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1 (Bulletin of FFPRI)Vol.7No.2(No.47) 81-89June28 論文 (Original article) 御嶽山における密なチマキザサ林床をもつ亜高山帯針葉樹林の構造と動態 1)* 2) 1) 3) 3) 3) 4) Structure and dynamics of a subalpine coniferous forest Hisashi SUGITA 1)*, Kojiro IWAMOTO 2), Takeshi MORISAWA 1), Tomoyuki SAITO 3), Daisuke KABEYA 3),, Toru OKAMOTO 3) and Hisao SAKAI 4) Abstract Stand structure and its change during recent 8-years were investigated in a subalpine coniferous forest composed of, Picea jezoensis var. hondoensis, Abies veitchii and Abies mariesii with dense undergrowth of Sasa palmata in a 5 5 m plot on the southeast side of Mt. Ontake, central Japan. Canopy layer contained abundant openings (32 % in relative area). Abies veitchii and A. mariesii showed the L-shaped DBH distribution, while T. diversifolia and P. jezoensis var. hondoensis showed bimodal distribution: two peaks of canopy trees and suppressed trees. Most of T. diversifolia and P. jezoensis var. hondoensis trees had stilt-rooted growth form, or were established on uprooted mounds and buttresses, and few of these species were established on the ground. Two Abies species were often found on the ground more than Tsuga and Picea: however, the relative frequency was only 15% for A. veitchii and 35% for A. mariesii. The majority of Abies trees was established on uprooted mounds, buttresses, and rocks, or they were stilt-rooted. The preference for substrata other than the ground for the establishment microsites of Abies species was more conspicuous as compared to those reported for other subalpine stands with undergrowth without Sasa. The difference suggests that the inhibition of conifer establishment on the ground by dense undergrowth of S. palmata affects the stand structure in the study site. The mortality and recruitment rate for the stand during an 8-year period ( ) was.6 %/yr and 1.44 %/yr, respectively, indicating that recruitment was higher than mortality. The loss and gain in total basal area was.91 %/yr and.96 %/yr, respectively, indicating almost balanced. As for values according to species, only P. jezoensis var. hondoensis showed higher mortality rate and loss in basal area than recruitment rate and gain in basal area, while the other species demonstrated an opposite relationship between these values. Key words : Abies mariesii, Abies veitchii, dwarf bamboo, microsite, Picea jezoensis var. hondoensis, 5m 5m 8 32% L L 15% 35% ( 5cm ).6%/ 1.44%/.91%/.96%/ Received3 December Accepted 13 February 28 1) Tohoku Research Center, Forestry and Forest Products Research Institute(FFPRI) 2) Tama Forest Science Garden, Forestry and Forest Products Research Institute(FFPRI) 3) Kiso Experimental Station, Forestry and Forest Products Research Institute(FFPRI) 4) Hokkaido Research Center, Forestry and Forest Products Research Institute(FFPRI) * Tohoku Research Center, Forestry and Forest Products Research Institute(FFPRI), Morioka, Iwate 2-123, Japan; sugitah@ffpri.affrc.go.jp

2 82 SUGITA H. et al. ( ) (Oosting and Reed, 1952; Day, 1972; Fox, 1977; Peet, 1981; Veblen, 1986; Aplet et al., 1988; Kneeshaw and Bergeron, 1996; Takahashi, 1997; Antos and Parish, 22) (Kanzaki, 1984; Miyadokoro et al., 23) (Sugita and Tani, 21; Narukawa and Yamamoto, 21, 22; Mori et al, 24) (Kanzaki, 1984; Yamamoto, 1993, 1995) ha (Miyadokoro et al., 23; Mori and Takeda, 24a, b) (Miyadokoro et al., 24; 26 Mori et al., 27) (Miyadokoro et al, 23, 24; Mori et al, 27) ( 1976 Franklin et al, 1979; Sugita and Nagaike, 25) ( ) ( ) ( ) (Narukawa and Yamamoto, 22) (363m) (2256m) (219m) (1972) 15 2cm ( 2) 16 24m 22m ( 1951) ( ) ( 1958 Franklin et al, 1979) 213m S5 W 12 ( 24)

3 Structure and dynamics of a subalpine coniferous forest 83 (24 27 ) 9 18cm 128cm ( ) 1 1.5m 1% ( 24) (185m) 2ha (Miyadokoro et al, 23, 24) (2m) 1ha ( 1981; Mori and Takeda, 24b; Mori et al, 27) 1. 野外調査 ha 5m 5m ( 1.3m) 1m ( ) 動態パラメータの算出 5cm m r / T (Condit et al., 1999) m [ln{n /(N N d )}] / t1 r[ln{( N N d + N r ) / ( N N d )}] / t1 T2 / m r N N d N r t / (Hoshino et al., 22) (Loss in basal area) (Gain in basal area) {ln(ba /BA s )} /t1 [ln{(ba s1 + BA r ) /BA s }] /t1 BA BA s BA s1 BA r 1. 林分構造 1998 (DBH5cm ) Table /ha 46.8m 2 /ha 47% % 1% 2% 1998 Fig % ( 4.1%) 1 16m (14.3%) 22.7m (Table Table 1. Density, basal area and maxinum height of each component tree species * (ha -1 ) (m 2 /ha) (m) * (ha -1 ) (m 2 /ha) Density Basal area Maximum height Density Basal area 2 (29.1) (54.4) (27.7) (56.9) Picea jezoensis var. hondoensis 68 (9.9) (31.2) (8.2) (25.1) Abies veitchii 32 (46.5) (11.3) (48.4) (14.3) Abies mariesii 72 (1.5).832 (1.8) (1.9).965 (2.1) Betula ermanii 4 (.6).485 (1.) (.5).537 (1.1) Sorbus commixta 16 (2.3).86 (.2) 16 (2.2).119 (.3) Enkianthus campanulatus 8 (1.2).7 (.1) 16 (2.2).14 (.2) Euonymus macropterus (.).3 (.) (.).4 (.) Total (): (%) Relative value * 5cm Density of trees more than 5cm in DBH Bulletin of FFPRIVol.7No.228

4 84 SUGITA H. et al. 1) (12.%) 1998 Fig.2 L 1 4cm 15cm L 25 65cm 35 75cm 1cm 1 35cm 2. 定着マイクロサイト Fig.3 1 ( ) 15% 35% (33%) 85% ( 1998) (p<.1 2 ) ~ 26 年における死亡加入成長 Fig cm 21% Density (ha -1 ) Suppressed trees Suppressed trees in gaps Canopy trees in gaps Canopy trees Abies veitchii Abies mariesii 1 2 Fig. 1. Crown projection map Picea jezoensis var. hondoensis Betula ermanii DBH (cm) Fig. 2. DBH distribution

5 Structure and dynamics of a subalpine coniferous forest Abies mariesii Abies mariesii Density (ha -1 ) Abies veitchii Stilt-rooted trees Fallen logs Buttresses Uprooted mound Rocks Ground Density (ha -1 ) Abies veitchii Stem broken Standing dead Living Picea jezoensis var. hondoensis 4 Picea jezoensis var. hondoensis Betula ermanii Betula ermanii DBH (cm) DBH (cm) Fig. 3. Microsites for establishment Fig. 4. Occurrence of dead individuals during the 8-year period 5 1cm 1% 8 1cm (Fig.1) 26 Table (DBH5cm ) Table 2 Table 2. 5cm.6%/ 1.44%/.91%/.96%/ 2.43% 2.14% Demographic parameters during 8 years ( ) * * Mortality rate Recruitment rate Turnover (%/yr (%/yr (%/yr (%/yr yr Loss in basal area Gain in basal area Picea jezoensis var. hondoensis Abies veitchii Abies mariesii Total * DBH > 5 cm Bulletin of FFPRIVol.7No.228

6 86 SUGITA H. et al. Diameter growth during (cm/yr) Canopy trees (including in gaps) Fig.5 (Spearman rs=-.481 p=.5) ( p<.1p=.124 p=.97 Mann-Whitney U ) cm/ cm/ cm/ 2 ( p<.1 Mann-Whitney U ) 1. 林床型による林分構造のちがい L (Fig.2) ( 1981; Miyadokoro et al, 23; Mori and Takeda, 24b) Suppressed trees (including in gaps) Picea jezoensis var. hondoensis DBH in 1998 (cm) Fig. 5. Abies mariesii Abies veitchii Relationship between DBH and diameter growth (Fig.1) 31.8% 7.3% 8.5%(Yamamoto, 1993) 18.5%(Miyadokoro et al., 23) 5.6% 6.4% 8.9%( Yamamoto, 1995) 2. 樹種の定着場所に及ぼすチマキザサ林床の影響 (Christy and Mack, 1984; Harmon and Franklin, 1989) (Fig.3) (Sugita and Tani, 21) (Mori et al, 24) (Narukawa and Yamamoto, 22) (Sugita and Tani, 21) (Sugita and Nagaike, 25) (Knapp & Smith, 1982; Simard et al., 23) (Mori et al, 24) (Narukawa and Yamamoto, 22) (Sugita and Tani, 21) (Fig.3) 15% 35% (Narukawa & Yamamoto, 22) ( 1984)

7 Structure and dynamics of a subalpine coniferous forest 87 (Nakashizuka and Numata, 1982; Taylor and Qin, 1988) 31.8% (Fig.1) Takahashi(1997) (Nakashizuka, 1989) ( ) 3. 森林動態 (Fig.4).6%/ 1.44%/ (8 Table 2) 2.74% 1.2%(9 Miyadokoro et al, 24) 3.3% 1.42%(25 Mori et al, 27) (Miyadokoro et al, 23, 24; Mori et al, 27) (Table 2) (.8%) 3.2% 3.3% (Miyadokoro et al, 24) 3 (Fig.5) (Peet, 1981; Aplet et al., 1988) Mori et al.(27) 25 (Fig.1) ( ) (B-11 S-4-2(4)) Bulletin of FFPRIVol.7No.228

8 88 SUGITA H. et al. Antos, J.A. and Parish, R. (22) Structure and dynamics of a nearly steady-state subalpine forest in southcentral British Columbia, Canada, Oecologia, 13, Aplet, G.H., Laven, R.D., and Smith, F.W. (1988) Patterns of community dynamics in Colorado Engelmann spruce-subalpine fir forests, Ecology, 69, Christy, E.J. and Mack, R.N. (1984) Variation in demography of juvenile Tsuga heterophylla across the substratum mosaic, Journal of Ecology, 72, Condit, R., Aston, P.S., Manokaran, N., LaFrankie, J.V., Hubbell, S.P. and Foster, R.B. (1999) Dynamics of the communities at Pasoh and Barro Colorado: comparing two 5-ha plots, Phil. Trans. R. Soc. London, 354, Day, R.J. (1972) Stand structure, succession, and the use of southern Alberta's Rocky Mountains forest, Ecology, 53, Fox, J.F. (1977) Alternation and coexistence of tree species, American Naturalist, 111, Franklin, J.F., Maeda, T., Ohsumi, Y., Matsui, M., Yagi, H. and Hawk, G.M. (1979) Subalpine coniferous forests of Central Honshu, Japan, Ecological Monographs, 49, (1984) 26(1) Harmon, M.E. & Franklin, J.F. (1989) Tree seedlings on logs in Picea-Tsuga forest of Oregon and Washington, Ecology, 7, Hoshino, D., Nishimura, N. and Yamamoto, S. (22) Dynamics of major conifer and deciduous broadleaved tree species in an old-growth Chamaecyparis obtusa forest, central Japan, Forest Ecology and Management, 159, (1937) (1998) Kanzaki, M. (1984) Regeneration in subalpine coniferous forests I. Mosaic structure and regeneration process in a forest, Botanical Magazine, Tokyo, 97, Knapp, A.K. and Smith, W.K. (1982) Factors influencing understory seedling establishment of Engelmann Spruce (Picea engelmannii) and Subalpine Fir (Abies lasiocarpa) in southeast Wyoming, Canadian Journal of Botany, 6, Kneeshaw, D.D. and Bergeron, Y. (1996) Ecological factors affecting the abundance of advance regeneration in Quebec's southwestern boreal forest, Canadian Journal of Forest Research, 26, Miyadokoro,T., Nishimura,N., Hoshino,D. and Yamamoto,S. (24) Dynamics of forest canopy and major tree populations over nine years in a subalpine old-growth coniferous forest, central Japan, Ecoscience, 11, Miyadokoro, T., Nishimura, N. and Yamamoto, S. (23) Population structure and spatial patterns of major trees in a subalpine old-growth coniferous forest, central Japan, Forest Ecology and Management, 182, Mori, A., Mizumachi, E., and Komiyama, A. (27) Role of disturbance and demographic non-equilibrium in species coexistence, inferred from 25-year dynamics of a late-successional old-growth subalpine forest, Forest Ecology and Management, 241, Mori, A., Mizumachi, E., Osono, T. and Doi, Y. (24) Substrate-associated seedling recruitment and establishment of major conifer species in an oldgrowth subalpine forest in central Japan, Forest Ecology and Management, 196, Mori, A. and Takeda, H. (24a) Effects of mixedwood canopies on conifer advance regeneration in a subalpine old-growth forest in central Japan, Ecoscience, 11, Mori, A. and Takeda, H. (24b) Effects of undisturbed canopy structure on population structure and species coexistence in an old-growth subalpine forest in central Japan, Forest Ecology and Management, 2,

9 Structure and dynamics of a subalpine coniferous forest 89 Nakashizuka,T. (1989) Role of uprooting in composition and dynamics of an old-growth forest in Japan, Ecology, 7, Nakashizuka,T. and Numata,M. (1982) Regeneration process of climax beech forests I. Structure of a beech forest with the undergrowth of Sasa, Japanese Journal of Ecology, 32, Narukawa,Y. and Yamamoto,S. (21) Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan, Ecological Research, 16, Narukawa, Y. and Yamamoto, S. (22) Effects of dwarf bamboo (Sasa sp.) and forest floor microsites on conifer seedling recruitment in a subalpine forest, Japan, Forest Ecology and Management, 163, (1972) 24(1) Oosting, H.J. and Reed, J.F. (1952) Virgin spruce-fir forest of the Medicine Bow Mountains, Wyoming, Ecological Monographs, 22, Peet, R.K. (1981) Forest vegetation of the Colorado front range, Vegetatio, 45, Simard, M.J., Bergeron, Y andsirois, L. (23) Substrate and litterfall effects on conifer seedling survivorship in southern boreal stand of Canada, Canadian Journal of Forest Research, 33, (22) ( ) Sugita,H. and Nagaike,T. (25) Microsites for seedling establishment of subalpine conifers in a forest with moss-type undergrowth on Mt. Fuji, central Honshu, Japan, Ecological Research, 2, Sugita,H. and Tani,M. (21) Difference in microhabitatrelated regeneration patterns between two subalpine conifers, and Abies mariesii, on Mt. Hayachine, northern Honshu, Japan, Ecological Research, 16, Takahashi, K. (1997) Regeneration and coexistence of two subalpine conifer species in relation to dwarf bamboo in the understorey, Journal of Vegetation Science, 8, Taylor A. H. and Qin Z. (1988) Regeneration pattern in oldgrowth Abies Betula forests in the Wolong Natural Reserve, Sichuan, China, Journal of Ecology, 76, Veblen, T.T. (1986) Treefalls and the coexistence of conifers in subalpine forests of the central Rockies, Ecology, 67, Yamamoto, S. (1993) Gap characteristics and gap regeneration in a subalpine coniferous forest on Mt Ontake, central Japan, Ecological Research, 8, Yamamoto, S. (1995) Gap characteristics and gap regeneration in subalpine old-growth coniferous forests, central Japan, Ecological Research, 1, Bulletin of FFPRIVol.7No.228

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橡 PDF ( ) * Regeneration of many tree species by soil scarification under canopies -Relation between canopy openness and species richness- Hajime SATO ( )20 20 40 PAR30 0.5 1 (2) 1970 1993 1997 1996 1982 Hokkaido