全 文 ：东南亚地区五加科植物进化关系的初步研究
?
文 军1 , 2 , 朱昱苹3 , Chunghee LEE1 ,
Elizabeth WIDJAJA
4
, LENG Guan Saw
5
( 1 Department of Botany, National Museumof Natural History, MRC166 , Smithsonian Institution, Washington, DC 20013-7012 ,
U . S . A ; 2 中国科学院植物研究所系统与进化植物学国家重点实验室 , 北京 100093; 3 南京大学生命科学院 ,
江苏 南京 210093 ; 4“ Herbarium Bogoriense,” Puslitbang Biologi , Jalan Raya Juanda 22 , Bogor, Indonesia;
5 Forest Research Institute Malaysia ( FRIM) , Kepong, 52109 Kuala Lumpur, Malaysia)
摘要 : 东南亚五加科包含 14 个属约 500 种 , 本文应用 ITS 片段对该区五加科植物的进化关系作了初步研
究。研究显示该地区五加科植物具有复杂的起源 , 很多属属于亚洲掌状复叶类群或 Hedereae族的一支中。
该区特有类群 Harmsiopanax形态上非常特殊 , 但其系统位置尚未不明朗。在 Brassaiopsis属中 , 有几种形态
差异较大的种 , 但它们属同一单系 , 加之各种间 ITS序列差异较小 , 故应是新近起源于马来亚半岛和苏门
达腊岛的种类。 Wardenia simplex 聚类在 Brassaiopsis一支中 , 故不支持将 Wardenia 作为独立的属。东南亚地
区对于 Schefflera属的发育非常重要 , 已有的证据显示该区的 Schefflera属植物属于该属的 Heptapleurum类群。
马来亚与泰国南部的 Dendropanax lancifolius并没有与 Dendropanax属的核心类群聚在一起 , 其系统地位需进
一步研究。 Macropanax maingayi 是非常特殊的一个种 , 曾被独立分出 , 成立了单种属 Hederopsis。本文的分
析清楚表明它属于 Macropanax属。 Aralia merrillii 因为其不同寻常的攀缘特性而被独立出来 , 建立了单种属
Acanthophora, 但 ITS序列分析支持将它置于 Aralia属中。新增的取样继续支持 Arthrophyllum的单系性。 Os-
moxylon的原初分布范围在东南亚 , 它是五加科系统进化树上孤立的类群。
关键词 : 五加科 ; 系统进化 ; 东南亚 ; Wardenia; Hederopsis
中图分类号 : Q 949 文献标识码 : A 文章编号 : 0253 - 2700 (2008) 04 - 391 - 09
Evolutionary Relationships of Araliaceae in the Malesian
Region: a Preliminary Analysis
WEN Jun1 , 2 , ZHU Yu-Ping3 , ChungheeLEE1 , Elizabeth WIDJAJA4 , LENG Guan Saw5
( 1 Department of Botany, National Museumof Natural History, MRC166 , Smithsonian Institution, Washington, DC 20013-7012 ,
U . S . A ; 2 State Key Laboratory of Systematic and Evolutionary Botany, Instituteof Botany, Chinese Academy of Sciences,
Beijing 100093 , China; 3 Collegeof Life Sciences, Nanjing University, Nanjing 210093 , China;
4“ Herbarium Bogoriense,” Puslitbang Biologi , Jalan Raya Juanda 22 , Bogor , Indonesia;
5 Forest Research Institute Malaysia ( FRIM) , Kepong, 52109 Kuala Lumpur, Malaysia)
Abstract: We employ the nuclear ribosomal ITS sequences to assess the evolutionary relationships of Araliaceae in the
Malesian region . Malesian Araliaceae consist of 14 genera and about 500 species . Our analysis suggests a diffuse origin of
Araliaceaetaxa, withmany genera belongto the Asian palmatecladeor the tribe Hedereae . TheMalesian endemic Harm-
siopanax is morphologically unique and its phylogenetic position is not well resolvedat present . Several morphologically di-
verse species of Brassaiopsis perhaps have a relatively recent origin in the Malay Peninsula and Sumatra, as suggested by
their monophyly as well astheir low ITS sequencedivergence . Wardenia is not supportedas W. simplex ( = B. simplex) is
nested within Brassaiopsis . TheMalayan region is important for the development of Schefflera, andavailableevidence sug-
云 南 植 物 研 究 2008 , 30 (4) : 391～399
Acta Botanica Yunnanica DOI : 10 .3724?SP. J . 1143 .2008.07267
? ?Received date: 2007 - 11 - 09 , Accepted date: 2008 - 02 - 21
作者简介 : 文军 , 女 , 教授 , 主要从事分类学研究。
gests that Schefflera in the region forms a clade with the Heptapleurum group . Dendropanax lancifolius does not form a
clade with the core group of Dendropanax, and itsstatus needs to be further analyzed . Macropanax maingayi was consid-
ered to be a highly distinct member comprising the monotypic genus Hederopsis . Our analysis clearly places it in Macro-
panax . Aralia merrillii was onceconsideredto bethesolemember of thegenus Acanthophorabecauseof its unusual climb-
ing habit . The ITS data support its placement in Aralia . Our expanded sampling of Arthrophyllumcontinues to support its
monophyly . Osmoxylon has aprimarydistribution in the Malesianregion and it isa hylogenetically isolatedmember of Ara-
liaceae .
Key words: Araliaceae; Phylogeny; Malesia; Wardenia; Hederopsis
Araliaceae consist of approximately 45 genera and
1450 species with a cosmopolitan distribution, espe-
cially rich in species in the tropical regions . Three
centers of diversity may be recognizable: the eastern
and southeastern Asia, Australasia, and the neotro-
pics . The Araliaceae of eastern and southeastern Asia
have experiencedmany nomenclatural changes especial-
ly in the last decade, and at present they belong to 28
genera and consist of about 850 species (Table 1) .
The Asian Araliaceaehavebeen treated byvarious
workers, mostly with a regional focus . Linnaeus
(1753 ) described the first species from Asia: Aralia
chinensis L ., based on collections made by Pehr Os-
beck in southern China near Canton ( Guangzhou) in
1752 ( Wen and Reveal , 1992 ) . Philipp Franz von
Siebold, a physician fromthe Netherlands made exten-
Table 1 Generaof Araliaceae in Asia and their distribution
Genus Number of species Distribution
Anakasia Philipson 1 fNew Guinea
Aralia L . ( 1753 ?) 67 xAsia, North America to South America
ArthrophyllumBlume ( 1826 )) 30 xSE Asia to New Guinea and New Caledonia
Brassaiopsis Decne . & Planch . (1854 p) ca . 30 ?C China, Indochina to SE Asia, to the Himalayan region
Chengiopanax Shang & J . Y . Huang (1993 %) 2 fChina and J apan
Cromapanax A . J . C . Grierson (1991 ?) 1 fBhutan
Dendropanax Decne . & Planch . ( 1854 ?) ca . 70 E Asia to S Asia, & C & S America
Fatsia Decne . & Planch . ( 1854 ?) 3 fE Asia
Eleutherococcus Maxim . ( 1859 ?) ca . 35 E Asia & the Himalaya
Gamblea C . B . Clarke (1879 t) 3 fE Asia, the Himalayan region to SE Asia
Gastonia Comm . ex Lam . 9 fMadagascar, Mascarenes, Seychelles, Malesia, to theSolomon Islands
HarmsiopanaxWarb . ( 1897 9) 3 fJ ava to New Guinea
Hedera L . (1753 ?) 16 xTemperate Eurasia
Heteropanax Seem . ( 1865 ?) 8 fS & SW China, Indochina to SE Asia, & the Himalaya
Kalopanax Miq . ( 1863 ?) 1 fE Asia
Macropanax Miq . ( 1856 ?) ca . 15 C China to the Himalaya, to SE Asia
Merrilliopanax Li ( 1942 ?) 4 fW China & the Himalayan region
Metapanax J . Wen & Frodin ( 2001 &) 2 fC & SW China & N Vietnam
Oplopanax Miq . ( 1863 ?) 3 fE Asia & NW N America
Osmoxylon Miq . (1863 ?) 60 xTaiwan to Borneo, the Philippines, to New Guinea and nearby Pacific
islands
Panax L . (1753 ?) 18 xE Asia & E North America
Polyscias J . R . & G . Forster ( 1776 I) ca . 150 ?Old World tropics, only few in Malesia, but commonly cultivated
throughout Asia as ornamentals
Schefflera J . R . & G . Forster (1775 W) ca . 450 ?Asia, only the Asian group is referred to here
Sinopanax Li (1949 9) 1 fTaiwan
Tetrapanax C . Koch (1859 ) 1 fE Asia
Trevesia Vis . (1840 ?) ca . 10 S China to SE Asia
Tupidanthus J . D . Hooker & Thomson ( 1856 F) 1 fS Yunnan to NE India, N Thailand and N Indochina
Woodburnia Prain ( 1904 ?) 1 fN Myanmar
* Schefflera is the largest genus of Araliaceae, with 600 - 900 species . Recent studies have shown that Schefflera is polyphyletic ( Wen et al. , 2001 ;
Plunkett et al. , 2004a, 2005 ) . The 450 or so species in Table 1 refer to only a group of closely related Asian species of Schefflera .
293 云 南 植 物 研 究 30 卷
sive collections of Araliaceae in Japan . The Siebold′s
collections were studied by Zuccarini and published in
their Flora Japonica . Li ( 1942 ) and Hoo and Tseng
(1978 ) revised Araliaceae in China . The Araliaceae
from Vietnam were examined by several workers
(Grushvitzky et al. , 1990a, b; Wen and Lowry2002;
Wen et al. , 2003) . Miguel ( 1863) treated the Arali-
aceae from the Dutch East Indies ( now Indonesia) ,
where he recognized Aralia, Nothopanax Miq ., and
Panax . Philipson (1979) provided a treatment of Ara-
liaceae (except Schefflera) in the Malesian region and
revised several genera from the region ( Philipson,
1973 , 1977 , 1978 ) . Henderson ( 1933 ) , Stone
(e.g ., 1977 , 1978 , 1980) and Frodin (1978 ) pub-
lished a series of papers on Araliaceae fromMalaysia .
Araliaceae of the Malesian region are species-
rich, but the generic endemism is low . Fourteen gen-
era occur in the region and they are Anakasia ( 1
sp .) , Aralia ( 10 spp .) , Arthrophyllum ( 17 spp .) ,
Brassaiopsis ( 8 spp ., including Wardenia King) ,
Dendropanax ( 3 spp .) , Eleutherococcus ( 1 sp .) ,
Gamblea (1 sp .) , Gastonia ( 2 sp .) , Harmsiopanax
(3 spp .) , Macropanax ( 3 spp . including Hederopsis
C . B . Clarke) , Osmoxylon ( ca . 40 spp ., including
Boerlagiodendron Harms) , Polyscias ( ca . 25 spp .) ,
Schefflera ( ca . 450 spp .) , and Trevesia ( 4 spp .)
(Philipson, 1979 ) . Two of the 14 genera ( Anakasia
and Harmsiopanax) are endemic to the Malesian re-
gion, although Osmoxylon is nearly endemic there
(Philipson, 1979; Frodin and Govaerts, 2003) . Three
genera were also included in Araliaceae by Philipson
(1979 ) and they are now excluded from the family
based on recent phylogenetic evidence: Aralidium
Miq . ( now in Aralidiaceae) , Delarbrea Vieill . ( now
in Myodocarpaceae) and Mackinlaya F . Muell . ( now
in Apiaceae) (Plunkett et al. , 2004b) .
Objectivesof this study are to provide a prelimi-
nary analysis on the evolutionary relationships and di-
versification of Araliaceae in theMalesian region . Spe-
cifically we try to assess the taxonomic status of several
morphologically unusual members including Acan-
thophora Merr . (now in Aralia) , Hederopsis, which is
now included in Macropanax (Philipson, 1979 ) , and
Wardenia, which is now recognized as an element of
Brassaiopsis .
Materials and Methods
Sequences of 107 species representing 31 species from the
Malesian region and 76 species of Araliaceae from the other re-
gions throughout the distribution range of the family ( Table 2 ) .
Pittosporumresiniferumof the closely related Pittosporaceae was
selected as the outgroup . We selected the nuclear ribosomal ITS
region as our molecular marker because a large number of se-
quences arealready availablefor Araliaceae (Wen and Zimmer,
1996; Wen et al. , 2001 , 2003; Plunkett et al. , 2004a) . Fifty-
one sequences were newly generated in this study, and 57 se-
quenceswere obtained fromGenBank .
Total DNAs were extracted following Doyle and Doyle
(1987) . Theamplification and sequencing of the nuclear riboso-
mal ITS regions followed Wen andZimmer ( 1996) . Phylogenetic
analyseswereperformedwith PAUP* ver . 4 . 0 (Swofford, 2002)
usingmaximum parsimony in which alignment gaps were treated
as missing data or as new characters . The maximum parsimony
analysis was performed using a heuristic search with MULPARS
and furthest-addition sequenceoptions . The amount of support for
monophyletic groups revealed in the maximally parsimonious tree
(s) (MPTs) was also examined with 500 bootstrap replicates
(Felsenstein, 1985 ) .
Nucleotide substitution model parameters were determined
for the ITS data set using MODELTEST ver . 3 .0 ( Posada and
Crandall , 1998) . Bayesian analyses (Rannala and Yang, 1996;
Mau et al. , 1999) were carried out using MrBayes ver . 3 .0b3
(Huelsenbeck and Ronquist, 2001) with the model parameters .
Bayesian analyses started from random trees and employed four
Markov chains Monte Carlo ( mcmc) runs, monitored over one
million generations, re-sampling trees every 100 generations .
Runswere repeated twice to confirm results . The resulting log
likelihood and number of generations were plotted to determine
the point after whichthe log likelihoods had stabilized . After dis-
carding thetrees savedprior to this point as burn-in, the remain-
ing treeswere imported into PAUP anda50 % majority-rule con-
sensus tree was produced to obtain posterior probabilities of the
clades .
Results
The ITS data set had 734 aligned positions with
405 variable sites, 298 of which are parsimony-infor-
mative . Parsimony analysis of the ITS data set generat-
ed 13782 most parsimonious trees ( MPTs) with a
length of 1510 steps, a consistency index ( CI) of 0 .44
3934 期 WEN Jun et al . : Evolutionary Relationships of Araliaceae in the Malesian Region: a Preliminary Analysis
(0 . 39 excludinguninformativecharacters) and a reten- tion index ( RI ) of 0 .68 ( Fig . 1) .
Table 2 Accessions of Araliaceae sampled in the study (The 51 new sequences generated in this study are boldfaced)
Taxon Voucher Origin
GenBank
accession
Aralia bipinnata Blanco Wen 8276 3-1 ( F ) Philippines, Luzon DQ007354 ?
A ?. californica S . Watson Ornduff 8931 ( CS) USA , California AF551720
A ?. dasyphylla Miq . Widjaja 7574 ( F) Indonesia, Java DQ007355
A ?. excelsa (Griseb .) J . Wen Chiang s ?.n . ( CS) Mexico AF242231
A ?. foliolosa Wall . ex C . B . Clarke Pandey 5009 ( F ) India, West Bengal AY233312
A ?. gigantea J . Wen Pandey 5002 ( F ) India, West Bengal AY233313
A ?. humilis Cav . Wen 4974 3( F ) USA , Arizona AF242230
A ?. leschenaultii ( DC .) J . Wen Wen 4907 3( F ) Nepal AY394569
A ?. merrillii Shang Wen 8393 3( F ) Malaysia, Selangor DQ007356
A ?. nudicaulis L . Wen 849 ?( A ) USA , North Carolina U41674 Z
A ?. scopulorumBrandegee Wen 565 ?( OS) Mexico, Baja California U66927 Z
A ?. spinosa L . Wen & Dong 976 ?( A ) USA , Georgia U66928 Z
A ?. subcordata ( Don ) J . Wen Wen 5818 3( F ) India, Shillong DQ007357
A ?. thomsonii Seem . Wen 6168 3( F ) Vietnam, Ninh Binh DQ007358
Arthrophyllumahernianum Merr . Wen 8308 3( F ) Philippines, Luzon DQ007359 ?
A ?. diversifoliumBlume Wen 8372 3( F ) Malaysia, Perak DQ007361
A ?. mackeei Lowry, ined . Lowry 4670 ](MO) New Caledonia U63182 Z
A ?. montanum Ridley Wen 8365 3( F ) Malaysia, Pahang DQ007362
A ?. ovalifoliumJungh . & de Vriese Wen 8339 3( F ) Malaysia, Pahang DQ007360
Astrotricha pterocarpa Benth . Plunkett 1527 ?(MO) Australia, Queensland U63189 Z
Brassaiopsis aculeata Seem . Pathak & Bhaumik 4447 ?(CAL ) India, Arunachal Pradesh AY725110
B ?. elegans Ridley Wen 8408 3( F ) Malaysia, Selangor DQ007363
B ?. ficifolia Dunn Wen 7453 3( F ) Thailand, Chiangmai DQ007403
B ?. glomerulata ( Blume) Regel Lowry 4839 ](MO) Vietnam AY256901
B ?. hainla (Buch .-Ham .) Seem Wen 4905 3( F ) Nepal AY389028
B ?. malayana J . Wen & Frodin , ined . Wen 8378 3( F ) Malaysia, Malay Peninsula DQ007367
B ?. mitis C . B . Clarke Pandey 5005E (BHAG) India, West Bengal AF551726
B ?. polyacantha ( Wall .) R . N . Banerjee Wen 8367 3( F ) Malaysia, Pahang DQ007364
B ?. simplex ( King ) B . C . Stone Wen 8357 3-4 ( F ) Malaysia, Malay Peninsula DQ007366
B ?. sumatrana Ridley Wen 8406 3( F ) Malaysia, Selangor DQ007365
Cephalaralia cephalobotrys ( F . Muell .) Harms Plunkett 1519 ?(MO) Australia, Queensland AF229762
Cheirodendron trigynum ( Gaudich .) A . Heller Lammers & Holody 5677 ?(CHR) Hawaii , Maui U63183 Z
Cuphocarpus briquetianus Bernardi Lowry 5329 ](MO) Madagascar AY430378 ?
Cussonia paniculata Eckl . & Zeyh . Wen 8480 3( F ) Cult . in Chelsea Physic Garden, London DQ007368 ?
Dendropanax dentiger (Harms ) Merr . Wen 5539 3( F ) China, J iangxi DQ007369 ?
D ?. hainanensis ( Merr . & Chun) Chun Deng s. n . ( IBSC) China, Hunan DQ007402
D ?. lancifolius ( Ridley) Ridley Wen 8362 3( F ) Malaysia, Pahang DQ007370
D ?. maingayi King Wen 8364 3( F ) Malaysia, Pahang DQ007371
D ?. sessiliflorus (Standl . & A . C . Sm .) A . C . Sm . Wen 7002 3( F ) Costa Rica, Puntarenas Prov . DQ007401
D ?. trifidus (Thunb .) Makino ex Hara Wen 2461 3( F ) J apan, Tokyo AF242238
Eleutherococcus sieboldianus ( Makino) Koidz . Wen 7149 3( F ) Cult . in Oak Park, Illinois DQ007372 ?
E ?. trifoliatus ( L .) S . Y . Hu Wen 8301 3( F ) Philippines, Luzon DQ007373
Fatsia japonica ( Thunb .) Decne . & Planch . Mitchell s *. n . ( CHR 502463 ) Cult . in New Zealand U63163 Z
Gamblea malayana ( M . R . Henderson ) Shang ,
Lowry & Frodin
Wen 8361 3( F ) Malaysia, Pahang DQ007374 ?
Gastonia curtispongia Lam . Lowry 4976 ](MO) Cult . in Réunion AF229722 ?
Harmsiopanax aculeata K . Schum . Widjaja 7562 ?( F) Cult . in Bali DQ007376
H ?. harmsii K . Schum . Widjaja 7563 ( F) Cult . in Bali DQ007375
H ?. ingens Philipson Lowry 5266 ](MO) Irian Jaya AY389038
Hedera sinensis (Tobler) Hand .-Mazz . Wen 6083 3( F ) Vietnam, Laocai AY304817 ?
Heteropanax fragrans ( Roxb .) Seem . Wen 7414 3( F ) Thailand, Mae Hong Son Prov . DQ007377 ?
Kalopanax septemlobus (Thunb .) Koidz . Wen 3093 3( F ) China, Sichuan AY256899 ?
Macropanax dispermus ( Blume) Kuntze Wen 6058 3( F ) Vietnam, Laocai DQ007378 ?
493 云 南 植 物 研 究 30 卷
Continue table 2
Taxon Voucher Origin
GenBank
accession
M ?. maingayi ( C . B . Clarke) Philipson Wen 8355 3( F ) Malaysia, Selangor DQ007379
M ?. rosthornii ( Harms) C . Y . Wu ex G . Hoo Wen 5087 3( F ) China, J iangxi AF551738
M ?. undulatus ( Wall . ex G . Don) Seem . Lowry 4947 ](MO) Vietnam AY389039
Meryta denhamii Seem . Wen s ?.n . ( F ) Cult . in Belgium DQ007380
Metapanax davidii ( Franch .) J . Wen & Frodin Wen 1409 3( CS) China, Hubei AF242233 ?
M ?. delavayi ( Franch .) J . Wen & Frodin Wen 1217 3( CS) China, Yunnan AF242232
Munroidendron racemosum (C . N . Forbes) Sherff Plunkett 1342 ?( WS) Cult . in Missouri Botanical Garden AF229738
Neopanax arboreus (L . f .) Allan RBGE19981866 ?Cult . in Royal Botanic Garden Edinburgh DQ007381
Oplopanax elatus ( Nakai) Nakai Wen 5407 3( F ) China, J ilin AY389043 ?
Oreopanax argentatus ( Kunth ) Decne . & Planch Wen 6224 3( F ) Cult . in Belgium DQ007400 ?
O ?. echinops ( Schltdl . & Cham .) Decne . & Planch . Chiang 1278 z( CS) Cult . in Mexico City, Mexico AF242229
Osmoxylon geelvinkianumBecc . Plunkett 1489 ?(MO) New Guinea AF229727
O ?. lineare ( Merr .) Philipson Wen 7064 3( F ) Cult . in Hawaii DQ007382
O ?. novoguineense ( Scheff .) Becc . National Tropical Botanical
Garden Acc . 810791002 ?
Cult . in National Tropical Botanical
Garden
AF229726
Panax assamicus R . N . Banerjee Pandey 5000 ?( F ) India, Darjeeling AY233320
P ?. ginseng C . A . Meyer Wen 1250 3( CS) China, Heilongjiang U41680 Z
P ?. pseudoginseng Wall . Wen 4900 3( F ) Nepal AY275936
P ?. stipuleanatus Tsai & Feng Wen 5631 3-5 ( F ) China, Yunnan AY275945
P ?. trifolius L . Kramer & Kramer s Z. n . (CS) USA , Ohio U41698
Pittosporum resiniferum Hemsl . Wen 8286 3( F ) Philippines, Benguet Prov . DQ007353 ?
Polyscias cissodendron (C . Moore& F . Muell .) Harms Lowry 4749 ](MO) Madagascar AF229693 ?
P ?. javanica Koord . & Valeton Widjaja 7564 ( F) Cult . in Bali DQ007383
P ?. nodosa (Blume) Seem . Lowry 5286 ](MO) Cult . in Bogor Bot . Gard . AY430387
P ?. samoensis ( A . Gray ) Harms Wen 7078 3( F ) Cult . in Hawaii DQ007384
Pseudopanax ferox Kirk RBGE 19992225 Cult . in Royal Botanic Garden Edinburgh DQ007385 ?
P ?. latevirens ( Gay) Franch . Wen 7428 3-1 ( F ) Chile, Valdivia DQ007386
Raukaua anomalus ( Hook .) A . D . Mitchell , Frodin
& M . Heads
Acc # 18225 ?( CHR 500664) New Zealand, Peel Forest Nursery U63164 Z
Reynoldsia sandwicensis A . Gray Plunkett 1359 ?( WS) Hawaii AF229739
Schefflera brenesii A . C . Smith Wen 6998 3( F ) Costa Rica, Puntarenas Prov . DQ007387 ?
S ?. delavayi (Franch .) Harms Li 13830 ?( F ) China, Yunnan DQ007391
S ?. digitata J . R . Forst . & G . Forst . Mitchell s *. n . ( CHR 485642 ) Cult . in Lincoln, New Zealand U63188 Z
S ?. elegantissima (Veitch ex Mast .) Lowry & Frodin Wen 2488 3( CS) Cult . in Fort Collins, Colorado AF242239 ?,
AF242240
S ?. heterophylla ( Wall . ex G . Don ) Harms Wen 8392 3( F ) Malaysia, Selangor DQ007388
S ?. hullettii ( King) R . Vig . Wen 8377 3( F ) Malaysia, Perak DQ007392
S ?. impressa (C . B . Clarke) Harms Pathak & Bhaumik 4495 (CAL ) India, Arunachal Pradesh AY725132
S ?. microphylla Merr . Wen 8280 3( F ) Philippines, Benguet Prov ., DQ007389
S ?. oblongifolia Merr . Wen 8323 3( F ) Philippines, Ifugao Prov . DQ007390
S ?. oxyphylla ( Miq .) R . Vig . Wen 8396 3( F ) Malaysia, Selangor DQ007393
S ?. sp . Wen 8360 3( F ) Malaysia, Pahang DQ007394
S ?. yunnanensis H . L . Li Wen 5028 3( F ) China, Yunnan AY389060
Sinopanax formosanus ( Hayata) H . L . Li Lowry 4967 ](MO) China, Taiwan AF229768 ?
Tetrapanax papyrifer ( Hook .) K . Koch Mitchell s *. n . ( CHR 502422 ) Cult . In New Zealand U63192 Z
Tetraplasandra hawaiiensis A . Gray Wen 7075 3( F ) Hawaii DQ007395 ?
T ?. oahuensis ( A . Gray) Harms Lorence 8158 ( PTBG) Cult . in National Tropical Botanical
Garden
AF229740
Trevesia beccarii Boerl . Widjaja 7575 ?( F) Cult . in Bogor Bot . Gard . DQ007396
T ?. burckii Boerl . Wen 8358 3( F ) Malaysia, Perak DQ007398
T ?. burckii Boerl . Wen 8399 3( F ) Malaysia, Selangor DQ007397
T ?. lateospina Jebb Wen 7480 3-2 ( F ) Thailand, Lampang Prov . DQ007399
T ?. palmata ( DC .) Vis . Hao 901 ( F) Cult . in South China Bot Gard AF242247
T ?. sundaica Miq . Lowry 5285 ](MO) Cult . Bogor Bot Garden AY389063
5934 期 WEN Jun et al . : Evolutionary Relationships of Araliaceae in the Malesian Region: a Preliminary Analysis
Fig . 1 The strict consensus tree of the Araliaceae from the Malesian region and the close relatives based on thenuclear ribosomal ITS
sequences ( bootstrap values above 50 % shown below the branches, and the Bayesian posterior probabilities above 95 % were
indicated by a“ * ”above the branches) . The Malesian species were in boldface
693 云 南 植 物 研 究 30 卷
Discussions
Araliaceae fromthe Malesian region had a diffuse
origin ( Fig . 1 ) . The Malesian taxa are scattered
throughout the phylogeny . The Malesian region is geo-
graphically located at the converging point of two major
centers of species diversity of Araliaceae: the eastern
and southeastern Asian center, and the Australasian
center .
Aralia consists of 69 species with most species in
the north temperate zone ( Wen, 1993 , 2002 , 2004 ) .
Aralia merrillii of Malaysia and the Philippines was
initially described to be amonotypic member of the ge-
nus Acanthophora Merr . as Acanthophora scandens
Merr . (Merrill , 1918) , because of its unusual climb-
ing habit in the tropical evergreen forests . Our analysis
clearly places it within Aralia (also see Wen, 2004) .
Brassaiopsis is composed of about 30 species with
themajority of the species in the Himalayas, western
China, Indochina and Malay Peninsula ( Mitchell and
Wen, 2005 ) . The Malay Peninsula has about seven
species, two of which ( B. minor and B. simplex) were
treated as membersof adistinct genus, Wardenia . Our
ITS sequence data support merging Wardenia with
Brassaiopsis, because the type species of Wardenia,
W. simplex King ( = Brassaiopsis simplex) is nested
within Brassaiopsis and in fact showed little sequence
divergence with B. elegans ( pairwise sequence diver-
gence of 0 .452% . Furthermore, the Malayan species
of Brassaiopsis seemto belong to two clades of the ge-
nus, one to the B. glomerulata - B. hainla - B. pol-
yacantha clade (clade A) , and theother to B. elegans
- B. simplex - B. malayana - B. sumatrana clade
(cladeB) . Taxaof clade B are fromsoutheastern Asia
(Malay Peninsula and Sumatra) and the sequence di-
vergence among themwas low ( ranging from 0 .437%
between B. malayana and B. elegans, to 0 .954% be-
tween B. simplex and B. sumatrana) , suggesting a re-
cent origin of these taxa .
Hederopsis was described as a monotypic genus
consisting of H. maingayi King ( 1898 ) endemic to
Malaya and Sumatra (Stone, 1977 ) . The genus was
accepted by Stone (1977 , 1980) when he was treating
theMalayanAraliaceae . Stone (1977) contrasted Hed-
eropsis with Macropanax and supported the separation
of the two genera based on the locular numbers in the
ovary (5 - 6 in Hederopsis vs . 2 - 3 in Macropanax) .
Philipson ( 1979 ) treated Hederopsis as a synonym of
Macropanax ( also see Frodin and Govaerts, 2003 ) .
Our ITS sequencedata suggest that Hederopsis is nested
within Macropanax and thus support merging Hederop-
sis with Macropanax .
Harmsiopanax is a Malesian genus consisting of
three species and is characterized by its prickly stems,
palmately lobed simple leaves, non-articulated pedi-
cels, bicarpellate bristly ovary, and its unusual fruits
consisting of two dry mericarps ( Philipson, 1973 ) .
Philipson ( 1979 ) noted that species of the genus are
monocarpic ( i . e ., fruiting only once) . This habit
needs better documentation . Philipson ( 1979 ) pointed
out that the genus shows many characters of Umbellif-
erae . Our recent studies place the genus in Araliaceae
(Plunkett et al. , 2004a) and the ITS sequence data
here put it in the large clade of Aralia, Panax, Ar-
throphyllum, Polyscias, and their close relatives (Fig .
1) , although the bootstrap support is low ( < 50% ) .
Harmsiopanax appears to be isolated phylogenetically .
It forms a monophyletic group with Aralia and Panax
in the 50% majority-rule consensus tree, but is unre-
solved in the strict consensus tree . Further analysis is
needed to resolve its position within Araliaceae .
Arthrophyllum, agenus of about 30 species, has
a current center of diversity inMalesia also extendingto
theAndaman islandsof India, Laos and NewCaledonia
( Philipson, 1977 , 1979; Frodin and Govaerts,
2003) . Philipson ( 1978 ) merged Eremopanax of New
Caledonia with smooth, wrinkled to ruminate endo-
spermand normally thick and stony endocarp with Ar-
throphyllum (with deeply ruminate endosperm and en-
docarp never thick or stone-like) . The genus as cir-
cumscribed by Philipson is highly distinct from other
generamorphologically by its umbellately arranged in-
florescences with opposite leaf-like bracts ( Lim,
1986) , 1-locular ovaries, and 1-seeded fruits . Our
study sampled the more species of Arthrophyllum than
previous analyses ( Wen et al. , 2001; Plunkett et al. ,
2004) and continues to support the monophyly of the
7934 期 WEN Jun et al . : Evolutionary Relationships of Araliaceae in the Malesian Region: a Preliminary Analysis
Malesian and New Caledonian Arthrophyllum ( Fig .
1 ) . The widely distributed A. diversifolium of the
western Malesian region is closely related to A. aher-
nianum from the Philippines . The morphologically
dwarf and divergent A. montanum forms a well-sup-
ported clade with A. ahernianum and A. ovalifolium .
Philipson ( 1979) treated A. ovalifolium as a synonym
of A. diversifolium . Our ITS sequencedata suggest that
A. diversifoliums. l . is not monophyletic .
Dendropanax is a genus of about 75 species with
an intercontinental disjunct distribution between tropi-
cal?subtropical Asia and theneotropical regions inCen-
tral and South America . Only three species occur in
Malesia: D. borneensis ( Philipson) Merr ., D. lan-
cifolius, and D. maingayi . We sampled D. lancifolius
and D. maingayi fromthe Cameron Highlands, Malay-
sia . The two species are actually sympatric there, with
D. lancifolius as a large tree of ca . 20 m tall , and
D. maingayi as a shrub of 2 - 3 mtall in the montane
cloud forest . Phylogenetically the two taxa are not
closely related . D. maingayi forms a clade with the
core Dendropanax clade . D. lancifolius is one of the
two unusual lineages ( the other being D. hainanensis
from China and Indochina) that we can unresolved
phylogenetic position . These two taxa are large trees
and have branched and more elaborate inflorescences .
The monophyly of Dendropanax is not supported in our
ITS analysis ( Fig . 1 ) . A broader sampling scheme is
needed totest thenon-monophyly of thegenus and con-
struct its biogeographic diversification history .
Schefflera is the largest genus of Araliaceae and
has been shown to be polyphyletic ( Wen et al. , 2001;
Plunkett et al. , 2004a, 2005) .TheMalesian region is
highly species-rich for the genus and possesses up to
450 species with many yet-to-be described and new to
science (Frodin and Govaerts, 2003 ) . In this study,
our samplingof Schefflera fromthe region are fromthe
Philippines and Malay Peninsula . We sampled the
Heptapleurum group ( S. hulletii , S. heterophylla,
S. microphylla and S. oxyphylla) and the Cephalos-
chefflera group ( S. oblongifolia Merr .) (sensu Frodin,
1975 , 1986 ) . The two groups form a well-supported
clade with other Heptapleurum taxa from eastern Asia
and the Himalayas ( e.g ., S. bengalensis, S. hy-
poleuca, S. impressa, and S. yunnanensis, as well as
Tupidanthus Hook . f . & Thomson .
Osmoxylon, a genus of 60 species, has a wide
distribution in Malesia, western Melanesia to Vanuatu
and is especially well developed in the Philippines and
Solomon Islands ( Frodin and Govaerts, 2003 ) . The
relationships within the genus are poorly understood .
Morphologically thegenus is characterized by the ligu-
late stipules and the marked petiole base with several
spiral or transversal crests . The inflorescence is a ter-
minal compound umbel with primary rays terminating
into three branches ( the lateral branches longer than
the central one) . The genus is a phylogenetically iso-
latedmember of Araliaceae and becomeswell developed
in Malesia and on the Pacific islands .
Acknowledgements : We thank the organizing committee of the
Sixth International Flora Malesiana Symposium for their efforts of
organizing the successful meeting, Richard Chung, Sam Yen
Yen, Ruth Kiew, Alang Bin Mahayu, Yee ChongChan, Kama-
rudin Saleh, EdwinoS . Fernando, and Leonardo Cofor herbari-
um and field assistance . The Pritzker Laboratory for Molecular
Systematics andEvolutionandtheBassFellowship Programof the
Field Museumsupported ElizabethWidjaja′s research visit to the
USA .
Reference :
Doyl ?e JJ , Doyle JL , 1987 . A rapid DNA isolation procedure for small
quantities of fresh leaf material [ J ] . Phytochem Bull , 19 : 11—15
Fels ?enstein J , 1985 . Confidence intervals on phylogenetics: an approch
using bootstrap [ J ] . Evolution, 39 : 783—791
Frod ?in DG, 1975 . Studies in Schefflera ( Araliaceae) : the Cephaloschef-
flera complex [ J ] . J Arnold Arbor, 56 : 427—448
Frod ?in DG, 1978 . Schefflera [ A ] . In: FSP Ng, Phil D eds, Tree Flora
of Malaya, vol . 3 [ M] . Singapore: Longman, 25—35
Frod ?in DG, 1986 . Studies in Schefflera ( Araliaceae) , II . Northern Lu-
zon ( Philippines) species of the Heptapleurumgroup [ J ] . Proc Acad
Nat Sci Philadelphia, 138: 403—425
Frod ?in DG, Govaerts R , 2003 . World Checklist and Bibliography of Ara-
liaceae [ M] . Kew: Royal Botanic Gardens
Grus ?hvitzky IV, Skvortsova NT , Ha TD, 1990a . Species in the tribe
PseudopanaceaeSeem . ( Araliaceae) in the flora of Neit Nam [ J ] .
Novost Sist Vyssh Rast, 27 : 104—113 ( in Russian)
Grus ?hvitzky IV , SkvortsovaNT , Ha TD et al. , 1990b . Conspectus of the
family Araliaceae Juss . in the flora of Vietnam [ J ] . Novost Sist
Vyssh Rast, 22 : 153—192 ( in Russian)
Hend ?erson MR , 1933 . Additions to the flora of the Malay peninsula [ J ] .
893 云 南 植 物 研 究 30 卷
Gard Bull Straits Settlem, 7 : 87—128
Hoo *G, Tseng CJ , 1978 . Angiospermae, dicotyledoneae, Araliaceae .
Flora ReipublicaePopularis Sinicae, Vol . 54 [M ] .Beijing: Science
Press
Huel senbeck JP , Ronquist F , 2001 . MrBayes: Bayesian inference of
phylogenetic trees [ J ] . Bioinformatics, 17 : 754—755
Li H L , 1942 . The Araliaceae of China [ J ] . Sargentia, 2 : 1—134
Lim AL , 1986 . The comparative inflorescence structure in five Malaysian
species of Arthrophyllum ( Araliaceae) [ J ] . Kew Bull , 41 : 769—
780
Linn ?aeus C , 1753 . Species plantarum, Stockholm
Mau -B , Newton M, Larget B , 1999 . Bayesian phylogenetic inferencevia
Markov chain Monte Carlo methods [ J ] . Biometrics, 55 : 1—12
Merr ?ill ED, 1918 . New or noteworthy Philippineplants, XIV [ J ] . Philip
J Sci , 13 : 263—333
Miqu +el FAW, 1863 . Araliaceae novae [ J ] . Ann Mus Bot Lugduno-Bata-
vum, 1 : 1—27
Mitc ?hell A , Wen J , 2005 . Phylogeny and evolutionary radiation of Bras-
saiopsis ( Araliaceae) in Asia, based on nuclear ( ITS and 5S-NTS)
DNA sequences [ J ] . Syst Bot, 30 : 872—886
Phil ?ipson WR , 1973 . A revision of Harmsiopanax ( Araliaceae) [ J ] .
Blumea, 21 : 81—86
Phil ?ipson WR , 1977 . A revision of the Malesian species of Arthrophyllum
Bl . ( Araliaceae) [ J ] . Gardens Bull Singapore, 30 : 299—312
Phil ?ipson WR, 1978 . The identity of Arthrophyllum and Eremopanax
( Araliaceae) [ J ] . Adansonia, 17 : 329—333
Phil ?ipson WR , 1979 . Araliaceae [ A ] . In: CGGJ van Steenis ed . Flora
Malesiana, Vol . 9 [ M ] . The Hague: Martinus Nijhoff Publishers,
1—105
Plun ?kett GM, Wen J , Lowry II PP, 2004a . Infrafamilial relationships and
characters in Araliaceae: insights from the phylogenetic analysis of
nuclear ( ITS) and plastid ( trnL-trnF ) sequence data [ J ] . Plant
Syst Evol , 245: 1—39
Plun ?kett GM, Chandler GT, Lowry II PP et al. , 2004b . Recent advances
in understanding Apiales and a revised classification [ J ] . S Africa J
Bot, 70 : 371—381
Plun ?kett GM , Lowry II PP, Frodin DG et al. , 2005 . Phylogeny and ge-
ography of Schefflera : pervasivepolyphyly in the largest genusof Ara-
liaceae [ J ] . Ann Missouri Bot Gard, 92 : 202—224
Posa ?da D, Crandall KA , 1998 . Modeltest: testing the model of DNA
substitution [ J ] . Bioinformatics, 14 : 817—818
Rann ?ala B, Yang ZH , 1996 . Probability distribution of molecular evolu-
tionary trees: A new method of phylogenetic inference [ J ] . J Mol
Evol , 43 : 304—311
Sieb ?old PF , Zuccarini JG, 1845 . Florae Japonicae Familiae Naturales,
pars prima: Plantae Dicotyledoneae Polypetalae [ J ] . Abh Math . -
PhysCl K?nigl Bayer Akad Wiss, 4 : 109—204
Ston ?e BC, 1977 . Notes on the systematy of Malayan phanerogams XXV .
Araliaceae [ J ] . Gard Bull Straits Settlem, 30 : 275—291
Ston ?e BC, 1978 . Araliaceae except Schefflera [ A ] . In: FSP Ng, Phil D
eds, . Tree Flora of Malaya . Vol . 3 [ M ] . Singapore: Longman,
11—25
Ston ?e BC, 1980 . A nomenclatural change in Malaysian Araliaceae:
Eleutherococcus malayanus ( Henderson) comb . nov . instead of Ac-
anthopanax malayanus Hend [ J ] . Malaysian For, 43 : 395
Swof ?ford DL , 2002 . PAUP : Phylogenetic Analysis UsingParsimony ( and
Other Methods) . Version 4 .0b10 [M ] . Sunderland, Massachusetts:
Sinauer
Wen J , 1993 . Generic delimitation of Aralia ( Araliaceae) [ J ] . Britton-
ia, 45 : 47—55
Wen ?J , 2001 . Evolution of the Aralia-Panax complex ( Araliaceae) as in-
ferred fromnuclear ITS ribosomal sequences [ J ] . Edinb J Bot, 58 :
243—257
Wen J , 2002 . Revision of Aralia sect . Pentapanax ( Seem .) In: J Wen
( Araliaceae) [ J ] . Cathaya, 13 - 14: 1—116
Wen J , 2004 . Systematics and biogeography of Aralia L . sect . Dimor-
phanthus (Miq .) Miq . ( Araliaceae) [ J ] . Cathaya, 15 - 16: 1—
188
Wen ?J , Reveal JL , 1992 . On the typification of the Linnean species of
Aralia ( Araliaceae) [ J ] . Taxon, 41 : 69—75
Wen J , Zimmer EA , 1996 . Phylogeny and biogeography of Panax L .
(the ginseng genus, Araliaceae) : Inferences from ITS sequences of
nuclear ribosomal DNA [ J ] . Mol Phylogen Evol , 6 : 167—177
Wen ?J , Lowry II PP, 2002 . Aralia hiepiana J Wen & Lowry, anewspe-
cies of Araliaceae fromVietnam [ J ] . Adansonia, 24 : 213—216
Wen ?J , Plunkett GM , Mitchell AD et al. , 2001 . The evolution of Arali-
aceae: aphylogenetic analysis based on ITS sequences of nuclear ri-
bosomal DNA [ J ] . Syst Bot, 26 : 144—167
Wen ?J , Lee C , Lowry II PP et al. , 2003 . Inclusion of the Vietnamese
endemic Grushvitzkya in Brassaiopsis ( Araliaceae) : Evidence from
nuclear ribosomal and chloroplast DNA [ J ] . Bot J Linn Soc, 142:
455—463
9934 期 WEN Jun et al . : Evolutionary Relationships of Araliaceae in the Malesian Region: a Preliminary Analysis