
Flora of China 24: 88–95. 2000. 9. PARIS Linnaeus, Sp. Pl. 1: 367. 1753. 重楼属 chong lou shu Liang Songyun (梁松筠 Liang Song-jun); Victor G. Soukup1 Daiswa Rafinesque; Euthyra Salisbury; Kinugasa Tatewaki & Suto. Herbs perennial. Rhizome slender or thickened. Stem erect, simple. Leaves 4 to many, very rarely 3, in a terminal whorl, petio- late, lanceolate to ovate, with 3 main veins and anastomosing veinlets. Flowers bisexual, solitary, terminal, pedunculate. Tepals 3–8, in 2 whorls, free; outer ones green, rarely ± white, ovate to lanceolate; inner ones linear or occasionally absent. Stamens 8–24 or more, 2–8 × as many as tepals; filaments narrow, flat; anthers basifixed, often with convex connective apically. Ovary subglobose, 1- loculed with parietal placentation or 4–10-loculed with axile placentation. Style short; stigma lobes 4–10. Fruit a berry or a berrylike capsule, indehiscent or loculicidal, several to many seeded. About 24 species: Bhutan, China, India, Japan, Korea, Laos, Mongolia, Myanmar, Nepal, Russia, Sikkim, Thailand, Vietnam; Europe; 22 species (12 endemic) in China. The rhizomes of many species are used medicinally in China. 1a. Ovary 4–10-loculed, with axile placentation; berry indehiscent; seeds sometimes with a spongy (never succulent) aril on 1 side. 2a. Rhizome creeping, elongate, slender, 2–5 mm thick; seeds without aril, funicle not swollen. 3a. Leaves usually 6–8; outer tepals oblong-lanceolate to ovate-lanceolate, 10–25(–30) mm wide ............... 20. P. verticillata 3b. Leaves usually 4; outer tepals narrowly lanceolate, 3–9 mm wide. 4a. Leaves oblong-lanceolate or ovate-elliptic; outer tepals reflexed, 3–5 mm wide .............................. 21. P. bashanensis 4b. Leaves broadly obovate or ovate; outer tepals spreading horizontally, 5–9 mm wide ....................... 22. P. quadrifolia 2b. Rhizome 8–45 mm thick; seeds ± enveloped by whitish green, spongy aril formed by swollen funicle. 5a. Leaves 9–10 ................................................................................................................................................ 19. P. dulongensis 5b. Leaves 4–6. 6a. Leaves wrinkled, obovate, 10–15 × 6.3–10 cm, basally rounded .............................................................. 18. P. rugosa 6b. Leaves not wrinkled. 7a. Leaves elliptic or oblanceolate, base subcuneate ................................................................................ 17. P. vaniotii 7b. Leaves oblong, ovate- or obovate-oblong, or oblong-lanceolate, base shallowly cordate or rounded. 8a. Seeds white or yellow-red, slightly enveloped by swollen funicle .............................................. 16. P. forrestii 8b. Seeds yellow-brown, 1/2 enveloped by greenish white or rarely orange, spongy aril formed by enlarged funicle ............................................................................................................................... 15. P. axialis 1b. Ovary 1-loculed with parietal placentation; berrylike capsule dehiscing irregularly; seeds with a succulent aril. 9a. Stamens 4–6 × as many as outer tepals; seeds enveloped by orange aril .............................................................. 1. P. dunniana 9b. Stamens 2 or 3 × as many as outer tepals; seeds enveloped by red aril. 10a. Leaves with white, yellowish, or purple markings. 11a. Leaves with purple markings; stamens ca. 3 × as many as leaves ................................................... 12. P. polyandra 11b. Leaves with white or yellowish markings; stamens 2 × as many as leaves. 12a. Outer tepals as many as leaves, shorter than inner ones ...................................................... 10. P. luquanensis 12b. Outer tepals fewer than leaves, longer than inner ones ......................................................... 11. P. marmorata 10b. Leaves without markings. 13a. Plants pubescent or papillose-pubescent. 14a. Free portion of connective 0.5–1 mm ............................................................................................. 8. P. mairei 14b. Free portion of connective 13–15 mm ................................................................................. 9. P. wenxianensis 13b. Plants glabrous or nearly so. 15a. Free portion of connective 6–15 mm. 16a. Inner tepals pendulous, margin undulate ........................................................................... 7. P. undulata 16b. Inner tepals (if present) not pendulous, margin not undulate .......................................... 14. P. thibetica 15b. Free portion of connective 1–4(–5) mm (except in P. polyphylla var. pseudothibetica). 17a. Filaments ca. 1 mm, shorter than free portion of connective ............................................ 5. P. daliensis 17b. Filaments more than 2 mm, longer than free portion of connective. 18a. Free portion of connective transversely ellipsoid, subglobose, or shortly conical, as wide as or wider than long .................................................................................... 13. P. fargesii 18b. Free portion of connective inconspicuous or longer than wide. 19a. Leaves (18–)23–26 × (11–)14–16 cm ....................................................... 3. P. vietnamensis 1 Herbarium, Department of Biological Sciences, University of Cincinnati, Cincinnati, Ohio 45221-0006, U.S.A. Flora of China 24: 88–95. 2000. 19b. Leaves 5–17 × 2–11 cm. 20a. Leaves ovate ........................................................................................ 2. P. cronquistii 20b. Leaves oblong to lanceolate. 21a. Inner tepals dark purple, usually much shorter than outer ones .... 4. P. delavayi 21b. Inner tepals yellow-green, usually slightly shorter or longer than outer ones ................................................................................... 6. P. polyphylla 1. Paris dunniana H. Léveillé, Repert. Spec. Nov. Regni Veg. • Ravine forests, mossy forests; 900–2100 m. SW Guangxi, Gui- 9: 78. 1910. zhou, Sichuan, SE Yunnan. 海南重楼 hai nan chong lou 2b. Paris cronquistii var. xichouensis H. Li, Bull. Bot. Res., Harbin 6(1): 113. 1986. Daiswa dunniana (H. Léveillé) Takhtajan; D. hainanensis (Merrill) Takhtajan; Paris hainanensis Merrill. 西畴重楼 xi chou chong lou Plants 1.5–3 m tall. Rhizome 2–5 cm thick. Leaves 4–8; Inner tepals ca. 3 cm × 5 mm. petiole 5–10 cm; leaf blade obovate-oblong 15–30 × 7.5–16 • cm. Peduncle 60–140 cm. Outer tepals (5 or)6–8, green, ob- Evergreen forests on limestone slopes; 1400–1500 m. SE Yun- nan (Xichou Xian). long-lanceolate, 5–10 × 1.5–2.4 cm; inner ones green, filiform- linear, longer than outer ones. Stamens 20 or more; filaments 3. Paris vietnamensis (Takhtajan) H. Li, Acta Bot. Yunnan. 6: (0.4–)0.8–1.5 cm; anthers 1.2–2.1 cm; free portion of connec- 357. 1984. tive 0.5–4 mm, apex acute. Ovary 1-loculed; stigma lobes (5 or) 南重楼 6–8, ca. 5 mm. Capsule berrylike, subglobose, ca. 4 cm in nan chong lou diam., dehiscent, pale green at maturity. Seeds white, wholly Daiswa hainanensis (Merrill) Takhtajan subsp. vietnamen- enveloped by orange, succulent, aril, ca. 4 mm wide. Fl. Mar– sis Takhtajan, Brittonia 35: 259. 1983. Apr, fr. Oct–Nov. 2n = 10*. Plants 1–1.5 m tall. Rhizome 1.5–3.5 cm thick. Leaves 4– • Forests; near sea level to 1100 m. Guizhou, Hainan, Yunnan. 6; petiole 4–10 cm; leaf blade obovate to obovate-oblong, (18–) Many plants of this species have been cultivated and closely 23–26 × (11–)14–16 cm, base shallowly cordate or subrounded. investigated by Li (Bull. Bot. Res., Harbin 6(1): 109–111. 1986). The Peduncle (5–)30–90 cm. Outer tepals 4–7, green, lanceolate to stamens are 20 (e.g., in Cavalerie 3652, the type gathering of Paris dunniana) or more, rather than 10–12, as described by Takhtajan oblong-lanceolate, usually 6–8 × 2–3 cm; inner ones narrowly (Brittonia 35: 257–258. 1983). linear, 5.5–11 × ca. 1 mm, distally slightly widened. Stamens 2(or 3) × as many as outer tepals; filaments 2–5 mm; anthers 2. Paris cronquistii (Takhtajan) H. Li, Acta Bot. Yunnan. 6: 0.6–1.6 cm; free portion of connective 1–5 mm, apex acute. 357. 1984. Ovary violet, conical-ovoid, 1-loculed. Style short; stigma lobes 6. Fl. Apr–Jul. 2n = 10*. 凌云重楼 ling yun chong lou Evergreen broad-leaved forests; 1200–1900 m. Guangxi, Yunnan Plants to 1 m tall. Rhizome 2–10 × 2–3 cm. Stem scabrous. [Vietnam]. Leaves 4–6; petiole 2.5–7.5 cm; leaf blade ovate, 11–17 × 6–11 4. Paris delavayi Franchet, J. Bot. (Morot) 12: 190. 1898. cm, adaxially sometimes with purple markings, abaxially purple or mottled with purple. Peduncle 12–60 cm. Outer tepals 5 or 6, 金线重楼 jin xian chong lou green, ovate-lanceolate or lanceolate, 3.5–9 × 1.3–2 cm; inner Daiswa delavayi (Franchet) Takhtajan; Paris henryi Diels. ones yellow-green, narrowly linear, 3–8 cm × 1–5 mm. Stamens usually 15–18; filaments 3–8 mm; anthers 0.7–1.5 cm; free Plants 0.5–1.5 m tall. Rhizome 1.5–5 × 0.8–1.5 cm. Stem portion of connective 1–2 mm, apex acute. Ovary green or pale 30–70 cm. Leaves 6–8; petiole 0.8–1.5 cm; leaf blade lanceo- purple, globose, 1-loculed, 5- or 6-ribbed. Style short; stigma late or oblong-lanceolate, 6–15 × 2–3 cm. Peduncle 16–75 cm. lobes 5 or 6. Capsule red at maturity, dehiscent. Seeds sub- Outer tepals (3 or)4 or 5, usually recurved, purple-green or globose, wholly enveloped by red aril. Fl. May–Jun, fr. Oct– green, narrowly lanceolate, 2–4 cm × 4–9 mm; inner ones usu- Nov. ally dark purple, filiform-linear, shorter than outer ones. Sta- mens usually 8 or 10; filaments 2–4 mm; anthers 0.6–1.8 cm; • Evergreen forests on limestone slopes, ravine forests, mossy free portion of connective
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