全 文 ：·快讯 Short Communication·
基于线粒体 matR基因序列初步探讨三白草科的系统发育
孟少武1　陈之端2　李德铢1　梁汉兴1
(1.中国科学院昆明植物研究所植物生物多样性与生物地理学开放研究室 , 昆明 650204;
2.中国科学院植物研究所系统与进化植物学开放研究实验室 , 北京 100093)
摘要:　首次报道了三白草科 3 个种及其外类群齐头绒(Zippelia begoniaefolia Bl.)的 matR 基因序列。研究材料包括
三白草科现存的所有 6个种和两个选定外类群 ,分析软件是 PAUP 4.0b 4a。分别采用最大简约法 、距离法和最大似
然法进行分析 ,都得到一个同样的系统发育树 , 只是支持率有所不同。结果显示三白草科是一个单系 , Anemopsis 是
Houttuynia 的姐妹群 ,并且 , 它们构成三白草科的第一个分支 , 是 Gymnotheca_Saururus 支的姐妹群 。Gymnotheca 和
Saururus 的姐妹群关系也被强烈支持。
关键词:　三白草科;系统发育;matR 基因序列
中图分类号:Q941+.2　　　文献标识码:A　　　文章编号:0577-7496(2001)06-0653-04
Phylogeny of Saururaceae Inferred from matR Sequence Data
MENG Shao_Wu1 , CHEN Zhi_Duan2 , LI De_Zhu1 , LIANG Han_Xing1
(1.Laboratory for Plant Biodiversity and Biogeography , Kunming Inst itute of Botany , The Chinese Academy of Sciences , Heilongtan , Kunming 650204 , China;
2.Laboratory of Systematic and Evolutionary Botany , Institute of Botany , The Chinese Academy of Sciences , Beijing 100093, China)
Key words:　Saururaceae;phylogeny;matR gene sequences
　　Saururaceae , a member of paleoherbs , is an ancient
and relic family with 6 species in 4 genera , i.e.Sauru-
rus , Gymnotheca , Anemopsis and Houttuynia[ 1] .They
are perennial herbs with simple flowers.The flowers have
bracts and no perianths.Saururaceae is typically East
Asian_ North American disjuncted , with Anemopsis and
Saururus cernuus in North America , Houttuynia , Gym-
notheca and Saururus chinensis in East Asia.Due to its
importance of systematic position and interesting geo-
graphical distribution pattern , Saururaceae has been a key
family although it only includes several species.Howev-
er , the viewpoints on the phylogeny of Saururaceae are
very different based on morphology.Wu and Wang[ 2]
posed a systematic tree of Saururaceae based mainly on
the gross morphology and geography , including Saururus ,
Circaeocarpus , Anemopsis , Houttuynia and Gymnotheca
in Saururaceae.Later , however , Wu andWang[ 3] detect-
ed that the new genus , Circaeocarpus , was in fact a
member of Piperaceae , and Circaeocarpus saururoides and
Zippelia begoniaefolia were conspecific.Based on the ba-
sic chromosome numbers of some genera in Saururaceae ,
Okada[ 4] put forward that Anemopsis and Houttuynia were
derived from Saururus respectively , and they were at the
same advanced level.According to chromosome number ,
Lei et al[ 5] supported Okada s opinion , and thought that
Gymnotheca was the most advanced genus.On the basis
of a cladistic analysis for some conventional morphological
and ontogenetic characters , Tucker et al[ 6] regarded that
Saururus was originated from the ancestor of Saururaceae ,
and subsequently , Gymnotheca.Houttuynia was the sis-
ter group of Anemopsis.Gymnotheca was sister to Hout-
tuynia_Anemopsis clade , while Saururus to Gymnotheca_
Houttuynia_Anemopsis clade.Combining the data from
gross morphology , anatomy , embryology , palynology , cy-
tology and flower development[ 7-11] , Liang[ 1] proposed
that the systematic order of Saururaceae should be Sauru-
rus , Gymnotheca , Anemopsis and Houttuynia.She
thought Saururaceae derived two branches at early times ,
one was Gymnotheca_Anemopsis branch , the other was
Saururus_Houttuynia.
An overall research on the molecular systematics of
Saururaceae is very much needed although some genera
were ever involved in current studies on high_rank classi-
fication of the angiosperms[ 12-14] .As a part of a compre-
hensive study of the Saururaceae , we reported the phy-
logeny of Saururaceae based on matR gene sequences.
Received:2000-12-19　Accepted:2001-03-15
Supported by the Special Grant of the National Natural Science Foundation of China(39725001 to LI De_Zhu), the National Natural Science Foundation of China
(39930020)and Hundreds of Talents Program of the Chinese Academy of Sciences [ 1998] 0054(LI De_Zhu).
植　物　学　报　2001 , 43(6):653-656 　 　 　 　 　 　 　
Acta Botanica Sinica
1　Materials and Methods
Four species , Zippelia begoniaefolia Bl., Gym-
notheca chinensis Decne., G. involucrata Pei and
Saururus chinensis (Lour.)Baill., were collected from
natural populations or cultivated plants.Vouchers are de-
posited in the herbarium of Kunming Institute of Botany ,
the Chinese Academy of Sciences , Kunming , Yunnan
Province (KUN).The sequences of other four species are
cited from GenBank (marked with brackets)(Table 1).
Our procedures for molecular techniques followed
Kong and Chen
[ 15] .The primers matR 26F(5′_GACCGC
TNACAGTAGTTCT_3′) and matR 1858R (5′_
TGCTTGTGGGCYRGGGTGAA_3′) were used for both
PCR amplification and sequencing , while matR 879F
(5′_ACTAGTTATCAGGTCAGAGA_3′) and 1002R(5′_
CACCCACGATTCCCAGTAGT_3′)only for sequencing.
The four primers were designed by Mark W.Chase s
Molecular Systematic Laboratory at Kew Botanic Garden ,
United Kingdom.
2　Results
The sequenced lengths of matR DNA sequences are
following:Saururus chinensis , 1735 bp;S .cernuus ,
1775 bp;Gymnotheca chinensis , 1778 bp;G.involu-
crata , 1751 bp;Houttuynia cordata , 1804 bp;Anemop-
sis californica , 1831 bp;Zippelia begoniaefolia , 1747
bp;Piper betle , 1831 bp.When gaps were treated as
missing data , there are 1771 total aligned sites for matR
DNA after cutting the uneven head and tail of the matrix ,
and 1697 sites were constant , 44 sites were variable but
parsimony_uninformative , 30 sites were parsimony_infor-
mative.The percentage of phylogenetic informative sites
was 1.69%.Sequence divergence ranged from 0.057%
to 1.753% among Saururaceae species and from 1.439%
to 2.206%between the two outgroups and ingroup taxa.
When all eight sequences were analyzed through the
maximum parsimony method with branch and bound
search , a single most parsimonious tree was obtained.
Saururaceae was supported as a monophyletic group , with
a bootstrap value of 97%.Anemopsis was sister to Hout-
tuynia with a bootstrap value of 84%, and Saururus to
Gymnotheca with a bootstrap value of 95%.Anemopsis
and Houttuynia consisted of the sister group of Gymnothe-
ca_Saururus clade(Fig.1).Generally speaking , the re-
sult is different from all the other phylogenetic opinions on
Saururaceae proposed by previous authors[ 1-6] to some
extent.
As the matrix was analyzed through the neighbor_
joining search of the distance method , and the branch and
bound search of the likelihood method , respectively , only
one tree with the same topology as Fig.1 was generated by
either method.Moreover , the sister group relationships
between Anemopsis and Houttuynia , and between Sauru-
rus and Gymnotheca were also supported.In distance
method , Anemopsis sister to Houttuynia was moderately
supported by a bootstrap value of 66%, Saururus sister to
Gymnotheca was also highly supported by a bootstrap val-
ue of 96%.In likelihood method , Anemopsis sister to
Houttuynia was moderately supported by a bootstrap value
of 60%, Saururus sister to Gymnotheca was also highly
supported by a bootstrap value of 83%.
3　Discussion
matR is a mitochondrial functional gene that encodes
a maturase_related protein[ 16] .Generally , the matR gene
sequences are used to compare the systematic relationships
among families or higher_rank taxonomic groups because it
is highly conservative.However , there were 44 variable
sites and 30 informative ones when gaps were treated as
missing data so that we could well distinguish genera and
even species in Saururaceae through the analysis of matR
gene sequences(Fig.1).Moreover , the bootstrap values
were high.Therefore , matR DNA sequence data might be
valuable for the phylogenetic reconstruction among genera
of some ancient and relic families such as Saururaceae.
Table 1　The species sampled in this study and the GenBank accession numbers of their matR sequences
Species Collection number GenBank accession number
Piper betle L. (From GenBank) AF197750
Zippelia begoniaefolia Bl. Meng 99715 AF332107
Anemopsis californica(Nutt.)Hook. (From GenBank) AF197747
Gymnotheca chinensis Decne. Meng 99003 AF332103
G.involucrata Pei Liang 97015 AF332105
Houttuynia cordata Thunb. (From GenBank) AF197749
Saururus cernuus L. (From GenBank) AF197748
S.chinensis(Lour.)Baill. Meng 99525 AF332106
654　 植　物　学　报　　Acta Botanica Sinica 43 卷
Fig.1.　The single most parsimonious tree of Saururaceae based on matR sequences.
Length = 84 , CI=0.9167 , RI=0.8158 , RC=0.7478.Number of bootstrap replicates = 1 000.Base substitutions are indicated above
branches , and bootstrap values (%)below branches.◆, without flower_bract stalk;□, with stalk.
　　The sister group relationships between Anemopsis
and Houttuynia , and between Saururus cernuus and S .
chinensis were both supported by matR sequences.These
are partly in agreement with Wu① , but different from
Okada
[ 4] , Lei et al[ 5] , Tucker et al[ 6] and Liang[ 1] .
Based on Fig.1 , Gymnotheca chinensis and G.involu-
crata were also a pair of sister group.They should be dif-
ferent species of the same genus as Pei[ 17] and Zhang and
Lei[ 18] proposed , although their morphological variation is
not very high.
The topology in this analysis shows that Anemopsis_
Houttuynia clade is the sister group of Gymnotheca_
Saururus clade.However , many researchers based on the
morphology of Saururaceae thought that Saururus was the
most primitive in Saururaceae because they paid more at-
tention to some characters such as the position of ovaries
or stamina , the number of stamina , but neglected the
great variation of flower organs.Tucker[ 19] observed the
high variation of the number of stamina and carpels in
Saururus cernuus.So did Liang[ 10] in Gymnotheca.The
flower_bract stalks are absent in Anemopsis , Houttuynia
and two outgroups , but present in Saururus and Gym-
notheca.It supports the topology of Saururaceae in the ar-
ticle.
Acknowledgments:　We express our hearty thanks to
Ms.Huang Cha_Cha , Ms.Sun Ying_Xue , Mr.Hong
Ya_Ping , Ms.Song Bao_Hua and Mr.Yang Jun_Bo for
their assistance during the experimental period;to Prof.
Wang Xiao_Quan for helpful discussion , and to Prof.Ge
Song for critical review on the manuscript.
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656　 植　物　学　报　　Acta Botanica Sinica 43 卷