JAJ (2012) Predation risk, elk, and aspen: tests of a behaviorally mediated Mountains: implication for forest management. Ann For Sci 62:175–181 trophic cascade in the Greater Yellowstone Ecosystems. Ecology 93:2600–2614 Iijima H, Nagaike T, Honda T (2013) Estimation of deer population dynamics by Yokoyama S, Maeji I, Ueda T, Ando M, Shibata E (2001) Impact of bark stripping Bayesian state-space model with multiple abundance indices. J Wildl Manage by sika deer, Cervus nippon, on subalpine coniferous forests in central Japan. 77:1038–1047 For Ecol Manage 140:93–99 Iijima H, Nagaike T (2015) Appropriate vegetation indices for measuring the impacts of deer on forest ecosystems. Ecol Ind 48:458–463 Izumiyama S, Mochizuki T (2008) Seasonal range use of sika deer, which inhabits the subalpine zone in the southern Japan Alps. Bull Shinshu Univ Alps Field Center 6:25–32 (in Japanese with English summary) Izumiyama S, Mochizuki T, Takii A (2009) GPS tracking of Sika deer which inhabits the sub-alpine zone in the Southern Japan Alps. Bull Shinshu Univ Alps Field Center 7:63–71 (in Japanese with English summary) Jiang Z, Ueda H, Kitahara M, Imaki H (2005) Bark stripping by sika deer on Veitch fir related to stand age, bark nutrition, and season in northern Mount Fuji district, central Japan. J For Res 10:359–365 Kamei T, Takeda K, Koh K, Izumiyama S, Watanabe O, Ohshima K (2010) Seasonal pasture utilization by wild sika deer (Cervus nippon) in a sown grassland. Grass Sci 56:65–70 Kay S (1993) Factors affecting severity of deer browsing damage within coppiced woodlands in the south of England. Biol Cons 63:217–222 Submit your manuscript to a Kiffner C, Rößiger E, Trisl O, Schulz R, Rühe F (2008) Probability of recent bark journal and benefi t from: stripping damage by red deer (Cervus elaphus) on Norway spruce (Picea abies) in a low mountain range in Germany: a preliminary analysis. Silva 7 Convenient online submission Fennica 42:125–134 7 Rigorous peer review Koda R, Fujita N (2011) Is deer herbivory directly proportional to deer population density? Comparison of deer feeding frequencies among six forests with 7 Immediate publication on acceptance different deer density. For Ecol Manage 262:432–439 7 Open access: articles freely available online Ligot G, Gheysen T, Lehaire F, Hébert J, Licoppe A, Lejeune P, Brostaux Y (2013) 7 High visibility within the fi eld Modeling recent bark stripping by red deer (Cervus elaphus) in South 7 Retaining the copyright to your article Belgium coniferous stands. Ann For Sci 70:309–318 McLaren BE, Mahoney SP, Porter TS, Oosenbrug SM (2000) Spatial and temporal patterns of use by moose of pre-commercially thinned, naturally-regeneration Submit your next manuscript at 7 springeropen.com stands of balsam fir in central Newfoundland. For Ecol Manage 133:179–196

Journal

"Forest Ecosystems"Springer Journals

Published: Dec 1, 2015

Keywords: Ecology; Ecosystems; Forestry

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