De-yuan Hong & Kai-yu Pan

A revision of the Paeonia suffruticosa complex (Paeoniaceae)


Hong, D.-Y. & Pan, K.-Y. 1999. A revision of the Paeonia suffruticosa complex (Paeoniaceae). -Nordic Journal of Botany 19: 289-299. Copenhagen. ISSN 0107-055X.


1a.

Paeonia suffruticosa ssp. suffruticosa

1b.

Paeonia suffruticosa Andrews ssp. yinpingmudan D.-Y. Hong, K.-Y. Pan et Z. W. Xie

2.

Paeonia jishanensis T. Hong et W. Z. Zhao

3.

Paeonia qiui Y. L. Pei et D.-Y. Hong

4.

Paeonia ostii T. Hong et J. X. Zhang

5.

Paeonia rockii (S. G. Haw et L. A. Lauener) T. Hong et J. J. Li

5a.

Paeonia rockii subsp. rockii

5b.

Paeonia rockii subsp. taibaishanica Hong

x

Paeonia x papaveracea Andrews

x

Paeonia x baokangensis Z. L. Dai et T. Hong


The taxonomical concept of the Paeonia suffruticosa complex i.e. Sect. Moutan Subsect. Vaginatae, has changed greatly since 1990. Six species and four subspecies have been described as new and two subspecies raised to specific level. Five species and two subspecies are recognized in the present revision, viz. P. suffruticosa subsp. suffruticosa and subsp. yinpingmudan, P. jishanensis, P. qiui, P. ostii, P. rockii subsp. rockii and subsp. taibaishanica. P. yananensis, P. ridleyi, P. spontanea, P. moutan subsp. atava, P. suffruticosa subsp. atava, P. rockii subsp. linyanshanii and P. ostii var. lishizhenii are treated as synonyms. P. papaveracea and P. baokangensis are proposed to be interspecific hybrids. A key to the recognized species and subspecies is provided. Biological features of the species are described and their distributions are mapped. The relationships between species are inferred and the origins of commonly cultivated tree peonies (P. suffruticosa and P. ostii) are discussed.

D.-Y. Hong and K.-Y. Pan, Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan, Beijing 100093, China.


Introduction

Paeonia suffruticosa Andrews (tree peony, mudan in Chinese) has been cultivated in China for more than two thousand years since the Han Dynasty, and now throughout the temperate region in the world. It is named as "King of Flowers" in China and is one of four candidates for the national flower of China. As one of the most famous ornamental flowers and Chinese medicinal plants, however, its circumscription as a species and its relationships with other woody species in the genus are sometimes ambiguous. Andrews' Bot. Rep. 6: t. 373 (1804) (the type of P. suffruticosa} shows that P. suffruticosa is a cultivated plant with double and red flowers and lobed leaflets. It is the first species described in the woody group of Paeonia. Sims' Curtis' Bot. Mag. t. 1154 (1807) (the type of P. moutan) also shows a cultivated plant with double and red flowers. They are apparently conspecific and match well the tree peony commonly cultivated as an ornament in gardens. Rehder (1920) described for the first time a variety of P.suffruticosa from the wild, viz. var. spontanea. This variety was recognized by Stem (1946), Fang (1958), Anonymous (1972) and Pan (1979). Stem (1946) divided the section Moutan DC. into two subsections: Subsect. Vaginatae and Subsect. Delavayana. In the former he included only one species, P. suffruticosa and its variety, var. spontanea. However, Stem's (1946) description of P. suffruticosa is mainly based on a specimen from Choni Lamasery, Gansu (Kansu) (J. F. Rock s. n.) with bipinnate leaves, rose-pink to white petals with a dark purple blotch at the base and yellowish white floral disc. Thus, Stern's "P. suffruticosa" is different from the one commonly cultivated in gardens (Andrews' (1804) P. suffruticosa) and must belong to another species. Rock's plant was later called "Rock's Variety" (Haw & Lauener 1990; Stem 1946). Fang (1958) , following Stem, also included specimens from the wild in Gansu and Shaanxi in his P. suffruticosa. He described P. yunnanensis as new, but it was apparently a cultivated plant with a double flower, and has been reduced to a synonym of P. suffruticosa by Pan(1979). Anonymous (1972) used "P. papaveracea Andrews" for the species from the wild in Gansu and Shaanxi (Rock's Variety"). Pan (1979) also realized that the group from Gansu and Shaanxi was different from the typical mudan in gardens and treated it as P. suffruticosa var. papaveracea (Andr.) Kerner. So, her P. suffruticosa includes three varieties, var. spontanea, var. papaveracea in addition to the typical one. Haw & Lauener (1990) recognized three subspecies in P. suffruticosa: the typical one (cultivated), subsp. Spontanea and subsp. rockii (i.e. Stern's suffruticosa or Pan's papaveracea), because they found that Andrews' papaveracea was also cultivated and could not be applied to "Rock's Variety". Therefore, only one species was recognized in the complex, P. suffruticosa, until early 1990's, although the first author of the present paper and his co-workers (Hong et al. 1988) realized that tree peony cultivated for Chinese traditional medicine, “mudanpi”, was different from the one commonly cultivated as an omament in gardens in having single and pure white flowers, red-purple disc and long and often entire leaflets. However, they did not present a taxonomnic treatment but just used the name "P. aff suffruticosa" for it.

T. Hong and his coauthors (Hong, T. et al. 1992, 1994, 1997) have recently described five new species and one new subspecies, and raised Haw and Lauener's two subspecies to specific level, i.e. P. ostii T. Hong et J.X. Zhang, P. jishanensis T. Hong et W. Z. Zhao, P. yananensis T. Hong et M. R. Li, P. ridleyi Z. L. Dai et T. Hong, P. baokangensis Z. L, Dai et T. Hong, P. rockii (S. G. Haw et L. A. Lauener) T. Hong et J. J.Li, P. spontanea (Rehder) T. Hong et W. Z. Zhao and P. rockii subsp. linyanshanii T. Hong et G. L. Osti. In late 1980's and early 1990's the first author of the present paper, Hong, D.-Y. and his students, Qiu, J.-Z. and Pei, Y.-L. made several expeditions to the Shennongjia Mountains in W. Hubei Province, the Qinling Range in Gansu Province, Shaanxi Province, and the Zhongtiao Mountains in S. Shanxi Province, and W. Henan Province. As a result of the expeditions, one new species, P. qiui (Pei & Hong 1995) and two new subspecies, P. rockii subsp. taibaishanica (Hong 1998) and P. suffruticosa subsp. yinpingmudan (Hong et al. 1998) were described. Summarizing what we have mentioned above, there nine valid specific names in the P. suffruticosa group (Stern's (1946) Subsect. Vaginatae), eight of wich were described in 1990's from a relatively small area, W. Hubei, W. Henan, S. Shanxi, Shaanxi and SE. Gansu. The questions arise whether there are really so many species in the group and which names are appropriate for the tree peony commonly seen in gardens and allies; a taxonomical revision is badly needed for this very famous group of plants.


Observation and discussion

With the above questions in mind, the first author of the present paper, D.-Y. Hong recently made several field trips with assistance of his students or colleagues to the distribution range ofthe group, to S. Shanxi in 1993, to W. Henan and Yan'an of Shaanxi in 1994, to W. Henan, W. Hubei and Shaanxi in 1997, supported by the National Geographie Society (Grant 5515-95) and the National Natural Science Foundation of China (NSFC 39391500 and 39630030), in addition to a number of expeditions to this area made by his students, Qiu J. Z. and Pei Y. L. from 1990 to 1993. He has visited in flowering seasons all the type localities except for that of P. rockii.

The Paeonia suffruticosa complex (P. suffruticosa and its allies) forms a natural group in Paeonia Sect. Moutan DC. (tree peonies), which is characterized by floral disc entirely enveloping 5 (very occasionally 6) tomentose carpels, while the other three species of the section, P. decomposita (Hong, D.-Y. 1996, 1997b), P. delavayi (Hong, D.-Y. et al. 1998a) and P. ludlowii (Hong, D.-Y. 1997a), have 1-4 (5) glabrous carpels which are enveloped only at the base or lower half by a much shorter floral disc.

D.-Y. Hong and his co-workers (Hong, D.-Y. et al. 1998b) have shown that the tree peony commonly cultivated in gardens, P. suffruticosa (Plate l), may have been derived from the wild one in Anhui and Henan provinces (Plates 2 and 3, Fig. l) , not from Rehder's spontanea (P. jishanensis){Plate 4, Fig. 2) as previously proposed (Stern 1946). They indicated the differences between P. suffruticosa and P. jishanensis and recognized the latter as an independent species, while treated the wild one in Anhui and Henan as a subspecies, P. suffruticosa subsp. yinpingmudan.

Haw & Lauener's (1990) "rockii", i.e. Anonymous' (1972) "papaveracea" or Pan's (1979) "P. suffruticosa var. papaveracea" is rather widely but scatteredly distributed in the Qinling Range in SE. Gansu and Shaanxi, the Funiu Mountains in W. Henan and the Shennongjia Mountains in W. Hubei. There are some differences between populations, but they form a natural group with leaves tri- (or bi-) pinnate; leaflets mostly 19-33, very occasionally less than 19; petals white but with a dark purple blotch at the base; filaments yellow; floral disc and Stigmata yellowish (Figs 5, 6; Plate 7), in which they differ from all other plants in the complex. The taxon has been raised to specific level by T. Hong and his coworker, P. rockii (S. G. Haw et L. A. Lauener) T. Hong et W. Z. Zhao(1994). The taxonomic history and infraspecific division of the species has been discussed and treated by D.-Y. Hong (1998). We have examined the type of P. papaveracea Andrews (Bot. Rep. 7: t. 463. 1807) and found that it has a single flower, white petals with a purple-red blotch at the base, dark purple disc, purple-red filaments, red Stigmata and deeply lobed leaflets. This colour plate is extremely similar to the plant we found in the garden and the adjacent field, and in secondary forests on Wanhua Shan, Yan'an, Shaanxi (Plate 8). It is considered as a hybrid between P. rockii and P. spontanea and identical with P. yananensis T. Hong et M. R. Li (see the explanation under P. x papaveracea). The application of this name by Anonymous (1972) and Pan (1979) to the wild plant in Gansu and Shaanxi (=P. rockii), and by Haw & Lauener (1990) to a cultivar in P. suffruticosa subsp. suffruticosa is unreasonable.

Paeonia ostii T. Hong et J. X. Zhang is, according to the present authors, an independent species, which is widely cultivated mostly in villages and fields, but very rarely in gardens, and is just the source of "mudanpi", a famous Chinese traditional medicine. It has now become clear that the medicinal plant commonly cultivated in villages and fields with single flowers is P. ostii (Plate 6), whereas the omamental one commonly cultivated in gardens with double flowers is P. suffruticosa subsp. suffruticosa. Their differences are not only in whether flowers are double or single, but also in leaves which are bipinnate with most leaflets entire in P. ostii (Fig. 4), while biternate with all or most. leaflets lobed in P. suffruticosa subsp. suffruticosa.

Paeonia qiui Y. L. Pei et Hong was found only in one population in the Shennongjia Mountains (Songbai Town) when it was described as new. It was also found in 1997 in Baokang County, about 50 km E. of the type locality, Songbai Town. It is the most closely related to P. suffruticosa, from which it differs only by leaflets mostly entire (Fig. 3) and usually purple above and petals with a red blotch at the base (Plate 5).

Paeonia ridleyi Z. L. Dai et T. Hong is considered as a synonym of P. qiui because no any distinct difference has been found between them and their types are from the same mountain range; P. spontanea is a synonym of P. jishanensis T. Hong et W. Z. Zhao. P. yananensis T. Hong et M. R. Li is a hybrid and is a synonym of P. x papaveracea as indicated under P. x papaveracea, and P. baokangensis Z. L. Dai et T. Hong (Plate 9) is also proposed to be a hybrid (see below). As Hong, D.-Y. et al. (1988) pointed out, in the section Moutan (woody group) the chromosome number is always 2n=10, and no obvious difference in karyotype was detected between species. Therefore, there should be no reproductive barrier between species in this section as inferred from the karyotype.


Taxonomical treatment Key to the species and subspecies

1. The best-developed leaves biternate, with 9 leaflets

... 2

1. The best-developed leaves bi- or tri-pinnate with more than 9 leaflets

... 5

2. Petals with no blotch at base; leaflets always or nearly always lobed, green above; flowers double or single

... 3

2. Petals with a red blotch at base; leaflets mostly entire, often purple above; flowers single

3. P. qiui

3. Leaves pubescent at least along veins on lower surface; lateral leaflets rounded or broadly ovate, 3-lobed, lobes usually again 2-lobed

2. P. jishanensis

3. Leaves glabrous; lateral leaflets ovate, 2-or 3-lobed, rarely entire

... 4

4. Flowers double

1a.P.suffruticosa ssp. suffruticosa

4. Flowers single

1b.P.suffruticosa ssp. yinpingmudan

5. Petals entirely white; floral disc purple; best-developed leaves with 11-15 leaflets

4. P. ostii

5. Petals white but with a large dark-purple blotch at base; floral disc yellowish; best developed leaves with leaflets (17)19-33

... 6

6. Leaflets lanccolate or ovate-lanceolate, mostly entire

5a. P. rockii ssp. rockii

6. Leaflets ovate or ovate-rounded, mostly lobed

5b. P. rockii ssp. taibaishanica


l. Paeonia suffruticosa Andrews

Bot. Rep. 6:t. 373. 1804; Rehder, Man. Cult. Trees and Shrubs, 214. 1927; Chen, 111. Man. Chin. Trees & Shrubs, Ist ed. 261. Fig. 193. 1937, 2nd ed. 261. Fig. 193. 1953; Stern, Stud. Gen. Paeonia, 40. 1946, quoad nom.; Fang in Acta Phytotax. Sin. 7(4); 313, 1958 p.p. excl. syn. P. papaveracea Andrews and P. decomposita Hand.-Mazz.; K. Y. Pan in Fl. Reip. Pop. Sin. 27: 41. 1979, p. p. excl. syn. P. decomposita Hand.-Mazz. -Type: Andrews' plate. Bot. Rep. 6: t. 373. 1804(!)

The species is characterized by biternate leaves with no more than 9 leaflets and leaflets mostly lobed. It includes one of the most famous omaments, mudan, commonly cultivated in gardens, and its wild relative, here treated as two subspecies.

None of the specimens cited under “P. suffruticosa" by Stern (1946) belongs to this taxon; those from Kansu (Gansu), Tibet and Bhutan belong to P. rockii, while that from Tatsien-lu (Kangding), Sichuan, is P. decomposita. D.-Y. Hong (1997a, b) has made it clear that in NW. Sichuan (including Kangding) there is only woody species of Paeonia, P. decomposita, whereas the plants in S. Tibet and Bhutan are P. rockii which was introduced there.

1a. Paeonia suffruticosa ssp. suffruticosa top

Paeonia arborea Donn, Hortus Cantabrig., 3rd ed., 102. 1804, nom. nud.

P. suffruticosa var. purpurea Andrews, Bot. Rep. 7. t. 448, 1807. - Type: the plate cited(!)

P. moutan Sims, Curtis' Bot. Mag. 29. t. 1154. 1808. - Type: the plate cited (!)

P. yunnanensis Fang in Acta Phytotax. Sin. 7(4): 306. Pl. 61-2. 1958. - Type: Yunnan, Lijiang, T. T. Yu 8143 (KUN!)

Flowers always double, various in colour (Plates 1-3). Only known in cultivation. It has been cultivated for more than two thousand years and has hundreds of cultivars. The subspecies is considered to have been domesticated from its closest relative, the subspecies yinpingmudan (Hong, D.-Y. et al. 1998b).

1b. Paeonia suffruticosa Andrews ssp. yinpingmudan D.-Y. Hong, K.-Y. Pan et Z. W. Xie top

Acta Phytotax. Sin. 36 (6): 515-520. 1998. - Type: China, Anhui, Caohu, Yinping Shan, on cliff, 1997,04,28, Pan, K. Y. and Xie, Z. W. 9701 (PE!)

This subspecies has only two extant individuals according to our extensive field work. The type individual is said to have existed on the cliff for hundreds of years. The flowers are purely white and single (Plate 2). Its filaments and Stigmata are purple-red and floral disc is red-purple. One leaf we obtained is bitemate and glabrous; leaflets 9, with terminal ones triangular, segmented, lateral ones ovate, 2-or 3-lobed, rarely entire. Thus its leaf is extremely similar to P. suffruticosa but differs remarkably from P. jishanensis (Figs l and 2).

The individual cultivated by Mr Yang in Henan also has single but pale purple-red flowers, with leaves similar to P. suffruticosa, but different obviously from P. jishanensis (Plates 3 and 4). Mr Yang told us that he introduced the shrub from a nearby mountain in early 1960's, where it cannot be found at present.

Specimens examined. Henan, Songxian, Muzhijie Township, Shigunping, Secaogou, at side of Yang Hui-fang's house, 1997,04,28, Hong, Ye and Feng H97010 (PE, MO).

2. Paeonia jishanensis T. Hong et W. Z. Zhao top

Bull. Bot. Res. 12(3): 225. Fig. 2. 1992. - Type: China, Shanxi, Jishan, Xiqiu, alt. 1200 m. 1991,05,10. Hong, T.915010(CAF,n.v.)

P. spontanea (Rehder) T. Hong et W. Z. Zhao Bull. Bot. Res. 14(3): 238, 1994.

P. suffructicosa ssp. spontanea (Rehder) S. G. Haw & L. A. Lauener in Edinb. J. Bot. 47(3): 278. 1990.

P. suffruticosa var. spontanea Rehder in J. Arn. Arb. l. 193. 1920, p.p., excl. specim. Tai-pei-shan (Mt. Taibai), Purdom s.n.(syntype of var. spontanea. A!); Stern, Stud. Gen. Paeonia, 43. 1946; K. Y. Pan, Fl. Reip. Pop. Sin. 27: 45. 1979. - Type (lectotype): China, Shaanxi, "50 li W. of Yenanfu" [Yan'an!], 1910, Purdom 338 (here designated A! isolectotypes E, K).

The group has been treated at three different levels, as variety by Rehder (1920), Stern (1946), Anonymous (1972) and Pan (1979), as subspecies by Haw & Lauener (1990) and as independent species by Hong, T. & Zhao (1994). The major differences between "suffruticosa" and "spontanea" are leaflet shape (particularly lateral ones), leaflet division and leaf indumentum in addition to whether there exist plantlets from root turions (spontanea) or not (suffruticosa).

The entity is scatteredly found in the Zhongtiao Mountains (Yongji County in Shanxi and Jiyuan County in Henan), the Luliang Mountains (Jishan County, Shanxi), the Huashan Mountains (Huayin County, Shaanxi), and Tongchuan and Yan'an both in Shaanxi (Fig. 7). All these populations except the one in Yan'an grow in thickets or in secondary deciduous forests at altitudes from 900 to 1700 m. The population in Yan'an consists of several individuals at the western side of the peony garden behind Zhaojun Temple in Wanhua Shan, and may well be introduced and naturalized there. Because the garden has been established probably for hundreds of years, and no one (including local people) knows its exact history, it is hard to give a definite answer about the history of P. jishanensis in Yan'an. All the populations mentioned above have purely white petals which are occasionally pinkish at the periphery (Plate 4).

Specimens examined. Henan: Jiyuan, 1050 m alt. 1997,04,26 Wang. S. Y. H97001 (PE, MO); Huadong-shu Forest Farm, 1200 m alt., 1994,05,22, Wang, S.Y. and Zhang, Y.J. 940220 (PE).

Shaanxi, Yan'an, Wanhua Shan, 36.6N, 109.4E, near the peony garden, 1997,05,09, Hong, D. Y. and Feng Y. X. H97066 (PE); eodem loc., Hsia, W. Y. 3519 (PE); Shaanxi-Gansu Basin, Le, T. Y. s. n. (PE).

Shanxi, no precise locality, 1390 m alt 1916,06.16, P. Licent 1909 (PE); Jishan: Xishe, 1550 m alt. 1989,05, Qiu, J. Z. PB89201 (PE); eodem loc. 1440 m alt. Qiu, J. Z. PB89501 (PE) ; Majiagou Village, 35.7N, 110.9 E. 1400 m alt. 1996,05,14, Zhou, S. L. H96053 (PE, A, K, MO, S); Majiagou Forest Farm, Chayuangou, 1450 m alt. 1982,05,14, Liu T. W. and Zeng Z. F. 165 (PE); eodem loc. Pei, Y. L. 9002 (PE); eodem loc., Pei, Y. L. and Hong, D. Y. 93003 (PE), 93006 (PE), 93008 (PE), 93009 (PE); Yongji, Shuiyukou Village, Pei, Y. L. 9170 (PE), 9180 (PE). 9201(PE); eodem loc., Pei, Y. L. and Hong, D. Y. 93011 (PE), 93427 (PE).

3. Paeonia qiui Y. L. Pei et D.-Y. Hong top

Acta Phytotax. Sin. 33(1): 91. Fig. l. 1995. -Type: China, Hubei, Shennongjia, Sunbai (Songbai) Town, 1650-2010 m alt. on steep precipice and overhanging rocks, 1988, 05, 20, Qiu J. Z. PB88034 (PE!).

P. ridleyi Z. L. Dai et T. Hong in Bull. Bot. Res. 17(1):1. Fig.l(p.4). 1997, syn. nov. - Type: China, Hubei, Baokang County, Laoyashan, Chongchongya (Changchongya), 1400 m alt. 1994,05,03, Dai, Z. L. 94053 (Baokang Forestry Research Institute Hubei Province!; photo, PE!).

The species is characterized by bitemate leaves with 9 leaflets, which are ovate or broad-ovate, mostly entire and often purple above (Fig, 3), and whitish pink or pink petals with a red blotch at the base (Plate 5). Apparently it is the most closely retated to P. suffruticosa. It sometimes reproduces vegetatively by root turions.

P. qiui is so far found only in four localities in W. Hubei (Songbai Town in the Shennongjia Nature Reserve, and Houping Town and Laoyashan in Baokang County) and W. Henan (Xixia) (Fig. 7). Three of four remaining populations are found on cliffs with a few individuals. Therefore, it is the most endangered species in Paeonia, on the verge of extinction. It is very urgent to take effective measures to conserve the species.

Specimens examined. Hubei, Shennongjia, Songbai Town: 1988,04,25, Qiu, J.Z. PB88018 (PE) and PB88021 (PE); eodem loc. 2000 m alt. 1988,05, 06 Qiu, J.Z. PB88022 (PE), PB88023(PE), PB88024(PE), PB88025(PE) and PB88026(PE); eodem loc.,1991,05,04, Pei, Y.L. 9110 (PH); Songbai Town, sunny grassy slope, 2200 m alt. 1986,05,19, Chen, T. and Ma, L. M. PB86013 (PE); Baokang, Houping Town, Hongjiayuan, in front of Mr. SU Yuan-zhi's house, 1997,05,02, Hong, Ye and Feng H97023 (PE, A, K, MO, S); eodem loc. Yanghu Shan, 1000 m alt. limestone, in secondary forests, 1997,05,02, Hong, Ye and Feng H97027 (PE, MO) and H97028(PE, MO); Houping Town, Chefongping, 1300 m alt., E slope, limestone, on cliffs. 1997,05,03, Hong, Ye and Feng H97029 (PE, A, K, MO, S); eodem loc., Wudaoxia, 1000 m alt., 1997,05,04, Hong, Ye and Feng H97045 (PE).

Henan, Xixia, in forests, 1600 m, 1988,05, Qiu, J. Z. 8305(PE).

4. Paeonia ostii T. Hong et J. X. Zhang top

Bot. Res. 12(3): 223. Fig. l (p.231). 1992. - Type:

Zhengzhou Institute of Aeronautical Industrial Management, Arboretum of Rare and Endangered Trees, introduced from Mt. Yangshan of Songxian, 1990,05,10, Hong,T. 905010(CAF).

P. ostii T. Hong et J. X. Zhang var. lishizhenii B. A. Shen in Acta Phytotax. Sin. 35(4): 360. 1997. syn. nov.

- Type: Anhui, Nanling, Yashan 200-250 m alt. road-side on slope, 1984,04,18, Shen B. A. PB1018 (holotype in Wuhu Municipal Drug Bureau of Anhui Province; isotype, PE!).

The diagnostic characters of this species are leaves bipinnate; leaflets 11-15 on the best developed leaves, ovate, ovate-lanceolate, mostly entire (Fig. 4); flowers single and purely white; filaments red-purple; floral disc dark purple and Stigmata red (Plate 6).

Although we did not examine the type specimen, we have seen the shrub in the campus of Zhengzhou Institute of Aeronautical Industrial Management, from which the type specimen was collected.

It is a distinct species, but had been confused with P. suffruticosa subsp. suffruticosa (Pan 1979) until D.-Y. Hong et al. (1988) who realized their differences, though they did not formally describe it as new. They just used the name P. aff. suffruticosa for it.

We have compared the type and protologue of this species to plants commonly cultivated for "mudanpi" (a famous Chinese traditional medicine) in Tongling, Anhui Province, the famous cultivation locality of "mudanpi". They resemble each other very well and thus the plant commonly cultivated for "mudanpi" should be identified as P. ostii.

P. ostii is cultivated on a large scale in Tongling, Anhui Province and also widely but scatteredly in other provinces, Henan, Hubei, Shaanxi, Sichuan, etc. So far we have not found it in the wild. Hong, T. et al. (1992) stated that the shrub, from which the type was collected, was introduced from "Songxian, Yangshan, alt. 1200 m, in thickets on slopes". We visited the place in Henan Province twice, in 1994 and 1997, and searched extensively for the plant in the mountains. We did find the species cultivated in Secaogou Village, but not in wild in the mountains. However, we found in 1998 its existence in wild in W. Henan (Lushi Co) (Hong et al. H98005).

Specimens examined. Anhui: Nanling, Yashan, 1984,04,18, Sheng, B. A. 1018 (the type of P. ostii var. lishizhenii, PE); Mt. Jiuhua in cultivation, 1986,04,28, Hong, D. Y. and Chen, T. PB86007(PE).

Henan, Neixiang, Baotianman Nature Reserve, Mu2huliu Village, 800 m alt., cultivated. 1997,04,30. Hong, Ye and Feng H97021 (PE); Xixia, 1600 m alt. in forest. 1988,05,14, Qiu, J.Z. PB88302 (PE); Songxian, Jiulongdong, 1200 m alt. 1994,05,07, Jia H.Y. 001 (PE) and 002 (PE); Muzhuwa, 1150 m alt. 1994,05,20, Jia, H.Y. 004 (PE) and 005 (PE); Nuzhaihuai, 1150 m alt. 1994,05,24, Jia, H.Y. 011 (PE); Luanshitou, 1250 m alt. 1994,05,25 Jia, H.Y. 034 (PE); Bailigou, 1200 m alt. 1994,05,27, Jia H.Y. 036 (PE) and 038 (PE); Bailigou, Dongling, 1200 m alt. 1994,05,18, Jia, H.Y. 042(PE) and 043 (PE).

Hubei, Baokang, Siping Town, Jiejiaping Village, cultivated, 1997,05,05, Hong, Ye and Feng H97052(PE).

Shaanxi, Mt. Taibai, 1350 m alt. in cultivation, 1985,05,23. Hong, D. Y. and Zhu, X. Y. PB85052 (PE); Huangling, Huangdi Tomb, cultivated, 1994, 05, 04, Hong, Pan and Zhang 94007(PE); eodem loc. 1997, 05, 10, Hong, D. Y. and Feng, Y. X. H97069 (PE).

5. Paeonia rockii (S. G. Haw et L. A. Lauener) T. Hong et J. J. Li top

Bull. Bot. Res. 12(3): 227. Fig. 4. 1992; D. Y. Hong in Acta Phytotax. Sin. 37: 1998.

P. suffruticosa Andrews subsp. rockii S. G. Haw et L. A. Lauener in Edinb. J. Bot. 47(3): 279. Fig. l a (p.276). 1990. - Type: China, Kansu (Gansu), "probably near Wutu (Wudu), (Farrer's Chieh Jo)", Farrer no 8(?) (E, photo PE!).

P. papaveracea auct. non Andrews: Anonymous, Icon. Cormophyt. Sin. l: 652. Fig. 1303. 1972.

P. suffruticosa Andr. var. papaveracea auct. non. Andrews: K. Y. Pan in Wang, W. T. (ed.): Fl. Reip. Pop. Sin. 27: 45. pl. 3(p.43). 1979.

P. suffruticosa auct. non Andrews: Stern, Stud. Gen. Paeonia, 40. 1946, p. p.; Fang, in Acta Phytotax Sin. 7(4): 313. 1958, p.p.

P. moutan Sims subsp. atava Brühl in Ann. Bot. Gard. Calcutta 5(2): 114. pl. 126. 1896, syn. nov. -Type: Tibet (Xizang), Chumbi, Tuk Chang, 1884, 06, King's Collector 549 (K).

P. suffruticosa Andr. subsp. atava (Brühl) S. G. Haw et L. A. Lauener, in Edinb. J. Bot. 47(3): 280. 1990. syn.nov.

P. suffruticosa Andrews, "Rock's Variety' ('Joseph Rock') hort.

This species is characterized by leaves tri- (or bi-) pinnate; leaflets usually 19-33, very rarely less than 19; filaments yellow; floral disc yellowish white; Stigmata yellow and petals white but with a dark purple blotch at the base. Thus, the species differs distinctly from the other four in the complex. And its flowers are the largest in the whole genus.

D.-Y. Hong (1997a) has described bis searching for P. moutan subsp. atava in its type locality, Yadong, S. Tibet, in 1996 with failure, and explained his consideration that so-called "atava" was actually "P. rockii" introduced by lamas.

It is relatively widely distributed in four provinces, Gansu, Shaanxi, Henan and Hubei (Fig.8). Two subspecies can be recognized.

5a. Paeonia rockii subsp. rockii top

Paeonia rockii subsp. linyanshanii T. Hong et G. L. Osti in Bull. Bot. Res. 14(3):237, Figs l & 2. 1994. -Type: Gansu, Wenxian, Baimahegou, alt. 1570 m, 1993,04,28, Zhang, Q. R. 19930428 (CAF).

This subspecies (Fig. 5; Plate 7) is relatively widely distributed and usually grows in deciduous forests at an altitude of 1100-2800 m .

Specimens examined. Gansu, Zhugqu, Taozhou Forest Farm, on edges of Pinus armendii forests, 2800 m alt. 1959,05,22, Jiang S. and Jin C. L. 423 (PE); no precise locality, Pei, Y.L. 9116 (PE); Wudu, Lanshan, 2500 m alt., 1930,06,21, Hao K. S. 501 (PE); Wenxian, no precise locality, Pei Y. L. 9115 (PE); Zhilong Shan, Shihuigou, 1956,05,14, Yellow River Exped. 3698 (PE).

Shaanxi, Lueyang, Baishuijiang Town, Siping Village, Pei, Y. L. 9140 (PE).

Henan: Songxian, Muzhijie Township, Mt. Yangshan, Yangjiaohao, W. slope, limestone, in Ulmus forest, 1450 m alt. 1994,05,02, Hong, D.Y. and Ye Y. Z. 94003 (PE); eodem loc. Daxigou, under limestone rock, 1994,05,02, Wang, S. Y. and Wang, Y. Z. 94002 (PE); Xisangou, 1080 m alt., 1994,05,07, Jia, H. Y. 023 (PE) and 024 (PE); Neixiang, Baotiaman Nature Reserve, Muzhuliu, Mudanduo, 1100 m alt. in secondary deciduous forest, 1997,04,30, Hong, Ye and Feng H97015 (PE, MO); eodem loc, Muzhuliu Village, in cultivation, 1997,04,30, Hong, Ye and Feng H97016 (PE, A, CAS,K, MO, S).

Hubei. Shennongjia, Songbai, 1900 m alt. 1988,05,06, Qiu, J.Z. PB88027 (PE), PB88028 (PE) and PB88029 (PE); eodem loc., 2000 m alt. 1988,05,06, Qiu, J. Z. PB88030 (PE); eodem loc. 2100 m alt.

1988,05,06, Qiu, S.Z. PB88031(PE) and PB88032 (PE); eodem loc. 1900 m alt. 1988,05,20, Qiu, S.Z. PB88033 (PE) ; eodem loc“ 1600 m alt. 1988,05,20, Qiu, J.Z. PB88035 (PE). Songbai, Shantunya, 1988,05 Qiu, J.Z. PB3601 (PE); Songbai, shady slope, in forests, 1400-1800 m alt. 1986,05,18, Chen, T. and Ma, L. M. PB86008 (PE) & PB86075 (PE); Baokang, Houping Town, Hongjiayuan Village, in front of Mr. SU Yuan-zhi's house, in cultivation, 1997,05,02, Hong, Ye and Feng H97024 (PE); Houping Town, Zhanjiapo Village, in deciuotis forest, on limestone rocks, 1360 m alt. 1997,05,05, Hong, Ye and Feng H97051( PE, A, CAS, K, MO, S).

5b. Paeonia rockii subsp. taibaishanica Hong top

Acta Phytotax. Sin. 36: 1998. - Type: Shaanxi, Mt. Taibai, Shangbaiyun, 1750 m alt. in deciduous forest, on cliff, 1985,05,24, Hong, D. Y. and Zhu, X. Y. PB85061 (PE!).

The subspecies is clearly different from the typical one in having leaflets ovate or ovate-rounded, mostly or totally lobed (Fig. 6). According to our present knowledge, it is confined to northem slope of the Qingling Range (Fig. 8) and grows in deciduous forests at an altitude of 1300-2000 m.

Specimens examined. Shaanxi, Mt. Taibai, 1985,10,13, Zhu, X. Y. and Wu, Z. H. PB85086 (PE); eodem loc., Dadian Lamasery, 2300 m alt. in cultivation, 1985,05,24, Hong, D. Y. and Zhu, X. Y. PB85066(PE); eodem loc., Shangbaiyun, in front ofthe temple, 1820 m alt. in cultivation, 1997,05,08, Hong, Ye and Feng H97058 (PE, A, CAS, K, MO,. S); eodem loc., Heihukuan, among bushes, 1939,05,05, Fu, K. T. 2584 (PE); Longxian , Pei, Y. L. 916001 (PE). Gansu, Tianshui, Baiyangling, Zhang, Z. W. 13(PE).


Paeonia x papaveracea Andrews (pro sp.) top

Bot. Rep. 7:t. 463. 1807. - Type: the plate cited (!).

P. yananensis T. Hong et M. R. Li in Bull. Bot. Res. 12(3): 226. Fig. 3(p. 233). 1992, syn. nov. - Type: China, Shaanxi, Yan'an, Wanhua Shan, in Plalycladus forests, 1991, 05,13, Hong, T. 915013 (CAF).

The specimen Hong, D. Y. and Feng, Y. X. H97068 (see below) extremely resembles Andrews' P. papaveracea in having lobed leaflets, single flowers, white petals with a purple-red blotch at the base, purple-red filaments, red-purple floral disc and red Stigmata (Plate 8). Hong, Pan and Zheng 94009 was collected exactiy at the type locality of P. yananensis, and Hong, Pan and Zheng 94011 was collected at the place ca. 400-500m E of the type locality. These threee Specimens are very similar to each other, and the latter two differ from Hong, D. Y. and Feng, Y. X. H97068 only in having petals pink instead of white. The number of their leaflets is mostly 11-15, in between P. rockii and P. jishanensis. All their characteristics can be found in these two species. Furthermore, P. jishanensis exists on Wanhua Shan, near the peony garden and P. rockii can also be found in the garden. Therefore, we have sufficient evidence to propose that P. papaveracea Andrews is a hybrid between P. rockii and P. jishanensis, and the hybrid escaped from the garden and have become naturalized.


Specimens examined. Shaanxi, Yan'an, Mt. Wanhua Shan, in Platycladus forest, 1994,05,05, Hong, Pan and Zheng 94009(PE); eodem loc. in the valleyofeast side, in Platycladus forest, Hong, Pan and Zheng 94011 (PE); Yan'an, Wanhua Township, Wangjiagou Village, in waste field, Hong, Pan and Zheng 94008 (PE); Yan'an, Mt. Wanhua Shan, in the valley of east side, in cultivation, 1997,05,09, Hong, D. Y. and Feng, Y. X. H97068 (PE).

Paeonia x baokangensis Z. L. Dai et T. Hong (pro sp.) top

Bull. Bot. Res. 17(1): 2. Fig. 2 (p.5). 1997. - Type: China, Hubei, Baokang County, Houping Township, 1996,05,02, Dai Z. L“ Ran, D. Y. & Li, Q. D. 96047 (Baokang Forestry Research Institute! photo PE!).

We consider that this is a hybrid between P. qiui and P. rockii, because all the characteristics of P. baokangensis can be found in these two species. Dai and Hong, T.'s protologue cited the type: "Baokang, Houpingzhen, Chefenggou, alt. 1600 m, in bush, May 2, 1996, Dai, Z.L“ Ran, D. Y. & Li Q. D. 96047". The first author of the present paper with assistance of bis Student, Miss Feng Y. X., and Prof. Ye Y. Z. at Henan Agricultural University vistited Mr Dai and examined the type.

Following Mr Dai's guidance we found the shrub from where the type was collected. The shrub was actually growing at the side of Mr Qi Xin-hua's house in Hongjiayuan village, Houping Town, 380 m alt. When we visited Mr Qi, there was a flower on the shrub just at anthesis, which is exactiy the same äs the type specimen in every aspect (Plate 9). Mr Qi told us that the shrub was introduced from a nearby mountain at least 20 years ago. In front of Mr Su Yuan-zhi's house in the same village but a couple of kilometers W. of Mr Qi's house, numerous individuais of Paeonia were growing. Among them, there were different forms, P. rockii, P. qiui and P. obovata, all of which were introduced from the nearby mountains according to Mr Su, and several hybrid individuals apparently came from hybridization between P. qiui and P. rockii. The story may well be: the villagers introduced peonies to their houses; P. qiui and P. rockii hybridized when they met in the village. In the nearby mountains in Houping Town we found both P. qiui (Mt. Chefongping, Hong, Ye and Feng H97029) and P. rockii (near Zhanjiapo village, Hong, Ye and Feng H97051), but not P. baokangensis in the mountains.

Specimens examined. Hubei, Baokang, Houping Town, Hongjiayuan Village, at the side of Mr. Qi Xin-hua's house. 380 m alt. 1997,05,02, Hong, Ye and Feng H97026 (PE, A, CAS, K, MO, S); the same village, in front of Mr. Su Yuan-zhi's house, 400 m alt. 1997,05,02, Hong, Ye and Feng H97025(PE).

Acknowledgements - The field work was supported by the National Geographie Society (Grant 5515-95) and the National Natural Science Foundation of China (NSFC 39391500 and 39630030). We are gratefül to Dr Qiu Jun-zhuan, Dr Pei Ye-long, Dr Zhang Shou-zhou, Mr Xie Zhong-wen and Miss Feng Yu-xin at our labora-tory, Professors Ye Yong-zhong, Wang Sui-yi and Dr Wang Ying-zheng at the Henan Agricultural University and Zheng Hong-chun at the Yan'an University for their assistance in the field work. We are also indebted to Mr Dai Zhen-lun at Baokang Forestry Research Institute of Hubei Province for his kind help. Our thanks are extended to the Harvard University Herbaria for sending us the syntypes of Paeonia suffruticosa var. spontanea on loan and to the Edinburgh Botanical Garden for sending us a photo of the type of P. suffruticosa subsp. rockii.

References top

Anonymous, 1972. Icon. Cormophyt. Sin. l. 652. Fig. 1303. - Science Press, Beijing.

Fang, W.- P. 1958. Notes on Chinese paeonies. - Acta Phytotax. Sin. 7(4): 297-323. Haw, S. G. & Lauener, L. A. 1990. A revision of the infraspecific taxa of Paeonia suffruticosa Andrews. - Edinb. J.Bot. 47(3): 273-281. Hong, D.-Y. 1996. On the identity of Paeonia decomposita Handel-Mazzetti. - Taxon 45: 67-69.

- 1997a. Paeonia (Paeoniaceae) in Xizang (Tibet). - Novon 7: 156-161.

- 1997b. Notes on Paeonia decomposita Hand.-Mazz. - Kew Bull. 52(4): 957-963.

- 1998. Paeonia rockii and its one new subspecies from Taibai Shan, Shaanxi. - Acta Phytotax. Sin. 36: 538-543.

- , Pan K.-Y. & Yu, H, 1998a. Taxonomy of Paeonia delavayi (Paeoniaceae).- Ann. Missouri Bot. Gard. 85(4): 554-564.

- , Pan, K.-Y. & Xie, Z.-W. 1998b. Yingpingmudan - the wild relative of the king of flowers, Paeonia suffruticosa Andrews. - Acta Phytotax. Sin. 36: 515-520.

- , Zhang, Z.-X. & Zhu, X.-Y. 1988. Studies on the genus Paeonia (l) - Report ofkaryotypes ofsome wild species in China. - Acta Phytotax. Sin. 26(1): 33-43. Hong, T. & Osti, G. L. 1994. Study on the Chinese wild woody Peonies(II): new taxa of Paeonia L. Sect. Moutan DC. - Bull. Bot. Res. 14(3): 237-240.

- & Dai, Z.-L. 1997. Study on the Chinese wild woody Peonies (III): new taxa of Paeonia L. Sect. Moutan DC. - Bull. Bot. Res. 17(1); 1-5.

-, Zhang, J.-X., Li, J.-J., Zhao, W.-Z. & Li, M.-R. 1992. Study on the Chinese wild woody peonies (l): new taxa of Paeonia L. Sect. Moutan DC. - Bull. Bot. Res. 12(3):223-234. Pan, K.-Y. 1979. Paeonia. - In: Wang. W.-T. (ed.), Flora Reipublicae Popularis Sinicae. 27: 37-59. Science Press. Beijing.

Pei, Y.-L. & Hong, D.-Y. 1995. Paeonia qiui - a new woody species of Paeonia from Hubei, China. - Acta Phytotax. Sin. 33(1): 91-93.

Stern, F. C. 1946. A Study ofthe Genus Paeonia. - Roy. Hort. Soc., London.


Figures top

Fig. 1 A well-developed leaf of Paeonia suffruticosa subsp. yinpingmudan.

Fig. 2. The second (from below) (the best developed) leaf of Paeonia jishanensis.

Fig. 3. The second (from below) (the best developed) leaf of Paeonia qiui.

Fig. 4. The second (from below) (the best developed) leaf of Paeonia ostii

Fig. 5. The second (from below) (the best developed) leaf of Paeonia rockii subsp. rockii.

6. The second (from below) (the best developed) leaf of Paeonia rockii subsp. taibaishanica.

Fig. 7. Distribution map of: squares Paeonia suffruticosa subsp. yingpingmudan. - dots P. spontanea. - triangle P. qiui.

Fig. 8. Distribution map of Paeonia rockii: black dots subsp. rockii. - circles subsp. taibaishanica.