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Euphorbiaceae EUPHORBIACEAE 大戟科 da ji ke Li Bingtao (李秉滔 Li Ping-tao)1, Qiu Huaxing (丘华兴 Chiu Hua-hsing, Kiu Hua-shing, Kiu Hua-xing)2, Ma Jinshuang (马金双)3, Zhu Hua (朱华)4; Michael G. Gilbert5, Hans-Joachim Esser6, Stefan Dressler7, Petra Hoffmann8, Lynn J. Gillespie9, Maria Vorontsova10, Gordon D. McPherson11 Trees, shrubs, or herbs, rarely woody or herbaceous lianas, monoecious or dioecious, indumentum of simple, branched, stellate, or gland-tipped hairs, peltate or glandular scales or stinging hairs, latex often present, clear, white, or colored; roots woody, rarely roots tuberous and stems succulent, sometimes spiny. Leaves alternate or opposite, rarely whorled; stipules usually present, often free, sometimes modified into spines or glands, deciduous or persistent; petioles long to short, sometimes with glands at apex or base; leaf blade simple, sometimes palmately lobed, rarely compound, or reduced to scales, margins entire or toothed, sometimes with distinct glands along margin and/or on surface, venation pinnate or palmate. Inflorescences axillary or terminal, flowers in cymes or fascicles, these often arranged along an elongated axis, branched or unbranched, forming a thyrse, in congested heads, or in a flowerlike cyathium with very reduced flowers enclosed within a ± cupular involucre; bracts sometimes petaloid. Flowers unisexual, within same inflorescence or in separate inflorescences, actinomorphic. Sepals (1–)3–6(–8), free or connate into calyx tube, valvate or imbricate, rarely absent (Euphorbia). Petals free, often reduced or absent. Disk present or absent. Male flowers with disk intrastaminal or extrastaminal, entire to dissected. Stamens one to very many, hypogynous; filaments free or connate; anthers 2(–4)-locular, mostly dehiscing longitudinally, rarely transversely or by pores, introrse or extrorse; rudimentary ovary sometimes present. Female flowers rarely with staminodes; ovary superior, (1–)2–5(–20)-locular; placentation axile; ovules 1 or 2 per locule, anatropous or hemitropous; styles free or connate, entire or lobed, or multifid, lobes erect, horizontal or curved; stigma capitate, linear, fimbriate, fan-shaped or pinnatilobate. Fruit typically a capsule elastically dehiscent into 2-valved cocci from a persistent columella, sometimes a berry or drupe. Seeds 1 or 2 per locule; seed coat thin to indurate, sometimes fleshy to form a sarcotesta; caruncle sometimes present; aril sometimes present; endosperm present or absent; embryo straight to curved or folded; cotyledons usually broader than radical. x = 6–14. About 322 genera and 8910 species: widespread throughout the world, primarily in the tropics and subtropics, more poorly represented in tem- perate regions; 75 genera (one endemic, nine introduced) and 406 species (99 endemic, 27 introduced) in China, nearly 95% of which are found in the S and SW parts of the country. The Euphorbiaceae as treated here include the following families that have been proposed for segregation: Androstachydaceae, Antidesmataceae, Bischofiaceae, Hymenocardiaceae, Phyllanthaceae, Pedilanthaceae, Picrodendraceae, Porantheraceae, Putranjivaceae, Ricinocarpaceae, Scepaceae, Stilaginaceae, Trewiaceae, and Uapacaceae. The Pandaceae and Buxaceae, formerly included here, are now well established as separate families. Molecular data has shown that the traditional concept of Euphorbiaceae includes three major lineages that are relatively distantly related to each other: the Phyllanthoids (genera 1–16 in this account), the Putranjivoids (genera 17 and 18), and the Euphorbioids (genera 19–75). Many species of Euphorbiaceae are of economic importance, probably most importantly as the main source of rubber (Hevea) but also as sources of medicine; foods, both as a staple starch source (Manihot) and fruits (e.g., Phyllanthus emblica); seed oils (Ricinus, Vernicia); and insecticides. Li Pingt’ao. 1994. Euphorbiaceae. In: Li Pingt’ao, ed., Fl. Reipubl. Popularis Sin. 44(1): ii–viii, 1–217; Kiu Huashing, Hwang Shumei & Chang Yongtian. 1996. Euphorbiaceae (2). In: Kiu Huashing, ed., Fl. Reipubl. Popularis Sin. 44(2): ii–ix, 1–212; Ma Jinshuang & Tseng Yungchien. 1997. Euphorbiaceae (3). In: Ma Jinshuang, ed., Fl. Reipubl. Popularis Sin. 44(3): ii–vi, 1–150. 1a. Ovules 1 per locule; indumentum of simple, stellate, scalelike, glandular, or stinging hairs; latex present or absent; leaves mostly spiral, less often distichous, blade sometimes lobed (palmately compound in Hevea), margin entire or variously toothed, often with glands/extra-floral nectaries on petiole apex/leaf base and/or margins; inflorescence axillary or terminal, often thyrsoid or cymose, sometimes pseudanthial, rarely sessile clusters; seeds sometimes carunculate ......................................................... Key 3. Euphorbioid genera 1b. Ovules 2 per locule; indumentum of simple, very rarely branched, hairs; latex absent; leaves often strongly distichous, blade never lobed, margin entire or finely serrulate (3-foliolate and toothed in Bischofia), usually without glands (present in Aporosa); inflorescence mostly axillary, often without visible axis; seeds not carunculate, sometimes arillate. 1 Department of Forestry, South China Agricultural University, Guangzhou, Guangdong 510642, People’s Republic of China. 2 South China Botanical Garden, Chinese Academy of Sciences, Longdong, Shahe, Guangzhou, Guangdong 510650, People’s Republic of China. 3 Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 11225-1099, U.S.A. 4 Laboratory of Plant Phylogenetics and Conservation Biology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, 88 Xuefu Road, Kunming, Yunnan 650223, People’s Republic of China. 5 Missouri Botanical Garden c/o Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom. 6 Botanische Staatssammlung, Menzinger Straße 67, D-80638 München, Germany. 7 Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany. 8 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom. 9 Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada. 10 Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. 11 Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A. 163 164 EUPHORBIACEAE 2a. Stigmas usually bifid, not dilated; fruit a capsule or berry or if a drupe then relatively small and/or with 2 or more seeds and without persistent flaplike stigmas; base of leaf blade symmetrical ....... Key 1. Phyllanthoid genera 2b. Stigmas dilated, peltate or reniform; fruit a relatively large 1-seeded drupe, usually crowned by persistent flaplike stigmas; base of leaf blade often asymmetrical .................................................... Key 2. Putranjivoid genera Key 1. Phyllanthoid genera Trees, shrubs, or herbs, usually without latex (present in Bischofia); indumentum of simple hairs (branched in Phyllanthus reticulatus), often absent. Leaves alternate, often distichous, sometimes scalelike on main stems; petiole usually short, usually without glands (present in Aporosa); leaf blade simple, margin entire or minutely serrulate (long petioles, 3(–5)-foliolate with toothed margins in Bischofia); venation pinnate, rarely obscurely 3-veined from base. Inflorescences mostly axillary, without visible axis (present in Antidesma, Aporosa, Baccaurea, Bischofia, Richeriella). Male flowers with 2–8 stamens, anthers longitudinally dehiscent (variable in Phyllanthus); female flowers with 2 ovules per locule. Seeds without caruncle, sometimes with fleshy aril or fleshy testa. Fifty-nine genera and over 1700 species: mostly tropical, the greatest diversity in SE Asia; 16 genera and 138 species (41 endemic, four intro- duced) in China. 1a. Leaves 3-foliolate, margin distinctly serrate; plant with red or reddish juice; styles entire ........................................... 16. Bischofia 1b. Leaves simple, margin entire or minutely serrulate; plant without red juice; styles usually bifid, sometimes connate (Glochidion). 2a. Inflorescences in spikes, racemes, or pseudoracemes, with visible inflorescence axis; plants dioecious; petals absent. 3a. Disk present; ovary and fruit 1- or 3-celled; petioles and leaf margins without glands; stamens longer than sepals. 4a. Anthers U-shaped with an enlarged connective; fruit indehiscent; ovary and fruit 1-celled, often laterally flattened; male disk annular, in distinct parts, or cushion-shaped, enclosing stamens and pistillode, never lobed .................................................................................................................................................................... 13. Antidesma 4b. Anthers with parallel thecae and without an enlarged connective; fruit dehiscent, never flattened; ovary and fruit 3-celled; male disk 5-lobed .......................................................................................................................... 6. Richeriella 3b. Disk absent; ovary and fruit 2–4-celled; petioles and leaf margins sometimes glandular; stamens shorter to longer than sepals. 5a. Leaves evenly distributed along branches, usually distichously arranged; leaf margin undulate to denticulate; petiole apex usually glandular ................................................................................................................................. 14. Aporosa 5b. Leaves clustered toward branch tips, usually spirally arranged; leaf margin entire; petiole
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