全 文 ：Received: 2014–04–30　　　　Accepted: 2014–06–15
The project was supported by the National Natural Science Foundation of China (30770156, 31070177, 31110103911), and Main Direction Program of
Knowledge Innovation of Chinese Academy of Sciences (KSCX2-EW-Z-1).
* Corresponding author. E-mail: wangrj@scbg.ac.cn
热带亚热带植物学报　2014， 22(5)： 431 ~ 442
Journal of Tropical and Subtropical Botany
基于分子和形态学证据确定传统中药白花蛇舌草
及其替代品的学名
王瑞江*, 邓淑君, 廖琦
(中国科学院华南植物园，中国科学院植物资源保护与可持续利用重点实验室，广州 510650)
摘要： 白花蛇舌草是我国重要的传统中药，主要是指茜草科(Rubiaceae)钮扣草族(Spermacoceae)的 Oldenlandia diffusa，但伞房
花耳草(O. corymbosa)在民间或中药市场也常被作为替代品使用。由于长期以来 Hedyotis-Oldenlandia 复合群的分类存在许多
争论，因此白花蛇舌草有时被归入非洲耳草属(Oldenlandia L.)，有时又作为广义耳草属(Hedyotis L. s. l.)的成员。为了澄清白花
蛇舌草命名上的问题，基于 7 个叶绿体片段和 2 个核基因片段对钮扣草族 85 个分类群进行了系统发育分析。结果表明，白花
蛇舌草不属于以上两属中的任何一属，而应为蛇舌草属[Scleromitrion (Wight & Arn.) Meisn.]的成员。依此结果，对 5 种植物进
行了新组合，并提供了白花蛇舌草和伞房花耳草的形态学比较，以有助于在实践中更好地进行区分。
关键词： 白花蛇舌草； 分子系统学； 非洲蛇舌草属； 蛇舌草属； 中药
doi: 10.3969/j.issn.1005–3395.2014.05.001
Nomenclature Clarification of the Traditional Chinese Medicine Baihua-
sheshecao and Its Adulterants Based on Molecular and Morphological
Evidence
WANG Rui-jiang*, DENG Shu-jun, LIAO Qi
(Key Laboratory of Plant Resources Conservation and Sustainable Uitlization, South China Botanical Garden, Chinese Academy of Sciences,
Guangzhou 510650, China)
Abstract: Baihuasheshecao is one of the important traditional Chinese medicines and mainly refers to Oldenlandia
diffusa but can be adulterated with O. corymbosa in commercial and folk herbal medicine. Because of the long-
time taxonomic debate and confusion about the taxonomy of Hedyotis-Oldenlandia complex, the scientific names
of Baihuasheshecao components were treated under the genus of either Oldenlandia or Hedyotis in practice. In
order to clarify the nomenclatural confusion, a comprehensive molecular phylogenetic analysis was conducted
on basis of seven plastid loci and two nuclear gene regions from the selected 85 taxa in the tribe Spermacoceae
(Rubiaceae). The results demonstrated that O. diffusa should not be a member of Oldenlandia L. but Scleromitrion
(Wight & Arn.) Meisn. Five new combinations are accordingly made and a detailed comparison between the two
species is provided for identification in future practice.
Key words: Baihuasheshecao; Molecular phylogeny; Oldenlandia; Scleromitrion; Traditional Chinese medicine
432 第22卷热带亚热带植物学报
Baihuasheshecao is a popular traditional medicine
for anti-inflammatory, hepatoprotective, neuroprotective,
antioxidant, antipyretic, diuretic, blood-stimulant, anti-
carbuncular effects and enormous potential in the
therapy of cancer and tumor in many Asian countries[1–3].
It is also an important ingredient of herbal tea for
health maintenance in the Eastern and tropical Asia[4].
According to Pharmacopoeia of the People’s Republic
of China (2005), Baihuasheshecao mainly refers to
the species Oldenlandia diffusa, but can be substituted
by and interchanged with O. corymbosa in the official
medication[5]. However, it is also usually adulterated
with O. pinifolia and O. verticillata etc. in folk.
It was reported that Oldenlandia corymbosa has
antipyretic purposes in traditional Thai medicine[6]
and hepatoprotective effect known as “Parppatakapullu”
in India[7]. And many recent studies revealed that
O. diffusa and O. corymbosa had different chemical
compositions and bioactivities[4,8–9].
Hedyotis-Oldenlandia complex belongs to the
tribe Spermacoceae of the family Rubiaceae. The
centuries’ taxonomic debate and confusion about
the congeneric or separate between Hedyotis L. and
Oldenlandia L. bewildered not only the plant taxonomists
for species description but the traditional Chinese
physicians for medicinal practice. Consequently, most
species described under Hedyotis almost certainly has
a synonym under Oldenlandia, vice versa. This caused
almost arbitrary or random nomenclature choices[10].
Hao et al.[11] doubted the unrelated phylogenetic
relationship between O. diffusa and O. corymbosa,
but they did not uncover it explicitly due to the lack
of enough research materials at that time. Recent
phylogenetic study based on large scale samples
from Asian countries verified the previous doubt
and revealed that these two species belonged to two
different clades representing different evolutionary
lineages[10,12]. In the present study, a comprehensive
molecular phylogenetic analysis was conducted on
basis of seven plastid loci (petD, rps16, trnL-trnF,
atpB-rbcL, matK, rbcL, trnH-psbA) and two nuclear
gene regions (ETS, ITS) of 85 selected taxa from the
newly defined tribe Spermacoceae[13]. This study
not only is an integration of previous series results
but will present a deep analysis for this complicate
complex to reveal the phylogenetic relationship of
different components of Baihuasheshecao and clarify
their taxonomic and nomenclature confusions.
1 Methods
1.1 Taxon sampling
Referring to the several recent publications[10,12–14],
54 ingroup taxa related to Oldenlandia diffusa or O.
corymbosa, as well as 31 associated taxa, were selected
for the present phylogenetic analysis. The generic
name Oldenlandia, rather than Hedyotis, was chosen
for the ingroup taxa (if have) so as to keep consistency
with the previous phylogenetic analysis. Detailed
information about species names used currently and
previously, vouchers, and GenBank accession No. is
given in Table 1.
1.2 Pre-treatment for the taxon names used in this
　paper
Some scientific names of the selected taxa were
pre-treated in order to address the taxonomic issues
explicitly.
The name Oldenlandia erecta of Li Heng 11298
(A) from China and Neupane 2 (ODU) from Nepal[10]
was replaced by O. corymbosa var. linearis according
to the treatments by Deb and Dutta[15] and Dutta and
Deb[16].
Oldenlandia tenelliflora has ever been doubted to
be possibly indistinct from Scleromitrion angustifolium
(Cham. & Schlechtd.) Benth.[17]. The detailed examination
about the type materials of O. tenelliflora from Java
showed that it has ovate and 1–1.7 cm wide (vs.
linear or linear-lanceolate, 0.3–0.9 cm wide in S.
angustifolium) leaves, 7–9-bristled (vs. 1–4-bristled in
S. angustifolium) stipules, and septicidally from base
to apex and then loculicidally (vs. loculicidally from
apex only in S. angustifolium) dehiscent capsules. The
Chinese samples are in agreement with the characters
of S. angustifolium.
Moreover, the samples of YU05, YU06, and
第5期 433
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王瑞江等：基于分子和形态学证据确定传统中药白花蛇舌草及其替代品的学名
434 第22卷热带亚热带植物学报
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68
/
JX
11
10
80
JX
11
11
55
JX
11
11
90
28
O
ld
en
la
nd
ia
c
ap
en
si
s
L.

f.
Za
m
bi
a:
D
es
se
in
e
t a
l.
84
3
(B
R
)
A
M
93
29
74
A
M
93
94
96
EU
55
77
37
EU
54
30
48
EU
54
31
33
EU
54
29
80
/
/
/
29
O
. c
ap
en
si
s
L.
f.
v
ar
.
pl
ei
os
ep
al
a
B
re
m
ek
.
Ta
nz
an
ia
: K
ay
om
be
e
t
al
. s
.n
. (
B
R
)
A
M
93
29
75
A
M
93
94
97
EU
55
77
38
EU
54
30
49
EU
54
31
34
EU
54
29
81
/
/
/
30
O
. c
or
ym
bo
sa
L
.
A
us
tra
lia
: A
nd
er
ss
on

22
60
(G
B
)
A
M
93
29
77
A
M
93
95
00
/
/
/
/
/
/
/
31
O
. c
or
ym
bo
sa
L
.
C
hi
na
: M
ac
au
M
A
25

(C
U
H
K
)
/
H
Q
14
88
02
/
H
M
75
29
60
H
M
75
28
75
/
H
M
75
31
30
H
M
75
30
45
H
M
64
03
60
32
O
. c
or
ym
bo
sa
L
.
C
hi
na
: R
ui
jia
ng
W
an
g
14
27
(I
B
SC
)
/
JF
69
99
25
JF
70
00
74
/
/
/
JF
69
99
99
JF
69
98
49
JF
69
97
84
33
O
. c
or
ym
bo
sa
L
.
C
hi
na
: T
ai
w
an
T
A
01

(C
U
H
K
)
/
H
Q
14
87
57
/
H
M
75
29
15
H
M
75
28
30
/
H
M
75
30
85
H
M
75
30
00
H
M
64
03
15
34
O
. c
or
ym
bo
sa
L
.
C
hi
na
: X
in
g
G
uo
5
0
(I
B
SC
)
/
JF
69
99
27
JF
70
00
76
JX
11
13
05
JX
11
13
79
/
JF
70
00
01
JF
69
98
51
JX
11
11
94
35
O
. c
or
ym
bo
sa
L
.
G
ab
on
: A
nd
er
ss
on
&

N
ils
so
n
22
63
(G
B
)
A
M
93
29
78
A
M
93
95
01
/
/
/
/
/
/
/
续
表
(C
on
tin
ue
d)
第5期 435
N
o.
N
am
e
us
ed
in
th
is
p
ap
er
N
am
e
us
ed
w
hi
le

se
qu
en
ce
d*
Vo
uc
he
r
(H
er
ba
riu
m
)
ET
S
IT
S
pe
tD
rp
s1
6
tr
nL
- tr
nF
at
pB
- r
bc
L
m
at
K
rb
cL
tr
nH
- p
sb
A
36
O
. c
or
ym
bo
sa
L
.
Si
ng
ap
or
e:
R
ui
jia
ng

W
an
g
SI
N
02
(I
B
SC
)
/
JX
11
12
39
JX
11
11
21
JX
11
13
06
JX
11
13
80
/
JX
11
10
83
JX
11
11
58
JX
11
11
93
37
O
. c
or
ym
bo
sa
L
.
Za
m
bi
a:
D
es
se
in
e
t a
l.
48
7
(B
R
)
A
M
93
29
79
A
M
93
95
02
EU
55
77
39
EU
54
30
50
EU
54
31
35
EU
54
29
82
/
/
/
38
O
. c
or
ym
bo
sa
L
. v
ar
.
li
ne
ar
is
(D
C
.)
Ve
rd
c.
O
ld
en
la
nd
ia
e
re
ct
a
(M
an
ila
l &
S
iv
ar
.)
R
. R
. M
ill
C
hi
na
: L
i H
en
g
11
29
8
(A
)
H
E6
81
55
4
H
E6
57
75
8
H
E6
57
64
3
H
E6
49
89
7
/
/
/
/
/
39
O
. c
or
ym
bo
sa
L
. v
ar
.
li
ne
ar
is
(D
C
.)
Ve
rd
c.
O
ld
en
la
nd
ia
e
re
ct
a
(M
an
ila
l &
S
iv
ar
.)
R
. R
. M
ill
N
ep
al
: S
. N
eu
pa
ne
2

(O
D
U
)
H
E6
81
54
1
H
E6
57
74
5
H
E6
57
63
0
H
E6
49
88
2
/
/
/
/
/
40
H
ed
yo
ti
s
co
ry
m
bo
sa
L
.
va
r.
te
re
ti
ca
ul
is
K
o
C
hi
na
: X
in
g
G
uo
e
t a
l.
16
(I
B
SC
)
/
JF
69
99
26
JF
70
00
75
JX
11
13
07
JX
11
13
78
/
JF
70
00
00
JF
69
98
50
JX
11
11
95
41
O
ld
en
la
nd
ia
d
en
sa

(in
ed
.)
Za
m
bi
a:
D
es
se
in
e
t a
l.
34
6
(B
R
)
A
M
93
29
80
A
M
93
95
03
EU
55
77
51
EU
54
30
61
EU
54
31
47
/
/
/
/
42
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
C
hi
na
: H
on
g
K
on
g
TY
01
(C
U
H
K
)
/
H
Q
14
87
59
/
H
M
75
29
17
H
M
75
28
32
/
H
M
75
30
87
H
M
75
30
02
H
M
64
03
17
43
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
C
hi
na
: M
ac
au
M
A
19

(C
U
H
K
)
/
H
Q
14
87
96
/
H
M
75
29
54
H
M
75
28
69
/
H
M
75
31
24
H
M
75
30
39
H
M
64
03
54
44
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
C
hi
na
: R
ui
jia
ng
W
an
g
14
49
(I
B
SC
)
/
JF
69
99
31
JF
70
00
80
/
/
/
JF
70
00
05
JF
69
98
55
JF
69
97
88
45
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
C
hi
na
: T
an
C
e-
m
in
g
95
67
0
(U
PS
)
H
E6
81
53
9
H
E6
57
74
3
H
E6
57
62
8
H
E6
49
88
0
/
/
/
/
/
46
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
C
hi
na
: X
in
g
G
uo
5
1
(I
B
SC
)
/
JF
69
99
32
JF
70
00
81
JX
11
13
08
JX
11
13
81
/
JF
70
00
06
JF
69
98
56
JF
69
97
89
47
O
. d
if
fu
sa
(W
ill
d.
) R
ox
b.
O
ld
en
la
nd
ia

br
ac
hy
po
da
D
C
.
N
ep
al
: S
. N
eu
pa
ne
8
8
(O
D
U
)
/
H
E6
57
73
5
H
E6
57
62
0
H
E6
49
87
1
/
/
/
/
/
48
O
. d
ue
m
m
er
i S
. M
oo
re
U
ga
nd
a:
W
. H
. L
ew
is

60
18
(G
H
)
H
E6
81
54
0
H
E6
57
74
4
H
E6
57
62
9
H
E6
49
88
1
/
/
/
/
/
49
O
. g
al
io
id
es
(F
. M
ue
ll.
)
F.
M
ue
ll.
A
us
tra
lia
: H
ar
w
oo
d
15
11
(B
R
)
/
A
M
93
95
07
EU
55
77
43
EU
54
30
53
EU
54
31
39
EU
54
29
86
/
/
/
50
O
. g
ra
ci
li
pe
s
C
ra
ib
Th
ai
la
nd
: J
. F
. M
ax
w
el
l
01
-5
91
(L
)
H
E6
81
54
2
H
E6
57
74
6
/
H
E6
49
88
3
/
/
/
/
/
51
O
. h
er
ba
ce
a
(L
.)
R
ox
b
C
hi
na
: H
on
g
K
on
g
H
u
&
B
ut
2
24
91
(A
)
/
H
E6
57
74
9
H
E6
57
63
3
H
E6
49
88
6
/
/
/
/
/
52
O
. l
an
ci
fo
li
a
(S
ch
um
ac
h.
) D
C
.
Za
m
bi
a:
D
es
se
in
e
t a
l.
12
56
(B
R
)
A
M
93
29
76
A
M
93
94
99
/
/
/
/
/
/
/
53
O
. l
an
ci
fo
li
a
(S
ch
um
ac
h.
) D
C
.
Za
m
bi
a:
D
es
se
in
e
t a
l.
13
56
(B
R
)
A
M
93
29
90
A
M
93
95
12
/
EU
54
30
58
EU
54
31
44
EU
54
29
91
/
/
/
续
表
(C
on
tin
ue
d)
王瑞江等：基于分子和形态学证据确定传统中药白花蛇舌草及其替代品的学名
436 第22卷热带亚热带植物学报
N
o.
N
am
e
us
ed
in
th
is
p
ap
er
N
am
e
us
ed
w
hi
le

se
qu
en
ce
d*
Vo
uc
he
r
(H
er
ba
riu
m
)
ET
S
IT
S
pe
tD
rp
s1
6
tr
nL
- tr
nF
at
pB
- r
bc
L
m
at
K
rb
cL
tr
nH
- p
sb
A
54
O
. n
em
at
oc
au
li
s
B
re
m
ek
.
Za
m
bi
a:
D
es
se
in
e
t a
l.
92
4
(B
R
)
A
M
93
29
94
A
M
93
95
17
/
EU
54
30
60
/
EU
54
29
94
/
/
/
55
O
. o
va
ti
fo
li
a
(C
av
.)
D
C
.
H
ed
yo
ti
s
ov
at
if
ol
ia

C
av
.
C
hi
na
: X
in
g
G
uo
e
t a
l.
20
-1
(I
B
SC
)
/
JF
69
99
40
JF
70
00
90
JX
11
13
09
JX
11
13
82
/
JF
70
00
15
JF
69
98
65
JF
69
97
95
56
O
. p
in
if
ol
ia
(W
al
l.
ex
G
.
D
on
) K
un
tz
e
H
ed
yo
ti
s
pi
ni
fo
li
a
W
al
l.
ex
G
. D
on
C
hi
na
: H
on
g
K
on
g
Y
U
19
(C
U
H
K
)
/
H
Q
14
88
21
/
H
M
75
29
79
H
M
75
28
94
/
H
M
75
31
49
H
M
75
30
64
H
M
64
03
79
57
O
. p
in
if
ol
ia
(W
al
l.
ex
G
.
D
on
) K
un
tz
e
H
ed
yo
ti
s
pi
ni
fo
li
a
W
al
l.
ex
G
. D
on
C
hi
na
: M
ac
au
M
A
11

(C
U
H
K
)
/
H
Q
14
87
88
/
H
M
75
29
46
H
M
75
28
61
/
H
M
75
31
16
H
M
75
30
31
H
M
64
03
46
58
O
. p
in
if
ol
ia
(W
al
l.
ex
G
.
D
on
) K
un
tz
e
C
hi
na
: R
ui
jia
ng
W
an
g
12
31
(I
B
SC
)
/
JX
11
12
40
JX
11
11
22
JX
11
13
11
JX
11
13
84
/
JX
11
10
84
JX
11
11
59
JX
11
11
96
59
O
. s
to
ck
si
i H
oo
k.
f.
In
di
a:
K
la
ck
en
be
rg
&

Lu
nd
in
3
26
(S
)
H
E6
81
55
8
H
E6
57
76
3
H
E6
49
90
1
60
O
. t
ab
or
en
si
s
B
re
m
ek
.
Ta
nz
an
ia
: B
id
go
od
e
t a
l.
40
15
(B
R
)
/
A
M
93
95
22
EU
55
77
53
/
EU
54
31
49
EU
54
29
96
/
/
/
61
O
. u
m
be
ll
at
a
L.
In
di
a:
S
. N
eu
pa
ne
8
4
(O
D
U
)
H
E6
81
56
5
H
E6
57
77
0
H
E6
57
65
3
H
E6
49
90
8
/
/
/
/
/
62
O
. u
m
be
ll
at
a
L.
Sr
i L
an
ka
: F
. F
ag
er
lin
d
33
20
(S
)
H
E6
81
46
9
H
E6
57
67
4
H
E6
57
56
9
H
E6
49
80
6
/
/
/
/
/
63
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: H
on
g
K
on
g
Y
U
12
(C
U
H
K
)
/
H
Q
14
88
15
/
H
M
75
29
73
H
M
75
28
88
/
H
M
75
31
43
H
M
75
30
58
H
M
64
03
73
64
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: M
ac
au
M
A
14

(C
U
H
K
)
/
H
Q
14
87
91
/
H
M
75
29
49
H
M
75
28
64
/
H
M
75
31
19
H
M
75
30
34
H
M
64
03
49
65
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: R
ui
jia
ng
W
an
g
14
38
(I
B
SC
)
/
JF
69
99
65
JF
70
01
15
/
/
/
JF
70
00
40
JF
6 9
98
90
JF
69
98
16
66
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: R
ui
jia
ng
W
an
g
40
9
(I
B
SC
)
/
JF
69
99
66
JF
70
01
16
JX
11
13
12
JX
11
13
85
/
/
JF
69
98
91
JF
69
98
17
67
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: X
in
g
G
uo
6
6
(I
B
SC
)
/
JF
69
99
69
JF
70
01
19
JX
11
13
13
JX
11
13
86
/
JF
70
00
43
JF
69
98
94
JF
69
98
19
68
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: X
in
g
G
uo
e
t a
l.
4-
1
(I
B
SC
)
/
JF
69
99
67
JF
70
01
17
/
/
/
JF
70
00
41
JF
69
98
92
/
69
O
. v
er
ti
ci
ll
at
a
L.
H
ed
yo
ti
s
ve
rt
ic
il
la
ta
(L
.)
La
m
.
C
hi
na
: X
in
g
G
uo
e
t a
l.
4-
2
(I
B
SC
)
/
JF
69
99
68
JF
70
01
18
/
/
/
JF
70
00
42
JF
69
98
93
JF
69
98
18
续
表
(C
on
tin
ue
d)
第5期 437
N
o.
N
am
e
us
ed
in
th
is
p
ap
er
N
am
e
us
ed
w
hi
le

se
qu
en
ce
d*
Vo
uc
he
r
(H
er
ba
riu
m
)
ET
S
IT
S
pe
tD
rp
s1
6
tr
nL
- tr
nF
at
pB
- r
bc
L
m
at
K
rb
cL
tr
nH
- p
sb
A
70
O
. w
au
en
si
s
Sc
hw
ei
nf
.
ex
H
ie
rn
T
he
co
rc
hu
s
w
au
en
si
s
(S
ch
w
ei
nf
. e
x
H
ie
rn
) B
re
m
ek
.
Et
hi
op
ia
: F
rii
s e
t a
l.
25
60
(U
PS
)
A
M
93
30
18
A
M
93
95
48
EU
55
77
74
EU
54
30
76
EU
54
31
68
EU
54
30
17
/
/
/
71
O
. w
ie
de
m
an
ni
i K
.
Sc
hu
m
.
K
en
ya
: L
uk
e
&
L
uk
e
83
62
(U
PS
)
A
M
93
30
01
A
M
93
95
25
EU
55
77
56
EU
54
30
63
EU
54
31
51
EU
54
30
00
/
/
/
72
P
ar
ak
no
xi
a
pa
rv
if
lo
ra

(S
ta
pf
e
x
Ve
rd
c.
) V
er
dc
.
ex
B
re
m
ek
.
Za
m
bi
a:
D
es
se
in
e
t a
l.
67
8
(B
R
)
/
/
EU
55
7 7
57
EU
54
30
64
EU
54
31
52
EU
54
30
01
/
/
/
73
P
en
to
do
n
pe
nt
an
dr
us

(S
ch
um
ac
h.
&
T
ho
nn
.)
Va
tk
e
Za
m
bi
a:
D
es
se
in
e
t a
l.
59
8
(B
R
)
A
M
93
30
03
A
M
93
95
28
EU
55
77
59
EU
54
30
66
EU
54
31
54
EU
54
30
02
/
/
/
74
Sc
le
ro
m
it
ri
on

an
gu
st
if
ol
iu
m
(C
ha
m
. &

Sc
hl
ec
ht
d.
) B
en
th
.
H
ed
yo
ti
s
an
gu
st
if
ol
ia
C
ha
m
.
&
S
ch
ltd
l.
C
hi
na
: S
hi
u-
Y
in
g
H
u
54
01
(A
)
H
E6
81
45
5
H
E6
57
66
2
/
H
E6
49
79
2
/
/
/
/
/
75
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: H
on
g
K
on
g
(T
M
05
)
/
H
Q
14
87
62
/
H
M
75
29
20
H
M
75
28
35
/
H
M
75
30
90
H
M
75
30
05
H
M
64
03
20
76
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: H
on
g
K
on
g
(T
M
06
)
/
H
Q
14
87
63
/
H
M
75
29
21
H
M
75
28
36
/
H
M
75
30
91
H
M
75
30
06
H
M
64
03
21
77
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: X
in
g
G
uo
e
t a
l.
12
(I
B
SC
)
/
JF
69
99
57
JF
70
01
08
JX
11
12
97
JX
11
13
70
/
JF
70
00
33
JF
69
98
82
JF
69
98
10
78
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: X
in
g
G
uo
e
t a
l.
19
(I
B
SC
)
/
JF
69
99
59
JF
70
01
09
/
/
/
JF
70
00
34
JF
69
98
83
/
79
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
U
nk
no
w
n:
c
ul
t.
at
B
R
A
M
93
29
49
A
M
93
94
51
EU
55
77
10
EU
54
30
62
EU
54
31
06
EU
54
29
97
/
/
/
80
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: L
iu
Z
he
ng
yu

21
84
0
(M
O
)
H
E6
81
56
0
H
E6
57
76
5
H
E6
57
64
8
H
E6
49
90
3
/
/
/
/
/
81
S.
a
ng
us
ti
fo
li
um
(C
ha
m
.
&
S
ch
le
ch
td
.)
B
en
th
.
O
ld
en
la
nd
ia

te
ne
ll
if
lo
ra
B
lu
m
e
C
hi
na
: R
ui
jia
ng
W
an
g
99
2
(I
B
SC
)
/
JF
69
99
58
JF
70
01
07
JX
11
12
96
JX
11
13
71
/
JF
70
00
32
JF
69
98
84
JF
69
98
11
82
St
en
ar
ia
n
ig
ri
ca
ns

(L
am
.)
Te
rr
el
l
U
SA
: Y
at
sk
ie
vy
ch
9
6-
92

(M
O
)
/
A
M
93
95
46
EU
55
77
72
A
F3
33
37
3
EU
54
31
66
EU
54
30
15
/
/
/
83
Sy
na
pt
ea
nt
ha
ti
ll
ae
ac
ea

(F
.M
ue
ll.
) H
oo
k.
f.
A
us
tra
lia
: L
az
ar
id
es
&

Pa
lm
er
2
72
(K
)
/
A
M
93
95
47
EU
55
77
73
EU
54
30
75
EU
54
31
67
EU
54
30
16
/
/
/
84
T
he
ca
go
nu
m
b
if
lo
ru
m

(L
.)
B
ab
u
H
ed
yo
ti
s
bi
fl
or
a
(L
.)
La
m
.
C
hi
na
: R
ui
jia
ng
W
an
g
11
46
(I
B
SC
)
/
JF
69
99
06
JF
70
00
55
JX
11
13
00
JX
11
13
74
/
JF
69
99
81
JF
69
98
30
JF
69
97
67
85
T
. p
te
ri
tu
m
(B
lu
m
e)

B
ab
u
H
ed
yo
ti
s
pt
er
it
a
B
lu
m
e
C
hi
na
: R
ui
jia
ng
W
an
g
14
23
(I
B
SC
)
/
JF
69
99
42
JF
70
00
92
/
/
/
JF
70
00
17
JF
69
98
67
JF
69
97
97
*:
T
he
b
la
nk
st
an
ds
fo
r t
ha
t t
he
n
am
e
is
th
e
sa
m
e
as
c
ur
re
nt
ly
u
se
d;
/:
M
is
si
ng
se
qu
en
ce
s.
续
表
(C
on
tin
ue
d)
王瑞江等：基于分子和形态学证据确定传统中药白花蛇舌草及其替代品的学名
438 第22卷热带亚热带植物学报
YU07 from Hong Kong, China, identified as “Hedyotis
herbacea” in NCBI database and as “Oldenlandia
diffusa” by Guo et al.[12], should be H. koana, which
has terminal and upper axillary inflorescence, on basis
of our comprehensive voucher examination.
In addition, Oldenlandia brachypoda of Nepalese
sample (Neupane 88, ODU) identified by Wikström
et al.[10], was merged under O. diffusa, following the
treatment of Dutta and Deb[16].
1.3 Sequence assembly, editing and phylogenetic
　analyses
Sequences for each region were prealigned with
the Clustal X Version 2.1[18] and ambiguously aligned
regions were manually corrected using BioEdit Version
7.1.11[19].
The Bayesian analyses (BI) were carried out using
MrBayes version 3.2.1[20]. Each of the nine regions
was assigned its own best-fit nucleotide substitution
model, as determined by the Akaike information criterion
(AIC) in Modeltest Version 3.7[21]. The best models
were GTR+I+G (ITS), TVM+I+G (ETS, atpB-rbcL),
TVM+G (petD, trnH-psbA, and trnL-trnF), GTR+G
(matK, rps16), TrN+I (rbcL). Searches were based
on 10000000 generations with four chains of the
Markov Chain Monte Carlo (MCMC, one cold and
three heated under default heating values) in each
of two parallel runs, with each chain starting with a
random tree. Trees were sampled and recorded every
1000 generations of the MCMC chain. Analyses were
set to automatically stop when the average standard
deviation of split frequencies under 0.01, which indicates
the convergence of two runs. A 50% majority-rule
consensus tree was constructed after removing the
“burn-in period” samples (the first 25% of sampled
trees).
2 Results
Phylogenetic relationships indicated by the MCMC
analyses are summarized as a 50% majority-rule
consensus tree in Figure 1 (CI=0.589, RI=0.794). The
posterior probability values are indicated above each
node. Nodes with posterior probability values equal to
or greater than 95% are considered well-supported[22].
The topology of phylogenetic tree is almost
correspondent to that revealed previously. All
investigated species recognized under Oldenlandia,
except O. ovatifolia, are resolved into two main
monophyletic clades with both robust support values
(BPP=100). And O. corymbosa and O. diffusa are
included in the clade I and the clade II, respectively
(Fig. 1).
The strongly supported clade I comprises 19
terminals, of which, eleven are O. corymbosa and its
varieties from different localities. Except for the probably
misidentified Zambian sample (Dessein et al. 487),
all other O. corymbosa samples from pantropical
areas of Africa, Asia and Australia are grouped in the
monophyletic lineage. In addition, O. corymbosa var.
linearis and H. corymbosa var. tereticaulis were also
included in the subclade IA with robust support too
(BPP=100). Other accompanied eight taxa are from
either Africa or South Asia.
The second clade II with robust support includes
nine samples under the name O. diffusa from Asia
and 26 samples representing other nine species from
Asia and Africa (only O. lancifolia). However,
O. herbaceae on the basis of the collection Hu & But
22491 (A) from Hong Kong, China[10], is nested within
the subclade IIA of O. diffusa from China and Nepal.
3 Discussion
3.1 Oldenlandia diffusa and O. corymbosa belong to
two different clades, representing two different
genera
The phylogenetic tree involving the two main
components of Baihuasheshecao indicated that O.
diffusa and O. corymbosa are actually two species
belonging to two different lineages with robust support
values (Fig. 1). As suggested previously[10,12–13,23],
these two lineages actually represent two different
genera, Oldenlandia L. (clade I) and Scleromitrion
(Wight & Arn.) Meisn. (clade II). Morphologically,
they can be distinguished by inflorescence and flower
第5期 439
Fig. 1 Phylogenetic relationships of Baihuasheshecao, Oldenlandia diffusa and its adulterant O. corymbosa, revealing that they belong to two distinct
clades in the tribe Spermacoceae. The tree is a 50% majority-rule consensus tree from a Bayesian Markov Chain Monte Carlo (MCMC) analysis of a
combined dataset of seven plastid (petD, rps16, trnL-trnF, atpB-rbcL, matK, rbcL, trnH-psbA) and two nuclear (ETS, ITS) data. Bayesian Posterior
Probability (BPP) is indicated above branches. The number following the species name refers to the collection number of each sample listed in Table 1.
王瑞江等：基于分子和形态学证据确定传统中药白花蛇舌草及其替代品的学名
440 第22卷热带亚热带植物学报
characters. Oldenlandia usually has small erect or
prostrate stems, terminal or axillary panicles with
obvious or very short peduncles and pedicelled flowers,
loculicidally dehiscent capsules and trigonous seeds.
On the contrary, Scleromitrion has erect stems, either
axillary clusters of sessile flowers or a single flower
with a long and slim pedicel, apically and loculicidally
dehiscent capsules, and many obconic seeds. And
the homostylous androecia and gynoecia are usually
inserted within the corolla tube in Oldenlandia, but
exserted in Scleromitrion[12].
3.2 Oldenlandia clade including the type O. corymbosa
Oldenlandia corymbosa is a common weed in
tropical continents and the western Pacific Islands and
was reported to origin from Africa[24]. Hitchcock[25]
designated O. corymbosa as the lectotype of the genus,
although it was usually treated as a member of Hedyotis
for a very long time. Moreover, the close relationship
between O. corymbosa and O. umbellata has also gotten
support due to their similarity of pollen characters[26].
Besides, Hedyotis corymbosa var. tereticaulis is involved
in this Oldenlandia clade and should be transferred
accordingly.
Oldenlandia corymbosa L. var. tereticaulis (Ko)
R. J. Wang, comb. nov.
Basionym: Hedyotis corymbosa (L.) Lam. var.
tereticaulis Ko, Fl. Hainan. 3: 308, 580, 1974.
3.3 The species in the clade II are accommodated
by the genus Scleromitrion
The monophyletic clade II including O. diffusa
represented the resurrected genus Scleromitrion with
the generic type S. angustifolium[12]. Wikström et al.[10]
indicated that the Chinese O. herbacea (Hu & But
22491, A) that is nested within the subclade IIA is
clearly a representative of some other species because
the true African O. herbacea was proved to be sisterhood
to the genus Conostomium (Stapf) Cufod. with strong
support[23].
In addition, the mixture of O. brachypoda (Neupane
88, ODU) with all O. diffusa samples is in agreement
with their merger[16], given that the former was correctly
identified.
Taxonomically, the name transfer from Hedyotis
or Oldenlandia to Scleromitrion leads to the following
four new combinations.
(1) Scleromitrion diffusum (Willd.) R. J. Wang,
comb. nov.
Basionym: Hedyotis diffusa Willd., Sp. Pl. 1:
566, 1797.
(2) Scleromitrion koanum (R. J. Wang) R. J.
Wang, comb. nov.
Basionym: Hedyotis koana R. J. Wang, Acta
Phytotax. Sin. 45: 696, 2007.
(3) Scleromitrion pinifolium (Wall. ex G. Don) R.
J. Wang, comb. nov.
Basionym: Hedyotis pinifolia Wall. ex G. Don,
Gen. Hist. 3: 526, 1834.
(4) Scleromitrion verticillatum (L.) R. J. Wang,
comb. nov.
Basionym: Oldenlandia verticillata L., Mant. Pl.
1: 40, 1767.
Sivarajan and Biju[27] proposed the concept of
“Hedyotis corymbosa-diffusa complex” because of
their morphological similarity. In order to give a clear
elucidation between these two species, a comparison
was prepared on basis of their morphology, chromosome
numbers, palynology, chemical constituents, and their
bioactivity (Table 2).
In summary, this study clarified the traditionally
misapplied scientific name of Baihuasheshecao based
on the molecular phylogenetic analysis. The results
demonstrated that the two main mixed components
of the herbal Baihuasheshecao, Oldenlandia diffusa
and its adulterant O. corymbosa, phylogenetically
belonged to two separate lineages that representing
two respective genera. We then suggested the application
of Scleromitrion diffusum to replace the misapplied
name Hedyotis diffusa or Oldenlandia diffusa for
future scientific research and medicinal practice. The
proper application of the scientific name for traditional
Chinese medicine not only can promote the quality of
the Chinese medicinal material, but can enhance the
standardization and utilization of genuine traditional
Chinese herbs during the internationalization and
第5期 441
modernization of Chinese herbology.
Acknowledgments　　We are grateful to Professor But Paul
Pay-Hay from the Chinese University of Hong Kong for sharing
his medicinal knowledge about Baihuasheshecao.
References
[1]　 Yu L, Li Y, Guo X J. Simultaneous determination of anthraquinones
in Hedyotis diffusa by LC coupled with UV detection [J].
Chromatographia, 2009, 70(1/2): 211–215.
[2]　 Lin J, Wei L, Xu W, et al. Effect of Hedyotis diffusa Willd. extract
on tumor angiogenesis [J]. Mol Med Rep, 2011, 4(6): 1283–1288.
[3]　 Niu Y, Meng Q X. Chemical and preclinical studies on Hedyotis
diffusa with anticancer potential [J]. J Asian Nat Prod Res, 2013,
15(5): 550–565.
[4]　 Li M, Jiang R W, Hon P M, et al. Authentication of the anti-tumor
herb Baihuasheshecao with bioactive marker compounds and
molecular sequences [J]. Food Chem, 2010, 119(3): 1239–1245.
[5]　 Lin C C, Ng L T, Yang J J. Antioxidant activity of extracts of Peh-
Hue-Juwa-Chi-Cao in a cell free system [J]. Amer J Chin Med,
2004, 32(3): 339–349.
[6]　 Noiarsa P, Ruchirawat S, Otsuka H, et al. Chemical constituents
from Oldenlandia corymbosa L. of Thai origin [J]. J Nat Med,
2008, 62(2): 249–250.
[7]　 Sadasivan S, Latha P G, Sasikumar J M, et al. Hepatoprotective
studies on Hedyotis corymbosa (L.) Lam [J]. J Ethnopharmacol,
2006, 106(2): 245–249.
[8]　 Liang Z T, He M F, Fong W F, et al. A comparable, chemical and
pharmacological analysis of the traditional Chinese medicinal
herbs Oldenlandia diffusa and O. corymbosa and a new valuation
of their biological potential [J]. Phytomedicine, 2008, 15(4):
259–267.
[9]　 Lee H Z, Bau D T, Kuo C L, et al. Clarification of the phenotypic
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Table 2 Comparison of Oldenlandia diffusa and O. corymbosa
Oldenlandia diffusa Oldenlandia corymbosa References
Habit Annual herbs Perennial herbs
Branch Diffuse at base, erect or decumbent Erect or prostrate
Stem transverse section Subrounded; covered with unicellular cell of mastoid
process
Quadrangular; covered with unicellular cell of
mastoid process
[28]
Leaf petiole Sessile or subsessile Sessile
Leaf shape Elliptic lanceolate, linear-lanceolate or linear Linear to narrowly elliptic
Stipule With 2–3 fimbriate at apex With 2–5 fimbriate at apex
Inflorescence Solitary, rarely 2–3 flowered Corymbs, umbels or racemose cymes, 2–5 flowers
Inflorescence position Axillary Axillary
Peduncle Present rarely Always present
Flower Homostylous Homo- or heterstylous
Stamens and style Exserted Included
Capsule shape Subglobose, truncate or flat at top Globose or ellipsoid; truncate or slightly raised at top
Capsule dehiscence Apically and loculicidally dehiscent Loculicidally dehiscent
Epidermal cell of pericarp Anticlinal wall straight Anticlinal wall undulated [28]
Fruit septal cell Protuberance Strip-shaped, arranged in inlaid shape [28]
Seed Trigonous, narrowly winged, reticulated Trigonous, reticulated [29]
Seed germination period Most seeds within two weeks and all by 21 weeks Most seeds within 36 weeks [30]
Chromosomal number n=27 n=9
2n=18, 36, 54
[31]
[32–36]
Pollen aperture 3– (4)-colporate 3–(4–5)-colporate [26,29]
Chemical constituent 2-Hydroxy-3-methyl anthraquinone Furanocoumarins hedyotiscones A [37]
Biological activity Immunomodulating activity; inhibiting the growth of
cancer cell lines and inducing apoptosis; antitumor
activities
Significant hepatoprotective effects [38]
[7]
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王瑞江等：基于分子和形态学证据确定传统中药白花蛇舌草及其替代品的学名
442 第22卷热带亚热带植物学报
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