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PAEONIACEAE 1. PAEONIA Linnaeus, Sp. Pl. 1:530. 1753

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PAEONIACEAE 1. PAEONIA Linnaeus, Sp. Pl. 1:530. 1753 Flora of China 6: 127–132. 2001. PAEONIACEAE 芍药科 shao yao ke Hong Deyuan (洪德元)1, Pan Kaiyu (潘开玉)1; Nicholas J. Turland2 Shrubs or perennial herbs, to 3.5 m tall. Roots fleshy, thick but attenuate toward tip, or tuberous. Stems (in herbs) or current year’s branchlets (in shrubs) terete, with several persistent scales at base. Leaves alternate, compound; proximal 1–3 best developed, 1–3-ternate or 1- or 2-pinnate; leaflets entire or divided. Flowers solitary and terminal, or 2 or more per shoot and both terminal and axillary, more than 4 cm in diam. Bracts 1–6, leaflike, varying in shape and size, grading into sepals. Sepals 2–9, varying in shape and size. Petals 4–13, varying in color. Stamens up to 230; filaments and anthers varying in color. Disc leathery or fleshy, annular (in herbs) or extended into a sheath and 1/3 to wholly enveloping carpels (in shrubs). Carpels 1–5(–8), free, glabrous or hairy; ovules numerous, borne in two rows along ventral suture. Styles present or absent; stigmas laterally flattened, recurved, crested. Fruit a follicle. Seeds black or dark brown, globose or ovoid-globose, to 1.3 cm in diam. One genus and ca. 30 species: NW Africa, temperate Asia and Europe, W North America; 15 species (ten endemic) in China. Pan Kai-yu. 1979. Ranunculaceae subfam. Paeonioideae. In: Fl. Reipubl. Popularis Sin. 27: 37–59. 1. PAEONIA Linnaeus, Sp. Pl. 1:530. 1753 芍药属 shao yao shu Morphological characters and geographical distribution are the same as those of the family. Paeonia is a very complex genus and many of the species are not yet well defined. A consistent taxonomic treatment will require further studies throughout the world distribution of the genus in order to resolve questions about the limits of, and relationships between, the species. 1a. Herbs perennial; disc not well developed, annular. 2a. Leaflets and segments up to 9, entire; carpels glabrous ................................................................ 9. P. obovata 2b. Leaflets and segments (of proximal leaves) more than 9, at least some of them segmented; carpels hairy or glabrous. 3a. Leaflets and segments of proximal leaves up to 20, some of them entire, mostly more than 2 cm wide. 4a. Carpel 1(or 2) ...................................................................................................................... 12. P. emodi 4b. Carpels 2–5. 5a. Flowers usually several per shoot, both terminal and axillary; leaflets and segments with bristles along veins adaxially ................................................................................................ 11. P. lactiflora 5b. Flowers usually solitary, terminal; leaflets and segments glabrous ............................. 10. P. mairei 3b. Leaflets and segments of proximal leaves more than 20, all or nearly all of them segmented, mostly less than 2 cm wide. 6a. Plants glabrous throughout ........................................................................................... 13. P. sterniana 6b. Leaflets and segments usually with bristles along veins adaxially; carpels usually brown- yellow hirsute. 7a. All or most sepals caudate at apex; roots cylindric, attenuate toward tip ................. 14. P. anomala 7b. Most sepals (at least 2) not caudate at apex; roots obviously tuberous ................. 15. P. intermedia 1b. Shrubs; disc well developed, 1/3 to wholly enveloping carpels. 8a. Flowers usually 2 or 3 per shoot, both terminal and axillary, ± pendent; disc fleshy, enveloping only base of carpels. 9a. Carpels usually 2–5(–7); follicles to 4 × 1.5 cm; petals, filaments, and stigmas often not pure yellow; plants less than 2 m tall .............................................................................................. 7. P. delavayi 9b. Carpel nearly always 1; follicles 4.7–7 × 2–3.3 cm; petals, filaments, and stigmas always yellow; plants 1.5–3.5 m tall .......................................................................................................................... 8. P. ludlowii 8b. Flower 1 per shoot, terminal, erect; disc leathery, 1/2 to wholly enveloping carpels. 10a. Disc ca. 1/2 enveloping carpels at anthesis; carpels 2–4(or 5), glabrous; leaflets (30–)35–65, all lobed .............................................................................................................................. 6. P. decomposita 1 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. 10b. Disc wholly enveloping carpels at anthesis; carpels 5(–7), densely tomentose; leaflets usually less than 20 or, if more, at least some of them entire. 11a. Proximal leaves pinnate; leaflets more than 9. 12a. Proximal leaves 2-pinnate; leaflets up to 15, ovate-lanceolate to ovate, mostly entire ......................................................................................................................... 4. P. ostii 12b. Proximal leaves (2 or)3-pinnate; leaflets (17–)19–33, lanceolate or ovate-lanceolate and mostly entire, or ovate to ovate-orbicular and mostly lobed ........................... 5. P. rockii 11b. Proximal leaves 2-ternate; leaflets usually ca. 9. 13a. Leaflets adaxially often reddish, ovate or ovate-orbicular, mostly entire, abaxially densely tomentose at axils of major veins; petals often with a red spot at base ....... 3. P. qiui 13b. Leaflets adaxially green, long ovate, ovate, or ovate-orbicular, mostly divided, abaxially glabrous, villous along veins, or villous throughout; petals unspotted at base. 14a. Leaflets long ovate or ovate, abaxially glabrous; terminal leaflets deeply 3- lobed, with an additional 1 to several lobes; lateral leaflets 2- or 3-lobed, sometimes entire; lobes acute at apex ................................................................................ 1. P. suffruticosa 14b. Leaflets ovate-orbicular, abaxially villous along veins or throughout; all leaflets deeply 3-lobed; lobes again lobed, apex rounded to acute .................... 2. P. jishanensis 1. Paeonia suffruticosa Andrews, Bot. Repos. 6: t. 373. 1804. • Known only in cultivation in China; widely cultivated in temperate 牡丹 mu dan regions elsewhere. Shrubs to 1.5 m tall. Stems brown-gray. Proximal This subspecies has been cultivated for more than 2000 years and hundreds of cultivars are known. leaves 2-ternate; leaflets long ovate or ovate, 4.5–8 × 1b. Paeonia suffruticosa subsp. yinpingmudan D. Y. Hong 2.5–7 cm, both surfaces glabrous; terminal leaflets & al., Acta Phytotax. Sin. 36: 519. 1998. deeply 3-lobed, lobes again 2- or 3-lobed; some lateral 银屏牡丹 yin ping mu dan leaflets 2- or 3-lobed, others entire; all lobes acute at Flowers single; petals white or pale red-purple. Fl. Apr– apex. Flowers solitary, terminal, single or (in cultivated May. plants) double, 10–17 cm wide. Bracts 5, long elliptic, • Cliffs; ca. 300 m. C Anhui (Yinping Shan), W Henan (Song Xian). unequal. Sepals 5, green, broadly ovate, unequal. Petals At present only two individuals of this subspecies are known. The (in single flowers) 5–11, white, pink, red, or red-purple, plant in Anhui grows on a cliff and is said by local people to be at obovate, 5–8 × 4.2–6 cm, apex irregularly incised. least 500 years old; the plant in Henan is cultivated but is said by its Filaments pink or purple, white distally, ca. 1.3 cm; owner to have been collected on a nearby mountain during the early anthers long ellipsoid, ca. 4 mm. Disc wholly 1960s (Hong et al., Acta Phytotax. Sin. 36: 517. 1998). enveloping carpels at anthesis, purple-red, leathery, 2. Paeonia jishanensis T. Hong & W. Z. Zhao in T. Hong & al., Bull. Bot. Res., Harbin 12: 225. 1992. apex dentate or lobed. Carpels 5, rarely more, densely 矮牡丹 tomentose. Stigmas red. Follicles oblong, densely ai mu dan brown-yellow tomentose. Fl. Apr–May, fr. Aug. 2n = Paeonia spontanea (Rehder) T. Hong & W. Z. Zhao, 10*. nom. illeg. (included P. jishanensis); P. suffruticosa • Cliffs; ca. 300 m. Native in C Anhui and W Henan; widely Andrews var. jishanensis (T. Hong & W. Z. Zhao) cultivated in China and elsewhere. Halda; P. suffruticosa subsp. spontanea (Rehder) S. G. 1a. Flowers double; petals variously colored, Haw & Lauener; P. suffruticosa var. spontanea Rehder. white, pink, red, or red-purple 1a. subsp. suffruticosa Shrubs to 1.8 m tall. Roots attenuate toward tip. Turions 1b. Flowers single; petals white or pale present. Stems gray or gray-brown. Proximal leaves 2- red-purple ....................... 1b. subsp. yinpingmudan ternate; leaflets ovate-orbicular to orbicular, 4–6 × 3.5– 4.5 cm, villous along veins or throughout, deeply 3- 1a. Paeonia suffruticosa subsp. suffruticosa lobed; lobes again lobed, acute to rounded at apex. 牡丹(原亚种) mu dan (yuan ya zhong) Flowers solitary, terminal, single, 10–16 cm wide. Paeonia chinensis Oken (1841), not Vilmorin (1870), Bracts 2 or 3, long elliptic, unequal. Sepals 4 or 5, nor P. sinensis Steudel (1841); P. fruticosa Dumont de Courset; P. moutan Sims; P. suffruticosa var. purpurea green, broadly ovate, 2.5–5 × 1.8–2.5 cm, apex rounded. Andrews; P. yunnanensis W. P. Fang. Petals 5–11, white, occasionally pinkish at base or Flowers double; petals variously colored, white, pink, margin, obovate, 4.5–7.2 × 4–6 cm, apex irregularly red, or red-purple. Fl. Apr–May. incised. Filaments pink or purple, white distally, 8–10 mm; anthers yellow, linear, 8–11 mm. Disc wholly enveloping carpels, red-purple, leathery, apex dentate. wholly enveloping carpels, purple-red, leathery,
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