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(2) important temples. Understanding the bioclimatic and potential natural vegetation in this transition region of eastern China is thus based largely on the few forest remnants and on bioclimatic comparisons with other, potentially similar regions. One especially large and well protected area of Chinese evergreen broad-leaved forest remains around the Buddhist monastery a t Tiantong, an early center of Zen Buddhism (about 150 km south of Shanghai). This is an especially interesting location because it lies near the southern end of the transition region (and of the intensively cultivated lowlands) and is near the coast, where moderated winter temperatures permit some more southern plants to survive. The location of Tiantong (about 30°N) would correspond to southernmost Kyushu in Japan or to northeastern Florida in eastern North America. Both of these locations are areas of potential evergreen broad-leaved forest. Southernmost Kyushu represents the beginning of the Japanese subtropical region (cf. Fujiwara 1981-86, Miyawaki 1979, Kira 1949) , whereas northern Florida still is in the warm-temperate region of eastern North America (cf. Laessle and Monk 1961, Greller 1980, Christensen 1988, Box and Fujiwara 1988). From forest remnants such as Tiantong, Chinese botanists try to reconstruct pat terns of original vegetation and interpret the current potential natural vegetation and its dynamics in this especially long-settled transition region of eastern China (e.g. Han 1981, Zhou 1981, Liu and Huang 1982, Veg. of Anhui Comm. 1983, Zhao 1986, Forests of Shandong Comm. 1987). Study of such situations may also provide insight into vegetation dynamics and potentials in similar regions of other countries, including southern Japan and the southeastern USA. The purpose of this paper is to describe the environmental setting and forest vegetation a t Tiantong, attempt to interpret the local vegetation dynamics, and offer some comparison with potentially evergreen forests elsewhere. Study Site a Tianfong forest and monastery are located in low coastal hills a t about 29.7" N latitude, about 30 km southeast of Ningbo city and 10 km from the seacoast, on the south side of the wide Fuchun River estuary (Hangzhou Bay), in Zhejiang Province of eastern China (see Figure 1). The monastery a t Tiantong was founded about 1600 years ago, and its 99room temple complex still dominates the valley. The climate a t Tiantong is suggested by the climate diagram for Ningbo city, shown in Figure 2. The climate is humid and typical for an east-coast position a t 30" N latitude, with a hot, humid monsoonal summer and a drier winter with a short cold period. Winters are not completely without precipitation, due to the coastal location. Mean monthly temperatures are 7.0 "C for December, 4.1"C for January, and 5.3 "C for February, with frost possible from November to April. These mean temperatures are lower than a t the same latitude in Japan and eastern North America, but absolute minima (-8.8"C a t Ningbo) are lower in eastern North America (cf. Box 1988). Based on surrounding meteorological stations and normal extrapolation methods, a more detailed estimate of the climate a t Tiantong itself is attempted in Table 1. Absolute temperature extremes a t Tiantong (valley elevation about 100m) were estimated by local personnel as 36.7"C and -6.5"C, with an average annual precipitation of 1450 mm. Mean relative humidity a t Ningbo is above 80 % for every month except January (77%). Inside the closed forest, of course, wind and temperature extremes are ameliorated, a s are the lowest humidity values. The geologic substrate around Tiantong is given as acidic intrusive rock (Map Press.
(3) Figure 1.. Location of Tiantong forest and monastery, with surrounding cities and corresponding location in J a p a n and southeastern USA. '. Ningbo. (5rn.l. Figure 2.. Climate Diagram for Ningbo, near Tiantong.. Climatic d a t a for Tiantong a r e n o t available b u t can be approximated using d a t a from Ningbo, a b o u t 30 km northwest of Tiantong. The climate i s a typical eastcoast humid climate, a t a latitude (30°N) which normally represents the transition from a typical temperate climate with cold winters and deciduous forest t o a warm-temperate o r even subtropical climate with milder winters and evergreen broad-leaved forest. A t Tiantong and Ningbo, light f r o s t is possible from December t o April, and the J a n u a r y mean temperature of 4.1°Cis rather cold. The lowest temperatures ever recorded (18years of measurement), however, a r e only -8.8" for J a n u a r y , - 8.2"for December, and -7.2" for February. (Note t h a t the climate diagram represents the second column of d a t a under Ningbo in Table 1.).
(4) Table 1. Estimated Climatic Conditions a t Tiantong Forest, based on Extrapolations from nearby meteorological stations. Shanghai. Ningbo (25m). Tiantong forest 200m 400m. Temperature ("C) mean annual temperature mean July temperature mean January temperature absolute minimum temperature Precipitation (mm) average annual precipitation driest winter month driest summer month Water Balance (mm) annual potential evapotranspir. annual moisture surplus annual moisture index (MI) MI of driest summer month Climatic data were available for Shanghai and Ningbo cities (Song 1988) and from the Chinese Central Meteorological Bureau (1962, see second column of Ningbo values). Values for Tiantong forest were estimated using a n environmental lapse rate of 6.4"C/km, based on the 18-year record at Ningbo (first column of Ningbo values). Potential evapotranspiration was estimated by the method of Holdridge (1959), which seems to be more accurate in subtropical latitudes than the widely used Thornthwaite method (cf. Stephens and Stewart 1963, Box 1986). Moisture conditions a t Tiantong are somewhat better than in the cities, with significant water surplus and probably little deficit, even for short periods. Temperatures a t Tiantong are lower but not extreme in winter. Tiantong has over 200 fog days per year.. a. Edit. Dept. 1979, map 7), which corresponds to our finding of a granite substrate. Local personnel described the parent material as also involving mesozoic sediments. The lowland soil type ( op. cit , map 14) is given as " red type", i.e. subtropical red-yellow podzolic, which is the zonal soil. On the slopes a t Tiantong the soil is better, with a thick humus layer and several centimeters of leaf litter. The hills around the monastery rise to about 500 meters, or about 400 m above the valley floor, with slopes ranging from 20" to over 40" in places. Outcropping larger rocks and boulders could be found in some places. The flora of the Tiantong forest was described by local botanists a s including 349 woody species and 251 non-woody species, in a total of 112 families. Song ( 1988) describes finding 252 species (in 161 genera and 81 families) in just eight sample plots (usually 20 x 20m) in evergreen broad-leaved forests in Shanghai and Zhejiang Province. A national forest station was established a t Tiantong in 1957. Eventually, the entire temple complex and surrounding forest (watershed) were designated a national park, comprising about 700 hectares and including a hotel. The slopes on all sides are covered by relatively natural evergreen broad-leaved forest dominated mainly by Fagaceae (Figure 3). This type of lucidophyllous (Kira 1969) or "laurel" forest (as it is often called in Japan and Europe, cf. Miyawaki 1984, Hub1 1988) is quite similar to that of other parts of eastern China (Wang 1961, Wu 1980, Hou 1983) as well as much of southern Japan (Fujiwara 1981-86, Miyawaki 1980-89, Numata 1974, Numata et al. 1972) and southernmost Korea.
(5) Figure 3.. The Buddist Temple Complex a t Tiantong and the surrounding, protected Evergreen Broad-Leaved Forest.. (e.g. Uyeki 1941, Kim et al. 1984). Many of the forest trees a t Tiantong show the multistem growth form indicative of past coppicing, mainly for firewood and charcoal production. Sporadic bamboo patches also indicate disturbance within recent decades. Nevertheless, the forest is in a remarkably natural condition, considering the complete absence of forest over so much of the lowland East-Chinese landscape. Local personnel told us we were the first Japanese and American botanists to study the Tiantong forest.. Methodology The vegetation a t Tiantong was sampled by the Braun-Blanquet relev6 method, a well standardized methodology well suited to rapid, extensive sampling as well asgeographic comparisons. This methodology is described, in English, by Westhoff and van der Maarel (1973), by Mueller-Dombois and Ellenberg (1974), and in more complete detail by Fujiwara (1987). Species were identified conclusively by Song and other local botanists, but many could also be recognized by the Japanese authors a s common to southern Japan. Voucher specimens could not be taken. Nomenclature generally follows Wu ( 1980) and the Institute of Botany (1972-85). References for species ranges and foliation habits also included Wang (1961), Hayashi (1985), and Mabberley (1987). Evergreen Broad-Leaved Forest at Tiantong The forest (Figure 3) was sampled a t several elevations and locations on both sides of the valley, a s summarized in Table 2. Of the seven releves taken, all were in evergreen broad-leaved forest except one, in a deciduous area just above the bottom of a ravine. In.
(6) general, the slopes are covered primarily by Castanopsis forest, varying from about 20 to over 30 meters in height. On and near ridgetops, especially a t the higher elevations, the forest becomes shorter in stature and involves Quercus and Schima (Theaceae) in the canopy, in addition to Castanopsis. Species richness varied between 47 and 60 species in the slope-f orest relevbs but dropped to 30-39 species in the shorter, more exposed upper forests, a s summarized in Table 2. The greatest number of species was generally found in the shrub layer. The composition and structure of relatively mature forest a t Tiantong is suggested by the relev6 shown in Table 3. Overstorey and more important understorey species are summarized for all sites in Table 4. All four slope-forest relev&, on both sides of the valley, were dominated by Castanopsisfargesii and secondarily by C. carlesii, both species which Wang (1961) lists in his group of cosmopolitan Castanopsis species, i.e. occurring in both southeastern and southwestern China. C. carlesii was once considered to be a synonym of C. cuspiduta , a major canopy dominant in southern Japanese evergreen broadleaved forests. Other evergreen broad-leaved canopy trees included Persea thunbergii ( a dominant, character species of Japanese evergreen broad-leaved forests) , Myrica rubra Table 2. Relevb Sites, Vegetation, and Species Richness a t Tiantong.. Elev. Aspect. Veg . Slope Height. Vegetation type. Number of Species T I Tz S H Total. Location West Side 1. Mid-slope. 220m. E. 30". 25m. Castanopsis forest, with deciduous. 5. 3. 28 23 53. 2. Mid-slope. 230m. E. 35". 25m. Castanopsis-Pinus forest. 8. 9. 19 20 47 *. 3. upper slope. 390m. E. 42". 12m. Quercus-Schima forest. 6. Northeast Side 4. Mid-slope. 245m. S. 40". 32m. Castanopsis forest, with deciduous. 5. Rocky mid-slope 6. Below ridgetop. 290m 330m. W W. 32" 32". 22m 15m. Castanopsis forest 4 4 Quercus - Schima 9 Castanopsis forest. 38 14 53 24 11 39. 7. Ravine (low slope). 270m. SE. 22". 27m. Deciduous forest 2 4 with evergr. understorey. 40 30 72. 8. 20. 9. 30. 11 27 22 60. Relevbs are arranged in order of increasing elevation except for the deciduous ravine forest. Vegetation height and species richness both tend to decrease with increasing elevation (position on the slope), perhaps due to upwardly decreasing soil depth and increasing exposure to winter winds. The deciduous ravine forest was dominated by Choerospondias axillaris (Anacardiaceae) and Acer oliverianum but with mainly evergreen understorey elements and with evergreen forest on the slopes immediately above. All other relevbs had evergreen dominants (mainly Fagaceae, plus Schima) but also had a t least one deciduous tree in the T I layer (Liquidambar, Carpinus, Sassafras, Platycarya, etc.)..
(7) (eastern China plus southern Japan) , Distylium myricoides ( Hamamelidaceae, with a more restricted range), and a large (25m) Ilex individual which could not be identified conclusively (perhaps I. wilsonii) .. Table 3. Composition o f Relatively Mature Evergreen Broad-Leaved Forest at Tiantong, eastern China. TI Tz S H. 25m 13m 4m 0.8m. 80% 40% 30% 40%. Slope : 30" to E Elevation : about 220 m Relev6 size : 20 x 30 m Date : 8 November 1985 (KF, S Y , A M , EB). TI. 4 3 Castanopsis fargesii 1 1 Castanopsis carlesii 1 1 Schima superba. Tz. 2 3 Castanopsis fargesii 2 2 Lithocarpus henryi. Camellia fraterna Schima superba Symplocos caudata Ilex buergeri Symplocos laurina Cinnamomum nikkei Eurya nitida Myrica rubra Symplocos anomala Eurya loquiana Randia cochinchinensis Lithocarpus glabra Clerodendron trichotomum vines : 1 2 Stauntonia sp. + Vitaceae sp. Woodwardia japonica Carex sp. 1 (large) Trachelospermum jasminoides (vine) Arachniodes sp. Asplenium wrightii Ficus sarmentosa var. henryi (sdl.) Ardisia crenata (seedling) Lithocarpus henryi (seedling) Syzygium buxifolium (seedling) Paederia scandens (vine) Pyrola sp. Carex sp. 3 (medium). 2 1 Liquidambar formosana (decid.) 1 1 Diospyros lotus (decid.). +. Styrax fabri (=S. confusa) (decid.). Castanopsis fargesii Neolitsea aurata Meliosma rigida Maesa japonica Daphniphyllum oldhamii Litsea elongata Quercus glauca Cleyera japonica Symplocos stellaria Halesia cochinchinensis Elaeocarpus japonica Ilex wilsonii. +. Smilax sp.. Lophaterum gracile Schima superba (seedlings) Colysis elliptica Dryopteris erythrosora (bissetiana?) Quercus myrsinaefolia (seedl.) Carex sp. 2 (small) (C. conica?) Acer sp. (seedling) Persea thunbergii (seedling) Styrax fabri (seedling) Vitaceae sp. (vine) Diplazium subsinuatum.
(8) Table 4. Summary of Evergreen Forest Relevks a t Tiantong. W e s t s i d e Tall forests 1 2. N - E a s t Short forests 3 6. s i d e Tall forests. 4. 5. T I trees Castanopsis fargesii Castanopsis carlesii Schima superba Quercus stewardii Quercus gracilis Distylium myricoides Persea thunbergii Cleyera japanica Lithocarpus henryi Myrica rubra Ilex sp. Liquidambar formosana (decid.) Diospyros lotus (decid.) Carpinus fargesii (decid.) Platycarya strobilacea (decid.) Sassafras tsumu (decid.) Albizzia kalkora (decid.) Fraxinus sp. (decid.) Pinus massoniana (30m) Cunninghamia lanceolata. Tz trees Eurya loquaiana Meliosma rigida Cinnamomum subavenium Quercus myrsinaef o lia Quercus glauca Styrax fabri (decid.). (+>. (1.2). .. (+> (+>. +. 1.1 1.2. . .. 1.1 (+>. . . (1.2) . .. .. (1.2). .. 2.2 (1.1) (+>. .. 1.1 1.1 2.2. .. 1-2. (+I. . . . . (+> ..
(9) Shrub layer (common spp. only) Litsea elongata Neolitsea aurata v. chekiangensis Eurya rubiginosa v. attenuata Symplocos stellaris Ardisia crenata Camellia fraterna. e. (+>. (+>. 2-3. 2-2. . . . . .. 1.2. +-2 1.2. (+>. +. (+>. . .. 2-3. 2-2. . .. 3.3. .. 3.3 1-2. 2 SPP.. 1-2. 2-2. + 1-2. + +. 1 1 2.2. + +. 2.3. 1.1. +. 1.1 1-2 (+>. .. 2.3 2 3. .. 2.2 1.1. .. +*2 +* 2 2.2 1.2. +. 3-3. Possible differential species Rhododendron ovatum Syzygium buxifolium Symplocos botryantha Daphniphyllum oldhamii Symplocos caudata Rubus buergeri. Vines (shrub layer, common t a x a only) Trachelospermum jasminoides Stauntonia leucantha Smilax sp.. Herb layer (common spp. only) Woodwardia japonica Dryopteris erythrosora Lophaterum gracile Carex spp.. (+>. +. 2-3. +. 1.2 3 SPP.. + +. 1.2. . .. +. 2.2 1.2. +. 1-2. +. . .. +*2 1-2 1.2. Species a r e listed in the highest layer in which they occur in any relevb-at Tiantong. (Values given in refer to occurrences in lower layers.) Tall-forest relevbs were from mid-slope, while the short-forest relev6s came f r o m ridgetops a t higher elevations (see Table 2 for site descriptions). Castanopsis fargesii i s the dominant o r a co-dominant species in a l l relev6s except the higher-elevation samples (relev6s 3 and 6), in which Schima superba and Fagaceae other t h a n C. fargesii a r e more important. All sites have some deciduous trees in the canopy, though site 5 had only one and seems t o be the m o s t m a t u r e site..
(10) In addition to the evergreen trees, each releve had a t least one broad-leaved deciduous tree in the canopy (e.g. Figure 4), the most prominent being Liquidambar f ormosana, a common successional tree in Taiwan, southeastern mainland China, and north a s f a r a s Hunan. Some relev& also contained conifers in the canopy, Pinus massoniana ( a Chinese heliophilic pine with a wide range) or Cunninghamia lanceolata (Taxodiaceae, a Chinese endemic widely used for afforestations and a s an ornamental abroad). The occurrence of these deciduous and coniferous trees in the canopy suggests that the Tiantong forest is not completely mature but rather still recovering from disturbance earlier in this century.. Figure 4.. Castanopsis Forest a t Tiantong, with Deciduous Trees also in the Canopy..
(11) i. 73. The woody understoreys include a variety of smaller trees and shrubs characteristic of evergreen broad-leaved forests throughout various parts of East Asia, in particular Eurya and Camellia spp. (Theaceae) , Lit sea elongata and Neolit sea aurata (both Lauraceae common throughout southern China), Ardisia crenata (Myrsinaceae, from NE India t o Japan), and Symplocos spp. (Symplocaceae, important throughout East and SE Asia). In addition, two evergreen Quercus spp. (Qu. myrsinaefolia and Qu. glauca), both canopy dominants in evergreen forests in Japan, occurred in five of the six evergreen forest relevks, though only in understoreys. The most common vines were Stauntonia spp. (Lardizabalaceae) a s climbers and Trachelospermum jasminoides mainly in ground patches. The herb layer of all relevks was dominated or co-dominated by ferns, mainly Woodwardia japonica but also species of Dryopteris, Dicranopteris, and Arachniodes. Various forest Carex spp. were also common. The composition of the upper-slope forest, just under the ridgetops, is suggested by the relev& shown in Table 5. In the shorter ridge forests (Figure 5), dominance shifts to Schima superba, also a relatively cosmopolitan species, plus Quercus stewardii or Qu. gracilis (both evergreen). Both Castanopsis species occur also in the ridge forests but sometimes only in understoreys. Lithocarpus henryi (Fagaceae) and Cleyera japonica (Theaceae) occurred in the canopy of the ridge forests but only in the understoreys of the slope-f orest relev&. 01. Figure 5.. Castanopsis Forest a t a b o u t 400 m Elevation a t Tiantong. (near relev6 # 3)..
(12) Table 5.. Composition of T w o Relatively Mature but Shorter Stands of Evergreen Broad -Leaved Forest near Ridgetops. Upper slope at 330m Slope:32" to W Relevi! size: 15 x 15 m. Upper slope at 390m Slope:42" to E Relevi! size :15x 15m. T: 4 3 1 1 (1 1. 3 2 1 1 1) 1. Quercus stewardii Schima superba Distylium myricoides Cleyera japonica Lithocarpus henryi Carpinus fargesii (decid.). T: 3 - 2 3.2 3.2 2.2 1.2 1.1 1.1 2.1 1.1. Neolitsea aurata v. chekiangensis Castanopsis fargesii Rhododendron ovatum Lithocarpus henryi Schima superba Symplocos anomala Distylium myricoides Ilex buergeri Daphniphyllum oldhamii Syzygium buxifolium Cinnamomum nikkei Photinia serrulata Castanopsis carlesii Lithocarpus glabra Damnacanthus indica Ilex purpurea Symplocos botryantha Rhaphiolepis indica Schopfia jasminodora Vitis itaefolium (vine). S:. Woodwardia japonica Dichranopteris dichotoma Carex sp. (big) Schima superba (seedlings) Carex sp. 2 (medium) Stauntonia leucantha (vine) Stroemia sp. (seedling) Syzygium sp. (seedling) Dal bergia hancei (seedling). 2 2 2.2 1-2 1.2 1-2 1.2 1-2 1.1 1-1 1.1 1-1 2 +* 2 + a. + + + + + + + -I+ + +. Quercus gracilis Schima superba Castanopsis carlesii Symplocos botryantha Lithocarpus henryi Persea thunbergii Cleyera japonica Fraxinus sp. (decid.) Carpinus fargesii (decid.) Lithocarpus henryi Rhododendron ovatum Quercus rubina Eurya loquiana Symplocos stellaria Cinnamomum subavenium Camellia sp. Persea thunbergii Litsea elongata Eurya rubiginosa var. attenuata Ilex ficuoides Quercus gracilis Eurya rubiginosa Castanopsis fargesii Viburnum symplocoides Quercus glauca Neolitsea aurata Syzygium buxifolium Photinia serrulata Cleyera japonica Cunninghamia lanceolata Lorapetalum chinensis Rhus (sylvestris?) Stauntonia leucantha (vine) Woodwardia japonica Dichranopteris dichotoma Dryopteris pacifica Carex sp. (big) Tylophora japonica Stauntonia leucantha (vine) Lithocarpus henryi Ardisia crenata Prunus sp. (seedling) Styrax fabri (seedling) Lophaterum gracile.
(13) The herb layers of the slope-forest relevks contain a number of woody seedlings which may be useful indicators of forest type and future structure. These include Schima superba, Persea t hunbergii, Lit hocarpus henryi, evergreen Quercus spp., and Cinnamomum subavenium (all evergreen), which already occur in higher strata. In addition to these, though, there were also seedlings of Ficus sarmentosa var. henryi (a viny, evergreen arborescent), Styrax fabri (deciduous), Syzygium buxifolium and Ardisia crenata (evergreen understorey arborescents) , Paederia scandens (Rubiaceae, a common liana) , Dalbergia hancei (Leguminosae, a mainly subtropical-tropical genus) , Daphniphyllum oldhamii (Euphorbiaceae, an evergreen understorey arborescent ) , Maesa japonica, Damnacanthus sp., and Osmanthus fragrans (widespread evergreen understorey shrubs), Rubus spp., an occasional Acer seedling, and seedlings or young individuals of the vine taxa Smilax, Tetrast igma hemsleyanum (Vit aceae) , and Mussaenda scandens (Rubiaceae) . A1t hough the number of such seedlings was small, Fujiwara (Miyawaki et al. 1986), based on experience with evergreen broad-leaved forests in Japan, suggested several species a s possible differential species for identifying different communities within the more general evergreen broad-leaved forest type (see Table 4). Further subdivision, however, would require many more relev&.. a. The Deciduous Ravine Forest In a t least one ravine (and probably others), where groundwater is high, the evergreen forest was replaced by forest with a deciduous overstorey and evergreen broad-leaved understorey, a s shown by the relevl: in Table 6. The canopy is dominated by Choerospondias axillaris (Anacardiaceae; earlier Spondias axillaris, cf. Inst. of Botany 1972, vol. 2), a deciduous tree occurring from northeastern India to southern China (Mabberley 1987) but which also occurs in Japan (Hayashi 1985) and is mentioned by Wang ( 1961) from mixed mesophytic forests of the upper Changjiang (Yangtze) valley a s well a s Guangdong and Hainan. The only other tree species are the deciduous Acer oliverianum and four understorey tree species, including deciduous Aleurites fordii (Euphorbiaceae). The other three understorey trees are evergreen species more common to East Asian evergreen forests, and the shrub layer is quite diverse, with over half its 40 species being evergreen and typical of evergreen forests. The herb layer is also diverse (30 species), giving this relevb more species than any of the evergreen forest samples. For the ravine forest releve a s a whole, 39 of the 72 species did not occur in any of the evergreen forest relevbs a t Tiantong (19 of 40 shrub-layer species and 17 of 30 herb-layer species, plus the three deciduous tree species). At least some of these species do, however, occur in laurel forest. These include (in addition to the deciduous trees) : Cudrania cochinchinensis (Moraceae) , Ligust rum obt usif olium, Thea sinensis, Illicium lanceolat um, Persea lat ophylla, Phoebe sheareri, and Cephalotaxus fortunei (all evergreen shrubs or potential trees), plus Liriope minor and Ophiopogon japonicus, the ferns Neolepisorus ovatus and Diplazium met t enianum. Deciduous species include Celt is biondii, Ulmus parvif olia, Tilia sp., and Hovenia dulcis, and vines Wisteria sinensis, Clematis henryi and Clematis sp., Cayratia japonica (Vit aceae) , and Kadsura longipedunculata (Schisandraceae) . Above the saturated bottom and lowermost slopes of the ravine, the forest reverts to the evergreen Castanopsis-type forest which covers most of the Tiantong slopes. This suggests that the ravine is too wet (perhaps with anaerobic soil conditions) for larger evergreen broad-leaved trees and corresponds to a pattern found also in floodplains of southern.
(14) Table 6. Relev6 in a Deciduous Ravine Forest at Tiantong. Elevation: 270m Slope: 22" to SE (lower slope o f ravine, high groundwater) Relev6 size: 10m across x 20m down. TI :. T2 :. 4 3. 2 2. Choerospondias axillaris (decid.) Acer oliverianum (decid.). 2 3 2 3. Litsea elongata Aleurites fordii (Euph., decid.). 1 1 1 1. Lithocarpus henryi Cinnamomum subavenium. Lithocarpus henryi Persea latophylla Litsea elongata Phoebe sheareri Quercus gracilis ( E G ) Thea sinensis Illicium lanceolatum Camellia fraterna Castanopsis fargesii Stauntonia leucantha Acer oliverianum Persea thunbergii Cephalotaxus fortunei Pleioblastus sp. (bamboo) Quercus myrsinaefolia ( E G ) Eurya muricata Quercus glauca ( E G ) Ardisia crenata (Myrsinac.) Trachelospermum jasminoides Wisteria sinensis. Trema cannabina var. dielsiana Lonicera macrantha (climber) Symplocos caudata Lindera rubronervia Ficus pumila Cudrania cochinchinensis (Morac.) Ligustrum obtusifolium Prunus spinulosa Tilia sp. Ulmus parvifolia Hovenia dulcis (Rhamnac.) Castanopsis carlesii Celtis biondii Lonicera japonica Diospyros lotus Fraxinus retusa Carpinus fargesii Clerodendrum cyrtophyllum (Verbenac,) Euonymus streptopterus Paederia scandens (Rubiac.). Trachelospermum jasminoides Arachniodes pseudaristata Ficus sarmentosa var. henryi Diplazium mettenianum Woodwardia japonica Oplismenus undulatifolia Arachniodes rhomboidea Dryopteris fuscipes Ophiopogon japonicus Carex sp. 1 (large) Clematis henryi Rubus buergeri Rubus pacificus v. ningpoensis Zanthoxylum scandens Arachniodes simplicior. Kadsura longipedunculata (Schisandr.) Oreocnida frutescens Dryopteris bissetiana Liriope minor Solanaceae sp. ( f r ) Clematis sp. Cayratia japonica (Vitac.) Smilax arisanensis Aster ageratoides Carex sp. 2 (small) Neolepisorus ovatus Platycarya stro bilacea (Juglandac.) Phaenosperma globosa (Graminae) Rhus sylvestris Dalbergia hancei. This forest, with summergreen canopy but largely evergreen understoreys, occurred near the bottom o f a moist ravine, where high groundwater appeared to preclude growth o f theadjacent evergreen broad-leaved canopy dominants. Neither canopy species nor about half the other species o f the ravine relev6 occurred i n any sample o f evergreen forest at Tiantong, though some o f these are known to be evergreen forest species. (EG=evergreen, decid.=deciduous)..
(15) Japan (e.g. Ohno 1979). Shallower-rooted shrubs and arborescents, including evergreens, apparently are not excluded by the high groundwater, presumably finding enough aeration near the soil surface. Celtis, Ulmus, Fraxinus (see Table 6), and evergreen Ligustrum, on the other hand, are common floodplain species throughout the northern temperate zone.. Bioclimatic Terminology Discussion of vegetation and bioclimatic zonation in China is complicated immediately by the different usage in China of basic terminology, a s shown in Table 7. Thus, the northernmost zone of evergreen broad-leaved forest potential, which many scientists would call warm-temperate (e.g. Walter and Lieth 1960-67, Suzuki 1963, Greller 1980, Ovington 1983, Miyawaki 1984) is called subtropical in Chinese literature (cf. Wu 1980, Hou 1983, Song 1983, China Nat . Geogr. Comm. 1984) . This fundamental difference of opinion is explained by Song (1988) and derives mainly from the question whether the "subtropics" should represent a narrow transition surrounding the tropics o r the broader transitional zone between the tropics and the true temperate zone of higher latitudes (cf. Alisov 1954). Table 7. Forest Vegetation Zonation and Terminology on Continental East Sides in the Northern Hemisphere. ". Terminology in China w a r m - temperate. northern subtropical. middle subtropical. southern subtropical. tropical. tropical. Terminology in Japan/ eastern N. America. Vegetation Structure. (typical) temperate. Summergreen deciduous forest with deciduous understoreys. 1. Summergreen deciduous canopy with EG/ semi-EG understoreys 2. Semi-evergreen BL forest (mixed) 3. Summergreen-evergreen forest mosaic of unmixed stands Evergreen broad-leaved forest (temp. & subtropical taxa; secondary forest deciduous ; lianas & buttresses not large) Evergreen broad-leaved forest (mainly tropical taxa; larger lianas & buttresses; secondary forests mainly evergreen). >. temperate-warm temperate transition. warm-temperate. subtropical. Evergreen seasonal forest (mostly tropical taxa; large lianas & buttresses; some deciduous trees) Tropical rainforest (evergreen) (only tropical taxa; large lianas & buttresses; cauliflory; many epiphytes). tropical.
(16) The Chinese (e.g. Jiang 1960, Chen and Song 1963, Li et al. 1964, Zhu 1985) take the latter view, supported in a practical way by the fact that many tropical or subtropical crops can be grown further north a s annuals. The narrower view of the subtropics focuses more on the climate of the entire year (not only the agricultural growing season) and reflects the idea that the most obvious climatic feature of the tropics is the complete absence of frost, a t least in the lowlands. Thus, regions are considered subtropical if they have little o r no frost (but may have temperatures which approach freezing) and are considered temperate, including warm-temperate, if they have frost essentially every year (cf. Box, in press). M any scientists would thus find it difficult to accept a region a s subtropical when significant frost occurs many nights in winter and mean monthly winter temperatures are below 5 "C, a s is the case in eastern China a s f a r south a s Tiantong and other parts of Zhejiang Province. In this paper we use the somewhat more globally (but not universally) accepted terminology a s represented by most Japanese and American, a s well a s much European usage. Warm-temperate (in humid, continental east-side climates ) is taken t o mean climates which have freezing temperatures in most o r all years, though usually not below about -10°C (by which point evergreen broad-leaved trees may be damaged and often killed). Subtropical (humid, east-side) climates do not receive frost every year o r absolute low temperatures below about -2°C ever. These climatic criteria seem t o consign Tiantong and much of the East China transition region t o the warm-temperate zone (cf. southern Japan and the outer coastal plain of the southeastern USA), with Guangdong and other parts of southeastern China seemingly comparable t o subtropical Okinawa o r south Florida. Why then does the forest a t Tiantong have so many seemingly subtropical if not tropical species? At the very least, most Chinese a s well a s foreign treatments of eastern China seem to agree that there is a separate, independent vegetation zone in the transition region in which the potential forest would be dominated by evergreen broad-leaved trees but might also contain some summergreen canopy trees (cf. Han 1981, Zhou 1981, Song et al. 1982, He and Cai 1984, Zhou 1984, Wei and Deng 1986, a s well a s the general treatments by Wang 1961, Wu 1980, and Hou 1983).. Vegetation Dynamics In the perhumid evergreen "laurel forest" region of southern Japan, a t least on deep soils, the secondary forests which arise after forest cutting are mostly deciduous, composed mainly of temperate-zone taxa such a s deciduous Quercus, plus Castanea, Ulmaceae, and a few trees of tropical affinity, usually also deciduous (Miyawaki and Fujiwara 1983, cf. Miyawaki 1980-89). In less mesic situations (which may result from soil erosion), pine forests may be the main secondary vegetation and can then maintain themselves through their fire-tolerance (e.g. Nakagoshi 1987). Where mesic conditions remain, however, secondary deciduous forests and perhaps some pine forests can develop again into the stable, evergreen " laurel forests" considered t o be the potential natural vegetation of the region (Suzuki 1963, Numata 1974, Miyawaki 1984, Fujiwara 1981-86). This is due, presumably, t o the competitive advantages of evergreen broad-leaved trees over summergreen trees where conditions are continuously moist and winters relatively mild. This suggests, a s mentioned before, that the summergreen trees and conifers in the canopy a t Tiantong may not be components of the mature potential natural vegetation but rather remnants of past disturbance, which may disappear with time if no further disturbance occurs..
(17) On the other hand, under the influence of the monsoon climate of eastern China, with dry cold winters and dry conditions continuing into spring, the continuing presence of a certain number of deciduous canopy trees in otherwise evergreen broad-leaved forest has been noted as a characteristic of this vegetation type (Song, personal observation; cf . Wu 1980, and Song 1988: Table 4 ) . This suggests a similarity to forests of the southeastern US coastal plain, where "southern mixed hardwoods" (Quarterman and Keever 1962) are considered to be the natural vegetation and are mostly deciduous. Even in the most maritime, otherwise evergreen mesic forests of this region (Quercus-Magnolia-Ilex, see Wharton 1978, Box and Fujiwara 1988), persisting deciduous canopy trees (e.g. Carya glabra) can of ten still be found. In both regions, the persistence of summergreen canopy trees may be related to dryness. This dryness is seasonal in China but related more to the sandy substrate on much of the southeastern US coastal plain. Both situations would seem to reduce the competitive advantage of the evergreens, permitting coexistence of the two canopy forms. Of course, some canopy trees a t Tiantong could be recognized a s common successional species, such as deciduous Liquidambar f ormosana and coniferous Pinus massoniana. In general, the evergreen forests at Tiantong seemed to be recovering from disturbance and approaching a mature structure, with the relatively uniform physiognomy typical of East Asian " laurel forests" . Both the shorter ridgetop forests and the deciduous ravine forest seemed to be fairly stable communities related to their topographic situations. The ' Tiant ong forest is mostly evergreen, despite relatively low mean winter temperatures, because extreme temperature minima do not fall much lower (not below about -lO°C, perhaps -12°C a t highest elevations), humidity is high and fog frequent, and the forest microclimate reduces temperature extremes within the forest. The temperatures on exposed ridgetops a t even the highest elevations may not be much lower than elsewhere since these locations do not experience cold-air drainage and may experience better mixing with wetter oceanic air. The low winter temperatures in the East China transition region nevertheless suggest that this may be a marginal location for evergreen broad-leaved forest, as based on relationships in other countries. Low temperatures may explain why secondary vegetation in such climates, including southern Japan and the southeastern USA (except south Florida), is mainly deciduous (plus pines), whereas secondary vegetation further south is usually evergreen. This may be a characteristic for distinguishing warm-temperate from subtropical evergreen broad-leaved forest regions. In marginal climates, the microclimate established by forest structure is very important. Deforestation or other destruction of the evergreen forest structure leads to a southward expansion of the deciduous forest, perhaps even as the potential natural vegetation. Reforestation with evergreen species, including conifers, on the other hand, may lead to a northward expansion of the potential for evergreen broad-leaved forest, as shown by the invasion of Cryptomeria plantations in central Japan by evergreen broad-leaved tree species formerly found only further to the south (Fujiwara, personal observation). If evergreen broad-leaved forest existed further to the north across part of the long-def orested East China Plain, its necessary microclimatic conditions are now gone. Reestablishment of evergreen broad-leaved forest in this region would first require re-establishment of a forest microclimate through development of secondary, perhaps deciduous forests..
(18) Zonal Position of Tiantong Whether t o call Tiantong warm-temperate or subtropical, in the more global sense, w as not so obvious. Some seemingly subtropical genera such a s Schima and Syzygiurn, and the deciduous Choerospondias, could be found a t Tiantong and were sometimes important. The more moderate near-coastal location may permit this, but Tiantong, despite low winter temperatures, is further south than most of Japan, a s pointed out by Song (1988) in comparing the two regions. This plus the impression that there were no deciduous successional forests a t Tiantong suggests that Tiantong is subtropical, which agrees with the Chinese designation a s 'northern subtropical' (e.g. Song 1983, 1988) or 'transitional subtropical' (Hou 1983) . In the "Vegetation of China" (Wu 1980, pp. 3113141, the forest type which seemed to be most similar t o that at Tiantong is what is called the Castanopsis fargesii-C. fabri formation, which is clearly seen a s belonging to the (Chinese) subtropical zone. Another feature of subtropical forests, a t least in southern Florida, southern Yunnan, and generally in southern Japan, seems to be that taxa from more northerly regions tend to disappear rather completely (especially the typical summergreen genera, e.g. Acer, Carpinus, Fraxinus) or lose their dominance noticeably (mainly the evergreen Fagaceae) a s one goes from warm-temperate into subtropical forests. At the same time, a greater diversity of taxa with clearly tropical affinities seems to appear rather quickly. The low winter temperatures a t Tiantong, with significant frost a t least sometimes, and the continued importance a t Tiantong of taxa which seem to have temperate-zone affinities (e.g. the Fagaceae) seem t o suggest that Tiantong is still marginally warm-temperate rather than subtropical. This example of subtropical taxa occurring a t low winter minimum temperatures represents a major question for bioclimatic zonation on the humid east sides of continents. Even if Tiantong is warm-temperate, though, almost all of the warm-temperate zone, with its evergreen broad-leaved forest potential, must lie to the north. Further reconstruction of vegetation zonation is complicated by the region of drier climat e which separates the forest climates of southeastern and northeastern China. Acknowledgements We are greatly indebted t o the following people for helping with our work a t Tiantong: Mr. Wang Liang-yan (chief) and Mr. Ye Xiang-ming (vice-chief) of the Tiantong Forest Reserve; Liu Hang-gang (East China Normal University), who helped identify the plant species; and Zhang Li-quan (East China Normal University), who helped botanically and a s interpreter. In addition, two of us (EB and KF) are indebted to the organizers of the International Symposium on Development in Tropical and Subtropical Mountains, Chengdu, October 1985, which provided the opportunity for us to come t o China. Many other people helped us locally, whom we would like t o thank once again also. Finally, the non-Chinese authors would especially like t o thank again our friend and colleague, Prof. Song Yongchang, who provided the opportunity and arrangements for our visit to Tiantong..
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