BIOGEOGRAPHY AND THE LIMESTONE FLORA IN SOUTHERN …flora.huh.harvard.edu/china//novon/mobt-90-03-444.pdf · as the regional tropical seasonal rain forest off lime-stone, shares characteristics

ANN. MISSOURI BOT. GARD. 90: 444–465. 2003.

BIOGEOGRAPHY ANDFLORISTIC AFFINITIES OFTHE LIMESTONE FLORA INSOUTHERN YUNNAN, CHINA1

H. Zhu,2 H. Wang,2 B. Li,2 andP. Sirirugsa3

ABSTRACT

The forests on limestone in southern Yunnan, in tropical southwest China, were inventoried, and their floristiccomposition and biogeographical affinities are discussed. These limestone forests were characterized by phanerophytesmaking up ca. 78% of the total species and those with mesophyllous leaves comprising 75%. Ecological species groupsbased on their habitat preferences were discerned from field observations: the species exclusive to the limestone habitatsmake up 10% and the preferents make up ca. 12% of the total limestone flora. From these limestone forests, 1394vascular plant species belonging to 640 genera and 153 families were recorded. Based on their distributions, 12biogeographic elements at the generic level and nine at the specific level were recognized. About 90% of the seedplant 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 Asia, thelimestone flora of southern Yunnan revealed closer affinity to tropical floras than to temperate elements of eastern Asianfloras. This limestone flora is thus tropical and part of the tropical Asian flora at its northern margin.

Key words: biogeography, China, limestone forest, southern Yunnan.

Limestone in tropical China occurs mainly inYunnan and Guangxi Provinces of southern andcentral China. Because of the great diversity ofedaphic conditions and topography, vegetationtypes on limestone are extremely diverse and richin endemic taxa. Limestone vegetation in southernChina has been destroyed as much as other vege-tation types even though these limestone areas aremore difficult to access and to farm. Limestone veg-etation is also more vulnerable because it recoversmuch more slowly on usually thin soils. Our re-search was conducted mainly in the area of Xish-uangbanna, in the southern part of Yunnan, whereabout 19% (3600 km2) of the total area is limestone(Liu et al., 1990). Most of this limestone area isstill forested and is receiving increasing attentionfor its biodiversity and its urgent need of conser-vation. Primary floristic works in southern Yunnanhave been written (Zhu et al., 1996, 1997, 1998a,1998b; Wang et al., 1997). This paper represents asynthesis of its floristics, physiognomy, and biogeo-graphical affinities.

1 This project was funded by The National Natural Science Foundation of China (40271048), the Chinese Academyof Sciences (The Fund for Top One Hundred Young Scientists and KSCX2-1-06B), and the Yunnan Natural ScienceFoundation. 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 preparingthis paper during his visiting scholar’s year at the University of Cambridge. Finally, he thanks T. C. Whitmore, whohas greatly supported and helped him in his research, and two anonymous reviewers for their constructive commentson 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, Thailand.

GENERAL GEOGRAPHY

Xishuangbanna, the southern part of Yunnan,which borders Burma and Laos, is a mountainousarea at the northern margin of tropical SoutheastAsia (Fig. 1). Basically, the study area has a moun-tainous topography with the mountains runningnorth-south and decreasing in elevation southward.Altitude varies from 480 m in the lowest valley inthe south to 2400 m at the top of the highest moun-tain in the north. The limestone strata occur mainlyin southeastern Xishuangbanna and range in alti-tude from 600 to 1600 m.

The region of Xishuangbanna has a typical trop-ical monsoon climate with an annual mean tem-perature of 22ºC, annual temperature accumulation(the sum of daily temperature means where they are. 108C) of 80008C, and annual precipitation vary-ing from 1200 to 1556 mm, of which more than80% falls during the rainy season between May andthe end of October (Xu et al., 1987).

The rock substrate is hard limestone of Permianorigin with a rugged topography. The soil is mainly

Volume 90, Number 32003

445Zhu et al.Southern Yunnan Limestone Flora

Fig

ure

1.L

ocat

ions

ofth

ere

sear

char

eain

Xis

huan

gban

na,

sout

hern

Yun

nan,

Chi

na,

and

com

pare

dre

gion

alflo

ras

(see

Tabl

e6)

.—

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urre

sear

char

ea.

—2.

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ggan

,SW

Chi

na.

—3.

Daq

insh

anM

ount

ains

,SW

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na.

—4.

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inqi

ng,

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hina

.—

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n,C

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.—

6.D

ongy

ang

Mou

ntai

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na.

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phuo

ng,

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ietn

am.

—8.

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engd

ao,N

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ilan

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heM

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.—

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ing,

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ayP

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.

446 Annals of theMissouri Botanical Garden

brown, coarse in texture, and composed of loamylimestone with a pH of ca. 6.75 and ca. 3.56%organic matter (Liu et al., 1990).

METHODS

A complete floristic inventory was made basedon the identification of more than 5000 plant spec-imens collected from the limestone habitat insouthern Yunnan during 1985–1995 and depositedmainly at HITBC and SYS. The flora of the vege-tation on the limestone consisted of 153 families ofvascular plants, including 640 genera and 1394species. An initial floristic analysis was made basedon the inventory (Zhu et al., 1996). Three mainvegetation types occur on the limestone—tropicalseasonal rain forest, tropical seasonal moist forest,and tropical montane dwarf forest—which were se-lected for establishing plots. For the tropical sea-sonal rain forest, seven separate plots ranging insize from 2000 to 2500 m2 were established. Forthe tropical seasonal moist forest seven separateplots ranging in size from 500 to 2000 m2 were laidout. For the tropical montane dwarf forest, only twoplots of 10 by 10 m were made due to its restrictionto limestone summits. These different plot sizeswere used because of the differential coverage offorest type and site restrictions. The structure andspecies composition of the vegetation on the lime-stone were analyzed based on plot data alreadypublished (Zhu et al., 1998a). In the present paper,plant inventory lists of the two main forest types(excluding montane dwarf forest) were compiledfrom sample plots separately for the physiognomic(life form and leaf size) analysis. The criteria forlife form and leaf size classes suggested by Raun-kiaer (1934) and the importance value index (IVI)suggested by Curtis and McIntosh (1951) were usedin the physiognomic or ecological analysis. Ecolog-ical species groups were discriminated from fieldobservation and correspond to groups used in Shi-mizu (1964) and Chin (1977). Species-level bio-geographical affinities were assessed for the totalflora of the limestone vegetation. The floristic sim-ilarities between the limestone flora of southernYunnan and the floras on limestone and non-lime-stone habitats from southwest China, northern Vi-etnam, northern Thailand, and the Malay Peninsulawere also discussed.

CLASSIFICATION OF LIMESTONE VEGETATION

Based on plant physiognomy, forest profile, flo-ristic composition, and habitat, the primary lime-stone vegetation can be classified into three vege-tation types, i.e., tropical seasonal rain forest,

tropical seasonal moist forest, and tropical montanedwarf forest (Zhu et al., 1998a). Within these, sixformations, including nine communities, were rec-ognized:

(1) Ravine seasonal rain forest (including the Po-metia tomentosa–Alphonsea monogyna communityand Pometia tomentosa–Celtis philippensis var.wightii community);(2) Lower hill seasonal rain forest (including onlythe Celtis philippensis var. wightii–Lasiococca com-beri var. pseudoverticillata community);(3) Evergreen moist forest (including the Osmanthuspolyneurus–Dracaena cochinchinensis communityand Lasiococca comberi var. pseudoverticillata–Cleis-tanthus sumatranus community);(4) Semi-evergreen moist forest (including theBombax insignis–Colona floribunda community andBombax insignis–Garcinia bracteata community);(5) Evergreen dwarf forest (including only the Pho-tinia angusta–Pistacia weinmannifolia community);(6) Semi-evergreen dwarf forest (including only theFicus neriifolia–Dracaena cochinchinensis commu-nity).

Detailed descriptions and ecological analyses ofthe communities have been reported earlier (Zhu etal., 1998a). Here the classification of the limestonevegetation is concisely enumerated so that the bio-geographical components of the limestone vegeta-tion can be better understood.

TROPICAL SEASONAL RAIN FOREST

Tropical seasonal rain forest on limestone, justas the regional tropical seasonal rain forest off lime-stone, shares characteristics with the equatoriallowland rain forest. These forests are mainly ever-green, but there are some deciduous trees in theemergent layer. This is equivalent to the tropicalsemi-evergreen rain forest of Southeast Asia (Whit-more, 1984), or the tropical semi-evergreen forestof India–Burma (Champion, 1936), as well as theevergreen seasonal forest of tropical America(Beard, 1944, 1955). In southern Yunnan, theselimestone forests occur in wet valleys and on lowerslopes of hills or mountains below 1000 m altitude.This same forest type also occurs in northern Thai-land (Smitinand, 1966) and North Vietnam (Thin,1997), although different names were used. Thetropical seasonal rain forest represents SoutheastAsian tropical rain forest at its latitudinal and al-titudinal limits. The ecological structure of the trop-ical seasonal rain forest on limestone is almost ex-actly the same as the seasonal rain forest offlimestone in the Xishuangbanna region (Zhu, 1992,



Tabl

e1.

Lif

efo

rms

ofth

eli

mes

tone

fore

stin

sout

hern

Yun

nan.

Lif

efo

rm*

Par

aE

piph

Lia

na

Woo

dyH

erb

Pha

nero

phyt

es

Meg

aph

Mes

oph

Mic

roph

Nan

oph

Hph

All

Cha

mG

eoph

The

ro-

phyt

es

Lim

esto

nese

ason

alra

info

rest

(148

00m

2of

7pl

ots,

tota

l24

9sp

ecie

s)

Num

ber

ofsp

ecie

sP

erce

ntag

eof

tota

lsp

ecie

s1 0.

4%7 2.

8%48 19

.3%

3 1.2%

9 3.6%

84 33.7

%34 13

.7%

17 6.8%

12 4.8%

156 62

.7%

31 12.4

%3 1.

2%— —

Lim

esto

nese

ason

alm

oist

fore

st(9

650

m2

of7

plot

s,to

tal

211

spec

ies)

Num

ber

ofsp

ecie

sP

erce

ntag

eof

tota

lsp

ecie

s— —

18 8.5%

27 12.8

%5 2.

3%2 0.

9%62 29

%36 17

%21 9.

9%3 1.4%

124 58

.8%

28 13%

7 3.3%

2 0.9%

*L

ife

form

(Rau

nkia

er,1

934)

;Meg

aph

5M

egap

hane

roph

yte

(per

enni

als

over

30m

high

);M

esop

h5

Mes

opha

nero

phyt

e(p

eren

nial

s8

to30

mhi

gh);

Mic

roph

5M

icro

phan

erop

hyte

(per

enni

als

2to

8m

high

);N

anop

h5

Nan

opha

nero

phyt

e(p

eren

nial

s0.

25to

2m

high

);H

ph5

Her

bace

ous

phan

erop

hyte

(her

bace

ous

pere

nnia

lsov

er0.

25m

high

);C

ham

5C

ham

aeph

ytes

(per

enni

als

less

than

0.25

mhi

ghab

ove

grou

nd);

Geo

ph5

Geo

phyt

e(p

eren

nial

s,dy

ing

back

abov

egr

ound

);P

ara

5P

aras

itic

;E

piph

5E

piph

ytes

;T

hero

phyt

es(a

nnua

ls).

1997). Most species in the seasonal rain forest onlimestone are also found in the adjacent non-lime-stone seasonal rain forest, but the latter is morediverse with additional species, which are notfound on the limestone.

TROPICAL SEASONAL MOIST FOREST

Tropical seasonal moist forest occurs on the mid-dle and upper limestone slopes ranging from 650to 1300 m altitude. This vegetation type abuts theseasonal rain forest and was called monsoon forestby some Chinese authors (Liu, 1987; Wu, 1980).The term seasonal moist forest is preferred herebecause the forest is not equivalent to Schimper’smonsoon forest (Schimper, 1903), in spite of thefact that it is affected by seasonal dryness and con-tains a variable percentage of deciduous trees. Theseasonal dryness in the region is compensated tosome extent by dense fog accompanied by low tem-peratures in the same months (November to April)(Whitmore, 1984). Some deciduous trees, such asGmelina arborea Roxb., Anthocephalus chinensis(Lam.) Rich. ex Walp., and Homalium laoticumGagn. var. glabretum C. Y. Wu, shed leaves towardthe end of the dry season, while others, such asCratoxylon cochinchinensis (Lour.) Bl., Ficus reli-giosa L., and Elaeocarpus varunua Buch.-Ham. exMast., shed their old leaves as new ones develop.This suggests that deciduousness in the region ismore frequently associated with locally dry habitatsthan the seasonal dryness of climate. Therefore, us-ing the term monsoon forest for the evergreen orsemi-evergreen forest on limestone is confusing be-cause Schimper’s monsoon forest is more or lesscompletely leafless during the dry season.

MONTANE DWARF FOREST

Montane dwarf forest occurs only on the tops ofhills and summits of mountains at altitudes above900 m. There is only one dwarf tree layer with acanopy height of 7–15 m. Epiphytic orchids, suchas Eria hainanensis Rolfe and Bulbophyllum ni-grescens Rolfe, and non-vascular epiphytes (bryo-phytes and lichens) are abundant. In some sitessmall woody climbers, such as Derris caudatilimbaHow (Papilionaceae) and Pristimera arborea(Roxb.) A. C. Smith (Hippocrateaceae), are also fre-quent.

PLANT PHYSIOGNOMY OR ATTRIBUTES

From plot data, life form spectra (Raunkiaer,1934) of the two main forest types (seasonal rainforest and seasonal moist forest) are compiled in


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

Forest type

Leaf form

S C

Leaf texture

P L

Leaf size

Na Mi Me Ma

Limestone seasonalrain forest1

Percentage of speciesPercentage of Importance Value Index (IVI)3

72.376.3

27.723.7

47.952.8

52.147.2

00

13.83.8

76.691

9.65.3

Limestone seasonalmoist forest2

Percentage of speciesPercentage of Importance Value Index (IVI)

6874.8

3225.2

51.541.7

48.558.3

10.4

21.523.3

74.266.5

3.19.7

1 From 14800 m2 of 7 plots, total of 94 tree species . 5 cm DBH.2 From 9650 m2 of 7 plots, total of 97 tree species . 5 cm DBH.3 IVI 5 Relative dominant density 1 Relative frequency 1 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 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 Yunnannot endemic to southern Yunnan

Plants dominant on limestonePlants no restriction on limestonePlants found occasionally on limestone

24117170858225

1.78.4

12.261.616.1

Total 1394 100

Table 1. Leaf size spectra, leaf form, and leaf tex-ture are shown in Table 2. Both forest types weredominated by phanerophytes. Including lianas,these perennial phanerophytes make up 73.9–83.2% of the total species, while annual chamae-phytes account for only 12.4–13%. However, theseasonal moist forest shows lower percentages ofwoody lianas as well as megaphanerophytes andmesophanerophytes, but higher percentages of epi-phytes as well as microphanerophytes and nano-phanerophytes than the seasonal rain forest.

Both forest types have species with mesophyllousleaves making up ca. 75% of the total tree species,but the forests show clear differences if the speciesare weighted by importance value index (IVI). Thisincreases the percentage of mesophyllous perenni-als and decreases the percentage of micro- and ma-crophyllous trees in seasonal rain forest, while theopposite trend is seen in seasonal moist forest. Sea-sonal moist forest occupies much more rugged hab-itats with thinner and drier soils, and has more mi-crophyllous species. In weighting by IVI, theincrease in percentage of macrophyllous trees inseasonal moist forest is mainly due to the dominantevergreen species Dracaena cochinchinensis (Lour.)S. C. Chen (Agavaceae), with its long leathery lan-ceolate leaves, and the dominant deciduous treespecies Colona floribunda (Wall. ex Voigt) Craib(Tiliaceae) also with large leaves to 30 cm long. In

weighting the species by IVI, the percentage ofleathery leaves decreases in seasonal rain forest butincreases in seasonal moist forest due to the pres-ence of some species with these leathery leavessuch as Cleistanthus sumatranus (Miq.) Muell.-Arg.(Euphorbiaceae) and Dracaena cochinchinensis.

ECOLOGICAL SPECIES GROUP

Based on the study of the limestone floras of Ja-pan and Taiwan, Shimizu (1964) divided limestoneplants into five ecological groups:

(1) plants exclusive to limestone habitat;(2) plants selective for and found mainly in lime-

stone;(3) plants preferring and dominant on limestone;(4) taxa indifferent, with no special association

with limestone;(5) plants found only occasionally on limestone or

strangers to limestone.

To Shimizu, these first three groups were char-acteristic species for the limestone habitats and inparticular his exclusive and selective taxa were cal-cicoles. Chin (1977) accepted this classificationand similarly categorized plants on limestone in theMalay Peninsula into four groups, combining selec-tive and preferent plants. Similar ecological speciesgroups have been later recognized by Chinese bot-anists (Liang et al., 1985; Liu et al., 1994).



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

No. ofgenera

No. ofspecies %*

No. ofgenera

No. ofspecies %

OrchidaceaeRubiaceaeEuphorbiaceaePapilionaceaeMoraceaeVitaceaeAcanthaceaeRutaceaeAsclepiadaceaeUrticaceaeLauraceaeApocynaceaeMeliaceaeAnnonaceaeCucurbitaceaeRhamnaceaePiperaceae

3534272277

26111612101912129

193

8658585548383635353535333030242120

26.964.160.443.473.879.565.471.453.053.045.058.983.050.952.370.454.8

VerbenaceaeLabiataeGesneriaceaeSterculiaceaeDioscoriaceaeMenispermaceaeLiliaceaeAraceaeCompositaeMyrsinaceaeCommelinaceaeZingiberaceaeMyrtaceaeTiliaceaeMimosaceaeAnacardiaceaeConvolvulaceaeUlmaceae

6131371

1010108476136745

202019181817161515141414131212111111

43.530.163.346.864.360.764.044.114.638.260.942.445.857.166.664.737.4

100

*the no. of species on limestone

% 5 3 100the total no. of species in southern Yunnan

Following Shimizu and Chin’s classifications, wedivided the limestone flora of southern Yunnan intothese four ecological species groups (Table 3). Inour study, 141 vascular plant species are restrictedto limestone habitats and thus are exclusively foundhere. These include the following common speciesCeltis philippensis var. wightii, Amoora calcicola,Murraya tetramera, Pistacia weinmannifolia, aswell as species in Agapetes, Sageretia, Tupistra, andPristimera. Of these, 24 species are endemic tosouthern Yunnan. Taxa exclusive to limestone makeup about 10% of the total limestone flora, whichagrees with the results from Longgan limestone (ex-clusive taxa, 13%) (Liang et al., 1985) and Longhualimestone (exclusive taxa, 10%) (Liu et al., 1994)from Guangxi Province in China. Both the exclu-sive and preferent taxa make up 22.3% of the totalsum. They could be termed as characteristic spe-cies for limestone habitats (see Appendix 1). Thisis similar to the results from Longgan in Guangxi(with these characteristic species making up 20%of the total sum) (Liang et al., 1985) and from theMalay Peninsula (27.5%) (Chin, 1977).

THE FLORA AND ITS BIOGEOGRAPHY

In the limestone forests of southern Yunnan, Chi-na, 153 families of vascular plants including 640genera and 1394 species and varieties, were re-corded, of which seed plants compose 129 families,558 genera, and 1269 species (see Appendix 1).

More than 80% of the species also occur in thenon-limestone habitats of the Xishuangbanna re-gion.

The limestone flora makes up about one quarterof the total species of the regional flora. (The floraof the Xishuangbanna region was primarily docu-mented with 3336 native species of 1218 generaand 207 families of seed plants; see Li, 1996.)Some families show relative preference for lime-stone habitats (with more than 60% of the totalnumber of species in the region on limestone), forexample, Acanthaceae, Euphorbiaceae, Gesneri-aceae, Meliaceae, Menispermaceae, Moraceae,Rhamnaceae, Rubiaceae, Rutaceae, Vitaceae, andUlmaceae (Table 4). Other families, such as Hip-pocrateaceae, Icacinaceae, and Vacciniaceae, showan even stronger preference for limestone (foundalmost exclusively in limestone habitats in southernYunnan), although they are not among the predom-inant families in species richness.

The distribution types of Chinese seed plants atthe generic level were documented by Wu (1991).Based on Wu’s document, 544 of the 558 genera ofseed plants from the limestone forest of southernYunnan can be divided into 12 distribution typesor geographic elements (14 genera, which are cos-mopolitan in distribution, are not included in thegeographic statistics). One thousand two hundredforty-four of the 1269 species of seed plants fromthe limestone forest can be recognized in nine dis-


Tabl

e5.

Geo

grap

hic

affin

itie

sof

the

lim

esto

nefo

rest

sof

sout

hern

Yun

nan.

Geo

grap

hic

elem

ent

atth

ege

neri

cle

vel

(see

Wu,

1991

)P

erce

ntag

eof

gene

raG

eogr

aphi

cel

emen

tsat

the

spec

ific

leve

lP

erce

ntag

eof

spec

ies

1.P

antr

opic

2.Tr

opic

alA

sia–

Trop

ical

Am

eric

adi

sjun

ct3.

Old

Wor

ldTr

opic

s4.

Trop

ical

Asi

ato

Trop

ical

Aus

tral

ia5.

Trop

ical

Asi

ato

Trop

ical

Afr

ica

6.Tr

opic

alA

sia

7.N

orth

ern

Tem

pera

te8.

Tem

pera

teE

aste

rnA

sia

and

Nor

ther

nA

mer

ica

disj

unct

9.O

ldW

orld

Tem

pera

te10

.Te

mpe

rate

Med

iter

rane

an,

Wes

tern

Asi

ato

Cen

tral

Asi

a11

.E

aste

rnA

sia

12.

End

emic

toC

hina

21.1

%2.

9%13

.8%

9.0%

7.9%

35.3

%2.

8%2.

8%0.

9%0.

2%2.

9%0.

4%

1.P

antr

opic

2.Tr

opic

alA

sia–

Trop

ical

Am

eric

adi

sjun

ct3.

Old

Wor

ldTr

opic

s4.

Trop

ical

Asi

ato

Trop

ical

Aus

tral

ia5.

Trop

ical

Asi

ato

Trop

ical

Afr

ica

6.Tr

opic

alA

sia

6a.

Indi

a-M

alay

sia

6b.

Mai

nlan

dSo

uthe

aste

rnA

sia

toM

alay

sia

6c.

Sout

hern

Asi

ato

Mai

nlan

dSo

uthe

aste

rnA

sia

6d.

Mai

nlan

dSo

uthe

aste

rnA

sia

toSo

uthe

rnC

hina

7.E

aste

rnA

sia

8.So

uthe

rnC

hina

9.E

ndem

icto

Yun

nan

0.6%

0.2%

0.4%

3.2%

1.3%

(64.

5%)

17.0

%7.

3%19

.9%

20.3

%0.

5%10

.6%

18.6

%

Tota

lof

544

gene

ra10

0%To

tal

of12

44sp

ecie

s10

0%

tribution types based on their geographic distribu-tion (25 species of seed plants are not included dueto insufficient distribution references) (Table 5). Atthe generic level, the geographic elements of trop-ical distribution (1–6, Table 5) compose 90% of thetotal genera; the geographic elements of temperatedistribution (7–10, Table 5) make up only 6.7%. Atthe specific level, the species that are of typicaltropical distribution (1–6, Table 5) account for70.2% 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 fromlimestone forests in Xishuangbanna. If the speciesfrom the tropical areas adjacent to Xishuangbannafrom 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 tropicalmargin.

In a floristic comparison with nine similar floras,both limestone and non-limestone, from southwestChina, northern Vietnam (Thin, 1997), northernThailand (Smitinand, 1966), and the Malay Pen-insula (Chin, 1977, 1979; Burkill & Henderson,1925) (Table 6), the limestone flora of southernYunnan displays explicit taxonomic affinities to thetropical floras and shows a closer affinity to thefloras from the Malay Peninsula than to other florasfrom subtropical China (the floras of Huapin andDongyang, see Li et al., 1986; Xu, 1984), eventhough these Malaysian floras lie farther away geo-graphically from southern Yunnan. Our limestoneflora in southern Yunnan shares the most generawith the limestone flora of northern Vietnam (Cuc-phuong, see Thin, 1997) among those floras com-pared in this study. The similarity at the genericlevel between our limestone flora and the limestoneflora of northern Thailand (Doi Chiengdao) (Smitin-and, 1966) is less than would be expected from itsgeographic proximity. This lack of correspondencecould be because the plant list for Doi Chiengdaoused here for comparison is an incomplete one con-sisting of only 512 species, less than half reportedfor most other sites in Table 6. The limestone floraof Xishuangbanna did not show a higher floristicsimilarity to other regional limestone floras than tonon-limestone floras in our comparison. It appearsthat limestone floras develop from local or regionalfloras, supported also by the fact that only about10% of the total species of limestone floras (theexclusive group) are restricted to limestone habitatsin our study.

The floristic relationships between our limestoneflora in southern Yunnan and neighboring floras of



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.

Location HabitatSize of flora(Seed plants)

Shared taxaby both floras

Similaritycoefficients

2. Longgan, SW China22814–339N, 1068469E

limestone 149 families669 genera

1363 species

118371

91.266.5

3. Daqinshan Mountains, SW China228149N, 1078E

non-limestone 182 fam.871 gen.

1813 spp.

126389

97.469.8

4. Gulinqing, SW China228369N, 1048E

limestone 143 fam.496 gen.

1095 spp.

116261

89.952.6

5. Huapin, SW China25831–399N, 1098509E


1051 spp.

83150

72.833.2

6. Dongyang Mountains, SW China258149N, 1078569E

limestone 116 fam.367 gen.736 spp.

86153

73.741.6

7. Cucphuong, N Vietnam20814–249N, 105824–44 9E


1661 spp.

120428

9376.7

8. Chiendae, N Thailand19829N, 988549E

limestone 101 fam.342 gen.512 spp.

93181

92.152.9

9. Malay peninsula limestone1–68N, 100–1048E


1112 spp.

93244

81.651.6

10. Taiping, Malay peninsula48N, 1018E


1939 spp.

94243

82.551.6

Notes: The direct comparison of species composition between the different floras is not very significant before thelocal floras are updated and the taxa revised; therefore, the comparison of floristic similarities between the differentfloras 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 discussedby Zhu (1997). This limestone flora shares all fam-ilies and 88% of its genera with the flora of Indo-china (Lecomte, 1907–1951; Aubreville et al.,1960–1996), 96% of its families and 68% of itsgenera with the flora of the Malay Peninsula (Rid-ley, 1967; Keng, 1978), 73% of its woody plantgenera with Burma (Kurz, 1877), and more than97% of its families and more than 80% of its generawith other tropical floras of south China (includingHainan Island; see Wu, 1994). The limestone floraof Xishuangbanna demonstrates strong affinity toother tropical Asian floras.

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sine

nsis

(Hem

sl.)

G.

W.

Stap

les

Trid

ynam

iasi

nens

isva

r.de

lava

yi(G

agne

p.&

Cou

rche

t)G

.W

.St

aple

s

Ch1

170

(SY

S);

Wan

gH

ong

2655

(HIT

BC

)C

h469

(SY

S);

Zhu

Pei

-zhi

1052

7(H

ITB

C)

Ch1

232

(SY

S)C

h513

(SY

S)

CR

UC

IFE

RA

EC

arda

min

eca

lcic

ola

W.

W.

Smit

hLi

Yan-

hui

4217

(HIT

BC

)

DIO

SCO

RE

AC

EA

ED

iosc

orea

arac

hidn

aP

rain

&B

urki

llD

iosc

orea

aspe

rsa

Pra

in&

Bur

kill

Dio

scor

each

ingi

iP

rain

&B

urki

llD

iosc

orea

pulv

erea

Pra

in&

Bur

kill

Dio

scor

eate

ntac

ulig

era

Pra

in&

Bur

kill

Ch8

45(S

YS)

Ch9

21(S

YS)

Ch4

73(S

YS)

;Li

Yan-

hui

2036

(HIT

BC

)C

h791

(SY

S);

Wan

gH

ong

3147

(HIT

BC

)P

eiSh

eng-

ji99

85(H

ITB

C)

ER

ICA

CE

AE

Aga

pete

sbu

rman

ica

W.

E.

Eva

nsA

gape

tes

lobb

iiC

.B

.C

lark

eA

gape

tes

man

nii

Hem

sl.

LiYa

n-hu

i37

18(H

ITB

C),

2739

(HIT

BC

)Li

Yan-

hui

3719

(HIT

BC

)Li

Yan-

hui

3629

(HIT

BC

)

EU

PH

OR

BIA

CE

AE

Cle

ista

nthu

ssu

mat

ranu

s(M

iq.)

Mue

ll.-

Arg

.C

roto

nar

gyra

tus

Blu

me

Phy

llan

thus

clar

kei

Hoo

k.f.

Trig

onos

tem

onbo

nian

usG

agne

p.

LiYa

n-hu

i30

95(H

ITB

C),

2529

(HIT

BC

)C

h179

(SY

S)C

h563

(SY

S),

Ch8

12(S

YS)

Ch3

67(S

YS)

,C

h39

(SY

S),

Ch3

53(S

YS)


App

endi

x1.

Con

tinu

ed.

FAB

AC

EA

EA

lbiz

iaod

orat

issi

ma

(L.)

Ben

th.

Bau

hini

aca

rcin

ophy

lla

Mer

r.C

aesa

lpin

iats

oong

iiM

err.

Cal

lery

aeu

rybo

trya

(Dra

ke)

A.

M.

Scho

t

LiYa

n-hu

i14

11(H

ITB

C)

Ch1

097

(SY

S)C

h118

(SY

S)Li

Yan-

hui

4200

(HIT

BC

)D

erri

sca

udat

ilim

baH

owM

ilet

tia

tetr

apte

raK

urz

Soph

ora

praz

eri

Pra

inSo

phor

ato

nkin

ensi

sG

agne

p.

Ch3

98(S

YS)

,C

h504

(SY

S)Li

Yan-

hui

4100

(HIT

BC

)C

h394

(SY

S),

Ch5

59(S

YS)

;Li

Yan-

hui

2802

(HIT

BC

),42

77(H

ITB

C)

Ch6

19(S

YS)

FU

MA

RIA

CE

AE

Cor

ydal

ista

lien

sis

var.

siam

ensi

s(C

raib

)X

.Z

hang

LiYa

n-hu

i20

062

(HIT

BC

),37

27(H

ITB

C)

GE

SNE

RIA

CE

AE

Rap

hioc

arpu

sbe

goni

ifol

ius

(H.

Lev

.)B

.L

.B

urtt

Did

ymoc

arpu

sm

arga

rita

eW

.W

.Sm

.Ly

sion

otus

serr

atus

D.

Don

Orn

itho

boea

henr

yiC

raib

Ch9

47(S

YS)

;P

eiSh

eng-

ji98

92(H

ITB

C);

Tao

Guo

-da

1590

2(H

ITB

C)

Pei

Shen

g-ji

1054

(HIT

BC

)C

h624

(SY

S),

Ch8

06(S

YS)

;P

eiSh

eng-

ji59

-993

7(H

ITB

C);

Tao

Guo

-da

1575

0(H

ITB

C)

Exp

edit

ion

3262

5(H

ITB

C);

Pei

Shen

g-ji

59-1

0069

(HIT

BC

)

GU

TT

IFE

RA

EG

arci

nia

brac

teat

aC

.Y

.W

uex

Y.

H.

Li

LiYa

n-hu

i41

03(H

ITB

C),

3813

(HIT

BC

)

ICA

CIN

AC

EA

EG

omph

andr

ate

tran

dra

(Wal

l.)Sl

eum

.Ta

oG

uo-d

a11

182

(HIT

BC

)

LA

BIA

TA

EC

oleu

sxa

ntha

nthu

sC

.Y

.W

u&

Y.

C.

Hua

ngG

omph

oste

mm

apa

rvifl

orum

Wal

l.P

eiSh

eng-

ji10

138

(HIT

BC

)C

uiJi

ng-y

un14

659

(HIT

BC

);Ta

oG

uo-d

a15

798

(HIT

BC

)R

abdo

sia

erio

caly

xva

r.la

xiflo

raC

.Y

.W

u&

H.

W.

Li

LiB

ao-g

ui99

1120

9(H

ITB

C);

LiYa

n-hu

i37

59(H

ITB

C);

Wan

gZ

hong

-yu

s.n.

(HIT

BC

)

LA

MIA

CE

AE

Sym

phor

ema

invo

lucr

atum

Rox

b.Li

Yan-

hui

2988

(HIT

BC

),48

52(H

ITB

C)

LA

UR

AC

EA

EC

rypt

ocar

yaac

utif

olia

H.

W.

Li

LiYa

n-hu

i10

29(H

ITB

C),

3052

(HIT

BC

)

LIN

AC

EA

ER

einw

ardt

iain

dica

Dum

.E

xped

itio

n34

310

(HIT

BC

)

MA

LVA

CE

AE

Pte

rosp

erm

umch

ingt

unge

nse

C.

Y.

Wu

&H

sue

Pte

rosp

erm

umpr

oteu

sB

urki

llE

xped

itio

n34

533

(HIT

BC

)Li

Yan-

hui

3586

(HIT

BC

);W

ang

Hon

g83

1(H

ITB

C),

842

(HIT

BC

)



App

endi

x1.

Con

tinu

ed.

ME

LIA

CE

AE

Agl

aia

test

icul

aris

C.

Y.

Wu

Am

oora

tetr

apet

ala

(Pie

rre)

C.

Y.

Wu

Mun

roni

ahe

nryi

Har

ms

Ch3

39(S

YS)

Ch5

29(S

YS)

;W

ang

Hon

g17

10(H

ITB

C),

1715

(HIT

BC

)C

h351

(SY

S),

Ch1

92(S

YS)

;Li

Yan-

hui

3327

(HIT

BC

)

ME

NIS

PE

RM

AC

EA

EC

ycle

asu

tchu

enen

sis

Gag

nep.

Step

hani

ach

ingt

unge

nsis

H.

S.L

oSt

epha

nia

epig

aea

H.

S.L

o

Ch9

01(S

YS)

Tao

Guo

-da

4389

8(H

ITB

C)

Tao

Guo

-da

1567

2(H

ITB

C)

MY

RSI

NA

CE

AE

Myr

sine

sem

iser

rata

Wal

l.Li

Yan-

hui

2003

2(H

ITB

C);

Tao-

Guo

-da

1572

9(H

ITB

C);

Wan

gH

ong

1960

(HIT

BC

)

OL

AC

AC

EA

EN

atsi

atop

sis

thun

berg

iaef

olia

Kur

zTa

oG

uo-d

a42

370

(HIT

BC

)

OL

EA

CE

AE

Ligu

stru

msi

nens

eva

r.co

ryan

um(W

.W

.Sm

ith)

Han

d.-M

azz.

LiYa

n-hu

i42

9(H

ITB

C);

Tao

Guo

-da

1644

0(H

ITB

C);

Wan

gH

ong

2101

(HIT

BC

),20

91(H

ITB

C)

OR

CH

IDA

CE

AE

Bul

boph

yllu

mam

bros

ia(H

ance

)Sc

hltr

.B

ulbo

phyl

lum

kwan

gtun

gens

eSc

hltr

.C

h679

(SY

S)C

h22

(SY

S)B

ulbo

phyl

lum

nigr

esce

nsR

olfe

Den

drob

ium

sala

ccen

seL

indl

.E

ulop

hia

brac

teos

aL

indl

.P

holi

dota

chin

ensi

sL

indl

.

Zhu

Pei

-zhi

8683

(HIT

BC

)C

h158

(SY

S)Li

Yan-

hui

4150

(HIT

BC

)Ta

oG

uo-d

a44

107

(HIT

BC

)

PA

LM

AE

Car

yota

uren

sL

.C

hen

San-

yang

1895

9(H

ITB

C)

PIP

ER

AC

EA

EP

eper

omia

lept

osta

chya

var.

cam

bodi

ana

(C.

DC

.)M

err.

Ch9

50(S

YS)

PR

IMU

LA

CE

AE

Lysi

mac

hia

garr

etti

iF

letc

her

LiYa

n-hu

i33

02(H

ITB

C)

RH

AM

NA

CE

AE

Gou

ania

java

nica

Miq

.H

oven

iaac

erda

var.

kiuk

iang

ensi

s(H

u&

Che

ng)

C.

Y.

Wu

exY

.L

.C

hen

Sage

reti

ala

xiflo

raH

and.

-Maz

z.Ve

ntil

ago

caly

cula

tava

r.tr

icho

clad

aY

.L

.C

hen

&P.

K.

Cho

u

Ch7

98(S

YS)

,C

h850

(SY

S);

LiYa

n-hu

i37

14(H

ITB

C)

Tao

Guo

-da

3991

5(H

ITB

C)

Ch3

99(S

YS)

;Z

huP

ei-z

hi10

478

(HIT

BC

),10

479

(HIT

BC

)C

h550

(SY

S);

LiYa

n-hu

i52

30(H

ITB

C)


App

endi

x1.

Con

tinu

ed.

RU

BIA

CE

AE

Dam

naca

nthu

sin

dicu

sG

aert

n.f.

Hym

enop

ogon

para

siti

cus

var.

long

iflor

usH

owex

W.

C.

Che

nP

avet

tapo

lyan

tha

R.

Br.

Pav

etta

scab

rifo

lia

Bre

mek

.

Ch3

61(S

YS)

,C

h816

(SY

S)Ta

oG

uo-d

a13

651

(HIT

BC

)C

h368

(SY

S),

Ch2

70(S

YS)

;Li

Yan-

hui

1773

(HIT

BC

),41

11(H

ITB

C)

LiYa

n-hu

i42

40(H

ITB

C),

3871

(HIT

BC

)

RU

TA

CE

AE

Mur

raya

euch

rest

ifol

iaH

ayat

a

Mur

raya

tetr

amer

aH

uang

LiYa

n-hu

i37

16(H

ITB

C),

3309

(HIT

BC

);Ta

oG

uo-d

a15

722

(HIT

BC

);E

xped

itio

n32

673

(HIT

BC

)Li

Yan-

hui

3815

(HIT

BC

),42

69(H

ITB

C)

SCR

OP

HU

LA

RIA

CE

AE

Lind

enbe

rgia

phil

ippi

nens

is(C

ham

.&

Schl

tdl.)

Ben

th.

Ch9

6(S

YS)

;Li

Yan-

hui

363

(HIT

BC

),39

52(H

ITB

C)

SIM

AR

UB

AC

EA

EB

ruce

am

olli

sW

all.

LiYa

n-hu

i42

03(H

ITB

C)

STA

PH

YL

EA

CE

AE

Turp

inia

robu

sta

Cra

ibLi

Yan-

hui

393

(HIT

BC

);Z

hang

Jian

-hou

1370

7(H

ITB

C)

UL

MA

CE

AE

Cel

tis

wig

htii

var.

phil

ippe

nsis

(Pla

nch.

)E

.So

epad

mo

Ch3

2(S

YS)

,C

h371

(SY

S),

Ch1

089

(SY

S);

LiYa

n-hu

i39

1(H

ITB

C),

4211

(HIT

BC

)

UR

TIC

AC

EA

EE

lato

stem

ahe

rbac

eifo

lium

Hay

ata

Ela

tost

ema

salv

inio

ides

W.

T.W

ang

Lapo

rtea

uren

tiss

ima

Gag

nep.

Pel

lion

iasc

abra

Ben

th.

Pil

eaca

dier

eiG

agne

p.&

Gui

llau

min

Pro

cris

cren

ata

C.

B.

Rob

.

Ch8

77(S

YS)

Ch8

79(S

YS)

;Li

Yan-

hui

4105

(HIT

BC

);Z

huP

ei-z

hi86

49(H

ITB

C)

LiYa

n-hu

i39

6(H

ITB

C)

Ch9

17(S

YS)

Ch5

72(S

YS)

,C

h719

(SY

S)C

h514

(SY

S),

Ch6

32(S

YS)

;Li

Yan-

hui

3751

(HIT

BC

)

VE

RB

EN

AC

EA

EG

arre

ttia

siam

ensi

sF

letc

her

Ch3

58(S

YS)

,C

h710

(SY

S),

Ch9

41(S

YS)

VIT

AC

EA

ETe

tras

tigm

aca

mbo

dian

umP

ierr

eex

Gag

nep.

Tetr

asti

gma

dubi

um(L

awso

n)P

lanc

h.Te

tras

tigm

aru

pest

reP

lanc

h.

Ch9

1(S

YS)

,C

h925

(SY

S);

Zha

ngJi

an-h

ou13

602

(HIT

BC

)Li

Yan-

hui

3700

(HIT

BC

),27

37(H

ITB

C)

Ch1

39(S

YS)

ZIN

GIB

ER

AC

EA

EP

omm

eres

chea

lack

neri

Wit

tm.

Pei

Shen

g-ji

1007

3(H

ITB

C);

Tao

Guo

-da

4409

1(H

ITB

C)

Seed

Pla

ntTa

xapr

efer

ent

toli

mes

tone

habi

tats



App

endi

x1.

Con

tinu

ed.

AC

AN

TH

AC

EA

EA

ndro

grap

his

laxi

flora

(Blu

me)

Lin

dau

Era

nthe

mum

pulc

hell

umA

ndr.

Gol

dfus

sia

glom

erat

aN

ees

Pse

uder

anth

emum

pala

tife

rum

Rad

lk.

Ch3

76(S

YS)

,C

h808

(SY

S),

Ch9

74(S

YS)

;Li

Yan-

hui

3721

(HIT

BC

)Li

Yan-

hui

2898

(HIT

BC

)C

h119

1(S

YS)

;Ta

oG

uo-d

a37

75(H

ITB

C)

Ch1

37(S

YS)

;C

h172

(SY

S),

Ch1

80(S

YS)

,C

h310

(SY

S)P

seud

eran

them

umpo

lyan

thum

(C.

B.

Cla

rke)

Mer

r.R

hapi

dosp

ora

vaga

bund

a(R

.B

en)

C.

Y.

Wu

Sem

nost

achy

alo

ngis

pica

ta(H

ayat

a)C

.F.

Hsi

eh&

T.C

.H

uang

Ch9

1(S

YS)

;Li

Yan-

hui

1011

(HIT

BC

)C

h36

(SY

S),

Ch2

47(S

YS)

;Li

Yan-

hui

2829

(HIT

BC

)C

h118

2(S

YS)

AN

NO

NA

CE

AE

Mil

iusa

chun

iiW

.T.

Wan

gM

itre

phor

am

aing

ayi

Hoo

k.f.

&T

hom

s.M

itre

phor

ath

orel

iiP

ierr

e

Ch4

99(S

YS)

,C

h121

1(S

YS)

;U

nkno

wn

coll

.74

317

(HIT

BC

),74

318

(HIT

BC

)Li

Yan-

hui

3139

(HIT

BC

)C

h673

(SY

S),

Ch4

20(S

YS)

;Li

Yan-

hui

6749

3(H

ITB

C)

AP

OC

YN

AC

EA

EA

ntio

stel

ma

lant

sang

ensi

s(T

sian

g&

P.T.

Li)

P.T.

Li

Bid

aria

yunn

anen

se(T

sian

g)P.

T.L

iD

isch

idia

esqu

irol

ii(L

ev.)

Tsia

ngD

isch

idia

min

or(V

ahl)

Mer

r.G

ymne

ma

sylv

estr

e(R

etz.

)Sc

hult

.G

ymne

ma

lati

foli

umW

all.

exW

ight

Ch6

58(S

YS)

Ch2

42(S

YS)

,C

h764

(SY

S)C

h23

(SY

S)C

h729

(SY

S)E

xped

itio

n34

523

(HIT

BC

)C

h586

(SY

S),

Ch6

33(S

YS)

,C

h716

(SY

S);

Yang

Zhe

ng-h

ong

1097

6(H

ITB

C)

Hoy

aca

rnos

a(L

.f.)

R.

Br.

Hoy

aly

iL

ev.

Hoy

ane

rvos

aTs

iang

&P.

T.L

iH

oya

vill

osa

Cos

tani

n.Ja

smin

anth

essa

xati

lis

(Tsi

ang

&P.

T.L

i)W

.D

.St

even

s&

P.T.

Li

Mar

sden

iate

naci

ssim

a(R

oxb.

)M

oon

Toxo

carp

usvi

llos

us(B

lum

e)D

ecne

.

Ch6

51(S

YS)

Tao

Guo

-da

3547

4(H

ITB

C)

Zha

ngJi

an-h

ou18

562

(HIT

BC

)C

h960

(SY

S);

LiYa

n-hu

i25

74(H

ITB

C)

Tao

Guo

-da

4419

5(H

ITB

C)

Ch4

38(S

YS)

;E

xped

itio

n32

621

(HIT

BC

)C

h544

(SY

S),

Ch5

87(S

YS)

,C

h646

(SY

S)

AR

AC

EA

EA

glao

nem

api

erre

anum

Eng

l.C

oloc

asia

giga

ntea

Hoo

k.f.

Rha

phid

opho

rade

curs

iva

(Rox

b.)

Scho

ttR

haph

idop

hora

hong

kong

ensi

sSc

hott

Ch2

33(S

YS)

;C

h240

(SY

S);

Pei

Shen

g-ji

1027

4(H

ITB

C)

Ch1

169

(SY

S);

Tao

Guo

-da

9123

(HIT

BC

)C

h116

0(S

YS)

;Li

Jie

722

(HIT

BC

)C

h706

(SY

S),

Ch1

179

(SY

S)

BE

GO

NIA

CE

AE

Beg

onia

auga

stin

eiH

emsl

.B

egon

iadr

yadi

sIr

msc

h.B

egon

iapr

osta

taIr

msc

h.

Ch2

65(S

YS)

;Ta

oG

uo-d

a15

926

(HIT

BC

)Z

huP

ei-z

hi10

364

(HIT

BC

)C

h976

(SY

S)

BO

MB

AC

AC

EA

EB

omba

xce

iba

L.

LiYa

n-hu

i29

46(H

ITB

C)


App

endi

x1.

Con

tinu

ed.

BO

RA

GIN

AC

EA

EE

hret

iats

angi

iI.

M.

John

st.

LiYa

n-hu

i33

60(H

ITB

C),

3726

(HIT

BC

)

CA

PP

AR

IDA

CE

AE

Cap

pari

sm

embr

anif

olia

Kur

zC

appa

ris

urop

hyll

aF.

Chu

nC

h612

(SY

S);

Zha

oSh

i-w

ang

2258

2(H

ITB

C)

Ch8

53(S

YS)

,C

h136

(SY

S),

Ch1

53(S

YS)

,C

h195

(SY

S);

LiYa

n-hu

i21

12(H

ITB

C)

CA

PR

IFO

LIA

CE

AE

Vibu

rnum

tsan

gii

Reh

der

LiYa

n-hu

i42

35(H

ITB

C);

Zhu

Pei

-zhi

1047

0(H

ITB

C)

CA

RD

IOP

TE

RID

AC

EA

EP

erip

tery

gium

quin

quel

obum

Has

sk.

Exp

edit

ion

3476

1(H

ITB

C)

CO

MM

EL

INA

CE

AE

Por

andr

asc

ende

nsD

.Y

.H

ong

Ch9

02(S

YS)

;P

eiSh

eng-

ji99

47(H

ITB

C);

Unk

now

nco

ll.

262

(HIT

BC

)

CO

MP

OSI

TA

EYo

ungi

aja

poni

ca(L

.)D

C.

Ch1

201

(SY

S),

Ch3

26(S

YS)

;Ta

oG

uo-d

a17

42(H

ITB

C)

EB

EN

AC

EA

ED

iosp

yros

yunn

anen

sis

Reh

der

&W

ils.

Ch5

16(S

YS)

,C

h121

4(S

YS)

;C

uiJi

ng-y

un14

656

(HIT

BC

);P

eiSh

eng-

ji10

300

(HIT

BC

)

EU

PH

OR

BIA

CE

AE

Ade

noch

laen

asi

lhet

ensi

sB

enth

.A

ntid

esm

am

onta

num

var.

mic

roph

yllu

m(H

emsl

.)P.

Hof

fman

nC

leid

ion

brac

teos

umG

agne

p.C

leid

ion

brev

ipet

iola

tum

Pax

&K

.H

offm

.La

sioc

occa

com

beri

var.

pseu

dove

rtic

ella

ta(M

err.)

H.

S.K

uSu

mba

viop

sis

albi

cans

(Blu

me)

J.J.

Smit

h

Ch8

07(S

YS)

;Z

ouSh

uang

-yun

356

(HIT

BC

)Li

Yan-

hui

3575

(HIT

BC

)Li

Yan-

hui

4250

(HIT

BC

)C

h922

(SY

S);

LiYa

n-hu

i39

9(H

ITB

C)

Ch3

09(S

YS)

,C

h226

(SY

S),

Ch3

8(S

YS)

;Li

Yan-

hui

1694

(HIT

BC

)C

h131

(SY

S);

LiYa

n-hu

i24

23(H

ITB

C)

FAB

AC

EA

EB

auhi

nia

genu

flexa

Cra

ibM

ille

ttia

yunn

anen

sis

Pam

pan.

Whi

tfor

diod

endr

onfil

ipes

(Dun

n)D

unn

Ch2

5(S

YS)

,C

h97

(SY

S);

LiYa

n-hu

i15

00(H

ITB

C)

Tao

Guo

-da

1366

8(H

ITB

C)

Zha

oSh

i-w

ang

2249

2(H

ITB

C)

FL

AC

OU

RT

IAC

EA

EF

laco

urti

aru

kam

Zol

l.&

Mor

.C

h784

(SY

S)

GE

SNE

RIA

CE

AE

Chi

rita

dim

idia

taR

.B

r.P

arab

oea

rufe

scen

s(F

ranc

h.)

B.

L.

Bur

ttP

arab

oea

dict

yone

ura

(Han

ce)

B.

L.

Bur

ttP

arab

oea

sine

nsis

f.m

acro

phyl

la(S

tapf

)C

.Y

.W

u

Ch9

48(S

YS)

Ch3

79(S

YS)

;W

ang

Hon

g27

24(H

ITB

C)

Ch8

42(S

YS)

;W

ang

Hon

g27

07(H

ITB

C)

Ch2

72(S

YS)

;E

xped

itio

n34

359

(HIT

BC

);Ta

oG

uo-d

a16

748

(HIT

BC

),37

92(H

ITB

C)



App

endi

x1.

Con

tinu

ed.

GR

AM

INE

AE

Den

droc

alam

usst

rict

us(R

oxb.

)N

ees

Ch1

058

(SY

S);

Sun

Ji-l

iang

1815

1(H

ITB

C)

ICA

CIN

AC

EA

EA

pody

tes

dim

idia

taE

.M

eyer

exA

rn.

Tao

Guo

-da

1669

0(H

ITB

C)

LA

BIA

TA

EC

oleb

rook

eaop

posi

tifo

lia

Sm.

Els

holt

zia

stac

hyod

es(L

ink)

C.

Y.

Wu

Ch5

1(S

YS)

;Li

Yan-

hui

252

(HIT

BC

),Su

nJi

-lia

ng18

005

(HIT

BC

)Ta

oG

uo-d

a37

98(H

ITB

C)

LA

MIA

CE

AE

Cle

rode

ndro

nhe

nryi

Pei

Gar

rett

iasi

amen

sis

Fle

tche

rP

rem

naha

mil

toni

iJ.

L.

Ell

isSp

heno

desm

em

olli

sC

raib

Wan

gH

ong

2921

(HIT

BC

),29

22(H

ITB

C),

2940

(HIT

BC

)E

xped

itio

n32

276

(HIT

BC

);U

nkno

wn

coll

.243

4(H

ITB

C),

456

(HIT

BC

);W

ang

Hon

g27

27(H

ITB

C)

Ch1

77(S

YS)

,C

h278

(SY

S);

Exp

edit

ion

3261

7(H

ITB

C)

Ch8

64(S

YS)

LA

UR

AC

EA

ELi

tsea

dill

enii

foli

aP.

Y.

Bai

&P.

H.

Hua

ngLi

tsea

elon

gata

(Wal

l.ex

Nee

s)B

enth

.&

Hoo

k.f.

Pho

ebe

lanc

eola

ta(W

all.

exN

ees)

Nee

s

LiYa

n-hu

i30

53(H

ITB

C)

Ch2

69(S

YS)

,C

h119

(SY

S)C

h161

(SY

S),

Ch6

1(S

YS)

,C

h324

(SY

S);

LiYa

n-hu

i75

4(H

ITB

C)

LIL

IAC

EA

EP

olyg

onat

umpu

ncta

tum

Roy

leex

Kun

thTu

pist

raw

atti

iH

ook.

f.C

h711

(SY

S)E

xped

itio

n34

321

(HIT

BC

);Ya

ngZ

hen-

hong

6971

(HIT

BC

);W

ang

Hon

g31

02(H

ITB

C)

MA

LP

IGH

IAC

EA

EH

ipta

gebe

ngha

lens

isva

r.to

nkin

ensi

s(D

op)

S.K

.C

hen

Ch4

5(S

YS)

;Li

Yan-

hui

1051

(HIT

BC

)

MA

LVA

CE

AE

Ster

culi

avi

llos

aR

oxb.

Zou

Shua

ng-y

un32

4(H

ITB

C)

ME

LIA

CE

AE

Chu

kras

iata

bula

ria

var.

velu

tina

Kin

gD

ysox

ylum

lent

icel

latu

mC

.Y

.W

u&

H.

Li

Dys

oxyl

umsp

icat

umH

.L

.L

iTo

ona

cili

ata

var.

pube

scen

s(F

ranc

h.)

Han

d.-M

azz.

L.Ya

n-hu

i38

5(H

ITB

C)

Ch5

08(S

YS)

;C

uiJi

ng-y

un22

658

(HIT

BC

)U

nkno

wn

coll

.25

0(S

YS)

Ch6

80(S

YS)

,C

h566

(SY

S),

Ch5

81(S

YS)

MO

RA

CE

AE

Cud

rani

ajin

ghon

gens

isS.

S.C

hang

Fic

usor

thon

eura

Lev

.&

Vani

otF

icus

pros

tata

Wal

l.ex

Miq

.F

icus

pubi

gera

var.

mal

ifor

mis

(Kin

g)C

orne

r

LiYa

n-hu

i31

35(K

UN

)C

h41

(SY

S);

LiYa

n-hu

i38

02(H

ITB

C)

Zhu

Hua

&W

ang

Hon

g88

-11

(HIT

BC

)Li

Yan-

hui

3223

(HIT

BC

)

MY

RSI

NA

CE

AE

Ard

isia

garr

etti

iF

letc

her

Ch1

30(S

YS)

,C

h170

(SY

S),

Ch4

7(S

YS)

;Li

Yan-

hui

3867

(HIT

BC

)


App

endi

x1.

Con

tinu

ed.

MY

RT

AC

EA

ESy

zygi

umcl

avifl

orum

(Rox

b.)

Wal

lex

A.

M.

&J.

M.

Cow

anC

h501

(SY

S),

Ch4

92(S

YS)

OR

CH

IDA

CE

AE

Gas

troc

hilu

sob

liqu

us(L

indl

.)K

untz

eN

ervi

lia

plic

ata

(And

r.)Sc

hltr

.Tr

opid

iaan

gulo

sa(L

indl

.)B

lum

e

LiYa

n-hu

i27

34(H

ITB

C)

Ch7

13(S

YS)

;Li

Yan-

hui

3138

(HIT

BC

)Li

Yan-

hui

392

(HIT

BC

);W

ang

Pei

-qun

1111

5(H

ITB

C)

OR

OB

AN

CH

AC

EA

EA

egin

etia

indi

caL

.Z

hao

Shi-

xian

g16

5(H

ITB

C)

OX

AL

IDA

CE

AE

Aver

rhoa

cara

mbo

laL

.B

ioph

ytum

esqu

irol

iiL

ev.

LiYa

n-hu

i31

29(H

ITB

C)

LiYa

n-hu

i40

04(H

ITB

C)

PA

LM

AE

Car

yota

ochl

andr

aH

ance

Che

nYu

6207

9(H

ITB

C)

PIP

ER

AC

EA

EP

eper

omia

heyn

eana

Miq

.P

iper

arbo

rico

laC

.D

C.

Pip

ercu

rtip

edun

cula

tum

C.

DC

.

Ch8

04(S

YS)

;Li

Yan-

hui

2738

(HIT

BC

)C

h585

(SY

S),

Ch9

24(S

YS)

,C

h121

6(S

YS)

;U

nkno

wn

coll

.86

5(H

ITB

C)

Cui

jing-

yun

2281

1(H

ITB

C)

RH

AM

NA

CE

AE

Vent

ilag

ole

ioca

rpa

var.

pube

scen

sY

.L

.C

hen

&P.

K.

Cho

uZ

izyp

hus

atto

pens

isP

ierr

eC

h108

8(S

YS)

,C

h645

(SY

S)Li

Yan-

hui

1289

(HIT

BC

),C

h251

(SY

S)

RU

BIA

CE

AE

Ixor

ace

phal

opho

raM

err.

Pav

etta

aren

osa

Lou

r.P

avet

taho

ngko

ngen

sis

Bre

mek

.P

sych

otri

api

life

raH

utch

.P

sych

otri

asi

amic

a(C

raib

)H

utch

.Sp

irad

icli

sca

espi

tosa

f.su

bim

mer

saL

o

Ch1

218

(SY

S),

Ch1

229

(SY

S)Li

Yan-

hui

246

(HIT

BC

),28

14(H

ITB

C)

Ch3

55(S

YS)

;Z

huH

ua&

Wan

gH

ong

3004

(HIT

BC

)C

h712

(SY

S),

Ch1

65(S

YS)

;Li

Yan-

hui

1498

(HIT

BC

)C

h836

(SY

S);

LiYa

n-hu

i24

73(H

ITB

C),

258

(HIT

BC

)Li

Yan-

hui

1267

5(H

ITB

C)

RU

TA

CE

AE

Gly

cosm

isci

trif

olia

(Wil

ld.)

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dl.

Mur

raya

koen

igii

(L.)

Spre

ng.

Mur

raya

mic

roph

ylla

(Mer

r.&

Chu

n)Sw

ingl

eM

urra

yapa

nicu

lata

(L.)

Jack

.Z

anth

oxyl

umar

mat

umva

r.fe

rrug

ineu

m(R

ehde

r&

E.

H.

Wil

son)

C.

C.

Hua

ngZ

anth

oxyl

umla

etum

Dra

keZ

anth

oxyl

umut

ile

C.

C.

Hua

ng

Ch2

22(S

YS)

,C

h146

(SY

S);

Zou

Shua

ng-y

un33

3(H

ITB

C)

Ch1

76(S

YS)

,C

h330

(SY

S),

Ch3

52(S

YS)

,C

h313

(SY

S);

LiYa

n-hu

i18

59(H

ITB

C)

Exp

edit

ion

3426

7(H

ITB

C);

Tao

Guo

-da

4405

6(H

ITB

C)

LiYa

n-hu

i33

15(H

ITB

C)

Ch8

87(S

YS)

;U

nkno

wn

coll

.10

10(H

ITB

C)

Ch4

75(S

YS)

,C

h578

(SY

S)C

h636

(SY

S),

Ch7

77(S

YS)



App

endi

x1.

Con

tinu

ed.

STA

PH

YL

EA

CE

AE

Tapi

scia

yunn

anen

sis

W.

C.

Che

ng&

S.D

.C

huZ

huH

ua93

011

(HIT

BC

)

TE

TR

AM

EL

AC

EA

ETe

tram

eles

nudi

flora

R.

Br.

LiYa

n-hu

i85

41(H

ITB

C)

TIL

IAC

EA

EC

olon

aflo

ribu

nda

(Wal

l.)C

raib

Col

ona

thor

elii

(Gag

nep.

)B

urre

tC

h700

(SY

S),

Ch9

32(S

YS)

;P

eiSh

eng-

ji10

084

(HIT

BC

)Li

Yan-

hui

1696

(HIT

BC

)

UR

TIC

AC

EA

EB

oehm

eria

zoll

inge

rian

aW

edd.

Deb

rege

asia

edul

is(S

iebo

ld&

Zuc

c.)

Wed

d.E

lato

stem

aru

pest

re(B

uch.

-Ham

.)W

edd.

Pil

eam

onil

ifer

aH

and.

-Maz

z.P

ilea

plat

anifl

ora

C.

H.

Wri

ght

Pou

zolz

iasa

ngui

nea

(Blu

me)

Mer

r.

Ch1

064

(SY

S);

Cui

Jing

-yun

1941

6(H

ITB

C)

Ch3

11(S

YS)

;Ta

oG

uo-d

a41

641

(HIT

BC

)C

h168

(SY

S),

Ch3

88(S

YS)

,C

h359

(SY

S);

Wan

gH

ong

1494

(HIT

BC

)C

h984

(SY

S);

Wan

gH

ong

1484

(HIT

BC

)C

h381

(SY

S);

Ch6

57(S

YS)

;Li

Yan-

hui

4210

(HIT

BC

);Ta

oG

uo-d

a16

090

(HIT

BC

)C

uiJi

ng-y

un22

793

(HIT

BC

)

VIT

AC

EA

ELe

eaae

quat

aL

.Yu

aau

stro

-ori

enta

lis

(Met

calf

)C

.L

.L

iTe

tras

tigm

aca

ulifl

orum

Mer

r.Te

tras

tigm

ade

lava

yiG

agne

p.

Ch4

68(S

YS)

,C

h714

(SY

S);

Pei

Shen

g-ji

9359

(HIT

BC

)C

h844

(SY

S)C

h268

(SY

S);

LiYa

n-hu

i11

915

(HIT

BC

)C

h507

(SY

S);

Ch8

11(S

YS)

Tetr

asti

gma

erub

esce

nsva

r.m

onos

perm

umG

agne

p.Te

tras

tigm

apu

bine

rve

Mer

r.&

Chu

nVi

tis

bala

nsae

ana

Pla

nch.

Viti

sbe

tuli

foli

aD

iels

&G

ilg

Ch4

63(S

YS)

,C

h105

4(S

YS)

;Ta

oG

uo-d

a91

89(H

ITB

C)

Ch6

53(S

YS)

,C

h616

(SY

S)C

h668

(SY

S),

Tao

Guo

-da

1644

7(H

ITB

C)

Ch6

54(S

YS)

,C

h408

(SY

S)

ZIN

GIB

ER

AC

EA

EH

edyc

hium

sino

-aur

eum

Stap

fH

edyc

hium

vill

osum

var.

tenu

iflor

umW

all.

exB

aker

Pom

mer

esch

easp

ecta

bili

s(K

ing

&P

rain

)K

.Sc

hum

.

Ch1

240

(SY

S);

Wan

gH

ong

3066

(HIT

BC

);Ta

oG

uo-d

a16

049

(HIT

BC

)Li

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3471

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.)Ta

rdie

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Documents

BIOGEOGRAPHY AND THE LIMESTONE FLORA IN SOUTHERN …flora.huh.harvard.edu/china//novon/mobt-90-03-444.pdf · as the regional tropical seasonal rain forest off lime-stone, shares characteristics