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8. Phyllanthus Linnaeus, Sp. Pl. 2: 981. 1753.

叶下珠属 ye xia zhu shu

Authors: Bingtao Li & Michael G. Gilbert

Cathetus Loureiro; Cicca Linnaeus; Diasperus Kuntze; Emblica Gaertner; Epistylium Swartz; Eriococcus Hasskarl; Hemicicca Baillon; Kirganelia Jussieu; Macraea Wight; Niruri Adanson; Nymphanthus Loureiro.

Trees, shrubs, or herbs, mostly monoecious, less often dioecious; branching often "phyllanthoid": main stems with spiral phyllotaxy, ultimate branchlets sometimes clustered on short shoots, resembling pinnate leaves and often deciduous as a unit, less often stems all similar with spiral or distichous phyllotaxy; hairs simple, often absent, rarely branched. Leaves alternate, often reduced and scalelike on main stems, strongly distichous on leafy stems; stipules small, deciduous or persistent; petiole short; leaf blade simple, margin entire, venation pinnate. Inflorescences axillary, sometimes at leafless nodes, solitary or in fascicles, cymes, glomerules, racemes, or panicles; pedicels delicate. Male flowers: sepals (2 or)3-6, in 1 or 2 series, free, imbricate, margin entire, eroded, denticulate or fimbriate; petals absent; disk glands 3-6, usually free; stamens 2-6; filaments free or connate; anthers 2-locular, extrorse, thecae 2, connectives obscure, longitudinally or horizontally dehiscent, rarely obliquely so; pistillode absent. Female flowers: sepals as in male or more; disk glands usually small, free or connate into an annulus or urn-shape, surrounding ovary; ovary smooth or less commonly roughened, bullate, or hairy, 3(-12)-locular; ovules 2 per locule; styles 3(-12), apex 2-lobed or 2-branched, rarely entire, erect, spreading, or recurved. Fruit usually a capsule, globose or depressed globose, smooth or warty, dehiscent into 3 2-valved cocci when mature, less often a fleshy berry or drupe; columella persistent. Seeds without caruncle or aril, trigonous, surface smooth, sculptured or striate; seed-coat dry crustaceous, endosperm whitish, cartilaginous; embryo straight or slightly curved; cotyledons usually considerably broader than radical. x = 13.

About 750-800 species: primarily in the tropics and subtropics, poorly represented in temperate regions; 32 species (13 endemic, one introduced) in China.

Molecular studies have shown that several long-established genera nest within Phyllanthus, including Breynia, Phyllanthodendron, Glochidion, and Sauropus.

Phyllanthus acidus (Linnaeus) Skeel is recorded from Taiwan and Hong Kong, where it is presumably cultivated for its edible fruits.


1 Plant dioecious, with either male or female flowers; leaf blade 1-10 cm   (2)
+ Plant monoecious, with both male and female flowers; leaf blade often less than 1 cm   (3)
       
2 (1) Leaf blade 1-2 × 0.6-1.3 cm; inflorescences mostly 1-flowered; male sepals 6; female pedicel 2-3 mm.   24 P. cochinchinensis
+ Leaf blade 2.5-10 × 1.5-4 cm; inflorescences many flowered; male sepals 4; female pedicel 10-15 mm.   9 P. gracilipes
       
3 (1) Fruits indehiscent, fleshy   (4)
+ Fruit a dehiscent capsule, not fleshy, brown or brownish when mature   (9)
       
4 (3) Leaf base oblique; fruit a drupe, pale green or yellowish white, 10-13 mm in diam.; stamens 3 or 4   (5)
+ Leaf base symmetrical; fruit a berry, purple-black when mature, 6-10 mm in diam.; stamens (3-)5(or 6)   (6)
       
5 (4) Leaves linear-oblong, 0.8-2.3 × 0.15-0.6 cm; stamens 3, fused.   1 P. emblica
+ Leaves ovate or oblong-ovate, 2.4-10 × 1.9-4.5 cm; stamens 4, free. ..... P. acidus (see note above)   (4)
       
6 (4) Fruits 3-celled   (7)
+ Fruits 4-12-celled   (8)
       
7 (6) Fruiting sepals persistent; male sepals 6; fruit ca. 10 mm in diam..   2 P. glaucus
+ Fruiting sepals deciduous; male sepals 5; fruit ca. 6 mm in diam..   3 P. flexuosus
       
8 (6) Stamens 4 or 5(-7), filaments all free; fruit ca. 8 mm in diam., 4-seeded.   4 P. oligospermus
+ Stamens 5, filaments 3 connate and 2 free; fruit ca. 6 mm in diam., 8-16-seeded.   5 P. reticulatus
       
9 (3) Male sepal margins fimbriate, serrulate, or erose   (10)
+ Male sepal margins entire   (18)
       
10 (9) Ovary with dense, rugose-undulate or curved pubescence or densely setose   (11)
+ Ovary smooth or tuberculate   (12)
       
11 (10) Ovary with dense, rugose-undulate or curved pubescence; leaf blade 3-7 mm wide, base oblique.   6 P. chekiangensis
+ Ovary and fruits with very dense, soft prickles; leaf blade 6-10 mm wide, base symmetrical.   10 P. forrestii
       
12 (10) Female sepals 5   (13)
+ Female sepals 6   (14)
       
13 (12) Stipules 3-5 mm; female pedicel ca. 10 mm; stamens 4.   8 P. fimbricalyx
+ Stipules 1-1.5 mm; female pedicel 20-35 mm; stamens 2.   11 P. hainanensis
       
14 (12) Stamens 4   (15)
+ Stamens 2   (16)
       
15 (14) Branchlets glabrous; female sepals 1.2-2(-3.8) mm, obscurely erose or subentire; fruiting pedicel 1-1.5(-2.5) cm.   7 P. sootepensis
+ Branchlets rufous puberulent; female sepals ca. 3.5 mm; fruiting pedicel ca. 5 cm.   9 P. gracilipes
       
16 (14) Young branchlets puberulent; leaf blade 18-30 × 8-13 mm; inflorescence a bisexual axillary fascicle, usually with several male and 1 female flowers.   12 P. pulcher
+ Branchlets glabrous; leaf blade 3-13 × 3-7 mm; inflorescence 1-flowered   (17)
       
17 (16) Branchlets terete, glabrous; pedicels 2-8 mm; styles connate at base.   13 P. franchetianus
+ Branchlets angular, minutely papillate along upper side; pedicels 10-15 mm; styles free.   14 P. leptoclados
       
18 (9) Stamens 3, filaments ± free   (19)
+ Stamens 2-4, filaments 2 free, others connate   (21)
       
19 (18) Shrubs; leaf blades thinly papery or membranous; male disk cup-shaped.   17 P. clarkei
+ Herbs, branching mainly near base; leaf blades leathery or thinly leathery; male disk of separate glands   (20)
       
20 (19) Leaves ± distichous; stems usually ascending; female pedicel ca. 5 mm, disk orbicular, undivided; capsule with scalelike protuberances.   15 P. virgatus
+ Leaves spiral; stems spreading; female pedicel 1.5-2 mm, disk of separate glands; capsule smooth.   16 P. maderaspatensis
       
21 (18) Leaf blade margin hispidulous; ovary and fruits tuberculate.   18 P. urinaria
+ Leaf blade not hispidulous; ovary and fruits smooth   (22)
       
22 (21) Male sepals 5 or 6; stamens 3   (23)
+ Male sepals 4; stamens 2   (29)
       
23 (22) Herbs   (24)
+ Shrubs   (25)
       
24 (23) Perennial herbs; leaf blade elliptic or obovate, lateral veins 3 pairs; male sepals and disk glands 6.   19 P. arenarius
+ Annual herbs; leaf blade oblong-elliptic, lateral veins 4-7 pairs; male sepals and disk glands 5.   20 P. amarus
       
25 (23) Branchlets terete; male sepals and disk glands 5.   21 P. myrtifolius
+ Branchlets angular; male sepals and disk glands 6   (26)
       
26 (25) Branchlets usually clustered at apex of erect main stem; leaf blade base oblique; filaments connate at base only.   22 P. rheophyticus
+ Branchlets more evenly distributed along main stems; leaf blade base symmetrical; filaments connate into a column   (27)
       
27 (26) Leaf blade 3-14 × 1.5-3.5 cm; inflorescences on specialized racemelike leafless shoots, 2-10 cm; female disk cup-shaped, apex ± lacerate.   23 P. pachyphyllus
+ Leaf blade 0.2-2 × 0.1-1.3 mm; inflorescence 1-flowered or a 4-6-flowered fascicle, subtended by normal leaves; female disk urceolate or disk-shaped, apex entire   (28)
       
28 (27) Leaf blade obovate or spatulate, apex obtuse or rounded, lateral veins obscure; stipule margins ciliate; bract margins lacerate; flowers usually solitary.   24 P. cochinchinensis
+ Leaf blade oblong or lanceolate, apex apiculate, lateral veins prominent; stipule margins glabrous; bract margins entire; flowers 4-6-fascicled.   25 P. tsarongensis
       
29 (22) Branchlets, petioles, ovary, and fruits with stellate tufts of brown hairs; ovary 4-6-celled.   26 P. tsiangii
+ Branchlets, petioles, ovary, and fruits glabrous; ovary 3-celled   (30)
       
30 (29) Annual herbs; stamen filaments free.   27 P. ussuriensis
+ Shrubs; stamen filaments connate   (31)
       
31 (30) Leaf blade linear-oblong; flowers in glomerules.   28 P. taxodiifolius
+ Leaf blade not linear-oblong; flowers solitary or fascicled   (32)
       
32 (31) Leaf blade 4-5 × ca. 2 mm, base oblique; flowers solitary.   29 P. nanellus
+ Leaf blade 20-45 × 7-17 mm, base symmetrical; flowers fascicled   (33)
       
33 (32) Branchlets terete; leaf blade elliptic-ovate, lateral veins 6-9 pairs; male pedicels 5-7 mm.   30 P. guangdongensis
+ Branchlets angular-winged; leaf blade ovate-lanceolate or lanceolate, lateral veins 3-5 pairs; male pedicels 1-8 cm   (34)
       
34 (33) Sepals purple; male pedicels 0.3-0.8 cm; female pedicels 0.6-1 cm.   31 P. bodinieri
+ Sepals whitish; male pedicels 1-1.5 cm; female pedicels 2-2.3 cm.   32 P. fangchengensis

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