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Home , Aeginetia, Alphonsea, Begonia tonkinensis, Biophytum, Boehmeria, Celtis philippensis

BIOGEOGRAPHY AND H. Zhu,2 H. Wang,2 B. Li,2 and 3 FLORISTIC AFFINITIES OF P. Sirirugsa THE LIMESTONE FLORA IN SOUTHERN , CHINA1

ABSTRACT

The on limestone in southern Yunnan, in tropical southwest , were inventoried, and their floristic composition and biogeographical affinities are discussed. These limestone forests were characterized by phanerophytes making up ca. 78% of the total and those with mesophyllous comprising 75%. Ecological species groups based on their habitat preferences were discerned from field observations: the species exclusive to the limestone habitats make up 10% and the preferents make up ca. 12% of the total limestone flora. From these limestone forests, 1394 vascular species belonging to 640 genera and 153 families were recorded. Based on their distributions, 12 biogeographic elements at the generic level and nine at the specific level were recognized. About 90% of the plant genera (over 90% of the species) were tropical; furthermore, 35% of the seed plant genera (65% of the species) have tropical Asian affinities. In a comparison with other regional floras from southern China and tropical , the limestone flora of southern Yunnan revealed closer affinity to tropical floras than to temperate elements of eastern Asian floras. This limestone flora is thus tropical and part of the tropical Asian flora at its northern margin. Key words: biogeography, China, limestone , southern Yunnan.

Limestone in tropical China occurs mainly in GENERAL GEOGRAPHY Yunnan and Guangxi Provinces of southern and Xishuangbanna, the southern part of Yunnan, central China. Because of the great diversity of which borders Burma and Laos, is a mountainous edaphic conditions and topography, vegetation area at the northern margin of tropical Southeast types on limestone are extremely diverse and rich Asia (Fig. 1). Basically, the study area has a moun- in endemic taxa. Limestone vegetation in southern tainous topography with the mountains running China has been destroyed as much as other vege- north-south and decreasing in elevation southward. tation types even though these limestone areas are Altitude varies from 480 m in the lowest valley in more difficult to access and to farm. Limestone veg- the south to 2400 m at the top of the highest moun- etation is also more vulnerable because it recovers tain in the north. The limestone strata occur mainly much more slowly on usually thin soils. Our re- in southeastern Xishuangbanna and range in alti- search was conducted mainly in the area of Xish- tude from 600 to 1600 m. uangbanna, in the southern part of Yunnan, where The region of Xishuangbanna has a typical trop- about 19% (3600 km2) of the total area is limestone ical monsoon climate with an annual mean tem- (Liu et al., 1990). Most of this limestone area is perature of 22ºC, annual temperature accumulation still forested and is receiving increasing attention (the sum of daily temperature means where they are for its and its urgent need of conser- Ͼ 10ЊC) of 8000ЊC, and annual precipitation vary- vation. Primary floristic works in southern Yunnan ing from 1200 to 1556 mm, of which more than have been written (Zhu et al., 1996, 1997, 1998a, 80% falls during the rainy season between May and 1998b; Wang et al., 1997). This paper represents a the end of October (Xu et al., 1987). synthesis of its floristics, physiognomy, and biogeo- The rock substrate is hard limestone of Permian graphical affinities. origin with a rugged topography. The soil is mainly

1 This project was funded by The National Natural Science Foundation of China (40271048), the Chinese Academy of Sciences (The Fund for Top One Hundred Young Scientists and KSCX2-1-06B), and the Yunnan Natural Science Foundation. The senior author thanks Xu Zaifu for his great help with his research and Wu Zheng-yi and Zhang Hong- da, his academic advisors. He particularly thanks E. Tanner and P. Grubb for their help in analyzing data and preparing this paper during his visiting scholar’s year at the University of Cambridge. Finally, he thanks T. C. Whitmore, who has greatly supported and helped him in his research, and two anonymous reviewers for their constructive comments on this article. 2 Xishuangbanna Tropical Botanical Garden, The Chinese Academy of Science, Mengla, Yunnan 666303 P. R. China. E-mail address for correspondence: [email protected]; [email protected]. 3 Prince of Songkla University, Hatyai 90112, .

ANN.MISSOURI BOT.GARD. 90: 444–465. 2003. Volume 90, Number 3 Zhu et al. 445 2003 Southern Yunnan Limestone Flora iengdao, N 2. Longgan, SW Figure 1. Locations of the research area in Xishuangbanna, southern Yunnan, China, and compared regional floras (see Table 6). —1. Our research area. — China. —3. Daqinshan Mountains,Thailand. SW —9. China. The —4. Malay Gulinqing, Peninsula limestone. SW —10. China. Taiping, —5. Malay Huapin, Peninsula. China. —6. Dongyang Mountains, China. —7. Cucphuong, N Vietnam. —8. Ch 446 Annals of the Missouri Botanical Garden

brown, coarse in texture, and composed of loamy tropical seasonal moist forest, and tropical montane limestone with a pH of ca. 6.75 and ca. 3.56% dwarf forest (Zhu et al., 1998a). Within these, six organic matter (Liu et al., 1990). formations, including nine communities, were rec- ognized: METHODS (1) Ravine seasonal rain forest (including the Po- A complete floristic inventory was made based metia tomentosa–Alphonsea monogyna community on the identification of more than 5000 plant spec- and Pometia tomentosa– philippensis var. imens collected from the limestone habitat in wightii community); southern Yunnan during 1985–1995 and deposited (2) Lower hill seasonal rain forest (including only mainly at HITBC and SYS. The flora of the vege- the Celtis philippensis var. wightii–Lasiococca com- tation on the limestone consisted of 153 families of beri var. pseudoverticillata community); vascular , including 640 genera and 1394 (3) Evergreen moist forest (including the Osmanthus species. An initial floristic analysis was made based polyneurus–Dracaena cochinchinensis community on the inventory (Zhu et al., 1996). Three main and Lasiococca comberi var. pseudoverticillata–Cleis- vegetation types occur on the limestone—tropical tanthus sumatranus community); seasonal rain forest, tropical seasonal moist forest, (4) Semi-evergreen moist forest (including the and tropical montane dwarf forest—which were se- Bombax insignis–Colona floribunda community and lected for establishing plots. For the tropical sea- Bombax insignis–Garcinia bracteata community); sonal rain forest, seven separate plots ranging in (5) Evergreen dwarf forest (including only the Pho- size from 2000 to 2500 m2 were established. For tinia angusta–Pistacia weinmannifolia community); the tropical seasonal moist forest seven separate (6) Semi-evergreen dwarf forest (including only the plots ranging in size from 500 to 2000 m2 were laid neriifolia–Dracaena cochinchinensis commu- out. For the tropical montane dwarf forest, only two nity). plots of 10 by 10 m were made due to its restriction to limestone summits. These different plot sizes Detailed descriptions and ecological analyses of were used because of the differential coverage of the communities have been reported earlier (Zhu et forest type and site restrictions. The structure and al., 1998a). Here the classification of the limestone species composition of the vegetation on the lime- vegetation is concisely enumerated so that the bio- stone were analyzed based on plot data already geographical components of the limestone vegeta- published (Zhu et al., 1998a). In the present paper, tion can be better understood. plant inventory lists of the two main forest types (excluding montane dwarf forest) were compiled TROPICAL SEASONAL RAIN FOREST from sample plots separately for the physiognomic Tropical seasonal rain forest on limestone, just (life form and size) analysis. The criteria for as the regional tropical seasonal rain forest off lime- life form and leaf size classes suggested by Raun- stone, shares characteristics with the equatorial kiaer (1934) and the importance value index (IVI) lowland rain forest. These forests are mainly ever- suggested by Curtis and McIntosh (1951) were used green, but there are some deciduous in the in the physiognomic or ecological analysis. Ecolog- emergent layer. This is equivalent to the tropical ical species groups were discriminated from field semi-evergreen rain forest of Southeast Asia (Whit- observation and correspond to groups used in Shi- more, 1984), or the tropical semi-evergreen forest mizu (1964) and Chin (1977). Species-level bio- of –Burma (Champion, 1936), as well as the geographical affinities were assessed for the total evergreen seasonal forest of tropical America flora of the limestone vegetation. The floristic sim- (Beard, 1944, 1955). In southern Yunnan, these ilarities between the limestone flora of southern limestone forests occur in wet valleys and on lower Yunnan and the floras on limestone and non-lime- slopes of hills or mountains below 1000 m altitude. stone habitats from southwest China, northern Vi- This same forest type also occurs in northern Thai- etnam, northern Thailand, and the Malay Peninsula land (Smitinand, 1966) and North Vietnam (Thin, were also discussed. 1997), although different names were used. The tropical seasonal rain forest represents Southeast CLASSIFICATION OF LIMESTONE VEGETATION Asian tropical rain forest at its latitudinal and al- Based on plant physiognomy, forest profile, flo- titudinal limits. The ecological structure of the trop- ristic composition, and habitat, the primary lime- ical seasonal rain forest on limestone is almost ex- stone vegetation can be classified into three vege- actly the same as the seasonal rain forest off tation types, i.e., tropical seasonal rain forest, limestone in the Xishuangbanna region (Zhu, 1992, Volume 90, Number 3 Zhu et al. 447 2003 Southern Yunnan Limestone Flora

ϭ 1997). Most species in the seasonal rain forest on — — 2 0.9% phytes Thero- limestone are also found in the adjacent non-lime- stone seasonal rain forest, but the latter is more diverse with additional species, which are not 3 1.2% 7 3.3% found on the limestone. Microphanerophyte ϭ 31 12.4% 28 13%

Epiphytes; Therophytes TROPICAL SEASONAL MOIST FOREST

ϭ Tropical seasonal moist forest occurs on the mid- 62.7% 58.8% dle and upper limestone slopes ranging from 650 156 124 to 1300 m altitude. This vegetation type abuts the seasonal rain forest and was called monsoon forest 4.8% 3 12 1.4% by some Chinese authors (Liu, 1987; Wu, 1980). Parasitic; Epiph The term seasonal moist forest is preferred here ϭ because the forest is not equivalent to Schimper’s 6.8% 9.9% 17 21 monsoon forest (Schimper, 1903), in spite of the fact that it is affected by seasonal dryness and con-

Phanerophytes tains a variable percentage of deciduous trees. The 34 13.7% 36 17% seasonal dryness in the region is compensated to some extent by dense fog accompanied by low tem- 84 33.7% 62 29% peratures in the same months (November to April) (Whitmore, 1984). Some deciduous trees, such as Gmelina arborea Roxb., Anthocephalus chinensis 9 3.6% 2 0.9% Megaph Mesoph Microph Nanoph Hph All Cham Geoph Mesophanerophyte (perennials 8 to 30 m high); Microph (Lam.) Rich. ex Walp., and Homalium laoticum Herbaceous phanerophyte (herbaceous perennials over 0.25 m high); Cham ϭ Gagn. var. glabretum C. Y. Wu, shed leaves toward ϭ 3 1.2% 5 2.3% the end of the dry season, while others, such as Cratoxylon cochinchinensis (Lour.) Bl., Ficus reli- Liana giosa L., and Elaeocarpus varunua Buch.-Ham. ex Woody Herb 48 19.3% 27 12.8% Mast., shed their old leaves as new ones develop. This suggests that deciduousness in the region is 7 2.8% 8.5% more frequently associated with locally dry habitats 18 o 2 m high); Hph Geophyte (perennials, dying back above ground); Para than the seasonal dryness of climate. Therefore, us-

ϭ ing the term monsoon forest for the evergreen or — —

1 0.4% semi-evergreen forest on limestone is confusing be- cause Schimper’s monsoon forest is more or less completely leafless during the dry season.

MONTANE DWARF FOREST Montane dwarf forest occurs only on the tops of hills and summits of mountains at altitudes above 900 m. There is only one dwarf layer with a Megaphanerophyte (perennials over 30 m high); Mesoph Number of species Percentage of total species ϭ Number of species Percentage of total species canopy height of 7–15 m. Epiphytic orchids, such Nanophanerophyte (perennials 0.25 t as Eria hainanensis Rolfe and Bulbophyllum ni- ϭ grescens Rolfe, and non-vascular epiphytes (bryo- phytes and lichens) are abundant. In some sites small woody climbers, such as Derris caudatilimba How (Papilionaceae) and Pristimera arborea (Roxb.) A. C. Smith (), are also fre- quent. Life form* Para Epiph of 7 plots, total 249

of 7 plots, total 211 LANT HYSIOGNOMY OR TTRIBUTES 2 P P A 2 From plot data, life form spectra (Raunkiaer, species) species) 1934) of the two main forest types (seasonal rain Table 1. Life forms of the limestone forest in southern Yunnan. (14800 m * Life form (Raunkiaer, 1934); Megaph (9650 m (perennials 2 to 8 m high); Nanoph Limestone seasonal rain forest Chamaephytes (perennials less than 0.25 m high above ground); Geoph (annuals). Limestone seasonal moist forest forest and seasonal moist forest) are compiled in 448 Annals of the Missouri Botanical Garden

Table 2. Physiognomic characteristics of the limestone forest in southern Yunnan.

Leaf form Leaf texture Leaf size Forest type SC PLNa Mi Me Ma Limestone seasonal Percentage of species 72.3 27.7 47.9 52.1 0 13.8 76.6 9.6 rain forest1 Percentage of Importance Value Index (IVI)3 76.3 23.7 52.8 47.2 0 3.8 91 5.3 Limestone seasonal Percentage of species 68 32 51.5 48.5 1 21.5 74.2 3.1 moist forest2 Percentage of Importance Value Index (IVI) 74.8 25.2 41.7 58.3 0.4 23.3 66.5 9.7 1 From 14800 m2 of 7 plots, total of 94 tree species Ͼ 5 cm DBH. 2 From 9650 m2 of 7 plots, total of 97 tree species Ͼ 5 cm DBH. 3 IVI ϭ Relative dominant density ϩ Relative frequency ϩ Relative dominant breast area (Curtis & McIntosh, 1951). S: Simple leaves; C: Compound leaves; P: Papery leaves; L: Leathery leaves; Ma: Macrophyll (large to 164,025 mm2); Me: Mesophyll (to 18,222 mm2); Mi: Microphyll (to 2025 mm2); Na: Nanophyll (to 225 mm2) (Raunkiaer, 1934).

Table 1. Leaf size spectra, leaf form, and leaf tex- weighting the species by IVI, the percentage of ture are shown in Table 2. Both forest types were leathery leaves decreases in seasonal rain forest but dominated by phanerophytes. Including lianas, increases in seasonal moist forest due to the pres- these perennial phanerophytes make up 73.9– ence of some species with these leathery leaves 83.2% of the total species, while annual chamae- such as Cleistanthus sumatranus (Miq.) Muell.-Arg. phytes account for only 12.4–13%. However, the () and Dracaena cochinchinensis. seasonal moist forest shows lower percentages of woody lianas as well as megaphanerophytes and ECOLOGICAL SPECIES GROUP mesophanerophytes, but higher percentages of epi- Based on the study of the limestone floras of Ja- phytes as well as microphanerophytes and nano- pan and Taiwan, Shimizu (1964) divided limestone phanerophytes than the seasonal rain forest. plants into five ecological groups: Both forest types have species with mesophyllous leaves making up ca. 75% of the total tree species, (1) plants exclusive to limestone habitat; but the forests show clear differences if the species (2) plants selective for and found mainly in lime- are weighted by importance value index (IVI). This stone; increases the percentage of mesophyllous perenni- (3) plants preferring and dominant on limestone; als and decreases the percentage of micro- and ma- (4) taxa indifferent, with no special association crophyllous trees in seasonal rain forest, while the with limestone; opposite trend is seen in seasonal moist forest. Sea- (5) plants found only occasionally on limestone or sonal moist forest occupies much more rugged hab- strangers to limestone. itats with thinner and drier soils, and has more mi- To Shimizu, these first three groups were char- crophyllous species. In weighting by IVI, the acteristic species for the limestone habitats and in increase in percentage of macrophyllous trees in particular his exclusive and selective taxa were cal- seasonal moist forest is mainly due to the dominant cicoles. Chin (1977) accepted this classification evergreen species Dracaena cochinchinensis (Lour.) and similarly categorized plants on limestone in the S. C. Chen (Agavaceae), with its long leathery lan- Malay Peninsula into four groups, combining selec- ceolate leaves, and the dominant deciduous tree tive and preferent plants. Similar ecological species species Colona floribunda (Wall. ex Voigt) Craib groups have been later recognized by Chinese bot- () also with large leaves to 30 cm long. In anists (Liang et al., 1985; Liu et al., 1994).

Table 3. The ecological species groups of the limestone flora of southern Yunnan.

Ecological species groups (see Shimizu, 1964; Chin, 1977) Number of species % Plants found only on limestone: endemic to southern Yunnan 24 1.7 not endemic to southern Yunnan 117 8.4 Plants dominant on limestone 170 12.2 Plants no restriction on limestone 858 61.6 Plants found occasionally on limestone 225 16.1 Total 1394 100 Volume 90, Number 3 Zhu et al. 449 2003 Southern Yunnan Limestone Flora

Table 4. Predominant families found in limestone forests of southern Yunnan.

No. of No. of No. of No. of genera species %* genera species % Orchidaceae 35 86 26.9 6 20 43.5 Rubiaceae 34 58 64.1 Labiatae 13 20 30.1 Euphorbiaceae 27 58 60.4 Gesneriaceae 13 19 63.3 Papilionaceae 22 55 43.4 Sterculiaceae 7 18 46.8 7 48 73.8 Dioscoriaceae 1 18 64.3 Vitaceae 7 38 79.5 Menispermaceae 10 17 60.7 26 36 65.4 Liliaceae 10 16 64.0 Rutaceae 11 35 71.4 10 15 44.1 Asclepiadaceae 16 35 53.0 Compositae 8 15 14.6 12 35 53.0 Myrsinaceae 4 14 38.2 Lauraceae 10 35 45.0 Commelinaceae 7 14 60.9 19 33 58.9 Zingiberaceae 6 14 42.4 Meliaceae 12 30 83.0 Myrtaceae 1 13 45.8 12 30 50.9 Tiliaceae 3 12 57.1 Cucurbitaceae 9 24 52.3 Mimosaceae 6 12 66.6 Rhamnaceae 19 21 70.4 Anacardiaceae 7 11 64.7 3 20 54.8 Convolvulaceae 4 11 37.4 Ulmaceae 5 11 100 the no. of species on limestone * % ϭϫ100 the total no. of species in southern Yunnan

Following Shimizu and Chin’s classifications, we More than 80% of the species also occur in the divided the limestone flora of southern Yunnan into non-limestone habitats of the Xishuangbanna re- these four ecological species groups (Table 3). In gion. our study, 141 species are restricted The limestone flora makes up about one quarter to limestone habitats and thus are exclusively found of the total species of the regional flora. (The flora here. These include the following common species of the Xishuangbanna region was primarily docu- Celtis philippensis var. wightii, Amoora calcicola, mented with 3336 native species of 1218 genera Murraya tetramera, Pistacia weinmannifolia, as and 207 families of seed plants; see Li, 1996.) well as species in Agapetes, Sageretia, Tupistra, and Some families show relative preference for lime- Pristimera. Of these, 24 species are endemic to stone habitats (with more than 60% of the total southern Yunnan. Taxa exclusive to limestone make number of species in the region on limestone), for up about 10% of the total limestone flora, which example, Acanthaceae, Euphorbiaceae, Gesneri- agrees with the results from Longgan limestone (ex- aceae, Meliaceae, Menispermaceae, Moraceae, clusive taxa, 13%) (Liang et al., 1985) and Longhua Rhamnaceae, Rubiaceae, Rutaceae, Vitaceae, and limestone (exclusive taxa, 10%) (Liu et al., 1994) Ulmaceae (Table 4). Other families, such as Hip- from Guangxi Province in China. Both the exclu- pocrateaceae, Icacinaceae, and Vacciniaceae, show sive and preferent taxa make up 22.3% of the total an even stronger preference for limestone (found sum. They could be termed as characteristic spe- almost exclusively in limestone habitats in southern cies for limestone habitats (see Appendix 1). This Yunnan), although they are not among the predom- is similar to the results from Longgan in Guangxi inant families in species richness. (with these characteristic species making up 20% The distribution types of Chinese seed plants at of the total sum) (Liang et al., 1985) and from the the generic level were documented by Wu (1991). Malay Peninsula (27.5%) (Chin, 1977). Based on Wu’s document, 544 of the 558 genera of seed plants from the limestone forest of southern THE FLORA AND ITS BIOGEOGRAPHY Yunnan can be divided into 12 distribution types In the limestone forests of southern Yunnan, Chi- or geographic elements (14 genera, which are cos- na, 153 families of vascular plants including 640 mopolitan in distribution, are not included in the genera and 1394 species and varieties, were re- geographic statistics). One thousand two hundred corded, of which seed plants compose 129 families, forty-four of the 1269 species of seed plants from 558 genera, and 1269 species (see Appendix 1). the limestone forest can be recognized in nine dis- 450 Annals of the Missouri Botanical Garden

tribution types based on their geographic distribu- tion (25 species of seed plants are not included due to insufficient distribution references) (Table 5). At 7.3% 0.5% 0.6% 0.2% 0.4% 3.2% 1.3% the generic level, the geographic elements of trop- 17.0% 19.9% 20.3% 10.6% 18.6% (64.5%) of species

Percentage ical distribution (1–6, Table 5) compose 90% of the total genera; the geographic elements of temperate distribution (7–10, Table 5) make up only 6.7%. At the specific level, the species that are of typical tropical distribution (1–6, Table 5) account for 70.2% of the total species. Among these, the geo- graphic types that are considered to be from trop- ical Asia make up 64.5% of the total species from limestone forests in Xishuangbanna. If the species from the tropical areas adjacent to Xishuangbanna from southern China and Yunnan are included, these tropical species compose more than 90%. This indicates that the limestone flora at Xishuang- banna is principally tropical in nature and repre- sents the tropical Asian flora at its northern tropical margin. In a floristic comparison with nine similar floras, both limestone and non-limestone, from southwest China, northern Vietnam (Thin, 1997), northern Thailand (Smitinand, 1966), and the Malay Pen-

1. Pantropic 2. Tropical Asia–Tropical America3. disjunct Old World Tropics 4. Tropical Asia to5. Tropical Australia Tropical Asia to6. Tropical Tropical Asia 6a. India-Malaysia 6b. Mainland Southeastern Asia6c. to Southern Malaysia Asia to6d. Mainland Mainland Southeastern Southeastern Asia Asia7. to Eastern Southern Asia China 8. Southern China 9. Endemic to Yunnan insula (Chin, 1977, 1979; Burkill & Henderson, 1925) (Table 6), the limestone flora of southern Yunnan displays explicit taxonomic affinities to the tropical floras and shows a closer affinity to the floras from the Malay Peninsula than to other floras from subtropical China (the floras of Huapin and 2.9% 9.0% 7.9% 2.8% 2.8% 0.9% 0.2% 2.9% 0.4%

21.1% 13.8% 35.3% Dongyang, see Li et al., 1986; Xu, 1984), even of genera Geographic elements at the specific level Percentage though these Malaysian floras lie farther away geo- graphically from southern Yunnan. Our limestone flora in southern Yunnan shares the most genera with the limestone flora of northern Vietnam (Cuc- phuong, see Thin, 1997) among those floras com- pared in this study. The similarity at the generic level between our limestone flora and the limestone flora of northern Thailand (Doi Chiengdao) (Smitin- and, 1966) is less than would be expected from its geographic proximity. This lack of correspondence could be because for Doi Chiengdao used here for comparison is an incomplete one con- sisting of only 512 species, less than half reported for most other sites in Table 6. The limestone flora of Xishuangbanna did not show a higher floristic similarity to other regional limestone floras than to

(see Wu, 1991) non-limestone floras in our comparison. It appears that limestone floras develop from local or regional floras, supported also by the fact that only about 10% of the total species of limestone floras (the

Geographic element at the generic level exclusive group) are restricted to limestone habitats in our study. The floristic relationships between our limestone Table 5. Geographic affinities of the limestone forests of southern Yunnan. Total of 544 genera 100% Total of 1244 species 100% 1. Pantropic 2. Tropical Asia–Tropical America3. disjunct Old World Tropics 4. Tropical Asia to5. Tropical Australia Tropical Asia to6. Tropical Africa Tropical Asia 7. Northern Temperate 8. Temperate Eastern Asia9. and Old Northern World America Temperate disjunct

10. Temperate Mediterranean, Western11. Asia to Eastern Central Asia Asia 12. Endemic to China flora in southern Yunnan and neighboring floras of Volume 90, Number 3 Zhu et al. 451 2003 Southern Yunnan Limestone Flora

Table 6. Comparison of floristic similarities between the limestone habitats of Xishuangbanna, southern Yunnan, and the limestone and non-limestone habitats from southwestern China and southeastern Asia.

Size of flora Shared taxa Similarity Location Habitat (Seed plants) by both floras coefficients 2. Longgan, SW China limestone 149 families 118 91.2 22Њ14–33ЈN, 106Њ46ЈE 669 genera 371 66.5 1363 species 3. Daqinshan Mountains, SW China non-limestone 182 fam. 126 97.4 22Њ14ЈN, 107ЊE 871 gen. 389 69.8 1813 spp. 4. Gulinqing, SW China limestone 143 fam. 116 89.9 22Њ36ЈN, 104ЊE 496 gen. 261 52.6 1095 spp. 5. Huapin, SW China non-limestone 151 fam. 83 72.8 25Њ31–39ЈN, 109Њ50ЈE 475 gen. 150 33.2 1051 spp. 6. Dongyang Mountains, SW China limestone 116 fam. 86 73.7 25Њ14ЈN, 107Њ56ЈE 367 gen. 153 41.6 736 spp. 7. Cucphuong, N Vietnam limestone 167 fam. 120 93 20Њ14–24ЈN, 105Њ24–44 ЈE 860 gen. 428 76.7 1661 spp. 8. Chiendae, N Thailand limestone 101 fam. 93 92.1 19Њ2ЈN, 98Њ54ЈE 342 gen. 181 52.9 512 spp. 9. Malay peninsula limestone limestone 117 fam. 93 81.6 1–6ЊN, 100–104ЊE 535 gen. 244 51.6 1112 spp. 10. Taiping, Malay peninsula non-limestone 115 fam. 94 82.5 4ЊN, 101ЊE 682 gen. 243 51.6 1939 spp. Notes: The direct comparison of species composition between the different floras is not very significant before the local floras are updated and the taxa revised; therefore, the comparison of floristic similarities between the different floras at the specific level is not made. References: location 2 (Chen, 1985); 3 (Daqinshan Forest station of Guanxi Forestry Bureau, 1980); 4 (Li, 1987); 5 (Li et al., 1986); 6 (Xu, 1984); 7 (Thin, 1997); 8 (Smitinand, 1966); 9 (Chin, 1977, 1979); 10 (Burkill & Henderson, 1925). tropical Asia and southern China were discussed Beard, J. S. 1944. Climax vegetation in tropical America. by Zhu (1997). This limestone flora shares all fam- Ecology 25: 127–158. . 1955. The classification of tropical American ilies and 88% of its genera with the flora of Indo- vegetation types. Ecology 36: 359–412. china (Lecomte, 1907–1951; Aubre´ville et al., Burkill, I. H. & M. R. Henderson. 1925. The flowering 1960–1996), 96% of its families and 68% of its plants of Taiping, in the Malay Peninsula. Gard. Bull. genera with the flora of the Malay Peninsula (Rid- Straits Settlem. 3: 300–459. Champion, H. G. 1936. A preliminary survey of the forest ley, 1967; Keng, 1978), 73% of its woody plant types of India and Burma. Indian Forest Rec., n.s. 1: genera with Burma (Kurz, 1877), and more than 1–286. 97% of its families and more than 80% of its genera Chen, F. P. 1985. Limestone Flora in Longgang, Guangxi, with other tropical floras of south China (including China. Unpublished M.S. Thesis, Zhongshan University. [In Chinese.] Island; see Wu, 1994). The limestone flora Chin, S. C. 1977, 1979. The limestone hill flora of Malaya, of Xishuangbanna demonstrates strong affinity to I, II. Gard. Bull. 30: 165–219; 32: 64–203. other tropical Asian floras. Curtis, J. T. & R. P. McIntosh. 1951. An upland forest continuum in the prairie-forest border region of Wis- consin. Ecology 32: 467–496. Literature Cited Daqinshan Forest station of Guangxi Forestry Bureau. Aubre´ville, A., N. L. Tardieu-Blot & J. E. Vidal (editors). 1980. Plant list of Daqinshan, Guangxi. [Unpublished 1960–1996. Flora du Cambodge, du Laos et du Viet- manuscript, in Chinese.] nam. No. 1–28. Museum National d’Histoire Naturelle, Keng, H. 1978. Orders and Families of Malayan Seed Paris. Plants. Singapore Univ. Press, Singapore. 452 Annals of the Missouri Botanical Garden

Kurz, W. S. 1877. Forest Flora of British Burma 1 & 2. Wang, H., H. Zhu & B. G. Li. 1997. Vegetation on lime- Office of the Superintendent of Government Printing, stone in Xishuangbanna, southwest China. Guihaia 17: Calcutta. [Reprint 1974, International Book Distribu- 101–117. [In Chinese with English abstract.] tors.] Whitmore, T. C. 1984. Tropical Rain Forests of the Far Lecomte, H. (editor). 1907–1951. Flora generale de East, 2nd ed. Clarendon Press, Oxford. L’Indochine. Tome 1–7. Masson et Cie Editeurs, Paris. Wu, C. Y. (editor). 1980. Vegetation of China, pp. 363– Li, B. 1987. Flora of Gulinqing Nature Reserve in south- 397. Science Press, Beijing. [In Chinese.] eastern Yunnan. Unpublished M.S. Thesis, Zhongshan . 1991. The areal-types of Chinese genera of seed University. [In Chinese.] plants. Acta Bot. Yunnan Suppl. IV: 1–139. Li, S. K., S. F. Yuan, L. F. Liu & Z. Z. Chen. 1986. The Wu, T. L. (editor). 1994. A Checklist of Flowering Plants flora of Huapin. In: Reports on the Huapin Forest Area of Islands and Reefs of Hainan and Guangdong Prov- in Guangxi. Shandong Science Press, Jinan. [In Chi- ince. Science Press, Beijing. [In Chinese.] nese.] Xu, Y. C., H. Q. Jiang & F. Quan (editors). 1987. Reports Li, Y. H. (editor). 1996. List of Plants in Xishuangbanna. on the Nature Reserve of Xishuangbanna. Yunnan Sci. Yunnan National Press, Kunming. [In Chinese.] & Techn. Press, Kunming. [In Chinese.] Liang, C. F., J. Y. Liang & L. F. Liu. 1985. A report on Xu, Z. R. 1984. Limestone Flora of Dongyang Mountain the exploration of the flora of Longgang. Guihaia 5(3): in Libo, Guizhou, China. Unpublished M.S. Thesis, 191–209. [In Chinese with English abstract.] Zhongshan University. [In Chinese.] Liu, L. H. 1987. Rain forest. In: C. Y. Wu (editor), Veg- Zhu, H. 1992. Tropical rain forest vegetation in Xish- etation of Yunnan. Science Press, Beijing. [In Chinese.] uangbanna. Chin. Geogr. Sci. 2: 64–73. Liu, L., Y. T. Hu, Y. C. Yang, W. W. Liu & R. X. Guo . 1997. Ecological and biogeographical studies on (editors). 1990. Reports on Land and Economy of Xish- the tropical rain forest of south Yunnan, SW China with uangbanna. Yunnan People’s Press, Kunming. [In Chi- a special reference to its relationship with rain forests nese.] of tropical Asia. J. Biogeogr. 24: 647–662. , H. Ye, G. Zhang & H. Chen. 1994. Floristic anal- , H. Wang, B. G. Li & Z. F. Xu. 1996. A phyto- ysis of the Longhua natural reserve. Acta Bot. Austro- geographical research on forest flora of limestone hills sinica 9: 1–16. [In Chinese with English abstract.] in Xishuangbanna. Guihaia 16: 317–330. [In Chinese Raunkiaer, C. 1934. The Life Forms of Plants and Statis- with English abstract.] tical Plant Geography. Oxford Univ. Press, Oxford. , , & . 1997. Floristic re- Ridley, H. N. 1967. The Flora of Malay Peninsula, I–V. lationships between the limestone flora of Xishuangban- L. Reeve, London. [Reprint.] na and neighboring floras of tropical Asia and south Schimper, A. F. W. 1903. Plant-Geography upon a Phys- China. Acta Bot. Yunn. 19: 357–365. [In Chinese with iological Basis. Oxford Univ. Press, Oxford. English abstract.] Shimizu, T. 1964. Studies on the limestone flora of Japan , H. Wang & B. G. Li. 1998a. The Structure, Spe- and Taiwan. Part II. J. Fac. Textile Sci. Technol. Shinsu cies Composition and Diversity of the Limestone Veg- Univ., A 12: 1–88. etation in Xishuangbanna, SW China. Gard. Bull. Sin- Smitinand, T. 1966. The vegetation of Dao Chiengdao, a gapore 50: 5–33. limestone massive in Chiengmai, north Thailand. Nat. , H. Wang, B. G. Li & Z. F. Xu. 1998b. Research Hist. Bull. Siam Soc. 21(1–2): 93–128. on the tropical seasonal rainforest of Xishuangbanna, Thin, N. N. 1997. The vegetation of Cucphuong national south Yunnan. Guihaia 18: 371–384. [In Chinese with park, Vietnam. Sida 17: 719–759. English abstract.] Volume 90, Number 3 Zhu et al. 453 2003 Southern Yunnan Limestone Flora 7152 (HITBC), (holotype, KUN) (HITBC) of angiosperms are according Tao Guo-da 4177 Pei Sheng-ji 59-10292 (HITBC); (holotype, KUN) Tao Guo-da 13655 (HITBC); is the authoritative source for specific and infraspecific 2804 Li Yan-hui 3771 Li Yan-hui 1255 (holotype, KUN); (holotype, KUN; isotype, HITBC) (HITBC), (holotype, KUN) (holotype, KUN; isotype, HITBC) 3717 3814 386 (HITBC); (holotype, KUN) (isotype, HITBC) (holotype, YCE; isotype, KUN) (holotype, KUN) (holotype, KUN); (holotype, KUN) (holotype, KUN; isotype, HITBC) (HITBC), (holotype, KUN; isotype, HITBC) (HITBC), (holotype, KUN) (HITBC), (holotype, SYS) Li Yan-hui 03587 Li Yan-hui 4634 (SYS); Flora Reipublicae Popularis Sinicae (SYS); (HITBC) Chu Hua 1192 Ye Guang-zheng 9035 Zhang Shun-cheng 24027 Ch1050 Pei Sheng-ji 10347 Li Yan-hui 4215 Pei Sheng-ji 59-10072 Li Yan-hui 4231 Li Yan-hui 4576 Pei Sheng-ji 59-10040 Li Yan-hui 20078 Ch378 Li Yan-hui 33830 Li Yan-hui 3712 Cai C.T. 59-10784 W. T. Wang 10496 H. P. Tsui W.P.Fang&H.Y.Su H. T. Chang & H. Chu Y. L. Chen & P. K. Chou H. H. Hsue H. H. Hsue G. Z. Ye H. Li Y. C. Yang & P. H. Huang C. Y. Wu ex H. W. Li monglunensis pachyphyllum Y. C. Tseng H. W. Li cordiformis Huang Y. T. Chang C. Y. Wu & H. Li var. var. Y. T. Chang var. Appendix 1. Ecological species groups of the limestone vegetation in southern Yunnan. Baphicacanthus multibracteolata Dracaena menglaensis Alangium kurzii lii Dioscorea menglaensis Croton yanhuii Diospyros atrotricha Neolitsea menglaensis Ornithoboea calcicola Lindera nacusua Quercus yiwuensis Pterospermum menglunensis Pterospermum yunnanensis Amoora calcicola mischocarpum Sageretia thea names except for onesto recently the revised Angiosperm Phylogeny and Website in (http://www.mobot.org/MOBOT/research/APweb/). IPNI (www.ipni.org)Seed and Plant TROPICOS Taxa (http://mobot.mobot.org/W3T/Search/vast.html). exclusive The to familyACANTHACEAE limestone names habitats and endemic to southern Yunnan AGAVACEAE ALANGIACEAE DIOSCOREACEAE EUPHORBIACEAE EBENACEAE GESNERIACEAE LAURACEAE FAGACEAE MELIACEAE PIPERACEAE 454 Annals of the Missouri Botanical Garden (HITBC) (HITBC) (HITBC) (HITBC) 34923 (HITBC) (HITBC) Tao Guo-da 1212 2911 3726 (HITBC), (HITBC) (HITBC); (HITBC), Tao-Guo-da 3797 (HITBC), Zhu Hua & Wang Hong 2416 (SYS); (SYS); Expedition 32120 (HITBC) (HITBC) (HITBC) Ch1217 Li Yan-hui 5190 Tao Guo-da 13650 Ch985 3160 Pei Sheng-ji 10045 9008 Li Yan-hui 3300 4610 (HITBC); (HITBC) (SYS) (SYS), (SYS); (SYS) (SYS), (SYS); (holotype, KUN; isotype, HITBC) (HITBC); (HITBC) (HITBC), (HITBC); (HITBC) (HITBC) (HITBC), (HITBC) (HITBC), (HITBC) Ch1219 Ch1175 Ch1186 Ch715 Ch814 Ch373 (SYS), (SYS), (SYS), (SYS) (SYS) (SYS), (SYS) (SYS), (SYS) (SYS) (SYS), (SYS) Tao Guo-da 15933 Ch1193 Ch255 Li Yan-hui 20047 Li Yan-hui 4605 Ch454 Wang Hong 3159 Expedition 32613 Yang Zheng-hong 12178 Wang Hong 1792 Pei Shengji 10034 Ch29 Ch56 Li Yan-hui 1043 Ch870 Li Yan-hui 4225 Tao Guo-da et al.Expedition 16166 10201 Tao Guo-da 9007 Ch584 Ch802 Ch815 Ch386 Ch545 Ch909 (Bremek.) H. Chu Tsiang & P. T. Li C. Y. Wu & T. L. Ming Engl. T. Anders. glomeruliflora Tsiang C. B. Clarke ex W. W. Smith hirsutus Poiss. ex Franch. G. D. Tao pingpienense Pierre ex Gagnep. var. (Craib) C. Y. Wu & T. L. Ming (Roxb.) G. Don Engl. & K. Krause Wall. Finet & Gagnep. Diels ex Ast (Wall. ex Nees) Bremek. var. Roxb. ´v. Schott Merr. Finet & Gagnep. var. Le Dunn Hook. f. & Thoms. Wall. Appendix 1. Continued. Pyrenaria menglaensis Andrographis laxiflora Daedalacanthus tetragonus Pteracanthus alatus Strobilanthes psiostachys Thunbergia coccinea Dracaena cambodiana Dracaena terniflora Pistacia weinmannifolia Toxicodendron fulvum Spondias lakonensis Alphonsea squamosa Miliusa velutina Mitrephora calcarea Polyalthia petelotii Alphonsea boniana Alphonsea mollis Ehretia dunniana Remusatia vivipara Amorphophallus yunnanensis Gonatanthus pumilus Bombax insigne Cosmostigma hainanense Aganosma acuminata Anodendron affine THEACEAE Seed Plant Taxa exclusive toACANTHACEAE limestone habitats but not endemic to southern Yunnan AGAVACEAE ANACARDIACEAE ANNONACEAE BORAGINACEAE ARACEAE BOMBACACEAE APOCYNACEAE Volume 90, Number 3 Zhu et al. 455 2003 Southern Yunnan Limestone Flora (HITBC) Wang Hong 1573 (HITBC); (HITBC) (HITBC) (HITBC) (SYS) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Li Yan-hui 2736 13691 Ch353 Tao Guo-da 13664 2739 2529 Tao Guo-da 13672 (SYS); (HITBG) (SYS) (SYS), (HITBC), (HITBC) (HITBC); (HITBC) (HITBC), (HITBC) (HITBC) (HITBC) (HITBC), (HITBC); Ch1063 Wang Hong 2655 Zhu Pei-zhi 10527 Li Yan-hui 2036 Wang Hong 3147 Ch812 Ch39 Wang Hong 1575 (HITBC) (SYS), (SYS); (SYS) (SYS) (SYS); (SYS) (SYS) (SYS) (SYS); (SYS); (SYS) (SYS), (SYS), (SYS); Tao Guo-da 15739 Ch321 Li Yan-hui 455 YB004125 Li Yan-hui 3780 Ch1015 Ch1170 Ch469 Ch1232 Ch513 Li Yan-hui 4217 Ch845 Ch921 Ch473 Ch791 Pei Sheng-ji 9985 Li Yan-hui 3718 Tong Shao-quan 24960 Ch24 Li Yan-hui 3141 Li Yan-hui 3719 Li Yan-hui 3629 Li Yan-hui 3095 Ch179 Ch563 Ch367 (Gagnep. & Courchet) G. W. Staples (Hu) A. C. Smith A. C. Smith (Miq.) Muell.-Arg. delavayi Gagnep. Prain & Burkill (Pierre) A. C. Smith Wall. Prain & Burkill W. W. Smith W. E. Evans ´v. (Hemsl.) G. W. Staples var. Hook. f. (L.) C. Shih A. C. Smith Roxb. Stapf Prain & Burkill Le Prain & Burkill Roxb. Prain & Burkill Blume Hemsl. Craib & Hutchison C. B. Clarke Appendix 1. Continued. Sarcococca vagans Loeseneriella yunnanensis Pristimera cambodiana Loeseneriella merrilliana Pristimera setulosa Capparis bodinieri Pterocypsela indica Vernonia curtisii Dioscorea aspersa Dioscorea chingii Dioscorea pulverea Dioscorea arachidna Dioscorea tentaculigera Tridynamia sinensis Agapetes mannii Porana paniculata Porana racemosa Tridynamia sinensis Cardamine calcicola Agapetes burmanica Agapetes lobbii Anogeissus acuminata Croton argyratus Cleistanthus sumatranus Phyllanthus clarkei Trigonostemon bonianus BUXACEAE CAPPARIDACEAE COMPOSITAE DIOSCOREACEAE CONVOLVULACEAE CRUCIFERAE COMBRETACEAE EUPHORBIACEAE 456 Annals of the Missouri Botanical Garden (HITBC) (HITBC) (HITBC) (HITBC) Tao Guo-da 15750 (HITBC) Wang Zhong-yu s.n. 4277 842 (HITBC) (HITBC); (HITBC) (HITBC); (HITBC), (HITBC), Tao Guo-da 15902 (HITBC) (HITBC); (HITBC) (HITBC) (HITBC) Li Yan-hui 3759 Tao Guo-da 15798 Li Yan-hui 2802 Pei Sheng-ji 59-9937 3727 Pei Sheng-ji 59-10069 3813 4852 3052 Wang Hong 831 (SYS) (SYS); (SYS); (HITBC); (HITBC); (HITBC) (HITBC) (HITBC) (HITBC), (HITBC); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC), (HITBC), (HITBC), (HITBC); Ch504 Ch559 Pei Sheng-ji 9892 Ch806 (SYS) (SYS) (SYS), (SYS), (SYS); (SYS) (SYS), Li Yan-hui 1411 Ch1097 Ch118 Li Yan-hui 4200 Ch398 Li Yan-hui 4100 Ch394 Ch947 Ch619 Li Yan-hui 20062 Pei Sheng-ji 1054 Ch624 Expedition 32625 Li Yan-hui 4103 Tao Guo-da 11182 Pei Sheng-ji 10138 Cui Jing-yun 14659 Li Bao-gui 9911209 Li Yan-hui 2988 Li Yan-hui 1029 Expedition 34310 Expedition 34533 Li Yan-hui 3586 (Craib) X. Zhang C.Y.Wu&H.W.Li ´v.) B. L. Burtt C. Y. Wu & Hsue Wall. (H. Le Roxb. W. W. S m . Merr. laxiflora (Wall.) Sleum. siamensis H. W. Li Burkill Merr. (L.) Benth. How Gagnep. (Drake) A. M. Schot C. Y. Wu & Y. C. Huang Craib Dum. var. C. Y. Wu ex Y. H. Li D. Don var. Kurz Prain Appendix 1. Continued. Albizia odoratissima Bauhinia carcinophylla Caesalpinia tsoongii Callerya eurybotrya Milettia tetraptera Derris caudatilimba Sophora prazeri Sophora tonkinensis Lysionotus serratus Raphiocarpus begoniifolius Didymocarpus margaritae Ornithoboea henryi Corydalis taliensis Garcinia bracteata Gomphandra tetrandra Coleus xanthanthus Gomphostemma parviflorum Rabdosia eriocalyx Symphorema involucratum Cryptocarya acutifolia Reinwardtia indica Pterospermum chingtungense Pterospermum proteus FABACEAE GESNERIACEAE FUMARIACEAE GUTTIFERAE ICACINACEAE LABIATAE LAURACEAE LINACEAE MALVACEAE Volume 90, Number 3 Zhu et al. 457 2003 Southern Yunnan Limestone Flora (HITBC) 2091 (HITBC) (HITBC), Wang Hong 1960 Wang Hong 2101 (HITBC) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC); 10479 1715 (HITBC), (HITBC), (HITBC) Li Yan-hui 3714 Li Yan-hui 3327 Tao-Guo-da 15729 Tao Guo-da 16440 (HITBC) (SYS); (SYS); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC) (HITBC); Ch850 Wang Hong 1710 Zhu Pei-zhi 10478 Ch192 Li Yan-hui 5230 (SYS), (SYS) (SYS); (SYS); (SYS) (SYS) (SYS), (SYS); (SYS) (SYS) (SYS) Ch798 Ch339 Ch529 Tao Guo-da 39915 Ch399 Ch950 Li Yan-hui 3302 Li Yan-hui 429 Ch901 Tao Guo-da 43898 Tao Guo-da 15672 Li Yan-hui 20032 Ch351 Ch550 Chen San-yang 18959 Tao Guo-da 42370 Ch679 Ch22 Zhu Pei-zhi 8683 Ch158 Li Yan-hui 4150 Tao Guo-da 44107 (C. DC.) Merr. Y. L. Chen & P. K. Chou (Hu & Cheng) C. Y. Wu ex Y. L. Chen (W. W. Smith) Hand.-Mazz. Schltr. Kurz cambodiana H. S. Lo trichoclada Rolfe (Hance) Schltr. var. Lindl. coryanum Gagnep. var. Fletcher kiukiangensis Wall. Lindl. (Pierre) C. Y. Wu Lindl. Hand.-Mazz. H. S. Lo C. Y. Wu var. Miq. Harms var. L. Appendix 1. Continued. Sageretia laxiflora Ventilago calyculata Munronia henryi Aglaia testicularis Amoora tetrapetala Hovenia acerda Lysimachia garrettii Gouania javanica Cyclea sutchuenensis Stephania chingtungensis Myrsine semiserrata Stephania epigaea Peperomia leptostachya Natsiatopsis thunbergiaefolia Ligustrum sinense Bulbophyllum nigrescens Dendrobium salaccense Eulophia bracteosa Bulbophyllum kwangtungense Pholidota chinensis Bulbophyllum ambrosia Caryota urens MELIACEAE PRIMULACEAE RHAMNACEAE MENISPERMACEAE MYRSINACEAE PIPERACEAE OLACACEAE OLEACEAE ORCHIDACEAE PALMAE 458 Annals of the Missouri Botanical Garden (HITBC) Expedition 32673 4211 (HITBC); (HITBC) (HITBC) (HITBC), 4111 (HITBC) (HITBC) (HITBC) (HITBC), (HITBC) (HITBC) Li Yan-hui 391 Tao Guo-da 15722 Zhu Pei-zhi 8649 3952 (SYS); (SYS) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC); Tao Guo-da 44091 (HITBC), Li Yan-hui 1773 Ch941 Li Yan-hui 3751 Ch1089 Zhang Jian-hou 13602 3871 3309 4269 2737 Zhang Jian-hou 13707 (SYS); (SYS) (SYS), (SYS) (SYS); (SYS), (SYS); (HITBC); (HITBC) (HITBC), (HITBC), (HITBC), (HITBC) (HITBC), (HITBC); (HITBC) Ch270 Ch816 Ch710 Ch719 Ch632 Li Yan-hui 4105 Li Yan-hui 363 Ch371 Ch925 (SYS), (SYS), (SYS), (SYS) (SYS), (SYS), (SYS) (SYS); (SYS) (SYS); (SYS), (SYS), (HITBC) Tao Guo-da 13651 Ch368 Ch361 Li Yan-hui 4240 Li Yan-hui 3716 Li Yan-hui 3815 Ch96 Li Yan-hui 4203 Li Yan-hui 393 Ch32 Ch91 Li Yan-hui 3700 Ch358 Li Yan-hui 396 Ch917 Ch572 Ch514 Ch877 Ch879 Ch139 Pei Sheng-ji 10073 How ex W. C. Chen (Planch.) E. Soepadmo longiflorus (Cham. & Schltdl.) Benth. Pierre ex Gagnep. Hayata var. Wittm. W. T. Wang Gaertn. f. Hayata Gagnep. Planch. (Lawson) Planch. Fletcher Huang Bremek. R. Br. philippensis Craib Benth. C. B. Rob. Gagnep. & Guillaumin Wall. var. Appendix 1. Continued. Pavetta polyantha Pavetta scabrifolia Damnacanthus indicus Hymenopogon parasiticus Murraya euchrestifolia Murraya tetramera Lindenbergia philippinensis Brucea mollis Turpinia robusta Celtis wightii Tetrastigma cambodianum Tetrastigma dubium Tetrastigma rupestre Garrettia siamensis herbaceifolium Elatostema salvinioides Laportea urentissima Pellionia scabra Pilea cadierei Procris crenata Pommereschea lackneri RUBIACEAE RUTACEAE SCROPHULARIACEAE SIMARUBACEAE STAPHYLEACEAE ULMACEAE VITACEAE VERBENACEAE URTICACEAE ZINGIBERACEAE Seed Plant Taxa preferent to limestone habitats Volume 90, Number 3 Zhu et al. 459 2003 Southern Yunnan Limestone Flora (HITBC) (HITBC) 74318 (HITBC) (HITBC), (SYS) (HITBC) (HITBC) (HITBC) Li Yan-hui 3721 Ch310 Yang Zheng-hong 10976 (SYS); (SYS), (SYS) (SYS); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Unknown coll. 74317 Ch974 Ch180 Li Yan-hui 67493 Ch646 Pei Sheng-ji 10274 Ch716 Li Yan-hui 2829 (HITBC) (HITBC) (SYS); (SYS) (SYS), (SYS), (SYS); (SYS), (SYS); (SYS) (SYS), (SYS); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Tao Guo-da 3775 Tao Guo-da 9123 Li Jie 722 Ch808 Ch172 Ch1211 Ch1179 Tao Guo-da 15926 Ch420 Ch587 Ch240 Li Yan-hui 2574 Expedition 32621 Ch764 Ch633 Li Yan-hui 1011 Ch247 (SYS); (SYS) (SYS); (SYS); (SYS), (SYS); (SYS), (SYS), (SYS); (SYS) (SYS), (SYS), (SYS); (SYS) (SYS); (SYS); (SYS) (SYS), (SYS) (SYS), (SYS); (SYS), (SYS) Ch376 Li Yan-hui 2898 Ch91 Ch1191 Ch137 Ch36 Ch1182 Ch499 Ch1169 Ch1160 Ch706 Ch265 Zhu Pei-zhi 10364 Ch976 Li Yan-hui 2946 Li Yan-hui 3139 Ch673 Ch544 Ch233 Ch651 Tao Guo-da 35474 Zhang Jian-hou 18562 Ch960 Tao Guo-da 44195 Ch438 Ch658 Ch242 Ch23 Ch729 Expedition 34523 Ch586 (C. B. Clarke) Merr. Radlk. Schott (Hayata) C. F. Hsieh & T. C. Huang (Tsiang & P. T. Li) P. T. Li (R. Ben) C. Y. Wu (Roxb.) Schott Andr. Engl. (Roxb.) Moon (Blume) Lindau (Tsiang & P. T. Li) W. D. Stevens & P. T. Li ´v.) Tsiang Nees Hook. f. & Thoms. Wall. ex Wight (Tsiang) P. T. Li Hemsl. (Le Hook. f. (Blume) Decne. (Retz.) Schult. Pierre Irmsch. Irmsch. (Vahl) Merr. W. T. Wang L. (L. f.) R. Br. Tsiang & P. T. Li Costanin. ´v. Le Appendix 1. Continued. Andrographis laxiflora Goldfussia glomerata Pseuderanthemum palatiferum Pseuderanthemum polyanthum Rhapidospora vagabunda Semnostachya longispicata Miliusa chunii Mitrephora maingayi Mitrephora thorelii augastinei Begonia dryadis Bombax ceiba Begonia prostata Aglaonema pierreanum Colocasia gigantea decursiva Rhaphidophora hongkongensis Antiostelma lantsangensis Bidaria yunnanense Dischidia esquirolii Toxocarpus villosus nervosa Hoya villosa Jasminanthes saxatilis tenacissima Dischidia minor Gymnema latifolium Hoya carnosa Hoya lyi ACANTHACEAE ANNONACEAE BOMBACACEAE ARACEAE APOCYNACEAE 460 Annals of the Missouri Botanical Garden (HITBC) (HITBC) (HITBC) 3792 (HITBC) (HITBC), Pei Sheng-ji 10300 (HITBC) Li Yan-hui 2112 (HITBC); (SYS); (HITBC) (HITBC) (HITBC) Ch195 Unknown coll. 262 Tao Guo-da 16748 Li Yan-hui 1694 (HITBC) (HITBC) (SYS), (SYS); (HITBC); (HITBC) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC) Cui Jing-yun 14656 Tao Guo-da 1742 Ch153 Ch38 3726 Zhu Pei-zhi 10470 Li Yan-hui 1500 (HITBC) (SYS); (SYS); (SYS), (SYS), (HITBC) (SYS); (HITBC) (HITBC), (HITBC); (HITBC) (HITBC) Ch326 Zhao Shi-wang 22582 Ch136 Ch1214 Pei Sheng-ji 9947 Zou Shuang-yun 356 Li Yan-hui 399 Ch226 Li Yan-hui 2423 Wang Hong 2707 Expedition 34359 Wang Hong 2724 Ch97 (SYS), (SYS); (SYS), (SYS), (SYS); (SYS); (SYS); (SYS), (SYS); (SYS) (SYS); (SYS); (SYS) (SYS); (SYS), Li Yan-hui 3360 Ch612 Ch853 Ch516 Expedition 34761 Ch1201 Li Yan-hui 4235 Ch902 Ch807 Li Yan-hui 3575 Li Yan-hui 4250 Ch922 Ch309 Ch131 Ch25 Tao Guo-da 13668 Zhao Shi-wang 22492 Ch784 Ch842 Ch272 Ch948 Ch379 (Merr.) H. S. Ku (Hemsl.) P. Hoffmann (Stapf) C. Y. Wu Hassk. microphyllum (Dunn) Dunn Benth. Pax & K. Hoffm. Kurz pseudoverticellata Rehder & Wils. (Blume) J. J. Smith (Hance) B. L. Burtt var. Pampan. Gagnep. macrophylla Craib var. (Franch.) B. L. Burtt F. Chun D. Y. Hong f. Zoll. & Mor. R. Br. (L.) DC. Rehder I. M. Johnst. Appendix 1. Continued. Ehretia tsangii Capparis urophylla Capparis membranifolia Diospyros yunnanensis Porandra scendens Youngia japonica Viburnum tsangii Antidesma montanum Cleidion bracteosum Lasiococca comberi Sumbaviopsis albicans Adenochlaena silhetensis Cleidion brevipetiolatum Peripterygium quinquelobum Bauhinia genuflexa Millettia yunnanensis Whitfordiodendron filipes Flacourtia rukam Chirita dimidiata Paraboea rufescens Paraboea dictyoneura Paraboea sinensis BORAGINACEAE CAPPARIDACEAE EBENACEAE COMMELINACEAE COMPOSITAE CAPRIFOLIACEAE EUPHORBIACEAE CARDIOPTERIDACEAE FABACEAE FLACOURTIACEAE GESNERIACEAE Volume 90, Number 3 Zhu et al. 461 2003 Southern Yunnan Limestone Flora (HITBC) (HITBC) Wang Hong 2727 (HITBC); Wang Hong 3102 (HITBC) (HITBC) (HITBC) 456 (HITBC); (HITBC) (HITBC), (HITBC) 2940 Li Yan-hui 3867 Li Yan-hui 754 Sun Ji-liang 18005 (SYS) (HITBC) (HITBC) (SYS); (SYS); (HITBC), (HITBC) (HITBC) (HITBC) (HITBC), Ch47 Ch581 Expedition 32617 2922 Yang Zhen-hong 6971 Unknown coll. 2434 Ch324 (HITBC) (SYS), (SYS), (SYS) (SYS); (SYS), (SYS) (HITBC) (HITBC), (HITBC); (HITBC) (HITBC); (KUN) (HITBC) (HITBC) (HITBC) Sun Ji-liang 18151 Ch170 Cui Jing-yun 22658 Ch566 Ch119 Ch61 Ch278 Li Yan-hui 252 Li Yan-hui 3802 Li Yan-hui 1051 (SYS); (SYS), (SYS); (SYS), (SYS), (SYS), (SYS) (SYS), (SYS) (SYS); (SYS); (SYS); Ch1058 Tao Guo-da 16690 Ch51 Tao Guo-da 3798 Wang Hong 2921 Li Yan-hui 3135 Ch41 Expedition 32276 Ch130 Ch45 L. Yan-hui 385 Ch508 Unknown coll. 250 Ch680 Zhu Hua & WangLi Hong Yan-hui 88-11 3223 Zou Shuang-yun 324 Li Yan-hui 3053 Ch269 Ch161 Ch864 Ch177 Ch711 Expedition 34321 (Dop) S. K. Chen King (King) Corner (Franch.) Hand.-Mazz. tonkinensis Sm. velutina Royle ex Kunth C. Y. Wu & H. Li S. S. Chang (Roxb.) Nees var. (Link) C. Y. Wu H. L. Li Pei var. Craib maliformis E. Meyer ex Arn. Fletcher J. L. Ellis ´v. & Vaniot (Wall. ex Nees) Nees P. Y. Bai & P. H. Huang pubescens Le Roxb. Fletcher (Wall. ex Nees) Benth. & Hook. f. var. Hook. f. Wall. ex Miq. var. Appendix 1. Continued. Dendrocalamus strictus Apodytes dimidiata Colebrookea oppositifolia Elsholtzia stachyodes Sphenodesme mollis Clerodendron henryi Premna hamiltonii Garrettia siamensis Ficus pubigera Cudrania jinghongensis Ficus orthoneura Ficus prostata Ardisia garrettii Dysoxylum lenticellatum Dysoxylum spicatum Toona ciliata Chukrasia tabularia Sterculia villosa Phoebe lanceolata Litsea elongata Litsea dilleniifolia Polygonatum punctatum Tupistra wattii Hiptage benghalensis GRAMINEAE ICACINACEAE LABIATAE LAMIACEAE MORACEAE MYRSINACEAE MELIACEAE MALVACEAE LAURACEAE LILIACEAE MALPIGHIACEAE 462 Annals of the Missouri Botanical Garden (HITBC) (HITBC) Li Yan-hui 1859 (HITBC) (SYS); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Unknown coll. 865 Ch313 258 (SYS); (SYS), (HITBC) (SYS) (HITBC) (HITBC) (HITBC) (HITBC), Ch1216 Ch352 Zou Shuang-yun 333 Li Yan-hui 1498 Tao Guo-da 44056 Ch251 Wang Pei-qun 11115 2814 (SYS) (SYS) (SYS), (SYS) (SYS), (SYS); (SYS); (SYS) (SYS) (HITBC) (HITBC) (HITBC) (HITBC); (HITBC), (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC); (HITBC), Ch645 Ch1229 Li Yan-hui 2738 Ch924 Li Yan-hui 3138 Ch492 Ch330 Ch146 Ch165 Zhu Hua & Wang Hong 3004 Ch777 Ch578 Unknown coll. 1010 Li Yan-hui 2473 (SYS), (SYS), (SYS); (SYS), (SYS); (SYS), (SYS), (SYS), (SYS), (SYS); (SYS), (SYS), (SYS); (SYS); Cui jing-yun 22811 Ch1088 Li Yan-hui 1289 Ch804 Ch585 Li Yan-hui 2734 Ch713 Li Yan-hui 392 Zhao Shi-xiang 165 Li Yan-hui 3129 Li Yan-hui 4004 Chen Yu 62079 Ch1218 Li Yan-hui 246 Ch501 Ch176 Ch222 Ch712 Ch355 Ch636 Ch475 Li Yan-hui 3315 Ch887 Expedition 34267 Li Yan-hui 12675 Ch836 (Rehder & E. H. Wilson) C. C. Huang Lo Y. L. Chen & P. K. Chou ferrugineum C. DC. pubescens subimmersa var. Bremek. ´v. (Roxb.) Wall ex A. M. & J. M. Cowan (Merr. & Chun) Swingle f. (Lindl.) Kuntze Drake Miq. Le (L.) Jack. L. Merr. Pierre var. (Willd.) Lindl. (Craib) Hutch. Hance C. C. Huang Hutch. (Lindl.) Blume (L.) Spreng. L. C. DC. Lour. (Andr.) Schltr. Appendix 1. Continued. Ventilago leiocarpa Zizyphus attopensis Peperomia heyneana Piper arboricola Piper curtipedunculatum Ixora cephalophora Pavetta arenosa Pavetta hongkongensis Psychotria pilifera Psychotria siamica Spiradiclis caespitosa Caryota ochlandra Averrhoa carambola Biophytum esquirolii Nervilia plicata Tropidia angulosa Gastrochilus obliquus Syzygium claviflorum Zanthoxylum laetum Zanthoxylum utile Glycosmis citrifolia Murraya koenigii Murraya microphylla Murraya paniculata Zanthoxylum armatum RHAMNACEAE PIPERACEAE RUBIACEAE PALMAE ORCHIDACEAE MYRTACEAE RUTACEAE Volume 90, Number 3 Zhu et al. 463 2003 Southern Yunnan Limestone Flora (HITBC) (HITBC) (HITBC) Tao Guo-da 16090 (HITBC) (HITBC) (HITBC) (HITBC) (HITBC); Tao Guo-da 16049 Wang Hong 1494 (HITBC) (SYS); (HITBC); (HITBC) (HITBC) (HITBC) (HITBC) Tao Guo-da 9189 Zhu Pei-zhi 8642 Ch359 Li Yan-hui 4210 Pei Sheng-ji 10084 Pei Sheng-ji 9359 (SYS); (SYS), (SYS); (SYS); (SYS); (SYS) (SYS) (SYS) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Cui Jing-yun 19416 Wang Hong 3066 Ch388 Wang Hong 1484 Ch657 Tao Guo-da 41641 Ch932 Li Yan-hui 11915 Ch714 Ch811 Ch1054 Ch616 Tao Guo-da 16447 Ch408 (SYS); (SYS); (SYS), (SYS); (SYS); (SYS); (SYS), (SYS) (SYS); (SYS), (SYS); (SYS), (SYS), (SYS), (SYS), Ch168 Ch984 Ch381 Ch1064 Ch311 Li Yan-hui 1696 Zhu Hua 93011 Li Yan-hui 8541 Ch700 Ch1240 Li Yan-hui 4866 Tao Guo-da 13661 Expedition 34430 Li Yan-hui 2554 Li Bao-gui 734 Zhu Pei-zhi 8668 Li Bao-gui 803 Ch844 Ch268 Cui Jing-yun 22793 Ch468 Ch507 Ch463 Ch653 Ch668 Ch654 Gagnep. (Hook.) Bedd. Wall. ex Baker monospermum L. (King & Prain) K. Schum. edgeworthii tenuiflorum Wedd. Stapf Merr. Blume J. Ghatak var. Merr. & Chun W. C. Cheng & S. D. Chu Gagnep. (Metcalf) C. L. Li L. var. (Blume) Merr. R. Br. var. (Buch.-Ham.) Wedd. (Siebold & Zucc.) Wedd. (Wall.) Craib Planch. C. H. Wright Hand.-Mazz. Diels & Gilg (Gagnep.) Burret L. Appendix 1. Continued. Elatostema rupestre Pilea monilifera Debregeasia edulis Pilea plataniflora Pouzolzia sanguinea Boehmeria zollingeriana Colona floribunda Colona thorelii Tapiscia yunnanensis Tetrameles nudiflora Adiantum capillus-veneris Antrophyum callifolium Pommereschea spectabilis Adiantum caudatum Adiantum malesianum Adiantum caudatum Tetrastigma delavayi Yua austro-orientalis Tetrastigma cauliflorum Leea aequata Tetrastigma pubinerve Vitis balansaeana Hedychium sino-aureum Tetrastigma erubescens Vitis betulifolia Hedychium villosum URTICACEAE TILIACEAE STAPHYLEACEAE TETRAMELACEAE Pterydophyta preferent to limestone habitats ADIANTACEAE ANTROPHYACEAE VITACEAE ZINGIBERACEAE 464 Annals of the Missouri Botanical Garden (HITBC) Unknown coll. 9498 (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC); (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Li Yan-hui 3810 Li Bao-gui 737 Li Bao-gui 880 Li Yan-hui 3739 Li Yan-hui 2749 Tao Guo-da 43453 Li Qing-jun 42749 Zhu Pei-zhi 10487 Li Bao-gui 674 Tao Guo-da 43571 Tao Guo-da 17161 Li Bao-gui 756 Li Yan-hui 2735 Tao Guo-da 38772 Expedition 34292 Li Bao-gui 98085 Tao Guo-da 15730 Li Bao-gui 45749 Tao Guo-da 16010 Tao Guo-da 39738 Li Bao-gui 774 Li Bao-gui 600 Li Bao-gui 604 Li Yan-hui 3808 Expedition 34719 Tao Guo-da 38843 Tao Guo-da 38875 Li Qing-jun 42624 Li Bao-gui 9904152 Pei Sheng-ji 9933 (Hook.) Ching C. Presl (Kunze ex Mett.) Tagawa (L.) Link (H. Christ) Ching (Wall.) C. Presl Ching (Bedd.) Ching (Sw.) Ching (Bedd). Pic. Serm. Hook. (L.) Trimen (Hook.) Ching (D. Don) J. Sm. H. Christ (Baker) Diels C. Chr. ex Ching (D. Don) C. Chr. (Giesenh.) Ching Rosenst. C. Presl Wall. ex Hook. & Grev. (Sw.) Bedd. (Baker) Ching (Kunze ex Mett.) J. Sm. (Christ) Ching (Takeda) Ching (Kunze) Ching H. Christ (Giesenh.) Ching Appendix 1. Continued. Asplenium austrochinense Asplenium excisum Asplenium interjectum Asplenium prolongatum Neottopteris antrophyoides Asplenium saxicola Asplenium varians Hymenasplenium cheilosorum Allantodia pinnatifido-pinnata Allantodia alata Kuniwatsukia cuspidata Egenolfia tonkinensis Drynaria bonii Drynaria fortunei Phymatodes cuspidata Dryopteris cochlaeta Pityrogramma calomelanos Nephrolepis auriculata bicolor Lepidogrammitis rostrata carnosum Drynaria rigidula Polystichum deltodon Lemmaphyllum microphyllum Pyrrosia subfurfuracae Pyrrosia tonkinensis Pyrrosia assimilis Pyrrosia mollis Pyrrosia nuda Pyrrosia nummularifolia ASPLENIACEAE ATHYRIACEAE BOLBITIDACEAE DRYNARIACEAE HEMONITIDACEAE NEPHROLEPIDACEAE Volume 90, Number 3 Zhu et al. 465 2003 Southern Yunnan Limestone Flora (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) (HITBC) Li Bao-gui 921 Li Bao-gui 960463 Li Bao-gui 45178 Mengliandui 10242 Mengliandui 10140 Li Qing-jun 42687 Pei Sheng-ji 9277 Sun Ji-liang 18185 Li Bao-gui 98086 Li Yan-hui 1326 Ching & S. K. Wu H. Christ (Burn. f.) Trev. (Hook. & Grev.) Maxim. (L.) Tardieu (Sw.) Spring (Bedd.) Ching & C. H. Wang (C. Presl) Copel. (Kunze) Ching & C. H. Wang (Hope) Ching Appendix 1. Continued. Selaginella involvens Selaginella pulvinata Pteris actiniopteroides Aleuritopteris pseudofarinosa Ctenitopsis devexa Tectaria decurrens Cyclosorus papilio Cheilosoria tenuifolia Tectaria simaoensis Cyclosorus parasitica SELAGINELLACEAE PTERIDACEAE SINOPTERIDACEAE THELYPTERIDACEAE