Abstract The taxonomical history of the woody group( peonies) in the genus Paeonia is reviewed in the present paper. The group is endemic to China, but Paeonia suffruticosa is commonly cultivated throughout the north temperate region and has long been known as "the King of Flowers" in China. However, the taxonomy of the group had been neglected before the 1990's. Since 1990, a number of new species and subspecies have been published. With the support from the National Geographic Society, the senior author of the present paper and his coworkers have made expeditions to all parts of the distribution area of the group, visited all the type localities and sampled a total of 64 populations. Based on the character analysis and examination of the type specimens or photos, each taxon is reviewed with its name checked nomenclaturally, and finally the taxonomy of the whole group is revised. As a result, eight species, three of which each contains two subspecies, are recognized. They are Paeonia suffruticosa Andrews subsp. suffruticosa and subsp. yinpingmudan D. Y. Hong. K. Y. Pan et Z. W. Xie, P. jishanensis T. Hong et W. Z. Zhao, P. qiui Y. L. Pei et D. Y. Hong, P. ostii T. Hong et J. X. Zhang, P. rockii (S. G. Haw et L. A. Lauener) T. Hong et J. J. Li subsp. rockii and subsp. taibaishanica D. Y. Hong, P. decomposita Hand.-Mazz. subsp. decomposita and subsp. rotundiloba D. Y. Hong, P. delavayi Franch. and P. ludlowii (Stern et Taylor) D. Y. Hong. P. baokangensis Z. L. Dai et T. Hong and P. yananensis T. Hong et M. R. Li are considered as hybrids between P. rockii and P. qiui and between P. rockii and P. jishanensis respectively. In addition, the reduction of a number of names recently published to synonyms is explained.

Key words Paeonia; Section Moutan ; Taxonomical history; Revision Paeonia sect. Moutan DC.

pages [352][353][354][355][356][357][358][359][[360][361] in chinese only, help needed for translation

Key to species

1. Flowers usually 2 or 3, terminal and axillary, ± pendent; disc fleshy, enveloping only the base of carpels.

2. Carpels usually 2-5( -7) ; follicles <4 x 1.5 cm; petals, filaments and stigma often not purely yellow; plants less than 2 m tall

7. P. delavayi Franch

2. Carpels nearly always single; follicles 4.7-7 x 2-3.3 cm; petals, filaments and stigma always yellow; plants 1.5-3.5 m tall

6. P. ludlowii (Stern et Taylor)D. Y. Hong

1. Flowers solitary, erect; disc leathery, enveloping completely carpels or half of them.

3. Disc enveloping half of carpels at anthesis; carpels 2-4(5), glabrous; leaves decomposite; leaflets(29) 33 - 63 in number, all lobed

6. P. decomposita Hand.-Man.

4. Carpels nearly always 5; leaflets narrow, terminal ones with length/width (1. 5) 1.7-2.7 (-3. 3); lobes narrow, terminal ones with length/width (1.6-)2.4-3.7(-4.3)

6a. subsp. decomposita

4. Carpels 2-5, usually 3 or 4; leaflets broader, terminal ones with length/width (1.0-) 1.2-1.8 (-2.2); lobes broader, terminal ones with length/width (1.0 -) 1.3-2.4 (-3.0)

6b. subsp. rotundiloba D. Y. Hong

3. Disc completely enveloping carpels at anthesis; carpels 5 (-7), densely tomentose; leaves biternate or bi- or tripinnate; leaflets usually less than 20 in number, if more at least some of them entire.

5. Leaves biternate; leaflets usually 9 in number.

6. Leaflets ovate or ovate-rounded, mostly entire, often reddish above; petals with a red spot at base

3 P. qiui YL. Pei et D. Y. Hong

6. Leaflets long-ovate, ovate or nearly rounded, green above; petals without spot.

7. Leaflets long-ovate or ovate, terminal leaflets deeply 3-lobed, with additional 1 to several lobes, lateral leaflets 2- or 3-lobed, some leaflets entire; lobes acute at apex; leaves glabrous on lower surface

1. P. suffruticosa Andrews

8. Flowers double, cultivated

1a. subsp. suffruticosa

8. Flowers single, wild

1b. subsp. yinpingmudan D. Y. Hong, K. Y. Pan et Z. X. Xie

7. Leaflets ovate-rounded to rounded, all deeply 3-lobed; lobes again lobed, acute to rounded at apex; leaves villose along veins on lower surface

2 P. jishanensis T. Hong et W. Z. Zhao

5. Best developed leaves pinnate; leaflets more than 9 in number, long-ovate to lanceolate, mostly entire, less frequently ovate-rounded, mostly lobed.

9. Leaves bipinnate; leaflets no more than 15 in number, ovate to ovate-lanceolate, mostly entire; petals white, without spot

4. P. ostii T. Hong et J. X. Zhang

9. Leaves tri-(less bi-) pinnate; leaflets (17)19-33 in number, lanceolate or ovate-lanceolate, mostly entire, or ovate to ovate-rounded, mostly lobed

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

10. Leaflets lanceolate to ovate-lanceolate, mostly entire

5a. subsp. rockii

10. Leaflets ovate to ovate-lanceolate, mostly lobed

5b. subsp. taibaishanica D. Y. Hong

(1) .......

Paeonia suffruticosa Andrews, Bot. Rep. 61 t. 373. 1804; Rehder, Man. Cult. Trees and Shrubs. 214. 1927; Chen, 111. Man. Chin. Trees and Shrubs. 1st ed. 261, fig. 193. 1937; Stern, Stud. Gen. Paeonia 40. 1946, quoad nom. i Fang in Acta Phytotax. Sin. 7(4): 313. 1958, p. p. excl. syn. P. papaveracea Andrews et 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(1)

Stern(1946) P. suffruticosa ..............

362

............ P. decomposita,

(1a) .........

Paeonia suffruticosa subsp. suffruticosa

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. PI. 61:2. 1958. TYPE: China. Yunnan, Lijiang, T. T. Yu 8143(KUN! )

P. suffruticosa var. rubro-plena (hort. )Baily in Rhodora 18: 156. 1916. syn. nov. TYPE: not designated.

P. suffruticosa var. roseo-superba (hort. ) Baily, 1. c. syn. nov. TYPE: not designated.

P. suffruticosa var. vittata (van Houtte) Baily, I.e. syn. nov.

P. moutan var. vittata van Houtte, Fl. De Serr.7,5,747. TYPE: the table cited (!)

P. suffruticosa var. banksii (hort. ) Baily, I.e. syn. nov. TYPE: not designated.

P. suffruticosa var. rosea (Lodd. ) Baily, I.e. syn. nov. TYPE: not designated.

P. suffruticosa var. humei (hort. ) Baily, I.e. syn. nov. TYPE: not designated.

.............

(1b)..........

Paeonia suffruticosa subsp. yinpingmudan D. Y. Hong, K. Y. Pan et Z. W. Xie in Acta Phytotax. Sin. 36(6): 519, fig. 2, colour pi. 2 & 3. 1998. TYPE: China. Anhui, Caohu, YinpingShan, on cliff, 1997-04-28, K. Y. Pan and Z. W. Xie 9701 (holotype, PE!)

............

(2) .............

Paeonia jishanensis T. Hong et W. Z. Zhao in Bull. Bot. Res. (Harbin)12(3): 225, fig. 2. 1992. TYPE: China. Shanxi, Jishan, Xiqiu, alt. 1200 m, 1991-05-10, T. Hong 915010 (holotype, CAF, n.v.).

Paeonia suffruticosa var. spontanea Rehder in J. Am. Arb. 1: 193. 1920, p. p. excl. specim. Tai-pei-shan (Mt. Taibai), Purdom, s.n. (syntype of var. spontanea Rehder, A!); Stern, Stud. Gen. Paeonia, 43. 1946; K. Y. Pan in Fl. Reip. Pop. Sin. 27: 45. 1979. TYPE: China. Shaanxi, "50 li W of Yenanfu"(Yan'an! ), 1910, Purdom 338 (lectotype. A! isolectotype, E, K)

Paeonia suffruticosa subsp. spontanea (Rehder) S. G. Haw et L. A. Lauener in Edinb. J. Bot. 47(3): 278, fig. Ic. 1990.

Paeonia spontanea (Rehder) T. Hong et W. Z. Zhao in Bull. Bot. Res. (Harbin) 14 (3): 238. 1994.

............Haw & Lauener, 1990; Pan, 1979;

363

Stern. 1946).......... 900-1700 ............

(3) ...........

Paeonia qiui Y. L. Pei et D. Y. Hong in Acta Phytotax. Sin. 33(1): 91, fig. 1. 1995. TYPE: China. Hubei, Shennongjia, Songbai(Sunbai) Town, alt. 1650-2010 m, 1988-04-05, J. Z. Qiu PB88034 (holotype, PE! )

Paeonia ridleyi Z. L. Dai et T. Hong in Bull. Bot. Res. (Harbin) 17(1): 1, fig. 1. 1977. TYPE: China. Hubei, Baokang, Laoyashan, Chongchongya(Changchongya), alt. 1400 m, 1994-05-03, Z. L. Dai 94053(holotype in Baokang Forestry Research Institute of Hubei Province! Photo, PE! )

........... P. delavayi .................

.............

(4) .............

Paeonia ostii T. Hong et J. X. Zhang in Bull. Bot. Res. (Harbin) 12(3): 223, fig. 1. 1992. TYPE: China. Zhengzhou Institute of Aeronautical Industrial Management, Arboretum of Rare and Endangered Trees, introduced from Mt. Yangshan of Songxian, Henan Province, 1990-05-10, T. Hong 905010 (holotype, CAP)

Paeonia ostii T. Hong et J. X. Zhang var. lishizhenii B. A. Shen in Acta Phytotax. Sin. 35(4): 360. 1997. TYPE: China. Anhui. Nanling, Yashan, alt. 200-250 m, roadsides, on slope, 1984-04-18, B. A. Shen PB1018(holotype in Wuhu Municipal Drug Bureau of Anhui Province; isotype, PE!)

............ P. suffruticosa .............. P. ostii ..............

(5) .........

Paeonia rockii (S. G. Haw et L. A. Lauener ) T. Hong et J. J. Li in Bull. Bot. Res.(Harbin) 12(3): 227, fig. 4. 1992; D. Y. Hong in Acta Phytotax. Sin. 37(6); 539, fig. 1. 1998.

Paeonia suffruticosa Andrews subsp. rockii S. G. Haw et L. A. Lauener in Edinb. J. Bot. 47(3): 279, fig. la. 1990. TYPE: China. Kansu(Gansu), "Probably near Wutu [ 364 ] (Wudu), (Farrer's Chieh Jo)", Farrer no 8(?)(holotype, E; photo, PE! )

Paeonia papaveracea auct. non. Andrews: Anonymous, Icon. Cormophyt. Sin. 1: 652, fig. 1303. 1972.

P. suffruticosa Andrews var. papaveracea auct. non. Andrews: K. Y. Pan in Fl. Reip. Pop. Sin. 27: 45, pl. 3. 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.; Grierson et Long, Fl. Bhutan. Vol. 1. Part 2: 321. 1984.

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

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

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

...............

(5a) .............

Paeonia rockii subsp. rockii

Paeonia rockii subsp. linyanshanii T. Hong et G. L. Osti in Bull. Bot. Res. (Harbin) 14(3): 237, fig. 1 & 2. 1994. TYPE: China. Gansu, Wenxian, Baimahegou, alt. 1570m, 1993-04-28, Q. R. Zhang 19920428 (holotype, CAF, n.v.).

.............. 1100-2800 m ...........

(5b) ..............

Paeonia rockii subsp. taibaishanica D. Y. Hong in Acta Phytotax. Sin. 36(3): 542, fig. 2. 1998. TYPE: China. Shaanxi, Mt. Taibai, Shangbaiyun, alt. 1750 m, in broad-leaved deciduous forest on cliff, 1985-05-24, D. Y. Hong and X. Y. Zhu PB8506 (holotype, PE!)

....................

(6) .............

Paeonia decomposita Hand.-Mazz. in Acta Hort. Gothob. 13: 39. 1939; D. Y. Hong, K. Y. Pan & Y. L. Pel in Taxon 45: 68. 1996; D. Y. Hong in Kew Bull. 52(4): 957-963. 1997. TYPE: China. NW Sichuan, Chosodjo, H. Smith 4641 (holotype, UPS! Photo, PE!)

365

Paeonia szechuanica Fang in Acta Phytotax. Sin. 7: 315. 1958; K. Y. Pan in Fl. Reip. Pop. Sin. 27: 45, fig. 4. 1979; D. Y. Hong in L. K. Fued. China Plant Red Data Bookl: 536-537. 1992. TYPE; China. NW Sichuan, Barkam, X. Li (H. Lee)70316 (holotype. SZ! isotype, PE!)

(6a)..................

Paeonia decomposita subsp. decomposita D. Y. Hong in Kew Bull. 52(4): 958, fig. 1 B. 1997.

.............. 2050-3100 m ..................

(6b) ..............

Paeonia decomposita subsp. rotundiloba D. Y. Hong in Kew Bull. 52(4): 961, fig. 1A. 1997. TYPE: China. NW Sichuan, Lixian. D. Y. Hong, Y. B. Luo & Y. H. He H95033 (holotype, PE! isotype, GH/A, K, MO. US!).

.................... 1700 -2700 m...................

(7) ................

Paeonia delavayi Franch. in Bull. Soc. Bot. France 33: 382. 1886; Stern, Stud. Gen. Paeonia 44. 1946; K. Y. Pan in Fl. Reip. Pop. Sin. 27: 1979; D. Y. Hong, K. Y. Pan & H. Yu in Ann. Missouri Bot. Gard. 85: 554-564. 1998. TYPE: China. NW Yunnan, Lijiang(Likiang), Delavay 1142 (holotype, P; isotype, K).

Paeonia lutea Delavay ex Franch. in Bull. Soc. Bot. France 33: 382. 1886;

Paeonia delavayi var. lutea (Delavay ex Franch.) Finet & Gagnep. in Bull. Soc. Bot. France 51: 524. 1904. TYPE: China. NE Yunnan: Eryuan County, Hea Chan Men, 1883-05-25. Delavay s. n. (lectotype, P; isolectotype, K).

Paeonia potanini Kom. in Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 2: 7. 1921. TYPE: China. W Sichuan, Yajiang (Tachien-lu district in Yalung valley), Potanin s. n. (holotype, LE).

Paeonia trollioides Stapf ex Stern, J. Roy. Hon. Soc. 56: 77. 1931;

Paeonia potanini var. trollioides (Stapf ex Stern) Stern, Stud. Gen. Paeonia 50. 1946. TYPE: China. NW Yunnan, Deqen, Baima Shan, Mekong-Yangtse divide, 3300 m, open stony pastures, Forrest 13195 (holotype, E).

Paeonia delavayi var. lutea f. superba Lemoine, Rev. Holt. 14: cum tab., 1906;

Paeonia lutea var. superba (Lemoine )hort. ex Gard. Chron., Ser. 3, 44: 50, cum tab., 1908. TYPE: [pl. 14 in Lemoine. Rev. Hort. 1906]

Paeonia delavayi var. angustiloba Rehder & E. H. Wilson in Sarg., PI. Wilson. 1: 318. 1913. TYPE: China. W Sichuan, Yajiang(W of Tachien-lu, descent to Yalung river),

366

3000-3600 m, 1908-10, Wilson 1333(holotype, A).

Paeonia delavayi var. atropurpurea Schipcz. in Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 2: 47. 1921. TYPE: China. No locality given (holotype, ? LE).

Paeonia. delavayi var. alba Bean, Tree5 Shrubs 3: 265. 1933;

Paeonia potanini f. alba (Bean)Stern, Stud. Gen. Paeonia 49. 1946. TYPE: t. 49 in Stern, Stud. Gen. Paeonia, 1946" (holotype!).

(8).............................

Paeonia ludlowii (Stern et Taylor) D. Y. Hong in Novon 7: 157, fig. 1 & 2. 1997.

Paeonia lutea var. ludlowii Stern et Taylor in J. Roy. Hort. Soc. 76: 217. 1951. TYPE: China. SE Tibet[Kongbo Prov. Miling, Tsangpo Valley, Ludlow], Sherriff & Taylor 4540 (holotype, BM).

...........................

Paeonia x papaveracea Andrews ( pro. sp. ) in 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. TYPE: China. Shaanxi, Yan'an, Wanhua Shan, in Platycladus forests, 1991-05-13, T. Hong 915013(CAF)

.............P. rockii ....P. jishanensis, ............(Hong and Pan, 1999),.................

Paeonia x baokangensis Z. L. Dai et T. Hong (pro. sp.) in Bull. Bot. Res. (Harbin) 17(1): 2, fig. 2. 1997. TYPE: China. Hubei, Baokang County, Houping Township, 1996-05-02, Z. L. Dai, D. Y. Ran & Q. D. Li 96047(Baokang Forestry Research Institute! Photo, PE!)

...............Hong and Pan(1999), .............P. rockii ..P. qiui ..................

..............................

367

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