Papaveraceae (PDF)
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PAPAVERACEAE 罂粟科 ying su ke Zhang Mingli (张明理)1, Su Zhiyun (苏志云 Shu Tsi-yun)2; Magnus Lidén3, Christopher Grey-Wilson4 Herbs, annual, biennial, monocarpic perennial, perennial, or shrubby. Laticifers or elongated idioblasts present. Leaves alternate or in a basal rosette, rarely opposite or whorled, usually without stipules; leaf blade entire to compound. Inflorescences racemes, panicles, dichasia, pseudoumbels, or solitary flowers. Flowers actinomorphic, bisymmetric, or zygomorphic, always bisexual, usu- ally 2-merous, rarely 3- or 4-merous. Calyx caducous, green or petaloid. Corolla choripetalous or quasi-sympetalous, very rarely absent. Anthers opening by slits. Ovary superior, syncarpous with 2 to several carpels; placentation parietal. Forty genera and ca. 800 species: mainly in the N Hemisphere of both the Old and New Worlds, extending into Central and South America, a few in Africa; 19 genera (one endemic, two introduced) and 443 species (295 endemic, five introduced, one requiring verification) in China. Many species are used medicinally due to the high content of alkaloids. The two subfamilies recognized here are often treated as separate families. Wu Chengyi, Chuang Hsuan & Su Ziyun. 1999. Papaveraceae. In: Wu Chengyi, ed., Fl. Reipubl. Popularis Sin. 32: 1–483, 541–545. 1a. Laticifers present; sap watery or milky; sepals green, entire, enclosing corolla in bud, imbricate or (in Eschscholzia) fused into a caducous hood; corolla actinomorphic, with (0–)4–8 (rarely more) petals; petals all similar, flat, imbricate and crumpled in bud; stamens several with filiform filaments; nectaries absent .......................................................................................................................... 1. Subfam. Papaveroideae 1b. Laticifers lacking; sap watery, but sometimes colored; sepals usually petaloid, not enclosing petals in bud; corolla bisymmetric or zygomorphic; petals 4 in 2 very dissimilar whorls, with wings, crests, and/or spurs, pairwise valvate in bud; androecium of 8 thecae distributed on 2, 4, or 6 “stamens” with broad hyaline filaments; nectaries usually present ............................................................................. 2. Subfam. Fumarioideae 1. Subfam. PAPAVEROIDEAE 罂粟亚科 ying su ya ke Zhang Mingli (张明理); Christopher Grey-Wilson Herbs, annual, biennial, perennial, or shrubby, with milky or watery latex. Leaves alternate, rarely opposite or pseudowhorled. Inflorescences solitary flowers, or racemes, corymbs, panicles, or umbels. Flowers actinomorphic. Sepals in 1 whorl of 2 or 3(or 4), free, caducous. Petals 4–10, occasionally more, in 2 whorls, usually caducous, rarely absent. Stamens many, free. Ovule anatropous to subcampylotropous, bitegmic, crassinucellate. Fruit syncarpous, multicarpellate, many seeded. Twenty-three genera and ca. 230 species: mainly in the N Hemisphere, extending into Central and South America, a few in Africa; 12 genera (one endemic, two introduced) and 67 species (29 endemic, five introduced) in China. 1a. Perianth 3-merous; pistils at least 3-carpellate ................................................................................................................ 1. Argemone 1b. Perianth 2-merous or petals absent; pistils 2-carpellate, rarely more. 2a. Flowers small, apetalous, in large panicles ............................................................................................................. 12. Macleaya 2b. Flowers medium to large, with 4 petals or more. 3a. Inflorescences usually corymbs or panicles; seeds cristately carunculate. 4a. Stems scapose; leaves all basal, blade cordate, margin repand .................................................................. 11. Eomecon 4b. Stems not scapose; leaves cauline and basal, blade not cordate, margin with teeth to pinnatifid. 5a. Cauline leaves alternate; stems cymosely branched; fruit nearly moniliform ............................... 10. Chelidonium 5b. Leaves nearly opposite on stem apices; stems simple; fruit not moniliform. 6a. Flowers bracteate; ovary shortly pubescent; capsules 2–4-valvate from apex to base .............. 8. Stylophorum 6b. Flowers ebracteate; ovary glabrous; capsules 2-valvate from base to apex ................................ 9. Hylomecon 3b. Inflorescences simple or racemose; seeds without cristate caruncle. 7a. Pistils 3-carpellate to polycarpellate; inflorescence racemose or paniculate, flowers sometimes solitary. 8a. Styles generally obvious, stigmas claviform or capitate, free or united, actinomorphic, decurrent; plant yellow lactiferous ..................................................................................................... 2. Meconopsis 1 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 2 Kunming Institute of Botany, Chinese Academy of Sciences, 132 Lanhei Road, Kunming, Yunnan 650204, People’s Republic of China. 3 Botanic Garden, Uppsala University, Villavägen 8, SE-752 36, Uppsala, Sweden. 4 Red Lion Barn, East Church Street, Kenninghall, Norfolk NR16 2EP, United Kingdom. 261 262 PAPAVERACEAE 8b. Flowers astylous, stigmas actinomorphic, united into compressed or steepled disks; plant white lactiferous .................................................................................................................................................. 3. Papaver 7b. Pistils 2-carpellate (rarely 4-carpellate); flowers solitary, terminal. 9a. Petals purple or red; capsules (2–)4-valvate ........................................................................................... 7. Roemeria 9b. Petals yellow or orange, rarely red; capsules 2-valvate. 10a. Anthers linear, longer than filaments; cauline leaves and basal leaves similar, blade ternate, polypinnately lobed, lobes linear ............................................................................... 4. Eschscholzia 10b. Anthers not linear, shorter than filaments; basal leaves many, pinnatilobate or pinnatipartite, lobes with teeth and petiolules, cauline leaves fewer, obviously smaller than basal leaves, sessile. 11a. Ovary 1-loculed; seeds ovoid and tessellate; plant frequently yellow lactiferous ..... 5. Dicranostigma 11b. Ovary with 2 false locules caused by septas forming spongy cells; seeds ovoid, reniform, alveolate; plant orange- or red lactiferous .......................................................... 6. Glaucium 1. ARGEMONE Linnaeus, Sp. Pl. 1: 508. 1753. 蓟罂粟属 ji ying su shu Zhang Mingli (张明理); Christopher Grey-Wilson Herbs, annual, biennial, or perennial, usually stout, mostly branched, spiny, bitter yellow lactiferous. Stems mostly erect. Leaves pinnate, lobes repand dentate, teeth apically spiny. Flowers solitary, terminal or arranged in cyme, trimerous; receptacles narrowly conical. Flower buds erect. Sepals (2 or)3, caducous, apex with horned appendices. Petals (4–)6, in 2 whorls, contorted or imbricate in bud, orange, yellow, yellow-white, or white, occasionally pink or mauve. Stamens many, free; filaments filiform or slightly ampli- ate below middle, apically subulate; anthers linear, nearly basal, bifid, extrorse, curved after dehiscence. Ovary ovoid, conical-ovoid, or nearly elliptic; carpels (3 or)4–6, united; ovules many; styles very short or absent; stigmas as many as carpels, actinomorphic. Capsule spiny, rarely not spiny, apex divided for a short distance into 3–6 valves, rarely parting nearly to base. Seeds many, spheroidal; caruncle very small or absent, seed coat tessellate. Twenty-nine species: mainly in North America from United States to C Mexico and the West Indies, nine species in South America (three in Chile), one in Hawaii, and the others scattered along the NW and W coasts of the Americas; one species (introduced) in China. 1. Argemone mexicana Linnaeus, Sp. Pl. 1: 508. 1753. sparsely spiny. Petals 6, yellow or orange, broadly obovate, 1.7–3 cm, base broadly cuneate, apex rounded. Filaments ca. 7 蓟罂粟 ji ying su mm; anthers narrowly oblong, 1.5–2 mm, coiling after dehis- Herbs, annual or occasionally short-lived perennial, 30– cence. Ovary elliptic or oblong, 7–10 mm, with explanate ful- 100 cm tall, usually stout. Stems mostly short, branched, with vous spines; styles very short; stigmas dark red, 4–6-lobed. sparse explanate fulvous spines. Basal leaves dense; petiole 5– Capsule oblong to broadly elliptic, 2.5–5 × 1.5–3 cm, sparsely 10 mm; blade glaucous with blue-green markings on veins, fulvous spiny, 4–6-valved dehiscing from apex to 1/4–1/3 of paler abaxially, broadly oblanceolate or obovate to elliptic, 5– length. Seeds spherical, 1.5–2 mm in diam., obviously tessel- 20(–25) × 2.5–7.5(–8) cm, glabrous, sparsely sharply spiny on late. Fl. and fr. Mar–Oct. 2n = 28. veins, base cuneate, margin pinnatipartite, apex acute; lobes un- Naturalized along coasts, introduced in courtyards. Naturalized in dulate teeth, teeth apically spiny. Cauline leaves alternate, simi- Fujian, Guangdong, Taiwan, ?Yunnan; introduced in many provinces in lar to basal leaves, but upper leaves smaller and sessile, often China [native to Central and tropical America]. subamplexicaul. Flowers solitary, sometimes in few-flowered This is a widely cultivated plant which has become a naturalized cymes. Pedicel very short. Flower buds ovoid, ca. 1.5 cm. Se- weed in many parts of the world, particularly in subtropical regions. The pals 2(or 3), cymbiform, ca. 1 cm, apex spurred, glabrous or seeds are poisonous. 2. MECONOPSIS Viguier, Hist. Nat. Pavots Argémones, 48. 1814. 绿绒蒿属 lü rong hao shu Zhang Mingli