全 文 ：植 物 分 类 学 报 43(2): 169–177(2005)
Acta Phytotaxonomica Sinica
———————————
Received: 21 June 2004 Accepted: 9 December 2004
* Author for correspondence. E-mail: .
Notes on taxonomy of Paeonia sect. Moutan DC.
(Paeoniaceae)
HONG De-Yuan* PAN Kai-Yu
(Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences,
Beijing 100093, China)
Abstract In this paper we argue for the specific status of Paeonia suffruticosa Andrews, for
the rationality of P. suffruticosa ssp. yinpingmudan D. Y. Hong, K. Y. Pan & Z. W. Xie and its
wildness, and for the treatment of P. suffruticosa ssp. atava (Brühl) S. G. Haw & Lauener as a
subspecies of P. rockii, P. rockii ssp. atava (Brühl) D. Y. Hong & K. Y. Pan. P. jishanensis T.
Hong & W. Z. Zhao is justified as a legitimate name, while P. spontanea (Rehder) T. Hong &
W. Z. Zhao is recognized as a superfluous name. Halda’s four combinations and two new
hybrid names in sect. Moutan DC. were treated as synonymy. As a result, one new combination
is made and five new synonyms are proposed here in this paper.
Key words Paeonia, Paeonia suffruticosa, Paeonia suffruticosa ssp. yinpingmudan, Paeonia
rockii ssp. atava, new combination, new synonymy.
In the 1990s we published several papers dealing with taxonomy of tree peonies
(Hong, 1996, 1997a, b, 1998; Hong & Pan, 1999a, b; Hong et al., 1998a, b; Zhou et al.,
2003; Hong et al., 2004). Haw (2001) basically accepted our taxonomy of the eight species
in sect. Moutan, but he held different views in three points. He recognized Paeonia
suffruticosa Andrews as a hybrid, treated P. suffruticosa ssp. yinpingmudan D. Y. Hong, K.
Y. Pan & Z. W. Xie as a synonym of P. ostii T. Hong & J. X. Zhang, and he still
considered P. moutan Sims ssp. atava Brühl (=P. suffruticosa ssp. atava (Brühl) S. G. Haw
& Lauener) as a doubtful taxon. Halda (1997) made a number of new combinations in sect.
Moutan and described two new hybrids, which were unfortunately neglected by us. This
paper is to address Haw’s three questions, and deal with Halda’s combinations and hybrid
names.
1 On Paeonia suffruticosa Andrews
1.1 Subspecies yinpingmudan D. Y. Hong, K. Y. Pan & Z. W. Xie—a wild form of P.
suffruticosa distinct from P. ostii
Paeonia suffruticosa ssp. yinpingmudan (Hong et al., 1998a) was treated by Haw
(2001) as a synonym of P. ostii, and the two specimens cited in the protologue were
suspected by him to be escaped. However, P. ostii is characterized by having bipinnate
lower leaves with 15 leaflets, which are mostly entire, ovate or ovate-lanceolate, and
hirsute at the base above, but glabrous beneath, whereas the type specimen of ssp.
yinpingmudan is characterized by having biternate lower leaves with 9 leaflets, which are
oblong to ovate-orbicular, all divided or lobed, and glabrous on both sides. Anyone who
has compared the type specimen of ssp. yinpingmudan (Mt. Yinping, Chaohu, Anhui
Province, K. Y. Pan & Z. W. Xie 9701 (PE)) with the type of P. suffruticosa (Andrews’
plate) (Fig. 1), would say that the identification of the collection K. Y. Pan & Z. W. Xie
Acta Phytotaxonomica Sinica Vol. 43 170
9701 as an element of P. suffruticosa is justifiable. From a comparison between Hong and
Pan’s (1999a) figures 1 and 4 (Fig. 2), P. suffruticosa ssp. yinpingmudan cannot be
confused with P. ostii.

Fig. 1. A comparison between the type of Paeonia suffruticosa Andrews and that of P. suffruticosa ssp. yinpingmudan
D. Y. Hong, K. Y. Pan & Z. W. Xie. A, The type of P. suffruticosa (Bot. Rep. 6: t. 373. 1804). B, The type of P.
suffruticosa ssp. yinpingmudan (K. Y. Pan & Z. W. Xie 9701(PE)).

We are not yet sure whether the material from Henan of ssp. yinpingmudan (Hong et
al. H97010 (PE)) is cultivated or introduced from a nearby mountain as Mr. Yang, our local
guide in the field, told us, but the type specimen of this subspecies (K. Y. Pan & Z. W. Xie
9701 (PE)) is quite certainly a wild one, because, first, it grew 30-40 m above the ground
on a cliff nearly 100 m high; second, the cliff is in a hilly area, far from cities or towns, and
there was no park or garden there; and third, it was said by local people that the peony had
existed there at least for 500 years. Therefore, we are not convinced that the peony was
escaped from cultivation. Our experience in 1985 on the Taibai Mountain, the Qinling
Range, perhaps could give us an implication. On the mountain we spent two days searching
for P. rockii (S. G. Haw & Lauener) T. Hong & J. J. Li ex D. Y. Hong, but only three
individuals were found, with two adults on cliffs and one young in a forest. A local person
told us, however, that about 25 years ago (i.e. around 1960) the peony was so common that
30-50 kg of fresh “Dan Pi” (bark of roots and stems) could be harvested a day by a single
person. There is possibility that wild P. suffruticosa (the subspecies yinpingmudan) was
more frequent in ancient time in mountainous or hilly areas of Anhui, Henan and some
other provinces. Due to the loss of its habitats and random harvest for medicinal and
ornamental purposes, it became rarer and rarer, reaching an extent that only the plant on
cliffs has survived.
No. 2 HONG De-Yuan et al.: Notes on taxonomy of Paeonia sect. Moutan DC. (Paeoniaceae) 171


Fig. 2. The lower (the best developed) leaves of the five species in the Paeonia suffruticosa Andrews complex. A, P.
suffruticosa ssp. yinpingmudan D. Y. Hong, K. Y. Pan & Z. W. Xie (type). B, P. jishanensis T. Hong & W. Z. Zhao. C,
P. qiui Y. L. Pei & P. Y. Hong. D, P. ostii T. Hong & J. X. Zhang. E1, P. rockii (S. G. Haw & Lauener) T. Hong & J. J.
Li ex D. Y. Hong ssp. rockii. E2, P. rockii ssp. atava (Brühl) D. Y. Hong & K. Y. Pan (syn. P. rockii ssp. taibaishanica
D. Y. Hong). Assembled from the figures 1-6 of Hong & Pan (1999a).
Acta Phytotaxonomica Sinica Vol. 43 172
1.2 Paeonia suffruticosa is not a hybrid
Haw (2001) recognized P. suffruticosa Andrews as a hybrid, but he did not give
substantial evidence to support his point of view. As we have pointed out (Hong et al.,
2004), cultivated tree peonies are polyphyletic, comprising hundreds of cultivars.
Single-flowered tree peonies cultivated for medicinal purpose belong to P. ostii.
Ornamentally cultivated tree peonies have been derived from different sources; some from
wild P. jishanensis T. Hong & W. Z. Zhao (cv. “Yi Ping Zhu Yi ”, “Chun Hong Jiao Yan”
in Luoyang peony gardens), some from wild P. rockii (a cultivar in Pengzhou peony
gardens), some from wild P. suffruticosa, i.e. ssp. yinpingmudan (see below), and many
others seem to have originated from hybridization between them. We consider that
ornamentally cultivated tree peonies comprise at least four species, P. suffruticosa, P.
jishanensis, P. rockii, and P. ostii, and numerous hybrids, and that not all of cultivars are of
hybrid origin. For example, the cultivars “Jin Yao Lou”, “Hong Dan Lan” in Pengzhou
peony gardens, Sichuan, and “Ge Jin Zi”, “Wang Hong” in Luoyang peony gardens,
Henan, are extremely similar to both the type of P. suffruticosa and the type specimen of P.
suffruticosa ssp. yinpingmudan, with no indication of involvement of P. jishanensis, P.
rockii or P. ostii in their origin.
1.3 Molecular data
On our molecular tree of Paeonia sect. Moutan constructed using GPAT gene
sequences, P. suffruticosa ssp. yinpingmudan and P. ostii form a clade, which is separated
from the P. jishanensis-P. qiui Y. L. Pei & D. Y. Hong clade with the bootstrap value as
high as 100%, but within the clade, ssp. yinpingmudan and P. ostii form two separate
clades with the bootstrap value as high as 96% (Zhao et al., 2004). This fact clearly
indicates that P. suffruticosa ssp. yinpingmudan is distinct from P. ostii, P. jishanensis and
P. qiui. We have also constructed a reduce-medium (RM) network of wild tree peonies
using PCR-RFLP data of 12 restriction enzymes. In the network P. suffruticosa ssp.
yinpingmudan forms an independent branch, which also shows the distinctness of P.
suffruticosa ssp. yinpingmudan from the other species (Zhao et al., 2004).
Twelve cultivars identified as P. suffruticosa based on morphology were compared
with P. suffruticosa ssp. yinpingmudan using PCR-RFLP profiles of 12 restriction
enzymes. Six of the 12 cultivars were found very similar to the subspecies, but none of
them was identical with it. The result indicates that ssp. yinpingmudan is not an escape, but
is a wild peony, from which many cultivars of P. suffruticosa have been derived (Zhou et
al., unpublished).
All the lines of evidence indicate that P. suffruticosa is an independent species with
ssp. yinpingmudan as its wild form. The species has been involved in the formation of
many, but not all of ornamentally cultivated tree peonies. The other species, P. rockii, P.
jishanensis and P. ostii, have been also involved in the origins of ornamentally cultivated
tree peonies. Cultivated tree peonies are polyphyletic in origin, comprising at least four
species, P. suffruticosa, P. jishanensis, P. rockii and P. ostii, and their hybrids. Therefore,
P. suffruticosa is an important component of the P. suffruticosa complex, and it still has a
wild form, ssp. yinpingmudan, from which many cultivars have been derived, and it has
been involved in the origin of many hybrid cultivars.
2 Paeonia qiui Y. L. Pei & D. Y. Hong—an independent species
The specific status of P. qiui has been recognized since its description in 1995.
However, Halda (1997, 2004) treated P. qiui as a variety of P. suffruticosa ssp. spontanea
(Rehder) S. G. Haw & Lauener.
No. 2 HONG De-Yuan et al.: Notes on taxonomy of Paeonia sect. Moutan DC. (Paeoniaceae) 173
Paeonia qiui is characterized by having lower leaves biternate, leaflets 9, ovate or
broadly ovate, mostly entire (Fig. 2), often purple above, and densely villose in axils of
major veins beneath, and petals whitish pink or pink, with a red or reddish blotch at the
base (Hong & Pan, 1999a, fig. 3, pl. 5). These characters clearly distinguish it from all the
other species, and thus P. qiui is a distinct species. Halda’s (1997) combination is not
justifiable.
3 Paeonia jishanensis T. Hong & W. Z. Zhao—a legitimate name
Haw (2001) still used the name P. spontanea (Rehder) T. Hong & W. Z. Zhao (Hong
& Osti, 1994), while treated P. jishanensis T. Hong & W. Z. Zhao (Hong et al., 1992) as its
synonym. P. jishanensis T. Hong & W. Z. Zhao was considered by him as an illegitimate
name, because, according to him, the type of P. suffruticosa var. spontanea Rehder was
included in the concept of P. jishanensis T. Hong & W. Z. Zhao. However, T. Hong &
Zhao’s (Hong et al., 1992) P. jishanensis does not include the type of P. suffruticosa var.
spontanea, buy instead T. Hong & Zhao indicated in both Chinese and Latin that P.
jishanensis differs from P. suffruticosa var. spontanea by having white petals but no
petaloid stamens (Hong et al., 1992). These authors did state (in Chinese) that petaloid
stamens in ssp. spontanea is an important character induced by domestication of wild tree
peonies, and thus ssp. spontanea should be reduced into a cultivar, “cv. spontanea”, but
they did not say that “cv. spontanea” was an element of P. jishanensis. Therefore, we
cannot see the reason to recognize P. jishanensis T. Hong & W. Z. Zhao as an illegitimate
name. It was two years later that they raised ssp. spontanea (Rehder) S. G. Haw & Lauener
to specific rank and treated P. jishanensis as its synonym. Thus, they created a superfluous
name, Paeonia spontanea (Rehder) T. Hong & W. Z. Zhao.
4 Paeonia ostii T. Hong & J. X. Zhang—a species distinct from P. suffruticosa
As described by T. Hong & J. X. Zhang (Hong et al., 1992) in the protologue, and
shown in their figure (Fig. 1), P. ostii has lower leaves bipinnate and leaflets mostly entire,
lanceolate or ovate-lanceolate, which match those of widely cultivated tree peonies for the
medicinal purpose. Hong and Pan (1999a, Fig. 4 & pl. 6) described and illustrated the
characters of P. ostii clearly. However, P. suffruticosa, as shown in the type (see Hong &
Pan, 1999a, pl. 1.), has lower leaves biternate with 9 leaflets, which are all divided or
lobed. Therefore, Halda’s (1997) treatment of P. ostii as a subspecies of P. suffruticosa is
unacceptable.
5 Paeonia suffruticosa ssp. atava (Brühl) S. G. Haw & Lauener—a
subspecies of P. rockii
Haw (2001) insisted his opinion that P. moutan ssp. atava Brühl as a doubtful taxon
(Haw & Lauener, 1990). The first author of the present paper thought after his expedition
to Tibet that the peony belonged to P. rockii, which was also cultivated in the Tashihungpo
Temple in Xigazê City, Tibet, and might well be introduced by Buddhist monks from the
Qinling Range, Shaanxi Province (Hong, 1997a). He also examined the type specimen in
the Royal Botanic Gardens, Kew (K) and found no difference between it and the tree peony
on Mt. Taibai, the Qinling Range, except in the petal colour. Haw & Lauener (1990) and
Haw (2001) emphasized the character of petals of ssp. atava, “entirely unblotched”,
“unblotched…, quite unlike those of P. rockii”. However, the blotched colour fades with
time if the specimen is not well preserved, a phenomenon we observed both in the field and
Acta Phytotaxonomica Sinica Vol. 43 174
herbarium. The type specimen of ssp. atava (King’s collector 549) does not show
“blotched petals” now, but in the protologue of this subspecies the petals were indeed
described as “white with a rosy hue, with a large dark-purple spot near the base”. This
character is typical of P. rockii. Anyone who compares Brühl’s (1896) pl. 126 with Hong’s
(1998) figure 2 (based on the type of ssp. taibaishanica) will find that ssp. “atava” is
extremely similar to ssp. “taibaishanica”, without significant difference between them.
Therefore a new combination, P. rockii ssp. atava (Brühl) D. Y. Hong & K. Y. Pan, is
made below and ssp. taibaishanica D. Y. Hong should be treated as its synonym.
6 Paeonia delavayi Franch.—a highly variable species
Haw (2001) agreed with us (Hong et al., 1998b) that P. delavayi is a very variable
species, including “the several formally recognized species and varieties...”, and “only one,
variable, species can be recognized…” We agree with him that “ for horticultural purposes,
it would be useful and appropriate for clones within this species to be given forma or
cultivar names”.
Since we recognize P. delavayi as one variable species, with P. lutea Delavay ex
Franch. and P. potanini Kom. treated as its synonyms, we naturally do not recognize P.
×franchetii and P. ×handel-mazzettii described by Halda (1997) as real hybrids. They
are just two of the innumerable forms in P. delavayi.
7 Paeonia ludlowii (Stern & Taylor) D. Y. Hong—a distinct species
When Hong (1997a) separated “ludlowii” from P. lutea and treated it as a new species,
he clearly indicated the differences between P. ludlowii and P. lutea (a synonym of P.
delavayi). One year later Hong and his co-workers (Hong et al., 1998b) showed strong
evidence that P. delavayi was a highly variable species with surprising polymorphism in
petal colour, and P. lutea was just one of the innumerable petal colour forms of P. delavayi.
Halda (1999) surprisingly still treated the entity as a subspecies in P. lutea, P. lutea
ssp. ludlowii (Stern & Taylor) J. J. Halda.
However, Haw (2001) fully supported Hong’s (1997a) separation of var. ludlowii
Stern & Taylor from P. lutea and elevation of the entity to specific rank. The following
revised key clearly distinguishes P. ludlowii from P. lutea (=P. delavayi).
1a. Carpels 1, rarely 2; follicles 4.7-7 cm long, 2-3.3 cm broad; seeds ca. 1.3 cm diam.; petals, filaments
and stigmata always pure yellow; leaf segments and lobes with acuminate teeth; plants congested,
1.5-3.5 m tall …………………………………………….……………………………………P. ludlowii
1b. Carpels 2-5; follicles less than 3.5 cm long, 1.5 cm broad; seeds less than 1 cm diam., petals, filaments
and stigmata variable in colour; leaf segments and lobes mostly entire; plants scattered, less than 2 m
tall………………………………………………….………………………………...………...P. delavayi
8 Taxonomic treatments
1. Paeonia suffruticosa Andrews in Bot. Rep. 6: t. 373. 1804; Stern, Stud. Gen.
Paeonia, 40. 1946, quoad nom.; D. Y. Hong & K. Y. Pan in Nord. J. Bot. 19: 291, pl. 1.
1999; D. Y. Hong, K. Y. Pan & Z. Q. Zhou in Acta Phytotax. Sin. 42: 275-283. 2004.
Type: Andrews’ plate cited (!).
Paeonia×suffruticosa S. G. Haw in the New Plantsman, 8 (3): 166. 2001, syn. nov.
1a. ssp. suffruticosa
牡丹

No. 2 HONG De-Yuan et al.: Notes on taxonomy of Paeonia sect. Moutan DC. (Paeoniaceae) 175
1b. ssp. yinpingmudan D. Y. Hong, K. Y. Pan & Z. W. Xie in Acta Phytotax. Sin. 36: 515.
1998. ——P. yinpingmudan (D. Y. Hong, K. Y. Pan & Z. W. Xie) B. A. Shen in
Lishizhen Medic. Mater. Med. Res. 12: 330. 2001. Type: China. Anhui (安徽): Chaohu (巢
湖), Mt. Yinping, on cliff, 1997-04-28, K. Y. Pan & Z. W. Xie 9701 (holotype, PE!).
银屏牡丹
2. Paeonia jishanensis T. Hong & W. Z. Zhao in Bull. Bot. Res. (Harbin) 12: 225. 1992;
D. Y. Hong & K. Y. Pan in Nord. J. Bot. 19: 292. 1999; S. G. Haw in The New Plantsman
8 (3): 163. 2001, pro syn. ——P. suffruticosa Andrews ssp. spontanea (Rehder) S. G.
Haw & Lauener var. jishanensis (T. Hong & W. Z. Zhao) J. J. Halda in Acta Mus.
Richnov. Sect. Nat. 4 (2): 30. 1997. Type: China. Shanxi (山西): Jishan (稷山), Xiqiu, alt.
1200 m, 1991-05-10, T. Hong 915010 (CAF).
Paeonia spontanea (Rehder) T. Hong & W. Z. Zhao in Bull. Bot. Res. (Harbin) 14:
238. 1994; S. G. Haw in The New Plantsman 8 (3): 163. 2001; D. Y. Hong & K. Y. Pan in
Nord. J. Bot. 19: 292. 1999, pro syn. ——P. suffruticosa Andrews var. spontanea Rehder
in J. Arn. Arb. 1: 193. 1920, p.p. excl. specim. Tai-Pei-Shan, Purdom s.n. (A). ——P.
suffruticosa Andrews ssp. spontanea (Rehder) S. G. Haw & Lauener in Edinb. J. Bot. 47:
278. 1990. Type: China. Shaanxi (陕西): “50 li W of Yenanfu” (Yan’an, 延安), 1910,
Purdom 338 (holotype, A!; isotypes, E!, K!, P!).
矮牡丹
3. Paeonia rockii (S. G. Haw & Lauener) T. Hong & J. J. Li ex D. Y. Hong in Acta
Phytotax. Sin. 36: 539. 1998; T. Hong & J. J. Li in Bull. Bot. Res. (Harbin) 12: 227. fig. 4.
1992, comb. invalid. ——P. suffruticosa Andrews ssp. rockii S. G. Haw & Lauener, in
Edinb. J. Bot. 47: 279. fig. 1a. 1990. Type: China. Gansu (甘肃): Wudu (武都), Farrer no 8
(holotype, E!; photo, PE!).
Paeonia papaveracea auct. non Andrews: Anonymous in Icon. Cormophyt. Sin. 1:
652, fig. 1303. 1972.
3a. ssp. rockii
紫斑牡丹
Paeonia rockii ssp. linyanshanii T. Hong & G. L. Osti in Bull. Bot. Res. (Harbin) 14:
237. figs. 1, 2. 1994. ——P. suffruticosa ssp. rockii var. linyanshanii (T. Hong & G. L.
Osti) J. J. Halda in Acta Mus. Richnov., Sect. Nat. 4 (2): 30. 1997, syn. nov. Type: China.
Gansu (甘肃): Wen Xian (文县), Baimahegou (白马河沟), alt. 1570 m, Q. R. Zhang
19930428 (CAF).
3b. ssp. atava (Brühl) D. Y. Hong & K. Y. Pan, comb. nov. ——P. moutan Sims ssp.
atava Brühl in Ann. Bot. Gard., Calcutta, 5 (2): 114. t. 126. 1896. ——P. suffruticosa
Andrews ssp. atava (Brühl) S. G. Haw & Lauener in Edinb. J. Bot. 47: 280. 1990, syn. nov.
Type: China. Xizang (西藏 ): Chumbi, Tuk Chang, 1884-06, King’s Collector 549
(holotype, K!).
Paeonia rockii (S. G. Haw & Lauener) T. Hong & J. J. Li ex D. Y. Hong ssp.
taibaishanica D. Y. Hong in Acta Phytotax. Sin. 36: 542. fig. 2. 1998. syn. nov. Type:
China. Shaanxi (陕西): Mt. Taibai (太白山), Shangbaiyun, alt. 1750 m, 1985-05-24. D. Y.
Hong & X. Y. Zhu PB85061 (holotype, PE!).
Paeonia suffruticosa Andrews var. spontanea Rehder in J. Arn. Arb. 1: 193. 1920,
p.p., quoad specim. Mt. Taibai.
太白山紫斑牡丹

Acta Phytotaxonomica Sinica Vol. 43 176
4. Paeonia delavayi Franch. in Bull. Bot. Soc. Fr. 33: 382. 1886; Stern, Stud. Gen.
Paeonia, 44. 1946; D. Y. Hong, K. Y. Pan & H. Yu in Ann. Missouri Bot. Gard. 85: 561.
1998; S. G. Haw in The New Plantsman 8: 167. 2001.
Paeonia delavayi var. angustiloba Rehder & Wilson in Sarg., Pl. Wilson. 1: 318.
1913; S. G. Haw in The New Plantsman 8: 252. 2001.
Paeonia×francheti J. J. Halda (P. lutea Franch.×P. potanini Kom.) in Acta Mus.
Richnov., Sect. Nat. 4 (2): 31. 1997, syn. nov.
Paeonia×handel-mazzettii J. J. Halda (P. delavayi Franch.×P. lutea Franch.) in Acta
Mus. Richnov., Sect. Nat. 4 (2): 31. 1997, syn. nov.
滇牡丹
Acknowledgements We are grateful to Ms. MA Li-Ming for her assistance in
preparation of the manuscript, and to Mr. YANG Xue-Jian for developing photos.
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芍药属牡丹组分类新注
洪德元* 潘开玉
(中国科学院植物研究所系统与进化植物学重点实验室 北京 100093)
摘要 自我们近年发表一系列牡丹组分类文章以来, 国内外基本上赞同我们8个种的分类系统,但
对一些问题仍有不同见解。本文进一步申述牡丹Paeonia suffruticosa Andrews 是一个独立的种而不
是人工杂种, 以及银屏牡丹P. suffruticosa ssp. yinpingmudan是牡丹P. suffruticosa的野生类型而不是
逸生的P. ostii的理由。上述论点也得到了分子树的支持。P. jishanensis T. Hong & W. Z. Zhao是一个
合法名称, 而P. spontanea (Rehder) T. Hong & W. Z. Zhao则确实是一个多余名。太白山紫斑牡丹的学
名应是P. rockii ssp. atava (Brühl) D. Y. Hong & K. Y. Pan, 因此P. moutan Sims ssp. atava Brühl不应是
可疑的分类群。本文还对Halda的6个组合和两个杂交种名作了处理。结果, 本文包括了5个新异名和
一个新组合。
关键词 芍药属; 牡丹; 银屏牡丹; 太白山紫斑牡丹; 新组合; 新异名