PHYTOGEOGRAPHICAL Tsun-shen Ying,2 David E. Boufford,3 RELATIONSHIPS OF THE and Yuling Tu4 GENERA OF ANGIOSPERMS IN THE FANJING SHAN MOUNTAIN RANGE, NORTHEASTERN GUIZHOU, CHINA' ABSTRACT The Fanjing Shan mountain range, one of six Man and the Biosphere (MAB) Preserves in China, covers an area of ca. 567 km2 in northeastern Guizhou Province, China. The flora comprises ca. 1,400 species of flowering plants in 636 genera, 23 of which are endemic to China or barely extend into bordering areas. The importance of the 20 largest families, which together contain more than half of the total species, is evaluated, and the phytogeographical affinities of the genera of angiosperms in the flora are analyzed and briefly discussed. In the Fanjing Shan region, tropical and temperate genera are nearly equally represented. The distributional patterns of the genera are analyzed, and some comparisons are made with eight other major mountain regions in south-central and southern China that also contain rich floras of phytogeographical significance. Fanjing Shan is most similar floristically to the Wuyi Shan, Jinfo Shan, and Shennongjia regions, areas to the north and east, than it is to any of the others, which, except for one, are located to the south and west. Besides the tropical and temperate components of the flora, the Fanjing Shan region also contains a rich representation of both Sino-Himalayan and Sino-Japanese elements. China is particularly rich in numbers of plant species. The estimated 29,000 species of flowering plants and ferns within the country, including about 7,500 indigenous trees and shrubs (Pei, 1984), represent approximately 8-12% of ail vascular plants known worldwide. Part of the richness can be accounted for by the diverse geology, complex climatic patterns, and broad altitudinal and lati- tudinal ranges. Elevational differences in China span more than 9,000 m from the summit of Qomo- langma Feng (Mt. Everest) on the border with Nepal at 8,848 m above sea level to 154 m below sea level in the Turpan Depression in the Xinjiang Uygur Autonomous Region. China also occupies the only place on earth where an unbroken tran- sition of vegetational types exists, ranging from trop- ical rainforests in the south through subtropical, temperate, and boreal forests, to tundra and alpine vegetation in China's southwest and north. This vegetational continuum has resulted in associations of plants not seen in other parts of the world and includes taxa with diverse and widespread phyto- geographical relationships extending around the world. Many of the plants in these associations, or their ancestors, are among those considered to have had a much wider distribution during the Tertiary. Fieldwork in the Fanjing Shan mountain range in northeastern Guizhou by Chinese and American botanists in 1986, and fieldwork in other mountain ranges throughout China by Ying and his collab- We thank our colleagues, Bruce Bartholomew, Qianhai Chen, Sizhao Fang, Jingen Qi, Stephan A. Spongberg, Zhanhuo Tsi, Peishan Wang, and Yinghai Xiang, who accompanied us in the field in northeastern Guizhou in 1986, and Ying-chien Chien, Yenfeng Fu, Weilian Huang, and Xianpu Wang for logistic arrangements prior to our fieldwork. We are particularly grateful to the National Geographic Society, the Chinese Academy of Sciences, the Academy of Sciences of Guizhou, and the Institute of Botany, Beijing, for generous financial support. The local government officials of Tongren District, Jiangkou Xian, Yinjiang Xian, and Songtao Xian aided us in many ways and deserve special thanks for their efforts and hospitality. We are also grateful to two anonymous reviewers for their comments and useful information. 2 Institute of Botany, Chinese Academy of Sciences, 141 Xizhimenwai Dajie, Beijing, People's Republic of China. Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. 4 Department of Geography, Guizhou Normal University, Guiyang, Guizhou, People's Republic of China. ANN. MISSOURI BOT. GARD. 78: 338-358. 1991. Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan orators, has provided the basis for interpreting the phytogeographical relationships of one particularly rich area, the Fanjing Shan mountain range, lo- cated between subtropical and temperate China. HISTORICAL AND BOTANICAL SETTING Botanically, Guizhou is one of the least known provinces in China. The province, which occupies an area of about 174,000 km2 and reaches from ca. 24�50' to ca. 29�10'N and from ca. 103�46' to 10900'E, is located in a transitional position between the tropics in Guangxi and southern Yun- nan and the subtropical and temperate areas of Hunan, Sichuan, and Hubei. The province also occupies a transitional position between the Sino- Himalayan flora, which extends from Yunnan through the Himalayas, and the Sino-Japanese flora of central and eastern China. As such, Guizhou is vital for the analysis and understanding of vege- tational and floristic patterns and relationships in ail of eastern Asia. Yunnan Province, one of only a few provinces in China for which more or less accurate figures are available, and which borders Guizhou on the west, is reported to contain ca. 14,000 species of flowering plants (Wu, 1984), or nearly half of ail the angiosperms in China. Si- chuan, to the north, is also considered to be flo- ristically rich, but no accurate figures for the num- bers of indigenous plants there are available. Similarly, Guizhou has a rich flora but, again, exact figures are not known. Until relatively recent times Guizhou was one of the most backward and sparsely populated prov- inces in China (Geelan & Twitchett, 1974). It is largely mountainous with the major ranges in the north-central and south-central parts of the prov- ince. The highest peak, in the Fanjing Shan range in the Wuling Mountains, exceeds 2,550 m ele- vation (Deng, 1982). Extensive areas of flat land are scarce, and except for narrow areas along major streams, much of the land in cultivation has resulted from terracing. The two major roads in Guizhou roughly divide the province into quarters. One road runs from north to south through the center of the province and the other extends from Hunan Province in the east to Yunnan Province in the west. They intersect in the capital at Gui- yang. Minor roads reach most other parts of the province, but travel in Guizhou has always been difficult because of the rugged terrain; even now travel by highway is slow, averaging only about 40 km per hour on the best highways. Because of these factors and the difficulties involved in ex- ploiting the natural resources, the vegetation of Guizhou, although frequently heavily disturbed, is somewhat better preserved than in other parts of China. One exceptionally well conserved region in northeastern Guizhou, the Fanjing Shan mountain range, an area with an unusually rich assemblage of plants and animals, has been designated a Man and the Biosphere Preserve. Few vegetational studies have been conducted in Guizhou. The first was a general discussion on the flora and vegetation of northeastern Guizhou based on observations made in the late summer and fall of 1931 during a collecting trip to the Fanjing Shan region (Steward & Chiao, 1933). Photographs published at the time show the veg- etation to be more intact than at present, but signs of drastic disturbance are already evident in some of the photos. The more recent and most important and ex- tensive vegetational studies in Guizhou were also conducted in the Fanjing Shan mountain region and the findings published in Scientific Survey of the Fanjingshan Mountain Preserve by the Fan- jingshan Mountain Preserve Institute (1982). In- cluded in this publication are discussions on David- ia involucrata Baillon forests (Yang et al., 1982), Quercus stewardiana A. Camus (reported as Cy- clobalanopsis stewardiana (A. Camus) K. M. Lan) forests (Yang, 1983), the general vegetation in the Fanjing Shan region (Huang et al., 1982a), a dis- cussion of soils in the mountains (Zhang & Zhang, 1982), and geomorphoiogy and recent crustal movements (Yang, 1982). Another paper by Huang et al. (1982b) discusses the newly described Abies fanjingshanensis W. L. Huang, Y. L. Tu & S. Z. Fang and its forest associations and phytogeo- graphic relationships. The Davidia forests are composed of a mixture of deciduous trees with some broad-leaved evergreens and conifers at elevations between 1,000 and 1,800 m on the lower parts of moist slopes. These forests have been described as "complex," and from the descriptions are rem- iniscent of forests in which Davidia occurs in the Shennongjia Forest District in western Hubei Prov- ince (Bartholomew et al., 1983). The Davidia for- ests in the Fanjing Shan region have been consid- ered so significant that they have been given government protection. The Abiesfanjingshanen- sis forests are among the southernmost Abies for- ests in southeast Asia. According to Huang et al. (1982b) these forests, at 2,100-2,350 m eleva- tion, were first discovered in 1981. The trees grow on north-, northwest-, and northeast-facing slopes with gradients of 50-60�. The species composition and community structure of these forests are some- what similar to other Abies forests in southwest 339 Annals of the Missouri Botanical Garden China, but are reported to differ in a number of characteristics. The occurrence of five species of Fagus L. in Guizhou Province (Guizhou Flora Committee, 1982) is of particular interest and is perhaps the greatest number of species of the genus growing in close proximity anywhere. Three of these species were the subject of investigation in the early 1970s by Tsien et al. (1975; see discussion below), who found that they form distinct forests on Fanjing Shan. Guizhou is also located near the center of con- centration in China of many eastern Asian-eastern North American disjunct genera. Among the taxa belonging to this association in the Fanjing Shan region are Liriodendron L. (Magnoliaceae), several species of Asarum L. sensu lato, whose affinities appear to be somewhat intermediate between As- arum sensu stricto and the eastern North American Hexastylis Raf. In the Polygonaceae are several species of Polygonum L. sect. Echinocaulon Meis- ner, a section with species in Asia and North Amer- ica and one species, P. sagittatum L., attributed to the eastern part of both continents (Steward, 1930; Park, 1988). Another genus in the Polygo- naceae, Antenoron Raf. (Tovara Adanson in many references), with two species reported for Guizhou, is also frequent and abundant in thickets, moist woods, and along streams. The number of species in the genus is questionable, and the relationship of the Asian taxa to each other and to the single North American species, A. virginianum (L.) Rob- erty & Vautier, is uncertain. Two species of Liq- uidambar L. occur in the Fanjing Shan area, L. formosana Hance and L. acalycina Chang. Small forests and individual trees of either or both of these two species are rather common in north- eastern Guizhou. Liquidambar acalycina is con- sidered to be closely related to the North American L. styraciflua L. and has been placed in the same section by Chang (1979). Most recently, a treatment of the vegetation of the entire province has appeared (Huang et al., 1988). This work, which the authors state is aimed at university teachers and students, conservation- ists, geographers, ecologists, foresters, technicians, and farmers, discusses the physical environment, regionalization of the flora, and main vegetation types and their evolution, and utilization of the forest types in conjunction with farming, forestry, and animal husbandry. CONSERVATION Conservation in China has recently been given high priority, and it is particularly noteworthy that the Fanjing Shan mountain range and forests have been designated a Man and the Biosphere Pre- serve. Unfortunately, however, habitat destruction is taking place at a rapid rate throughout China and even Fanjing Shan is not being spared. Trees around the margins of the preserve are actively being cut for conversion to charcoal, which is then transported several kilometers to villages at lower elevations, where it is sold. Other selected trees are being cut throughout much of the easily ac- cessible parts of the preserve to build houses and for other construction purposes. The herbaceous flora suffers from the activities of overzealous dig- gers of medicinal plants who sell their harvests in ail of the larger cities and towns surrounding Fan- jing Shan. Tourism is being encouraged, and while most tourists remain close to the trails, their num- bers are becoming so great that they are having a very definite and negative impact on many of the areas they visit. Their presence has become even more noticeable with the introduction and spread of nonbiodegradable food and beverage containers throughout China over the past few years. The lack of adequate sanitary facilities will undoubtedly have a deleterious effect on water resources. Unless these threats to the integrity of the area are ad- dressed in the very near future, the Fanjing Shan region will ultimately resemble much of the rest of rural China, and many species of both plants and animals will be lost locally, if not completely. GEOGRAPHICAL LIMITS, SOIL TYPES AND CLIMATE The Fanjing Shan mountain range is generally known to include the highest peaks in Guizhou province. It is centered between 27�49'50" and 28�1'30"N latitude and 108045'55" and 108�48'30"E longitude in northeastern Guizhou. The area is deeply cut on all sides by five deep, V-shaped valleys, resulting in a steep topography. The highest peak is about 2,570 m above sea level, and the relative elevation over the surrounding countryside ranges from 1,000 to 2,000 m. Four major soil types have been described in this region: (1) yellow-red earth, below 700 m; (2) yellow earth, (500-)700-1,400 m; (3) yellow-brown earth, 1,400-2,000 m; (4) mountain brown podzolic soil, above 2,200 m. All exhibit an acid reaction, with pH values lower than 5.0. Zhang & Zhang (1982) also reported a soil type, which they call a dark, mountain thicket soil, different from the mountain yellow brown earth and the soil in shrub meadows, in a zone between 2,000 and 2,300 m. An analysis of the climatic features of the Fan- 340 Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan jing Shan mountain area was made by Wang (1982), who classified the vertical climatic zones as middle subtropical, northern subtropical, south- ern temperate, and middle temperate. On the basis of these floristic, vegetational, and environmental studies in the Fanjing Shan moun- tain range, we are now able to draw some conclu- sions on the relationships of its highly diverse and interesting flora and to discuss to some extent the phytogeography of an important segment of the Chinese flora. COMPOSITION OF THE VEGETATION IN THE FANJING SHAN MOUNTAIN RANGE There are five forest zones from the base to the crest of the Fanjing Shan mountain range: (1) evergreen broad-leaved forest zone (below 1,300 m); (2) mixed evergreen-deciduous broad-leaved forest zone (1,300-1,900 m); (3) deciduous broad- leaved forest zone (1,900-2,100 m); (4) subalpine coniferous forest zone (2,100-2,350 m); and (5) subalpine shrub and meadow zone (above 2,350 m). The dominant genera in the forests are Cas- tanopsis (D. Don) Spach, Quercus L., Fagus, Acer L., Rhododendron L., and the gymnosperm gen- era, Tsuga Carr. and Abies Miller, while the dom- inant members of the subalpine shrub-meadow zone are Rhododendron spp., Sinarundinaria chungii (Keng) P. C. Keng, and Arundinariafangiana C. Camus. Because the zone occupied by the latter association is so narrow, it is not an important floristic component of the Fanjing Shan range. The distribution of the dominant species in the other zones is as follows. EVERGREEN BROAD-LEAVED FORESTS The dominant species of this forest type are Castanopsis tibetana Hance, C. eyrei (Champion) Tutcher, C. fargesii Franchet, C. carlesii Hayata, Quercus stewardiana, Q. englergiana Seemann, and Q. dentata Thunberg var. oxyloba Franchet. Almost all of the dominant species have distribu- tions south of the Chang Jiang (Yangtze River) and are concentrated in central-southeastern China. Castanopsis fargesii, C. carlesii, and C. eyrei extend to Taiwan. DECIDUOUS BROAD-LEAVED FORESTS The deciduous broad-leaved forests contain the primary portion of the Fanjing Shan flora. The dominant members of the forests are Fagus lucida Rehder & Wilson, F. longipetiolata Seemann, F. engleriana Seemann, and Acer flabellatum Reh- der. The range of distribution of Acer flabellatum includes northeastern Yunnan, eastern Sichuan, western Hubei, northern Guangxi, northwestern Jiangxi, and Guizhou, an area where Acer is richly represented. The three species of Fagus were the subject of investigation in the early 1970s by Tsien et al. (1975), who found that they form distinct forests on Fanjing Shan. Each of the species is segregated according to slope exposure, with Fagus lucida on west- and east-facing slopes, F. engleriana on north- facing slopes, and F. longipetiolata on south-fac- ing slopes. Paleopalynological studies reveal that these Fagus communities have been in a contin- uous state of flux in response to changes in climatic conditions and that the three species have occupied various positions in relation to each other as cli- mates differed in the past (Kong et al., 1977). The present distribution of these three species of Fagus in the Fanjing Shan region corresponds to their latitudinal distribution, where F. longipetiolata is a more southern species, F. engleriana is northern, and F. lucida is a more centrally distributed plant. SUBALPINE CONIFEROUS FORESTS Among the important species in this forest type are Tsuga forrestii Downie, T. longibracteata Cheng, and Abies fanjingshanensis, with sporad- ically intermixed deciduous broad-leaved trees. Of the two species of Tsuga, one occurs in southern Sichuan, northwestern Yunnan, and western Hu- bei; the other extends from northeastern Guizhou through southern Hunan, northern Guangxi and Guangdong to Fujian. The distributional areas of these two species overlap in the Fanjing Shan and adjacent regions (Fig. 1). Abiesfanjingshanensis, undoubtedly a relict species, is endemic to Fanjing Shan at elevations between 2,100 and 2,350 m. As with the examples mentioned above, we find that nearly ail of the dominant species in the flora of Fanjing Shan are endemics concentrated in cen- tral and southeastern China. SURVEY OF LARGER FAMILIES OF ANGIOSPERMS The largest families of flowering plants in the Fanjing Shan mountain range are the Rosaceae (66 species), Poaceae (54 species), and Fabaceae (51 species) (Table 1). The Rosaceae are a major family in this region and are very characteristic of the flora and vegetation of temperate areas in Chi- na. Poaceae and Fabaceae also exhibit an extraor- dinary frequency in forests and elsewhere in this 341 Annals of the Missouri Botanical Garden *l g.oa Mi enimf flAM FIGURE 1. Distribution of Tsuga forrestii (A) and 7 longibracteata (S). Note overlap in their ranges in the Fanjing Shan region. region. The development of the Fabaceae is mainly in the Papilionoideae, which has 41 species, while the Mimosoideae and Caesalpinioideae have only 2 and 8 species, respectively. There are three families with between 40 and 50 species: the Liliaceae (48 species), Asteraceae (45 species), and Lauraceae (41 species). The Lil- iaceae and Orchidaceae form the major part of the wealth of the petaloid monocots, and together com- prise nearly 9% of the total flora. The Lauraceae, with some 41 native species, comprise an important element of the flora. This family is tropical in nature, and even most genera in temperate regions have their distribution centers in tropical or sub- tropical locations. There are five families with between 30 and 40 species. They are the Orchidaceae (38 species), Lamiaceae (36 species), Fagaceae (33 species). Cyperaceae (33 species), and Ericaceae (30 spe- cies). The 33 species in 6 genera in the Fagaceae are important elements in the flora. Many members of the family, especially species of Castanopsis, Lithocarpus, and Quercus, are the dominant forest species, and Fagus, with five species in China and three on Fanjing Shan, is centered and well de- veloped in this region. Two families, the Ericaceae and Lamiaceae, are relatively poorly represented, but three genera of Lamiaceae endemic to China (Bostrychanthera Bentham, Hanceola Kudo, and Rostrinucula Kudo) occur here. The above 11 families comprise nearly 34.8% of the species of flowering plants in the Fanjing Shan flora. There are nine families with between 20 and 30 species. In order of size they are: Ranunculaceae (29 species); Apiaceae (29 species); Theaceae (28 species); Saxifragaceae (28 species); Scrophulari- aceae (24 species); Rubiaceae (24 species); Vita- ceae (22 species); Caprifoliaceae (21 species); and Celastraceae (21 species). Except for the Thea- ceae, Rubiaceae, and Vitaceae, these families are north and south temperate, or only north temper- ate, in distribution. The large number of archaic families in the fora of Fanjing Shan should also be mentioned. Their presence may be explained by the antiquity of the flora, although they are always represented by only one or a few species. The polycarpic Magnoliidae, generally considered to comprise the most primitive families and genera of flowering plants (Cronquist, 1981), are distributed mainly in eastern and south- eastern Asia and in North, Central, and northern South America; there are 13 species in 8 genera 342 Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan TABLE 1. Ranking of families in the Fanjing Shan flora based on numbers of species. Number of Number of genera species Number of endemic Number of endemic genera' to China species2 to China 1. Rosaceae 14/48 66/855 51 2. Poaceae 41/228 54/1,202 8 3. Fabaceae 33/163 51/1,252 22 22.3% 4. Liliaceae 21/55 48/335 25 5. Asteraceae 41/227 1 45/2,323 8 6. Lauraceae 8/20 41/422 24 . 34.8% 7. Orchidaceae 29/165 38/998 13 8. Lamiaceae 20/99 3 36/808 16 9. Cyperaceae 9/31 33/668 6 10. Fagaceae 6/6 33/281 22 11. Ericaceae 7/14 30/718 21 12. Ranunculaceae 12/40 29/736 14 13. Apiaceae 15/95 29/525 11 14. Theaceae 6/7 28/397 18 15. Saxifragaceae 14/26 28/440 19 16. Scrophulariaceae 12/60 24/634 10 17. Rubiaceae 19/75 1 24/477 13 18. Vitaceae 6/7 22/109 12 19. Caprifoliaceae 5/12 21/207 17 20. Celastraceae 5/13 21/184 12 Total 322/1,398 701/13,571 342 Percentages 50.2% 50.1%/46.73% 48.9%" ' Number of genera in Fanjing Shan/number in China. 2Number of species in Fanjing Shan/number in China. 3 Percentage of total species of larger families in Fanjing Shan region. (Illicium L. (Illiciaceae) Liriodendron L., Mag- nolia L., Manglietia Blume, Michelia L. (Mag- noliaceae), Kadsura Jussieu and Schisandra Mi- chaux (Schisandraceae), and Tetracentron Oliver (Tetracentraceae)), in this category within the Fanjing Shan area. The Hamamelidaceae are also thought to be an archaic group; in the Fanjing Shan region the family is represented by 13 species in 6 genera. The main families of the old "Amentifer- ae," such as the Betulaceae, Fagaceae, Juglan- daceae, Moraceae, Salicaceae, and Ulmaceae, are found on Fanjing Shan. In addition, three families endemic or nearly endemic to China, and also con- sidered to be primitive or relictual, the Davidiaceae, Eucommiaceae, and Sargentodoxaceae (also in northern Laos and Vietnam), are also found here. The Fanjing Shan mountain range therefore har- bors a great number of archaic or primitive taxa regardless of the criteria on which they are based. The combined families mentioned above contain 701 species, or 51.1% of the total flora, and play an important role in shaping the characteristics of the forests of the Fanjing Shan mountain range. RELATIONSHIPS SHAN FLORA OF THE FANJING A. GEOGRAPHICAL AFFINITIES OF THE GENERA The Fanjing Shan mountain range is very rich in genera of flowering plants, with 636, but is not especially rich in numbers of species, with only 1,431 (2.25 species per genus). Based on the geo- graphical distribution given for each genus in Wil- lis's (1973) Dictionary of the Flowering Plants and Ferns (eighth edition), and Mabberley's (1987) The Plant-book, and supplemented by information gathered from specimens in the herbaria at A, GH, HGAS (Guizhou Academy of Sciences), Guizhou Normal University, and PE, we were able to classify the genera of flowering plants in the Fanjing Shan preserve into 14 distribution types (Table 2) as follows. 1. Cosmopolitan genera Some genera, mainly aquatics and mesophytes, are so widespread that they can be classified as 343 Annals of the Missouri Botanical Garden TABLE 2. Distribution types of genera of seed plants in the Fanjing Shan region. Percent- Number of age of ail genera in genera in Percent- China with China with Number of age of ail Number of Percentage this dis- this dis- genera in genera in species in of ail species tribution tribution Distribution type the region the region the region in the region pattern pattern Cosmopolitan 56 8.8 162 11.3 107 52.3 Pantropic 109 17.1 291 20.3 373 29.2 Tropical America and tropical Asia 11 1.7 34 2.4 96 11.5 Old World tropics 36 5.7 63 4.4 162 22.2 Tropical Asia and tropical Australia 29 4.6 48 3.4 152 19.1 Tropical Asia and tropical Africa 23 3.6 31 2.2 160 14.1 Tropical Southeast Asia 60 9.4 114 8.0 570 10.5 North Temperate 112 17.6 332 23.2 297 37.7 Eastern Asia and North America 46 7.2 114 8.0 120 38.3 Old World Temperate 31 4.9 54 3.8 163 19.0 Temperate Asia 6 0.9 13 0.9 58 10.3 Mediterranean, western Asia and central Asia 1 0.2 1 0.07 169 0.6 Eastern Asia 93 14.6 149 10.4 294 31.6 Chinese endemics 23 3.6 25 1.7 214 10.7 Total 636 99.9 1,431 100.07 2,935 21.7 cosmopolitan. Fifty-six genera fall into this cate- gory. These taxa belong to 31 of the 143 native families in the Fanjing Shan region and include 162 species. These cosmopolitans comprise about 8.8% of the flora of Fanjing Shan, and 21.7% of the total families. Many mesophytes such as Senecio L., Aster L., Geranium L., Ranunculus L., Carex L., Polyg- onum L., Galium L., and Cyperus L. are the dominant species of the herbaceous layer in forests and in montane grasslands. Because information derived from the distribution of cosmopolitan gen- era makes it impossible to determine the geograph- ical affinities of the flora of the Fanjing Shan range, they are excluded from the statistical data. 2. Pantropical genera Genera found in ail three tropical regions (the Americas, Africa-Madagascar, Asia-Australasia) may be considered pantropical. A typical example is Hypoxis L. Other pantropical genera are con- centrated in the tropics of one continent, but have few species in one or both of the other two regions. As an example, Nertera Banks & Solander ex Gaertner contains about 12 species, one common to New Zealand, South America, and Tristan da Cunha, one on Tristan da Cunha, one in Australia, one in Chile, one on Granada, two in the Philip- pines, one in south China, and eight in New Zealand (van Steenis, 1962; Wu & Li, 1965). Although the distribution of the genus might argue for the existence of Nertera before the breakup of the southern landmass, a more reasonable explanation is that members of the genus have been dispersed more recently by migrating birds that eat the berry fruits. In the Fanjing Shan region 109 genera, or ap- proximately 17.1% of the total genera (excluding the cosmopolitan ones), may be classified as pan- tropics. They contain about 291 species, which is 20.3% of ail species in the flora of Fanjing Shan. Of the 109 pantropical genera in Fanjing Shan, about 52 (47.7%) are herbaceous 42 (38.5%) are woody, 4 (3.7%) consist of lianas, and 11 others include both herbaceous and woody species. Al- though the distribution centers of these genera are primarily concentrated in the tropics of both hemi- spheres, a few of them, such as Ilex L., Cynan- chum L., Impatiens L., Buddleja L., Celastrus L., Cuscuta L., Euphorbia L., Indigofera L., and Grewia L., extend to subtropical or even temperate regions. 3. Tropical America and tropical Asia This distribution type is represented by propor- tionally few genera, especially when compared to those with a pantropical distribution. Only 11 gen- era in 10 families, representing only 1.7% of the total genera, are included in this category. These genera, ail of which are woody, contain 34 species, 344 Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan or 2.4% of the total flora of Fanjing Shan. In Australasia some of these genera may extend to northeast Australia or the southwest Pacific Is- lands, but their distribution centers are always lim- ited to the American and Asian tropics. In China, several genera extend into temperate regions, e.g., Picrasma Blume, Sageretia Brongniart, and Lin- dera Thunberg, the latter to Liaoning province at lower elevations. Ail three, however, are concen- trated in south-southwest China. Similar examples are found in Litsea Lamarck, but the genus attains its greatest diversity in southern and southwestern China, which has been considered to be its place of origin (Li, 1979). In North America, Lindera extends into the New England states and just reach- es southern Canada; Litsea is found as far north as southeast Virginia and Tennessee. The American members of these two genera may not, therefore, be considered strictly tropical. These genera are common in the Fanjing Shan region and many of their species are the main constituents of the ev- ergreen and deciduous broad-leaved forests. 4. Old World tropics This distribution type is difficult to separate from the subtropical and sometimes even temperate pat- tern. We have segregated these genera mainly on the degree of their concentration in the tropics of the Old World. An example is Dopatrium Bu- chanan-Hamilton ex Bentham, which has about 12 species in the Old World tropics (Mabberley, 1987), but only one species, which extends as far north as Henan province, in China. Within the flora of Fanjing Shan, 36 genera (5.7%) may be classified as Old World tropical. Of these, 14 (38.9%) are woody, 14 (38.9%) are herbaceous, 3 (8.3%) are lianoid and 5 genera have both woody and her- baceous species. Among them, Pittosporum Banks ex Gaertner, Albizia Durazzini, and Mallotus Loureiro are constituents of broad-leaved forests, while others such as Maesa Forsskal and Embelia Burman f. are also rather common, but only in the shrub layers of the forests. 5. Tropical Asia and tropical Australia In one sense this category is a subset of the Old World tropical pattern in that it includes genera with much the same kind of distribution, but with the main areas of concentration in the eastern and southeastern portion of the Old World tropics rath- er than in the west. The western extremity of this distribution type extends to Madagascar, but not to the African mainland. An example is found in Balanophora Forster & G. Forster in which 80 species occur in Australia, Polynesia, Malaysia, South China, Japan, and Madagascar, but none occur on the African continent. Twenty-nine genera (4.6% of ail genera) and 48 species (3.4% of ail species) can be placed in this category. Among the genera, 10 are woody, 14 are herbaceous, and 5 are lianoid; only one genus (Dunbaria Wight & Arn.) has both woody and herbaceous species. The genus Microcarpaea R. Brown, with only one species, is a typical rep- resentative of this distribution type. In China these genera occur predominantly in southern China, but a few of them enter the temperate zone. Some, such as Elaeocarpus L., Toona (Endlicher) M. Roemer, Wikstroemia Endlicher, and Nothopa- nax Miquel, are important constituents of sub- tropical forests and shrublands. Nothopanax has a disjunct distribution in southwest China and New Zealand. 6. Tropical Asia and tropical Africa This category is also a subcategory of the Old World tropical pattern in that it includes genera with much the same kind of distribution, but with their centers of concentration in the west and south- west rather than in the east. The eastern extremity of this distribution type extends to Fiji in the South Pacific, but not to Australia. A good example of this pattern is seen in Lecanthus Wedd., where three species occur in tropical Asia, one in tropical Africa, and one in Fiji, but none occur in Australia. A similar example is found in Girardinia Gaudi- chaud-Beaupre, which also extends into subtropical to temperate China. The genus Crassocephalum Moench contains 30 species, 29 of which are re- stricted to Africa and Madagascar, but one ag- gressive species occurs in eastern Asia. In the Fanjing Shan region, 23 genera (3.6% of the total) belong to this distribution type. These genera contain 31 species (2.2% of the total spe- cies). Among the genera, 6 are woody, 10 are herbaceous, 3 are lianoid, and 4 contain both woody and herbaceous members. Except for Debregeasia Gaudichaud-Beaupre and Miscanthus Andersson, ail of them are uncommon in this region. The monotypic genus Toddalia Jussieu is a typical rep- resentative of this distribution type. 7. Tropical southeast Asia The Indo-Malesian region has one of the richest floras in the world. The eastern portion of the Indian region, and the northwestern portion of the Malesian region, through the Malay and Indo- chinese peninsulas, are very closely linked floris- tically with the mountains of south-central China. 345 Annals of the Missouri Botanical Garden Thus, a fairly large number of "typical" Indo- Malesian genera occur in south China. In the Fan- jing Shan region, 60 of the 636 genera (9.4%) are included in this category. These genera contain about 114 species and constitute 8.0% of ail the species in the flora. Among these 60 genera, 33 (55%) are woody, 17 (17%) are herbaceous, 6 (10%) are lianoid, and only 4 genera contain both woody and herbaceous species. Some of the woody genera, such as Phoebe Nees, Machilus Nees, Neolitsea (Bentham) Merrill, Camellia L., Schima Reinwardt ex Blume, and Adinandra Jack, Daphniphyllum Blume and Carrierea Franchet are the main constituents of evergreen, deciduous, and mixed broad-leaved forests in the Fanjing Shan area. Ophiorrhiza L. and Dichroa Loureiro are rather common in the herb and shrub layers of these forests. 8. North Temperate Included under this heading are ail the widely distributed genera restricted to the temperate regions of Eurasia and America. There are also a few genera that are equally characteristic of North Temperate areas and parts of the tropics, especially in the more mountainous regions of the latter. For example, the genus Vaccinium L. is widespread in temperate Eurasia and America, but some species extend into tropical regions of Asia and America, especially into tropical mountainous regions. Other examples are Juglans L., Cotoneaster Medikus, Artemisia L., Rosa L., Thalictrum L., and Sam- bucus L. In the Fanjing Shan area the genera belonging to this category total 112, or 17.6% of the total genera enumerated. They represent the highest proportion of distribution types in the flora of Fanjing Shan. These genera contain over 330 species, nearly 25% of ail species in this area. Among them are 42 woody and 64 herbaceous genera, 8 genera with both woody and herbaceous species, and only 1 lianoid genus. This category forms the nucleus of the flora of the Fanjing Shan mountain range. Among the es- pecially prominent and significant genera in terms of their dominance in the vegetation of the region are Acer, Fagus, Betula L., Carpinus L., Alnus Miller, Cornus L., Castanea Miller, Juglans, Sor- bus L., Prunus L., Tilia L., Rhododendron, Ilex, and Fraxinus L. Most of these genera contain only deciduous trees and ail of them are typical North Temperate genera. Within this category, however, 17 of the 112 genera occur disjunctly in both North Temperate and South Temperate regions, while Coriaria L., a common shrub genus, shows a dis- continuous distribution in the Mediterranean area, eastern Asia from the Himalayas to Japan, New Zealand, Mexico, and Chile. 9. Eastern Asia and North America Among North Temperate disjunction patterns, the one between eastern Asia and North America is perhaps the most important biogeographically for the amount of interest it has generated. It has been one of the best known since Asa Gray (1846) first drew special attention to it, and it has been much studied and discussed (Boufford & Spong- berg, 1983; Graham, 1972a, b; Hara, 1952, 1956, 1972; Hu, 1935, 1936; Li, 1952; Wu, 1983). In the flora of Fanjing Shan 46 genera, which represent 7.2% of the total genera, belong to this distribution type. These genera contain 114 spe- cies, or 8.0% of ail species in the region. Among these genera, 23 (50% of this distribution type) are woody, 17 (37%) are herbaceous, 4 (8.7%) are lianoid, and only 2 genera (4.3%) have both woody and herbaceous species. Although genera with this distribution pattern are concentrated in eastern Asia and North America, some of them (such as Itea L., Nyssa L., Castanopsis, and Lith- ocarpus) extend into the Indo-Malesian region, or occasionally even into Central Asia (Veronicastrum Moench), or Australia (Lespedeza Michaux) in the Old World, and into tropical regions in the New World (Catalpa Scopoli, Muhlenbergia Schreber, and Illicium). Tsuga and Castanopsis are impor- tant constituents of hemlock forests and broad- leaved evergreen forests, respectively. Lithocar- pus, Liquidambar, Hydrangea L., Illicium, Sas- safras Nees & Ebermaier, Photinia Lindley, and Aralia L. are also major constituents of evergreen and broad-leaved deciduous forests in this region. Two additional genera, Abelia R. Brown and Cley- era Thunberg, are also rather common; they are disjunctly distributed in eastern Asia and Mexico. 10. Old World Temperate Discussed here are genera widely distributed over the North Temperate Zone of the Old World. At both ends of their range, and more especially in the west, many of these genera, such as Dip- sacus L., Inula L., Lactuca L., Lotus L., Oenanthe L., and Peucedanum L., often show a tendency to extend southward into Africa. Two additional gen- era, Ligustrum L. and Daphne L., although they range throughout the Old World Temperate Zone, may also be mentioned here because, respectively, they also have one or a few species in Australia and in the Indo-Malesian region. In the flora of Fanjing Shan, 31 genera, or 4.9% of ail genera, belong to this distribution type. These 346 Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan genera contain 54 species, or about 3.8% of the total. Among the genera, 6 are woody, 22 are herbaceous, and only 3 have both woody and her- baceous members. The woody genera are not the main constituents of the forests of Fanjing Shan, but the herbaceous genera are the main constitu- ents of the montane grasslands or of the herb layers of the forest. 11. Temperate Asia This type is represented by proportionally few genera, especially when compared with the North Temperate type. Included here are ail the genera mainly distributed in Temperate Asia. Their dis- tributional areas sometimes extend southward to the more elevated regions of the subtropical zone. For example, Campylotropis Bunge (Fabaceae) contains about 45 species, of which 29 are dis- tributed in China, and almost ail of which are concentrated in the subtropical regions of the coun- try (Fu, 1987). These species show their temperate nature, however, in their ecologically restricted occurrence to areas with elevations between 1,000 and 3,000 m. We have segregated this group of genera mainly on the basis of the ecological pref- erence of their subtropical representatives. In the flora of Fanjing Shan this distribution type is represented by only 6 genera, or 0.9% of ail genera. Together they contain about 13 species, which comprise only 0.9% of ail species. With the exception of one woody genus, Campylotropis, ail are herbaceous. 12. Mediterranean, western Asia to central Asia and China; North America to Central America In the fora of Fanjing Shan, only Pistacia L. belongs to this distribution type. 13. Eastern Asia This category is a large one calculated to contain about 93 genera (14.6% of the total) and is rather difficult to distinguish from the temperate Asian distribution type. The genera occupy smaller rang- es and are essentially distributed in China and Japan. These genera can be roughly divided into three subgroups based on their distribution centers in relation to Guizhou. One includes genera extending from the Himalayan region across China to Japan; the other two subgroups have eccentric distribution centers, either westward or northeastward. Mem- bers with a westward distribution show a Sino- Himalayan range, from China to the Himalayas; the northeastward group shows a Sino-Japanese pattern, from China to Japan. Among the primarily Eastern Asian genera, about one-fifth belong to the Sino-Himalayan-Japan subgroup, one-fifth to the Sino-Himalayan subgroup, and three-fifths to the Sino-Japanese subgroup. These genera sometimes extend southward into Burma, Indochina, and/or the Malay Archipelago. Among the genera distributed from the Hima- layan region to Japan are: Choerospondias B. L. Burtt & A. W. Hill, Dichocarpum W. T. Wang & Hsiao, Hovenia Thunberg, Neillia D. Don, Houttuynia Thunberg, Euscaphis Siebold & Zuc- carini, Pterostyrax Siebold & Zuccarini, Liriope Loureiro, Boenninghausenia Reichenbach ex Meisner, and Helwingia Willdenow. Of the 93 genera in this category in the flora of Fanjing Shan (14.6% of the total genera), 47 are herbaceous, 38 are woody, and 8 are lianoid. The woody genera, such as Cercidiphyllum Sie- bold & Zuccarini, Acanthopanax (Decaisne & Planchon) Miquel, Dendrobenthamia Hutchinson (= Cornus L. subg. Syncarpea (Nakai) Q. Y. Xi- ang), Enkianthus Loureiro, Idesia Maximowicz, Platycarya Siebold & Zuccarini, Pterocarya Kunth, and Tetracentron Oliver, are important constituents of the broad-leaved forests, while the herbaceous genera, such as Ophiopogon Ker-Gaw- ler, Liriope, Reineckea Kunth, Tripterospermum Blume, and Ainsliaea DC., play an important role in the herb layer of the same forests. Sinarundi- naria Nakai is the main constituent under trees in the broad-leaved deciduous forests and in subalpine scrublands. 14. Endemic genera For the Fanjing Shan mountain range as a whole, 23 of the 636 native genera (3.6%) are endemic to China, but ail of them are shared with other provinces. These endemic genera are listed with their ranges and number of species in Table 3. Twelve of these genera are monotypic and nine are oligotypic; the two remaining genera have mul- tiple species. Among the 23 are 11 arborescent genera, of which nine are deciduous and two are evergreen. Eleven genera are herbaceous and only one, Clematoclethra Maxim., is lianoid. Except for three genera, Eurycorymbus Handel-Mazzetti (Sapindaceae: 1 species), Dysosma Woodson (Ber- beridaceae: 7 species), and Cunninghamia R. Brown (Taxodiaceae: 2 species), which occur in Taiwan, all are restricted to central and south- eastern China. Based on the figures cited above we were able to conclude that among the native genera in the flora of Fanjing Shan, 268 (42.1%) are tropical, 347 Annals of the Missouri Botanical Garden TABLE 3. Chinese endemic or near endemic genera in the flora of Fanjing Shan with overall distribution of the genus. The second column indicates the number of species in Fanjing shan/number of species in China. Asteropyrum (Ranunculaceae) Bretschneidera (Bretschneideraceae) Bostrychanthera (Lamiaceae) Chimonanthus (Calycanthaceae) Camptotheca (Nyssaceae) Clematoclethera (Actinidiaceae) Cyclocarya (Juglandaceae) Cunninghamia (Taxodiaceae) Davidia (Nyssasaceae) Dysosma (Berberidaceae) Dickinsia (Apiaceae) Dipteronia (Aceraceae) Eucommia (Eucommiaceae) Eomecon (Papaveraceae) Emmenopterys (Rubiaceae) Eurycorymbus (Sapindaceae) Hanceola (Lamiaceae) Latouchea (Gentianaceae) Pteroceltis (Ulmaceae) Rostrinucula (Lamiaceae) Sinojohnstonia (Boraginaceae) Thyrocarpus (Boraginaceae) Whytockia (Gesneriaceae) 1/2 Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan. 1/1 Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Tai- wan, E Yunnan, Zhejiang. 1/1 Fujian, Guangdong, Guangxi, Guizhou, Hubei, Sichuan, Tai- wan. 1/2 Anhui, Fujian, Gansu, Guangxi, Guizhou, Henan, Hubei, Hun- an, Jiangsu, Jiangxi, Shaanxi, Sichuan. 1/1 Fujian, Hubei, Hunan, Guangdong, Guangxi, Guizhou, Jiangxi, Sichuan. 1/4 Guizhou, Hubei, Shaanxi, Sichuan, Yunnan 1/1 Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Zhejiang. 1/2 South of the Qinling range. 1/1 Guizhou, Hubei, Sichuan, N Yunnan. 3/7 South of the Qinling range. 1/1 Guizhou, Hubei, Hunan, Sichuan, Yunnan. 1/2 S Gansu, Guizhou, Henan, W Hubei, Shaanxi, Sichuan, SE Yunnan. 1/1 E to SW China. 1/1 Fujian, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan. 1/1 E to SW China (and India). 1/1 S and SW China, Taiwan. 1/8 Guangxi, Guizhou, Hunan, Sichuan, Yunnan. 1/1 Fujian, Guangdong, Guizhou. 1/1 N to S and SW China. 1/2 Guangxi, Guizhou, Hubei, Hunan, Shaanxi, Sichuan, Yunnan. 1/3 Guizhou, Hubei, Jiangxi, Shaanxi, Sichuan, Zhejiang. 1/3 Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, S Jiang- su, Jiangxi, E Yunnan. 1/3 Guizhou, Taiwan, SE Yunnan. 289 (45.4%) are temperate, and 23 (3.6%) are Chinese endemic genera. It is clear in the numbers and percentages that tropical and temperate genera are of about equal importance in the flora of Fanjing Shan. Taking a more general view, however, and looking for ultimate sources, it appears that the flora of Fanjing Shan can be thought of as com- prising two main elements, a northern and a south- ern one, disbursed, as would be expected, along an altitudinal gradient in five vegetational zones (as discussed above). B. RELATIONSHIPS WITH OTHER MOUNTAINOUS REGIONS IN CHINA To determine the area's floristic affinities within China, eight mountainous regions in the central and south-central part of the country (Fig. 2) were selected for comparison with the Fanjing Shan re- gion. Based on the worldwide distribution of the genera in the nine mountain regions, 15 distribution patterns can be recognized (Table 4). As expected, temperate genera increase and tropical genera de- crease in numbers with increasing latitude. Fanjing Shan, with 268 tropical and 289 temperate genera (excluding the Chinese endemic genera), has about an equal representation of temperate and tropical elements. Taibai Shan, located to the north of Fan- jing Shan in Shaanxi province, has 397 (68.0%) temperate and 105 (18.0%) tropical genera; cos- mopolitan genera (60, or 10.2%) and endemic genera (22, or 3.8%) make up the remaining 14%. Wuzhi Shan has the highest proportion of tropical genera (88.8%) of any of the areas examined, which would be expected from its southerly location on Hainan Island at about 19�N latitude. The num- bers of shared genera and the coefficients of sim- ilarity between these mountainous regions are given in Table 5. The higher coefficients of similarity between the Fanjing Shan region (A) and Wuyi Shan (B) (52.8%), the Fanjing Shan region and Shennongjia (D) (50.6%), and the Fanjing Shan region and Jinfo Shan (C) (50.7%) indicate closer phytogeographical links between those four areas 348 Volume 78, Number 2 1991 Ying et al. Phytogeography of the Fanjing Shan li O- Sa & - O0 o z \o-0vOLt0r-O Lrr O L0 � OO t- L O - L c�O\r-' un --4 l C- Il ag 0< o � -v Q Il I Uz o 2 < �aa Q cH H o d Q v U x t O V� c 3 `l) �o d � Il o-IO O s- t .