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2. Sapindaceae

无患子科 wu huan zi ke

Authors: Nianhe Xia & Paul A. Gadek

Trees or shrubs (or woody vines with tendrils in Cardiospermum and allied genera), rarely herbaceous climbers. Indumentum usually of simple hairs, often glandular on young parts, buds, and inflorescences. Leaves alternate, usually estipulate; leaf blade pinnate or digitate, rarely simple; leaflets alternate to opposite, entire or dentate to serrate. Inflorescence a terminal or axillary thyrse; bracts and bracteoles small. Flowers unisexual, rarely polygamous or bisexual, actinomorphic or zygomorphic, usually small. Sepals 4 or 5(or 6), equal or unequal, free or connate at base, imbricate or valvate. Petals 4 or 5(or 6), sometimes absent, free, imbricate, usually clawed, often with scales or hair-tufted basal appendages. Disk conspicuous, fleshy, complete or interrupted, lobed or annular, rarely absent. Stamens 5-10(-74), usually 8, rarely numerous, variously inserted but usually within disk, often exserted in male flowers; filaments free, rarely connate; anthers dorsifixed, longitudinally dehiscent, introrse; staminodes sometimes present in carpellate flowers, but filaments shorter and anthers with a thick wall, indehiscent. Ovary superior, (1-)3(or 4)-loculed; ovules 1 or 2(or several) per locule, placentation axile, rarely parietal, anatropous, campylotropous, or amphitropous; style usually apical (terminal), semigynobasic in Allophylus [gynobasic in Deinbollia Schumacher & Thonning]; stigma entire or 2 or 3(or 4)-lobed, usually rudimentary in male flowers. Fruit a loculicidal capsule, berry, or drupe, or consisting of 2 or 3 samaras, often 1-seeded and 1-loculed by abortion. Seeds 1(or 2 or more) per locule; testa black or brown, hard, often with a conspicuous fleshy aril or sarcotesta; embryo curved, plicate, or twisted, oily and starchy; endosperm usually absent. 2n = 20-36.

One hundred thirty-five genera and ca. 1500 species: widely distributed in tropical and subtropical regions, especially well represented in tropical SE Asia; 21 genera (one endemic) and 52 species (16 endemic, one introduced) in China.

There is some variation in the circumscription of Sapindaceae in taxonomic treatments, particularly with regard to the inclusion of genera from the closely related, predominately temperate families Aceraceae and Hippocastanaceae. Several studies including Müller and Leenhouts (in Ferguson & Müller, Evolutionary Significance Exine: 407-445. 1976), and more recently those based on molecular data (Stevens, Angiosperm Phylogeny Website, 2001 onward; Harrington et al., Syst. Bot. 30: 366-382. 2005), supported the recognition of a broadly defined Sapindaceae incorporating Aceraceae and Hippocastanaceae. Harrington et al. (loc. cit.) proposed four subfamilies or clades, comprising Sapindoideae (including Koelreuteria and Ungnadia Endlicher), Dodonaeoideae, Hippocastanoideae (including taxa previously referred to Aceraceae and Hippocastanaceae, plus Handeliodendron), and a monotypic "Xanthoceratoideae". Within Hippocastanoideae, Acer Linnaeus and Dipteronia Oliver comprise a monophyletic group and are treated in this Flora as Aceraceae. Similarly, Aesculus Linnaeus, Billia Peyritsch, and the Chinese endemic Handeliodendron Rehder form a monophyletic group and are treated here as Hippocastanaceae. There is some support for "Xanthoceratoideae" being the first lineage to diverge within the broadly defined Sapindaceae assemblage; consequently, Xanthoceras is treated separately from genera in Sapindoideae and Dodonaeoideae in the following account of Sapindaceae s.s. The sequence of genera reflects Müller and Leenhouts (loc. cit.) as modified by recent analyses based on molecular and morphological data, rather than following the order developed by Radlkofer (Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 20: 105-379. 1890; and in Engler, Pflanzenreich 98a-h(IV. 165): 1-1539. 1931-1934), which was previously followed in FRPS.

The main economic uses of this family include (1) timber: Amesiodendron chinense, Dimocarpus longan, D. confinis, Litchi chinensis, Pavieasia kwangsiensis, and Pometia pinnata; (2) fruit: Dimocarpus longan, Litchi chinensis, and Nephelium lappaceum; (3) medicine: Dimocarpus longan (arillode), Litchi chinensis (seeds), and Sapindus saponaria (roots); (4) oil: Amesiodendron chinense, Delavaya toxocarpa, and Xanthoceras sorbifolium. Saponins occur widely in the family, commonly used as a fish poison and for their detergent properties.

Lo Hsien-shui & Chen Te-chao. 1985. Sapindaceae (excluding Handeliodendron). In: Law Yuh-wu & Lo Hsien-shui, eds., Fl. Reipubl. Popularis Sin. 47(1): 1-72.


1 Herbaceous climbers; inflorescences with basal tendrils.   21 Cardiospermum
+ Trees or shrubs; inflorescences without tendrils   (2)
       
2 (1) Fruit indehiscent, drupaceous or berrylike   (3)
+ Fruit capsular, loculicidal   (12)
       
3 (2) Leaves pinnate, sessile, first pair leaflets inserted at base of leaf axis like a pair of stipules; fruit without spines and tubercles   (4)
+ Leaves petiolate   (5)
       
4 (3) Leaves imparipinnate, leaflets entire; fruit not lobed into schizocarps, 1-1.2 cm.   10 Lepisanthes
+ Leaves paripinnate, leaflets serrate; fruit lobed into 2 schizocarps, usually only 1 developed, longer than 2 cm.   13 Pometia
       
5 (3) Exocarp fleshy; arillode absent; petals with scales   (6)
+ Exocarp leathery or crustaceous   (8)
       
6 (5) Leaves digitate, leaflets 1-5; fruit shorter than 1 cm; sepals and petals 4; disk 4-sectioned.   20 Allophylus
+ Leaves pinnate; fruit longer than 1 cm; sepals 5; disk complete or lobed   (7)
       
7 (6) Testa osseous, hilum linear; petals 4 or 5, rarely 6, with 2 earlike scales or 1 large scale; deciduous trees.   9 Sapindus
+ Testa membranous or crustaceous, hilum orbicular; evergreen trees or shrubs   10 Lepisanthes
       
8 (5) Seeds without arillode; fruit not lobed into schizocarps, densely tomentose.   10 Lepisanthes
+ Seeds with arillode; fruit parted into schizocarps, usually only 1 or 2 developed, abaxially usually with various tubercles or spines, glabrous or sparsely hairy   (9)
       
9 (8) Arillode distinct from testa; fruit without spines, usually with tubercles or nearly smooth   (10)
+ Arillode adnate to testa   (11)
       
10 (9) Sepals imbricate; leaflets abaxially with naked glands in lateral vein axils, if without naked glands then inflorescences with stellate hairs.   11 Dimocarpus
+ Sepals valvate; leaflets abaxially without naked glands; inflorescences tomentose.   12 Litchi
       
11 (9) Petals and sepals 4.   14 Xerospermum
+ Petals absent; calyx 5- or 6-lobed; fruit with soft spines.   15 Nephelium
       
12 (2) Leaves simple; petals absent; branches, leaves, and inflorescences with sticky juice; fruit winged.   2 Dodonaea
+ Leaves compound; branches, leaves, and inflorescences without sticky juice   (13)
       
13 (12) Leaves digitate, leaflets 3; exocarp leathery or nearly woody; seeds without arillode; petals 5, with scales.   5 Delavaya
+ Leaves pinnate   (14)
       
14 (13) Fruit inflated, exocarp membranous or papery, veined   (15)
+ Fruit not inflated, exocarp leathery or woody   (16)
       
15 (14) Leaves imparipinnate; sepals valvate; filaments villous; fruit not winged.   6 Koelreuteria
+ Leaves paripinnate; sepals imbricate; filaments glabrous; fruit 3-winged.   7 Boniodendron
       
16 (14) Seeds with arillode; leaves paripinnate   (17)
+ Seeds without arillode   (19)
       
17 (16) Fruit parted into schizocarps; leaflets abaxially with orbicular naked glands at lateral vein axils.   16 Arytera
+ Fruit not parted into schizocarps   (18)
       
18 (17) Fruit pearlike or clavate; stamens 8; ovary 3-loculed; leaflets with naked glands in lateral vein axils.   17 Mischocarpus
+ Fruit transversely ellipsoid or subglobose; stamens 5; ovary 2-loculed; leaflets without naked glands in lateral vein axils.   3 Harpullia
       
19 (16) Leaves imparipinnate   (20)
+ Leaves paripinnate   (21)
       
20 (19) Fruit with dense, short spines, 1-loculed, 1-seeded; petals ca. 1 mm, white, scaly; disk lobes without appendages; evergreen trees.   8 Paranephelium
+ Fruit without spines, 3-loculed, with several seeds per locule; petals ca. 1.7 cm, base red or yellow, without scales; disk lobes with appendages; deciduous shrubs or small trees.   1 Xanthoceras
       
21 (19) Fruit 7-8 mm, exocarp leathery, densely tomentose, testa black, hilum small; petals without scales.   4 Eurycorymbus
+ Fruit longer than 1 cm, exocarp woody, glabrous, testa brown, hilum large; petals with scales   (22)
       
22 (21) Fruit not parted into schizocarps; leaf axis trigonous.   18 Pavieasia
+ Fruit parted into schizocarps; leaf axis terete.   19 Amesiodendron

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