accepted name (2005):
1997
original description of J.J. Halda
P. obovata Maximowicz subsp. japonica (Makino) J. J. Halda stat. nov.
[bas: P. obovata Maximowicz var. japonica Makino In Bot. Mag. Tokyo 12:302 (1898)].
Differs from subsp. obovata in having glabrous leaves and white flowers with short stigmas and being diploid (2n=10). Sachalin, Japan.
Hong 2001:
Hong D Y, Pan K Y & Rao G Y, 2001: Cytogeography and taxonomy of the Paeonia obovata polyploid complex. Pl. Syst. Evol. 227(3/4): 123—136.
Plant Syst. Evol. 227: 123-136 (2001)
Plant Systematics and Evolution
© Springer-Verlag 2001 Printed in Austria
Cytogeography and taxonomy of the Paeonia obovata polyploid complex (Paeoniaceae)
D.-Y. Hong1, K.-Y. Pan1, and G.-Y. Rao2
(1) Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing, China
(2) College of Life Science, Peking University, Beijing, China
Received October 30, 1998 Accepted June 21, 2000
Abstract. 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 below.
Key words: Paeoniaceae, Paeonia obovata polyploid complex, morphological variation, cytogeog-raphy, taxonomic revision.
The 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. obovata (Stern 1946). He followed Miyabe and Takeda (1910) in distinguishing P. japonica as an independent species, and thus recognized three taxa, P. obovata 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 var. 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. obovata 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).
Previous 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.
This 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.
Materials and methods
From 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).
Results and discussion
Leaf 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. H98026).
Table 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)
Leaf 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).
Fig. 1. Morphological variation in the Paenia obovata complex:
a Hong et al. H98026 (Mt. Changbai. China) with spreading (instead of ascending) leaves in a red-flowered form:
b Hong et al. H98006 (Lushi Co. Henan China) with petals white but pink at periphery, filaments purple at base but white above, and orange-red anthers;
c Hong D. Y. P1190011 (Uwa-dio Ehime, Japan) with pink petals;
d 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
Sepals. They exhibit the least variation, are nearly always 3, very occasionally 2 or 4 in number and mostly rounded at the top. Only one of the sepals sometimes is caudate in some individuals of some populations (e.g. Hong et al. H98026).
Petals: 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. H98006)(Fig. 1b-e).
The 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).
Petals 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.
Stamens: 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.
The 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. obovata.
Carpels 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.
According 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.
Style 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.
Fig. 2. Chromosomes of the Paeonia obovata Maxim, complex, all at mitotic metaphase except for B, which is at meiotic anaphase II.
A Hong et al. PR8507H (red flowers), 2n= 10;
B Zhu X. V. et al. PR86020 (white flowers), n = 5;
C Hong et al. H9803I (white flowers?), 2n=10;
D Hong and Zhu PB85062 (red flowers), 2n = 20;
F. Hong and Zhu PB85067 (white flowers), 2n = 20;
F Hong and Zhu PR85024 (white flowers), 2n = 20;
G Hong el al. H98014 (red flowers), 2n = 20.
All X 1250. only D X 1600
Chromosomes. 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).
Fig. 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
Furthermore, 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.
Conclusions and taxonomical treatment
It 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.
Paeonia obovata Maximowicz, Prim. Fl, Amur. 29 (1859)
Type: Amur, Kitsi; Maximowicz (holotype, LE; isotype, K!)
The species comprises two geographically allopatric subspecies which are keyed out below:
Leaves 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)
............. subsp. obovata
Usually densely, very occasionally sparsely, hirsute or pubescent on lower surface; tetraploid (the only diploid found in a mixed population in Lushi Co, W. Henan)
.................. subsp. willmottiae
Subsp. obovata
P. 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)
P. obovata var. typica Makino, Bot. Mag. (Tokyo) 12:301 (1898); J. Jap. Bot. 5(9): 33. 1928
P. 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).
P.vernalis Mandl in Bot. Kozl., 19:90. (1921); type: Russia, the Far East, Nikolsk-Ussurisk (holotype, LE).
P. obovala var. amurensis Schipczinsky, Not. Syst. Herb. Hort. Bot. Pctrop. 2: 44. (1921); type: not designated.
P. obovata var. australis Schipczinsky, Not. Syst. Herb. Hort. Bot- Petrop. 2: 44. (1921): type: not designated.
P. obovata var. glabra Makino in J. Jap. Bot. 5(9):33. (1928). type: Japan (not designated).
P. obovam var. alba Saunders, Nat. Hort. Mag. 13, tab., 227. (1934); type: Saunders' figure cited.
Paeonia japonica var. pilosa Nakai in J. Jap. Bot. 12: 394-395(1937); type: Korea, Heinan. ML Rorinsan (holotype. Til),
This 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.
Representative specimens examined:
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. G,37902(PE, 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).
JAPAN: 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:
Koguchi, Yuansongci, s.n. (KYO); Yamanashi: Fujiyoshida City, Araya, Konta, F. 11786 (KYO).
KOREA 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).
RUSSIA: 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).
Paeonia obovata Maxim, subsp. willmottiae (Stapf) D. Y. Hong et K. Y. Pan, stat. nov.
P. willmottiae Stapf in Curtis' Bot. Mag. 142: tab. 8667(1916); type: "Miss. Willmott May 22, 1915" (holotype K!)
P. obovata Maxim, var. willmottiae (Stapf) F. C. Stern in J. Roy. Hort. Soc. 68: 128 (1943) et Stud. Gen. Paeonia, 76(1946).
This 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.
Representative 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).
We 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
particularly 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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Addresses of the authors: Hong De-yuan and Pan Kai-yu, Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan, Beijing 100093, China. Rao Guang-yuan, College of Life Science, Peking University, Beijing 100871, China.
