Syst. Evol. 227: 123-136 (2001)
Systematics and Evolution
Springer-Verlag 2001 Printed in Austria
and taxonomy of the Paeonia obovata polyploid complex
Hong1, K.-Y. Pan1, and G.-Y. Rao2
Laboratory of Systematic and Evolutionary Botany, Institute of
Botany, Chinese Academy of Sciences, Beijing, China
College of Life Science, Peking University, Beijing, China
October 30, 1998 Accepted June 21, 2000
The taxonomy of the Paeonia obovata polyploid complex
in E-Asia has been controversial with treatments ranging from one
species and two varieties to five species and numerous
infraspecific taxa. Extensive observations in the field, analysis
of morphological characters based on 32 populations sampled, and
the examination of a large amount of herbarium specimens show that
all the characters used by previous authors for distinguishing the
species are variable between or even within populations: Leaf
position, petal colour, stamen number (from 21 to 110 in the
red-flowered and from 58 to 239 in the white-flowered form),
filament and anther colour, style length as well as follicle
number and position. Therefore, all these characters are of no
value for species delimitation. However, an extensive chromosome
survey and the analysis of morphological characters show that the
ploidy level is basically correlated with the geographical
distribution and with the indumentum on the lower leaf surface.
Thus, the recognition of one species with two subspecies is
proposed: Paeonia obovata subsp. obovata, mostly
diploid, widely distributed in the north, east and south of the
distribution range and glabrous to sparsely pubescent or hirsute
on the lower leaf surface, and subsp. willmottiae, tetraploid,
limited to the west and with leaves densely hirsute or pubescent
words: Paeoniaceae, Paeonia obovata polyploid complex,
morphological variation, cytogeog-raphy, taxonomic revision.
Paeonia obovata polyploid complex is a group of herbaceous
peonies, widely distributed in the forests of East Asia (Fig. 3).
It is distinguished from other groups (except the P. coriacea
group) by its ternate leaves with 9 entire leaflets, solitary
and terminal flowers, and glabrous carpels. From the P.
coriacea group it differs by its terminal leaflets being
obovate or broadly obovate.
The first species of
the complex described was P. obovata Maximowicz (1859)
based on specimens from the Amur area with rose-purple petals. The
second species, P. oreogeton S. Moore (1879) was based on
specimens from "Kwandien" (Kuandien), Liaoning Province,
China. Makino (1898) discovered a white-flowered form in Honsu,
Japan, and described it as a variety of P. obovata, i.e.
var. japonica, which was raised by Miyabe and Takeda (1910)
to specific level, P. japonica. The fourth species, P.
willmottiae Stapf (1916) was based on a plant in the garden of
Miss Willmott, Warley Place, Essex, England, which
was raised from seeds collected by E. H. Wilson from China.
Finally. Mandl (1921) described the fifth species of the complex
with white flowers, from the Far East of Russia (Nikolsk-Ussuria).
Schipczinski (1937) recognised only four species from Russian
territory: P. obovata, P. japonica. P. oreogeton, and
P. vernalis. The
first author revising the taxonomy of the complex was F. C. Stern
(1943). He reduced P. willmottiae to
a variety of P. obovata and
suggested to merge P. oreogeton and
P. vernalis with P.
(Stern 1946). He followed
Miyabe and Takeda (1910) in distinguishing P.
japonica as an
independent species, and thus recognized three taxa, P.
var. typica Makino
and var. willmottiae (Stapf)
F. C. Stern, and P. japonica.
Fang (1959) did not agree
with Stern, still recognizing P. willmottiae
as a separate species. Nearly
all Japanese botanists (e.g. Ohwi I97S) recognized two species in
Japan, each with two infraspecific taxa: P. obovata
and var. glabra
Makino. and P.
japonica var. japonica
and var. pilosa
Nakai. The second author of
the present paper, Pan (1979), recognized only one species, P.
with var. willmottiae.
but failed to mention P.
japonica. Following Pan's
treatment, nearly all Chinese authors (e.g. Wang 1987, Zhao 1990.
Lin 1992) recognized only one species, P. obovata.
However, Ding and Liu (1991)
argued for the status of P. japonica as
an independent species and listed additional differential
characters, a treatment which was also accepted by Li (1995).
chromosomal studies on the Paeonia obovata complex (Hong et
al. 1988: Nishikawa 19K5; Okado and Tamura 1979; Sokolovskaya
1960, 1966) have indicated differentiation in ploidy (with
diploids and tetraploids). In order to understand the geographical
distribution of 2x and 4x, and possible correlations between
ploidy and any morphological characters, we have conducted an
extensiv karyological survey in China and Japan.
present paper provides a taxonomical revision of the complex based
on extensive field work, examination of a large amount of
herbarium specimens, analyses of morphological characters, and
up-to-date cytogeographical data.
1985 to 1998 the first author and his assistents made a number of
expeditions visiting numerous sites throughout China and several
sites in Japan. .. populations in total were sampled during the
expeditons (of which 25 are listed in Tab.1). Besides, we examined
a great amount of herbarium specimens in the Herbaria of the
Institute of Botany. Chinese Academy of Sciences (PE). Shenyang
Institute of Applied Ecology, Chinese Academy of Sciences (IFP).
the Northwest Normal University (NWTC), the Northwest Institut
Botany in Shaanxi (WUK). the Hokkaido University (SAPS), and the
Kyoto University (KYO). More than 20 populations were sampled for
the karyological survey. Origins and vouchers are given in Table
1, all the latter are preserved in the Herbarium of Institute of
Botany. Chinese Academy of Sciences (PE). Root tips were collected
in the fields, prelrealed in an aqueous solution of 0.05%
colchicine or saturated p-diehlombenzol for 3-5 hrs and then fixed
in Carnoy's I fixative. For Zhu et al. PB86020 (Jinfushan of
Chong..) flower buds were directly fixed in the field by Carnoy II
fixative for meiotic observation. Chen Z. D. s.n. was collected
and transplanted to Beijing in 1996. and its root tips were
obtained in the greenhouse. Chromosome preparations followed Hong
ct al. (1988).
orientation. This was considered by Ding
and Liu (1991) as an important character for distinguishing P.
japonica (spreading) from P. obovata (ascending).
According to our field observations, however, this character is
unstable even within a population. In the populations Hong et al.
H98026, H98006, H98007, for example, both ascending spreading
leaves were found. Figure 1f ...nitely shows spreading instead of
ascending leaves in a red-flowered population (Hong el al.
1. Variation of leaf indumentum, floral parts, and chromosome
numbers in 25 populations of the Paeonia ohovata complex. Leaf
indumentum is expressed in grades, number of individuals are in
parentheses (see text)
indumentum. This character
was used by Japanese botanists for recognizing infraspecific taxa
both in Paeonia
obovata (var. typica
and var. glabra)
and P. japonica
(var. japonica and
var. pilosa). We
examined hundreds of specimens, both in the field and herbaria,
and found that leaf indumentum on the lower surface was an
extremely variable character, from entirely glabrous to densely
hirsute or pubescent. For roughly quantifying this character and
thus revealing the pattern of variation in the whole complex we
used a more or less arbitrary system of six grades to measure the
density of leaf indumentum with 0 indicating
entirely glabrous, 1 very sparsely hirsute or pubescent with some
hairs only along major veins, while 5 the most densely hirsute or
pubescent (nearly entirely covered with hairs), and 2, 3 and 4
indicating intermediate states. Our observations are shown
in Table 1. Among 20 sampled populations (with Hong et al. H98002
and H98025 excluded), only three (Hong et al. H98010, Hong D.Y.
PB90010 and PB90011) were
purely glabrous; in ten populations all individuals sampled were
hairy, and within each of the rest seven populations both hairy
and glabrous individuals were found. Two populations (Zhu X. Y. el
al. PB86020 and Qiu J. Z. PB88501) are worth mentioning because
they exhibit a great range of variation in leaf indumentum, from
entirely glabrous to densely hairy (grade 4). From that it is
rather clear that the recognition of Paeonia
obovata var. glabra
and P. japonica
var. pilosa based
on leaf indumentum is unreasonable. In addition, we examined
hundreds of herbarium specimens from China, Japan and the Korea
Peninsula and projected their indumentum states onto the
distribution map of the complex (Fig. 3). It can be seen that leaf
indumentum is correlated to some extent with the geographical
distribution and ploidy. The plants of this complex in W. Hubei,
NW. Sichuan. N. Chongqing, Shaanxi, Shanxi, and W. Henan are all
tetraploid and nearly always densely hirsute or pubescent
except those in Shanxi, which were variable from mostly
hairy (e.g. Hong et al. H98009) to mostly
glabrous or sparsely hairy (e.g. Hong et al. H98010).
1. Morphological variation
in the Paenia
Hong et al. H98026 (Mt. Changbai. China) with spreading (instead
of ascending) leaves in a red-flowered form:
Hong et al. H98006 (Lushi Co. Henan China) with petals white but
pink at periphery, filaments purple at base but white above, and
Hong D. Y. P1190011 (Uwa-dio Ehime, Japan) with pink petals;
Hong et al. H98007 (Lushi Co. Henan, China)
with purple-red petals, totally white filaments and yellow
anthers; e Hong and Zhu PB85024 (Wolong Nature Reserve. Sichuan,
China) with totally white and widely spreading petals; f Hong el
al. H9H026 (Mt. Changbai. Antu Co, Jilin) with red petals, purple
filaments, and yellow ovaries
They exhibit the least variation, are nearly always 3, very
occasionally 2 or 4 in number and mostly rounded at the top. Only
of the sepals sometimes is caudate in some individuals of some
populations (e.g. Hong et al. H98026).
colour, posture and number. The most important character used
for distinguishing P. japonica from P. obovata is
the colour of petals, rose-purple in P. obovata and white
in P. japonica. Our extensive observations revealed that
the colour pattern in this complex is not so simple, but actually
varies from purely white (Hong and Zhu PB85024), white with
pinkish shade at the base or periphery (Hong et al. H98006, in W.
Henan), rose (Hong D. Y. PB90011), pink-red (Hong ct al. H9S033),
red (Hong et al. H98010), purple-red (Hong et al. H98007) to
purple-red with two white stripes on the back (Hong et al.
orientation of petals was said by Ding and Liu (1991) to be a
valuable character for distinguishing white-flowered P.
japonica (ascending) from red-flowered P. obovata
(spreading). However, the two orientations could be found in
each species. For example, spreading petals were found in a
population with white flowers (Hong and Zhu 85024) (Fig. 1e).
are usually 5 or 6, but flowers with 4 (Hong et al. H98019) or 7
petals (Hong et al. H98006) occasionally occur. Even within a
population, for example in Hong el al. H98019, the number of
petals varied from 4 to 6.
number and colour. The number of stamens was used by Ding and
Liu (1991) as an important character in distinguishing P.
japonica (100-280) from P. obovata (14-75). Our
extensive observations did not confirm this. As shown in Table 1,
the number of stamens varied from (58) 70 to 230 in the
white-flowered form (P.japonica), and from 21 to 110 in the
red-flowered form (P. obovata). Although there is some
difference in number of stamens between the two colour forms, the
two ranges overlap much. Therefore, as number of stamens varies
continuously in the complex, this is not a diagnostic character
for distinguishing the two colour forms as separate species.
colour of filaments was found to vary greatly in the
complex from white, yellow-green, purple below and white above to
entirely purple (Fig. 1d. f). The two colour types of filaments
(white and purple) were found even within a single population
(Hong et al. H98007). Anthers were yellow in the red-flowered
populations, but yellow, orange-red or dark purple in the
white-flowered populations (Fig. 1). Within the population Hong et
al. H98006 both purple and orange-red anthers were found.Thus it
is obvious that the colour of filaments and anthers are of no
taxonomic value for separating P.japonica and P.
and follicles. The number of carpels was found to vary from 1
to 5, usually 2 or 3 in both the white-flowered and
red-flowered forms (Table 1). Flowers with a single carpel were
found in Hong et al. H98023 (Kuandian, Liaoning) and Hong et al.
H98015 (Qianshan, Liaoning), while those with 5 carpels were found
in Hong et al. H98004 (Xixia County, Hcnan) and Hong el al. H98009
(Yimnqu County, Shanxi). The number of carpels varies even within
a population, for example, from 1 to 3 in Hong et al. H98001 and
Hong el al. H98007. from 3 to 5 in Hong et al. H98009 and from 2
to 4 in Zhu et al. PB 86020.
to Ding and Liu (1991), the orientation of mature follicles can be
used for distinguishing P. obovata (strongly recurved) and
P. japonica (slightly recurved). However, follicles recurve
gradually during ripening and the two states can be found in both
P. obovata and P. japonica. For example, Tang. T.
963 (red-flowered) and Wang, W. T. 2531 (red-flowered) had mature
follicles also strongly reversed with the top attaching the stem
as stated by Ding and Liu (1991) for the white-flowered form.
and stigma. We discovered that styles change their length with
the development of the carpels. In bud stage styles are hardly
visible; they become apparent when the flower opens and elongate
with time. In open flowers style length varies from 1.0 to 5.0 mm
long in the white-flowered form, and from 0.5 to 3.0 mm long in
the red-flowered (Table 1). The longest style (5 mm) was
discovered in a population (Xic.
Z. W. H98035) with petals white but pink below (Table 1).
Therefore, differences in length of styles between the two forms
are hardly significant.
2. Chromosomes of the Paeonia obovata Maxim,
complex, all at mitotic metaphase except for B, which is at
meiotic anaphase II.
Hong et al. PR8507H (red flowers), 2n= 10;
Zhu X. V. et al. PR86020 (white flowers), n = 5;
Hong et al. H9803I (white flowers?), 2n=10;
Hong and Zhu PB85062 (red flowers), 2n = 20;
Hong and Zhu PB85067 (white flowers), 2n = 20;
Hong and Zhu PR85024 (white flowers), 2n = 20;
Hong el al. H98014 (red flowers), 2n = 20.
X 1250. only D X 1600
The results of our karyological survey are shown in Table 1
and Fig. 2. It is clear that ploidy level is not correlated with
the flower colour: diploids and tetraploids occur in both the
white-flowered (P. japonica) and the red-flowered
form (P. obovata). Therefore, the chromosome data do not
support the recognition of P. japonica as an independent
taxon. However, ploidy level is shown to be correlated with the
geographical distribution. Nearly all the populations sampled are
tetraploids in the western part of distribution area of the
complex: Shanxi, Shaanxi. NW. Sichuan. N. Chongqing, W. Ik-nan and
W. Hubei (Table 1; Fig. 3). But there are two exceptions: In Lushi
County, W. Henan, there is a mixed population (Honget al. H98006
and H98007: Ding 1988), in which both red-flowered and
white-flowered forms were found to grow together, and both 2n= 10
and 2n = 20 were counted (Table 1). In the Far East of Russia,
precisely in Sakhalin and the lower reaches of the Amur River, and
in NE China, both diploids and tetraploids were recorded
(Sokolovskaya 1966, 1960; Table 1).
3. Distribution map of the Paeonia obovata complex
with ploidy levels and the density of indumentum. White triangles
= diploids; black triangles = tetraploids; the columns indicate
line relative density of the lower leaf side indumentum: totally
white = glabrous; totally black = most densely hirsute (see text
for details). The chromosomal data are from Table 1 and in
addition from Ding 1988; Ding and Liu 1991; Haga and Ogata 1956;
Hong et al. 1988; Lee 1967; Nakamura and Nomoto 1981: Nakata and
Hong 1991; Nishikawa 1985; Okado and Taimira 1979; Sokolovskaya
I960. 1966; Starodubtsev 1985; Uspenskaya 1987; Yang and Zhu 1989
to a certain extent there is correlation between ploidy level and
indumentum on the lower surface of leaves (Fig. 3). In the diploid
area (Japan, the Korea Peninsula, the Far East of Russia and the
north, east and south parts of distribution range in China), the
plants are glabrous to sparsely hairy, with the densest hairiness
reaching the third or, very occasionally, the fourth grade, e.g.
in the population Zhu, X. Y. et al. PB86020 in Mt. Jinfu-shan, S.
Chongqing, but there glabrous to very sparsely hairy individuals
could also be found. On the contrary, in the distribution range of
the tetraploids nearly all the populations sampled are densely
hirsute or pubescent, except those in C. and NE. Shanxi of China
and the Far East of Russia. In C. and NE. Shanxi, precisely on Mt.
Kuandi of Jiaocheng County, Mt. Huoshan of Huoshan County, and Mt.
Wutai, most individuals observed were glabrous or sparsely
pubescent, though they were tetraploid.
and taxonomical treatment
is clear from what we have described above that the characters
used by previous authors for distinguishing species or
infraspecific taxa are not useful as diagnostic characters except
the indumentum on the lower leaf surface. Because this character
was frequently found to vary greatly from totally glabrous to
moderately hairy, even within a population, it is not reasonable
to base varieties on it. However, indumentum is correlated
generally with the geographical distribution and ploidy level.
Therefore, it is justified to recognize only one species in the
complex, P. obovata, with two subspecies: the typical
subspecies for the populations in the northern, eastern and
southern parts of the distribution range, which are diploid (very
occasionally tetraploid in the Far East of Russia and in the
Changbai Mountains, Jilin, China) and mostly glabrous to
moderately pubescent or hirsute, and the subspecies willmottiae
for the populations in the western part of the distribution
range, which are tetraploid (diploid and tetraploid only in one
mixed population in Lushi County, W. Henan Province), and mostly
densely (less frequently moderately) hirsute or pubescent. It is
likely that reticulate evolution and polyphyletic origins of
polyploids as documented for the whole genus (Sang et al. 1995,
1997) also play a role within the P. obovata complex.
obovata Maximowicz, Prim. Fl, Amur. 29 (1859)
Amur, Kitsi; Maximowicz (holotype, LE; isotype, K!)
species comprises two geographically allopatric subspecies which
are keyed out below:
mostly glabrous or sparsely, occasionally densly pubescent or
hirsute at lower surface, diploid (tetraploids rare in the Far
East of Russia and on Mt. Changbai, NE. China)
densely, very occasionally sparsely, hirsute or pubescent on lower
surface; tetraploid (the only diploid found in a mixed population
in Lushi Co, W. Henan)
oreogeton S. Moore in J. Linn. Soc. Lon don, 17:376. (1879);
type: China, Liaoning Kwandien (Kuandian) s. coll. (holotype, K! =
P. obovata Maxim, f. oreogeton (S. Moore Kitagawa in
Lneam. Fl. Mansh. 221(1939)
obovata var. typica Makino, Bot. Mag. (Tokyo) 12:301
(1898); J. Jap. Bot. 5(9): 33. 1928
obovata Maxim, var. japonica Makino in Bot. Mag.
(Tokyo) 12:302. (1898); type: Miyabe et Takeda's fig. 153, cited
(!) = P. japonica (Makino) Miyabe et Takeda in Gard. Chron.
Ser.3, 48:366. Fig. 153(1910).
Mandl in Bot. Kozl., 19:90. (1921); type: Russia, the Far
East, Nikolsk-Ussurisk (holotype, LE).
obovata var. amurensis
Schipczinsky, Not. Syst.
Herb. Hort. Bot. Pctrop. 2: 44. (1921); type: not
obovata var. australis Schipczinsky, Not. Syst. Herb.
Hort. Bot- Petrop. 2: 44. (1921): type: not designated.
obovata var. glabra Makino in J. Jap. Bot. 5(9):33.
(1928). type: Japan (not designated).
obovata var. alba Saunders, Nat. Hort. Mag. 13, tab.,
227. (1934); type: Saunders' figure cited.
japonica var. pilosa Nakai in J. Jap. Bot. 12:
394-395(1937); type: Korea, Heinan. ML Rorinsan (holotype. Til),
subspecies grows in deciduous broad-leaved, mixed broad-leaved and
needle-leaved or conifers forests at altitudes from ca. 200 to
2000 m. It is distributed in Japan, the Korea Peninsula, the Far
East of Russia and NE., N. and E. China, in addition to a small
area in SW. China.
Anhui: Huangshan: E of
Tungwopong, Tsoong, P. C. 4090 (PE); Jiuhuashan: Sliangmingyuan,
Hong, D. Y. & Chen, T., PBH<i()()3(PE); Yuexi: Yaoluoping,
Xic, Z. W. H98035 (PE, A, CAS. K, MO, US): Beijing: Baihuashan:
Wang. C. W. 60273(PE): Miyun: W slope of Wulingshan. Hong, D. Y. &
Rao. G. Y. H98033 (PE, A, CAS, MO, US); Chongqing:
Nanchuan: Jinfushan, Tiechangping. Liu. Z. Y. H98038(PE, A.
CAS. K, MO. US); Cuizhou: Zunyi City: Xianrenshan, Qianbei
Exp. 0115 (PE); Hebei: Chengde: Wenchanggou. Liu, X. Y.
1665(PF): Chicheng: Dahaituoshan. Hong. D. Y.. Clicn. Y. &
Song, S. Y. PBH507K(PE); Donglmgshan: Shidaozihc, Dongling Exp.
269(PE); Fuping: Long-quankuan, Liaodaobai, Fuping Exp. 3(PE);
Huailai: Tangying Temple Hot Spring, s. coll. 2117(PE); Laishui:
Siqu, Yang, C. G. 8B8(PE); Neiqiu: Duanmutao. Liu. X. Y. 433 (PE):
Ping-shan: Muehang, Tongdonggou, s. coll. 466(PE); Qianxi:
Xishuiling. Qianxi Exp. 205(PE); Qinlong: Wudaoling, Qinlong Exp.
972(PE); Weichang: HuangUik, Toudaochuan, Huang 489(PE);
Xiaowulai: Beitai, Licent, P. 2622(PE): Xinglong: Wuling Shan,
Lianhuachi, Lion, T. N. 4740 (IFP); Yixian; Kuanzuoling. Lion. K.
M. 2004(PE); Yuxian: Nanliangqing:
Zhangjiakou Exp. ll(PF); Zanhuang: Loudi, Zliangshiyan, Shijiazuo
Exp. 53 l(PE);Zrmliiia: Eastern Tombs, Liou.T.N. 1411 (PE):
Heilongjiang: Dailing: Lianshuijian, Sino-Germ. Exp.
7476(PE); Hailin: Hengdaohezi, Hou X. Y. 11759(PE); Hulin:
Huloajian, WANG, G. Z. 40H(PE; 1EP): Mudanjiang: Provincial Wild
Plant Exp. Mudanjiang Team 0621(1'E); Ning'An: Jiangbhuijiao
Forest Farm, Daqinggou s. col!, s.n. (IFP); Shangzi: Weihe Distr.
Datutouzi, s. coll. s. n. (IFF); Upper Ussuri: Maximowicz. her
sec. anno 1860 (PE, K); Yichun: near Wuying, Sino-Gcrm. Exp.
7903(PE); Henan: Shangcheng: Jinganlai. Qiu, J. Z.
PB8850I(PEJ; Hunan: Sang/hi: Shadiping, Baheshc. Liu. L. IF
9344(PF); jiangxi: Lushan: Huanglong'en. Wang, M. J, 285(PE);
Jilin: Antu: Changbai Shan. 1500m. Hong, D. Y., Pan. K. Y.,
Rao. G. Y. & Cao W. H98026(PE, A, CAS, K, MO, US): Fusong:
Changbai Shan, Hong. D. Y. PB86071(PE|; Helong: near Guangping,
Yanbicn Second Team. 570 (PE); Hunchun: Changbai Shan lo Erdao
Water Power Station. Fang. Z. F. 3564(IFP]; Ji'an: Renao Forest
Farm, Fang, Z. F. 2285(1FP]; Jingyu: no precise locality, Sato, J.
10005(PE); Linjiang: Sanchazi Longwan, No. 2 to No.l. Liou, T. N.
1077(PE, IFP): Liuhe: Yousong, Liao'angou, Sandaolian. Deng. Y. C.
798(1FP); IJaoning: Anshan: Qianshan, Miaotai, Fifth Group
Village, Hong, D. Y., Pan, K. Y., Rao. G. Y. & Cao, W.
H98015(PE. A, CAS, K, MO, US); Benxi: 10 km W of Lianshaguan Fort,
Hong, D. Y., Pan. K. Y., Rao, G. Y. & Cao, W. H980I9(PE. A,
CAS. K, MO. US); Dalian: Laolicshan, Lu, Y. C. 3372(IFP);
Fcngchong: Yuanyang, Zhang, Y. L. 2513(IFP); Huanren: Mumengzi
Township, Cui, S. C. etal. 270 (IFP); Kuandian: Baishilazi Nature
Reserve, Hong. D. Y.. Pan. K. Y.. Rao, G. Y. & Cao, W.
H98023(PE, A. CAS. K. MO, US); Lingyuan: Dabiengou, the mountain
in front of Linjia Dayuan Wang, W. 690(IFP) Qingyuan: Xiaojiabaozi
Township, Xinhlun, Hong, D. Y,, Pan, K. Y., Rao, G. Y. & Cao.
W. H98031(PE. A, CAS, K, MO. US); Xifcng: Helong, Jiuru, Lin, Y.
Y. 7(IFP); Xinbing: Laogangshan, General Exp. 210 & 223 (IFP);
Xiuyan: Tangchi, Wang, W. el al. 152I(IFP); Yingkou: Jianyi
Township, Luwang-zipu, Yingkou General Exp. 16M (IFP); Zhuanghe:
Buyunshan. Dayang-gou, Li, S. X. 151 (IFP); iNei Mongol:
Ningchcng: Dayingzi, Bagudao, Erdaogou, Li, S. X. 4710 (IFP):
Sichuan: Ending: Erlongshan, Daba, HouShan, Hu. W.
SZ); Yuexi: Dongshan, Dashuigou, Zhu, X. Y. & Song, S. Y.
PB86024 (PE); Zhejiang: Anji:
Longwang Shan, Chen, L. 086 (ZJFC); Tianmu Shan: Sanmuping, Xiwan,
Hong, D. Y., Pan, K. Y. & Lou, L. H. H98001(PE,A,CAS,K,MO,US).
Aichi: Toei-cho, Torii, K. s.n. (KYO); Aomori:
Higashitsugaru-gun, Imbetsu-cho, Ooka-wadai, Mimoro, K. et al.
3901 (KYO); Ehime: Uma-gun, Doichou-cho, Mt. Akaboshi,
Hong, D. Y. PB90010(PE); To-Uwa-gun, Uwa-cho, Hong, D. Y.
PB90011(PE); Fukui: Nyu-gun, Echizen-cho, Rokuroshi,
Kurosaki, N. 10731 (KYO); Fukushi-ma: Nishishiragawa-gun,
Nishisato-mura, Suzuki, Z., s.n. (KYO); Gifu: Ibi-gun,
Kasuga-mura, from Sasamata to Mt. Ibuki, Takaheshi, H. &
Takano, H. 7313(KYO); Gumma: Agatsuma-gun, Tsuma-goi-mura,
Nidoage, Yahara, T. et al. 7033 (KYO); Hiroshima: Saiki-gun,
Mizuda-mura, Kato, M., s.n. (KYO); Hokkaido: Sapporo, Hong,
D. Y. s. n. (PE); Kuriler Islands, hikotan Ohwi, J. 250 (KYO);
Hyogo: Yabu-gun, Yoka-cho, Tsubairo, Kurosaki, N. 15145
(KYO); Iwate: Shimohei-gun, Akka-mura, Takasuka, Shimizu,
T. 01937(KYO); Kaga-wa: Kagawa-gun, Shioe-mura, Mt. Ohtaki, Hiroe,
M. 15608(KYO); Kanagawa: Hakone, Nagaotoge, Togasi, M.
1288(KYO); Kochi: Tosa, Tomioka-mura, Otakijinjya to Mt.
Tsutsujosan, Murata G. & Shimizu, T. 817(KYO); Kumamoto:
Yatsushiro-gun, Tzumi-mura, from Kureko to Mt. Iwaudo-yama,
Hotta, M. 12062(KYO); Kyoto: Funai-gun, Yamamoto, H.
533(KYO); Mie: Ichisi-gun, Misag-i-mura, Murata, G. &
Fukuoka, N. 194(KYO); Miyagi: Oshika-gun, Kinkasan-to,
Naito, T. s.n. (KYO); Miyazaki: Higashiusuki-gun,
Shiba-mura. Mt. Shiraiwa, Hotta, M. 6438(KYO); Nagano:
Shimoina-gun, Oojika-mura, Tsubame-iwa, Shimizu, T. 5815(KYO);
Nara: Yoshino-gun, Mts. Oomine, from Kawai to Gyojagaeri,
Hotta, M. et al. 27(KYO); Niigata: Sado (island), between
Mt. Donden & Mt. Kongo, Okamoto, M. 2438(KYO); Oita: N.
Oita, Yufudake, Tashiro, Z. s.n. (KYO); Okayama: Honkayabe,
Kawakami-mura, Mt. Maru-yama and Asanabewashiga-sen, Fujii, S. 164
(KYO); Saga: Tachodake, N of Saga city, Tashiro, Z. s.n. (KYO);
Shiga: Along the north ridge of Mt. Ibuki-yama, Fukuoka, N.
8061(KYO); Shimane: Tsuwanomachi, Ohwi, J. s.n. (KYO);
Shizuoka: Shizuoka, from Magosajima to Ikawa pass, Miki, E.
213(KYO); Tokushima: Miuma-gun, Ichiu-mura, from Kumadaira
to myotoike, Hotta, M. 10805(KYO); Yamaguchi:
Yuansongci, s.n. (KYO); Yamanashi: Fujiyoshida City, Araya,
Konta, F. 11786 (KYO).
PENINSULA: North Korea: Chongyang, Mills, R. G. s.n. (PE);
Chongsong, Koidzumi, G. s.n. (KYO); Bor.-orient, Ohwi, J. 2709(b)
(KYO); Musa, Ohwi, J. 1043 (a) (KYO); Changjin, Nom-ura, N. s.n.
(KYO); near Hamhung, Ohwi, J. s.n. (KYO); South Korea:
Sanchong, Igarashi, F. 23(?)(KYO).
Sakhalin: E. coast, Mt. Nupuripo, Mou-ale, T. s. n. (SAPS); W.
coast, Notasan (Mauka), Miyake, T. s. n. (SAPS); Mt. Kasipo, S.
Sakhalin, Takee, G., Abumiya, H. & Hoshiro, Y. s.n. (SAPS).
obovata Maxim, subsp.
D. Y. Hong et K. Y. Pan, stat. nov.
willmottiae Stapf in Curtis' Bot. Mag. 142: tab. 8667(1916);
type: "Miss. Willmott May 22, 1915" (holotype K!)
obovata Maxim, var. willmottiae (Stapf) F. C. Stern in
J. Roy. Hort. Soc. 68: 128 (1943) et Stud. Gen. Paeonia, 76(1946).
subspecies is characterized by tetra-ploidy and leaves mostly
densely or very densely hirsute or pubescent. It is confined to
the mountain area around the Qinling Range and grows in deciduous
forests at altitudes of 800-2800 m.
specimens examined: CHINA: Chongqing: Chengkou: Nanxi, Gaopo,
Dai, T. L. 101441 (PE, SZ); Wushan: Chaoyangping, Yang, G. H.
57903(PE); Wuxi: Hongchiba, Liu, Z. Y. H98039 (PE,A,CAS,K,MO,US);
Gansu: Kangxian: Jia'an, Qinhe Forest Farm, 1450 m, He, Y.
Q. & Tang, C. L. 315(WUK); Chengxian:Zhaoba Forest Farm, 1450
m, Lian Y. S. & Wang J. Q. 791244 (NWTC); Tewo: Nanshan, 2500
m, Gansu Provincial Medical Herb Group s.n. (NWTC); Tianshui:
Beiyinshan, Zhang 59 (NWTC); Zhugqu: Jiao'er-qiao, 2000m, Wei, Z.
P. 2566(WUK); Henan: Lushi: Shiziping, Dakuaidi (Qihe
Forest Farm), Hong, D. Y., Wang, S. Y., Pan, K. Y. & Rao, G.
Y. H98006 (PE, A, CAS, K, MO, US); Songxian: Yangshan,
Yangjiaohao, Hong, D. Y, & Ye, Y. Z. H94004(PE); Xixia:
Erlangping Township, Wan-tancun, Waitoushan, Hong, D. Y., Wang, S.
Y., Pan, K. Y. & Rao, G. Y. H98004 (PE, A, CAS, K,MO,
US); Yiyang: Longmao, Kuan, L. C. & Dai, T. L. 2418(PE);
Hubei: Badong: Luoping, Chen, Q. L. et al. 1920(PE); Baokang:
Dashui Forest Farm, Hong, D. Y., Ye, Y. Z. & Feng, Y. X.
H97050 (PE, MO); Hefeng: Zourria Township, Guchengping, Peng, P.
S. H93 (PE); Lichuan: Ye, Y. C. 248(PE) & 511 (PE);
Shennongjia (Songbai): Shantunyan, Qiu, J. Z. PB88022(PE);
Xingshan: Longmenhe, Chen, W. L. H98040 (PE, A,K,MO,US); Ningxia:
Jingyuan: Ningxia Medical Herb Group 2-118 (WUK); Qinghai: Xunhua:
Mengda Forest Farm, 2300m, Guo B. C. & Wang, W. Y. 25194
(HNWP); Shaanxi: Danfeng: Yaozhuang, Qipan-mo, He, S. B. 216
(WUK); Fengxian: Huangniupu, Donggou, 1650m, Zhang Z. Y. 090(WUK);
Fuping: Damahe, Fuziling, Kuo, P. C. 1493(PE); Huanglong: Yaoxian,
1400 m, Yang J. X. & Huang S. L. 6120 (WUK); Huashan: Liou, T.
N. 10887(PE); Langao: Bijiashan, 2070 m, Wei, Z. P. & Luo S.
H. 0291 (WUK); Lantian: Qinggangping, 1600 m, Su, G. X. 131 (WUK);
Longxian: Kuan-shan (Kwanshan), Wang, T. P. 4227(PE); Nanzh-eng:
Xiaoba, 1250m, Hou, X.X. 478(WUK); Shangxian: Qinwangshan, Shegou,
Wang, T. P. 16287(PE); Taibai Shan: Shangbaiyun, Hong, D. Y. &
Zhu, X. Y. PB85067(PE); Shanxi: Huoxian: Mt. Huoshan, above
Yanjiapin, near Forest Farm, Hong, D. Y., Pan, K. Y. & Rao, G.
Y. H98010 (PE,A,K,MO,US); Jiaocheng: Kuandi Shan, Hengjian,
Tashangcun, Liu, X. Y. 21363(PE); Jishan: Xishe, Qiu, J. Z.
PB88501 (PE); Ningwu: Dongzhai, Majiazhuang, Yuanwading, Shanxi
Exp. 265(PE); Qinxian: Yushuiquan, Shigaoshan, Kuan, K. C. &
Chen, Y. L. 983(PE); Wutai Shan: Taihuei Town, Zhenhaisi Temple,
Hong, D. Y., Pan, K. Y. & Rao, G. Y. H98013(PE,A,CAS,K,MO,US);
Yangcheng: Zhangui, Bao, S. Y. 2148(PE); Yich-eng: Dahe,
Beishenyu, Huanghe Exp. 202(PE); Yuanqu: Lishan, Houwentangcun to
Shunwang-ping, Hong, D. Y., Pan, K. Y. & Rao, G. Y. H98009
(PE, A, CAS, K, MO, US); Sichuan: Heishui: Duifengshan, Li, X. &
Zhou, J. X. 73835 (PE, SZ); Maoxian: Maoxian to Lixian, He, Z. &
Zhou, Z. L. 13120(PE, SZ); Wenchuan: Wolong Nature Reserve, Hong,
D. Y. & Zhu, X. Y. PB85024 (PE).
are grateful to the National Geographic Society (Grant 6179-98)
for support of the field work. Sincere gratitude is due to Dr. Qiu
Jun-zhuan, Dr. Chen Zhi-duan, Dr. Zhu Xiang-yun, Prof. Chen
Wei-lie, Prof. Wang Shui-yi, Dr. Wang Ying-zhen, Dr. Feng Yu-xing,
Mr. Cao Wei, and
to Dr. M. Nakata at the Toyama Botanical Garden, Japan, for their
assistance in the field work. We should also thank Dr. Ding Kai-yu
for his help in is making chromosome preparations, Mr. Yang
Xue-jian for his assistance in the preparation of photographs, and
Miss Li Qiao-ling for typing the manuscript.
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Guang-yuan, College of Life Science, Peking University, Beijing