全 文 ：r1-／7
广 西 植 物 Guihaia 14(I)：Il—l7．1994
&g叼 ／
羊蹄甲属的系统与生物地理学：1．厚盘组的分支分析
张奠湘 陈德昭
酬鞭鞘蚴眺肫州n们 (57 ，．
H 摘要 羊蹄甲属厚盘组作为一个分类实体包含有约18个种．本组植物主要分布于中国南部硬中南半
岛等地．其中很多种类在其分布区内地理替代现象相当明显。对此组的分支系统学研究，一方 面
有助于对本组种类系统进化关系的理解，另一方面叉为对热带亚热带亚洲犬陆各区最之间关系 的
理解提供帮助．为本地区的分支生物地理学研究提供材料。
本文用3O个形态及叶脉脉序性I捩对厚盘组进行了分支分析．云南羊蹄甲被选作外类群． 内类
群包括龙须藤亚组及攀援羊蹄甲亚组的所有l8个种． 分析产生了8个最简妁的分支图， 分析还表
明攀援羊蹄甲单独作 一个亚组是不可取的．厚盘组的分类及生物地理学需要进一步的研究．
关键诲 兰堕旦墨 墼望；．坌塞坌堑
Systematics and biofleoflraphy of Bauhinia L．
(Leguminosae～Caesalpinioideae)： I．Cladistic analysis
Of sect．Lasiobema (KO1th．)Benth．
Zhang Dianxiang and Chen Toohao (T。Chen)
(South CMaa Institute of Botany，Academia Siaica．Guangzhou 510650，China)
Abst ract A clad．蝎tie analysis ol sect Lasiobema based on 30 m0rpho logical and leaf
venal}ionaI characters is presented． Bauhln ¨na e s插 Franeh． chosen as its out—
group． A11 18 species in subse ct． Scandehies and subsect． Champlonae are ineluded；
Eight equally most parsimonivu8 cladograms ate produced． Some clades， the (B．
cofFJosa，B esquirolii)clads．the(且 $canden$，B．delavayi)，(B．[1ava，(B oxysel~ala，
B．penieilloba)))cIade， the(B．concre~a．B．curtissii)clade．the(B．harmsiana，(且
calycina，B．oodfroyi))elude．are stable in all eladograms produced．The sabsectional
status of B．sceandens is not warranted． Furthe r studying is needed to make nlore
detailed account on the systematics and biogeography of sect． Lasiobema．
Key word s Bauhinia：sect． 工aSfobema；cladlstic analysis
Introduetion
Lasiobema as a taxonomio entity WaS firs~recognized by Korthals (1841)．
But he failed to design a status for it． Bontham (1865)ranked it a section．Do
W it 0956)reoognized it aS genus iEt which the Tsbieolyx speoies inoIuded,
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12 广 西 植 物 14卷
Lasiobema as a section is maintained by W underlin，Larsen and Lars~n (1987)．
The section Lasiobema is a group mainly confined to the tropical cOntinental
A缸 the geological and hiogeographical history of wbSch is very eomplicared．
Some authors ov~~l think that a part of 80uthe ast Asia had Gondwanic origin
(Audl~y-Charles，1987)．A be~ter understanding of the b~ogoography of the areRs
concerned has to be based on a bettor understanding of the phylogenetic relation-
ship of organiSm s occur in which．
Since the publication of Heunig’s 。Phylogenetic Systematics in English in
1 966，Cladistics (i㈨e Farris，1 970；Nelson ＆ Platnick，1 981：W iley，1981)has
become 8 routine in elucidating the organisma~relationship．
Organism and its physic environment evolve together (Croizat，1958；1 964)．
Historical biogeography is to find patterns in organi smal distribution and to
elucidate liypothe8is on the procesS that generated these patterns (Brooks， 1981，
1990； Nelson ＆ Platni ckJ 1 981； Humphries ＆ Paren ti， 1986； Zande ＆ Roos，
1987)． In a cladistic biogeography analysisJ cladograms of unrelated groups of
organisms occuring in the area stud~ed is needed if general areagram i8 t0 be
built．But no other group comparable to sect．Lasiobema in distributional pattern
has been cladisticaliy analysed vet． A cladistic analysis on Lasiob i目carrjed
out to add to our knowledge of the evelution of this group in one hand， and in
the other hand，as the first step in building 。 general areagram of the areas
concerned．
In this study sect．Lasiobema incl de8 subsections Championae and subsect．
andente*帮 defined by W uudorlin et a1．(1987)． 8ubsoct． 尸ullae is excluded
from the section．subsect Pullae is more closely related to sect．T．bicalyx and
set．Loxocalyx according to a oladistie analysis 0n the genus Bauhinia (Zhang，
in p ep．)．
The 目peties listed below ” e us ed as OTUs in this study．
B．soandens L．：Java，mainland ThailandJ VietnamJ Laos，Cambed ia，Ⅱaiuan，
Burma，India and Sri Lanka．
B．apertilobata Mort．＆ Mete．：China：Fujian，Jiangxi，Guangdong and
GuangXi，on the Nanling Mountain Range．
B．calyclna Piorre ex Gagn．：0ambodia：KompOng Spsu．
B．ohampionii(Benth．)Benth．：China：Hainan，Guangden g，Guangxi，I=[un anJ
Huboi，Fujian，Taiwan，Jiangxi， Ouizhou，Zhejiang；Vietnam：Lang Son，
Quang Ninh，Ha Son Binh；India． ’
． comosa Craib：China：Yunnan，Sichuan (Liangshan)．
B．conoreta Craib：Th ail~nd：Peninsula．
B．curlissii Prain：Cambodia：Pursat．Laes：Savannakhet．Vietnam：Binh Tri
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1期 张莫湘等：羊蹄甲属的系统与生物地理学I 1．厚盘组的分支分桁 ls
Thien，Phu Khauh，Thuau Hai，，Dong Nai． Malaysia：Malay Peninsula：Lang-~
awl Is．；Thailand．
B·delavayi Franch·：China：Yunnan(Binehuan，Luoh~an，Fengqing)．
B·esqutrolii Gagnep·：China：Yunnan (Yuanjiang，Xingping，Binohuan，
Eryuan)and Guizhou (Guiding)·
B．flava(De Wit)Cusse~：Malaysia：Malay Peninsula：Langkawi Is
B．godfroyi Gagn．：Cambodia：Puxsat．
B．harmsiana Hosseus：Cambodia：Battambang．
B．hypoglauca Tang et W ang ex T． Chen： China Yunnan (Guangnan，
Xianshan)．
B．Jingyuenensis T．Chen：Chiha：NW Guangxi．
B．1onglstlpes T．Chen：China：Yuunan Yongsheng)．
B．oxysepala Gagn．：Vietnam：Ha Son Binh，Thanh Hoa：
B．penioilliloba Pierre ex Gagn．：Cambodia：Stung Treng．Laos：Savannakhet．
Sithandone．Vietnam：Dac Lae．
B．veflusiuta T．Chen：China：Guangxi(Guixian，Yongning)．
The distribution al data are collected from specimen rogerds and several floras
and monographs(de Wit，1956；Larsen et al，1980，1984；Chen。1988)．
Methods and material
The data matrix (Table 1)wa8 analyzed using Paup 8．0 (Sworford- 1991)
0丑 a Macintosh Plus microcomputer．Settings：HeureStioJ accelerated transforma~
tion，all characters ordered．
．
A computerized cladistic analysis based up0n the prineipie of parsimony does
not require decisions about polarity to be made for characters beforehand．Includ—
ing an outgroup or ancestral taxon in the analysisJ the program decides character
polarities based on the same parsimony criterion used in the reCOnStructing of
phylogenetio tree(Andersen，1991)·
Palynological characters show that · “ 疗仰 口 括 Franch has the gan1e typ。
of pollen with sect． Lasiobema species(Larsen，1975)． The cladiscie analysis of
the genus Bauhinia 01 series iovo~also supported the sister group of sect．Lagio
bema is the Yunnanentes-Corymbosue clade． ． u ” n口ns was thus used as the
outgroup in building the oladogram of Lasiobema．
Characters
All characters are morphological or leaf venational characters． Characters
are recorded from herbarium and laboratory materia1． Venation data ar9 from B
t e札ment of leaf venation of the tribe Cereideae by the Same author(Zhang，in
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14 广 西 植 物 14卷
prep)· Only morphological characters are uged in the analysis
．
1．Leaf On flowering branch：free leaflets(a) bilobed (b) entire(o)
2． Leaf：glabron S (8) pubescent on lower Surface (b) pubesGon t on both
surface (c)
8． 1nfl~rescence：panicle (a) slender raceme(b)
4．Pedicel：much longer than inflorescence internod e length (a) nearly equal
(b)
Brac t：subulate (a) linear (b)
Posit2on of bracteoles on podiceh lower pa rt (a) upper pa rt(b)
Bud：fusiform ( ) subglobose(b) globose (c)
Bud：otherwise(a) 5-linear (b)
Bud：otherwise (8) 5-dentate(b)
Bud：dalyx lobes not covering petals before anthesis(8) ca1vx lobes dosed
(b)
Bud：longer than 0．8 om (a) 0．8-0．5 (b) shorter than 0．5 cm (c)
Calyx：split to 5 lob8S (a) 2-3 lobes(b)
Calyx：only uppe r pa rt split(a) split to the mouth of hypa nthium (b)
Petal：glabrous(8) partly hairy (b) wholly hairy (e)
Petal：margin smooth (a) otherwiSe (b)
petal：claw longer than blade (a) blade longer (b)
Petah obova七e (8) laneeolate(b)
Filament：glabrous (8) hairy (b)
Ovary：wholly hairy (a) hairy on Suture(b) glabrous (o)
Crynophore：longer than half length of ovary (a) not SO (b)
S~yls：stout(8) slen der (b)
S~igma：incon spion ous (a) capltate (b)
DiSO：not swollen (a) swollen (b)
Disc：glabrous (a) halry (b)
Pod：obovate(b) strap-shape d (b) eDiptic (o)
Pod：longer than 6 em (a) 6-4 cm (b) shorter than 4 cm (c)
Pod：wider than 2．5 cm (a) less than 2 cm (b)
Venation：veinlets simple Or on ce-branched (a) twi ce-branched (b)
Venation：veinl et in most areoles (a) not SO (b)
Venation ：areoles moderate (a) very big (b)
Table 1． Data-matrix of Lasiobema
日·yunnaaensis aabab?babaabbaabaacaab??aaaaab
日．1ongistipe bbbabbbababbbbabaacaabaa???aab
日．COmOsa bobabbbaabeabbabaa~baabababbab
执 m 地 轨 执 峨弧
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l期 张奠湘等：羊蹄甲属的系统与生物地理学：1．厚盘组的分支分析
B．esqulrolii
B．seandens
B．delavayi
日．apert~lobata
bebabbbbabbaabaaebaabababbab
cabbaaeaaaeba~baacababbecbleab
bbbaaabaaabaababaabbbaaaeebaab
bbaababaabbabbabaal~aabaaaaaba
yun／1snensis
Iongistipes
esquiro】 i
delavayi
penicilliIoba
oxysepata
flava
apertldbata
CaiqUe]na
godfroyi
harmsiana
hypogIauca
cbamplonii
venustata
lingyuene~
c 临 8
B． c0n删 &
F ， I One of the eight most parsimonious cladograms·
ylelal~en$is
longlatlpes
esquirolli
scandens
deIavayi
penicilliloba
oxysepa【a
nava
apertilokata
champiomi
hypog1a11ca
1ing eⅡeⅡ8i昏
yelustata
caly~ilm f
gocfroyi
harmsia~a
eurti~ii
Fig． 2 The strict consenstl$ tree of eight trees
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l6 广 西 植 物
B．championii
B．hypoglauea
丑．1ingyuenensis
B． Pn sl0f0
B．ealyeina
B．harmsiana
B．penieilliloba
B．c“ ssii
B．]lava
B．c0ncrP Ⅱ
B．oxysepala
B．godfroyi
ebbaaabaaababbabaabbaabaaaaaba
bbbaaabazababba娃 abbaabaaaaaba
ebbaaaba~ababbbbaabbaaba?97 aba
cbbaaabaaababbabaaabaababababa
ebbaabazabebbbabbacababa?99997
bbbbaabaabcbbabbaababbbbaaaaba
bbbabaababaaaeabbaebbababhbabb
ebbaabbazabbbbaaaaeb~ababbbaba
bbbbnbHabb∞ 砒 a8c如 ababababa
ebba~acazaebbaabaaobaababbbaba
bbbaababahbabeabhaebhahag??999
bbbaabbaabebbb~ aacbbaba??????
14卷
Results and discussion
Eight trees with length of 85 step (Fig．1，one of．them )and a strict COIl-
senses tree(Fig．2)wore produced．
Some cls~es~ pear in all~ladog ramS produced．But the interposition of soD28
olades need further studying．
In the ingroup Bauhinia IongisHpes i8 the sister taxcra of a11 others together
all cladogramS produced． The next one to produce is always the(丑． eomo~,a
B．esqulrolii)elade．
The ((B．seandens，B．delavay
))) elade is another stable clads．
the(日．curtissiiJ B．eonereta)elade
calycina))elade．
)，(B．~lava，(B．oxysepala，B．penieillilo-
But the position of it is changeable． So are
and the(日．harmsiana， (日．godfrey~， B．
The positions of other species on eladograms are very unstable·Only if more
data were avaiIable can a general conclusion be made．
Species in a clade are usually biog eographicaflly closely related- It is shown
by the (B．comosa，B．esqairolii)clado in Southwestern China，the (B．cq issii，
B．Ooflcrefa)e~ade and the((日．penieilliloba，B．oxysepala) B．~lava)elade in
Indoehina and the Th ai-Malay Peninsular，the(B．harmsiana，(日．godfroyi， B．
calycina))clads in Cambodge．
One c2ade of the outgroup，B．yunnanensis Franoh．ocouring in W est China，
another dade of the outgroup，set．Corymbosae，having ahnost the same distri-
bufional area as the ingroup． The earliest elades of the ingroup are also from
SouthweStern China． W hether it is a general biogoographie pattern can only be
tested by other groups with a distributional a ea comparable． More detailed ac-
count of the evolution and biogeog raphy of the present group need further study‘—
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l圳 张真湘等 羊黼甲属的系统与生物地理学：1
． 厚盘姐的分支分析 1 7
Acknowledgements
This work is based Orl a draft wr~ien in Chinese
． The Pregent Version i日
partly finlshod in the Department of System atic Botany
， Aarhus University
where I (the first author)worked for o13e vear as a fellowshil>-holder supl：orted
by the Danish Research Academy． Prof．Kai L~rSen and Mrs
． Supoo 8．Larson，
M ·SoJ supervised my work and kindly helped me in so many ways．Dr．Susanne
S．Refiner permitted me to use PAUP． Dr．Anders Barfod and other mel~tbor8
in our Cladistlcs Discussion Group inspired me directly or indirectly in my works
in cladistics． 1 wish to ~hank 8ll the jnstjtu 0ng and indivJduaJs me．Mened
above．
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