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FOC | Family List | FOC Vol. 7 | Capparaceae

3. Capparis Linnaeus, Sp. Pl. 1: 503. 1753.

山柑属 shan gan shu

Shrubs, small trees, or vines, evergreen, erect, climbing, or sometimes prostrate. New branches often with branched or simple trichomes, glabrescent or sometimes with persistent trichomes; branchlet base sometimes with subulate scales (cataphylls). Stipular spines straight or curved, sometimes absent. Leaves alternate, spirally arranged or distichous [or secund], simple, petiolate or sometimes subsessile; leaf blade herbaceous to leathery, margin entire. Inflorescences superaxillary, axillary, or terminal, racemes, corymbs, umbels, or panicles, sometimes reduced to a single flower but usually 2-10-flowered; bract usually present at base of pedicels but often caducous. Sepals 4, in 2 whorls; sepals of outer whorl often thick, often inwardly concave or becoming navicular, covering other flower parts, sometimes basal one becoming saccate; sepals of inner whorl often thin, ± equal. Petals 4, imbricate, often 2 pairs roughly similar; anterior pair of petals distinct, clawless or sometimes clawed; posterior pair of petals with lateral margins adhering, base asymmetric, margin revolute or rarely inner lateral margin involute from base to near middle, base fully covering disk; petals sometimes all 4 ± equal, distinct. Stamens 7-120. Gynophore ± as long as filaments, scarcely lengthening in fruit but often becoming thicker, sclerified; ovary 1-loculed; placentae 2-6(-8); ovules few to many; style short or obsolete; stigma sessile. Fruit baccate, globose or ellipsoid, often with different color when mature or dry, usually not dehiscent. Seeds 1 to numerous per fruit, reniform to nearly polygonal; embryo bent.

Between 250 and 400 species: mostly in tropical and subtropical regions worldwide but some in temperate regions; 37 species (ten endemic) in China.

1 Inflorescences composed of superaxillary rows of (1-)2-10 flowers   (2)
+ Inflorescences composed of racemes, terminal corymbs, spikes, umbels, axillary fascicles, or sometimes solitary flowers in axils of leaves   (16)
2 (1) Flower-bearing portion of twig leafless, situated between two flushes of leaves, often with 50-100 flowers in groups of 4-10.   3 C. multiflora
+ Flower-bearing portion of twig leafy, each flower series subtended by a normally developed leaf   (3)
3 (2) Sepals 5 mm or shorter   (4)
+ Sepals 6 mm or longer   (7)
4 (3) Leaves subsessile with a petiole of 1-2 mm; leaf blade base cordate.   14 C. subsessilis
+ Leaves with a petiole longer than 3 mm; leaf blade base not cordate   (5)
5 (4) Spines straight (endemic to Taiwan).   6 C. henryi
+ Spines recurved or absent (mainland of S and SW China)   (6)
6 (5) Leaf blade 2-4 cm wide, apex acute to subacuminate, tip 0.3-0.7(-1) cm; twigs with strongly recurved spines.   15 C. tenera
+ Leaf blade 1-2(-3) cm wide, apex caudate, tip (1-)1.5-2.5 cm; twigs often without spines or spines small, ascending.   16 C. urophylla
7 (3) Spines present and recurved   (8)
+ Spines absent or if present straight   (10)
8 (7) Flowering gynophore base white tomentose; leaf blade apex leathery and mucronulate.   4 C. zeylanica
+ Flowering gynophore glabrous; leaf blade apex not leathery or mucronulate   (9)
9 (8) Flowers large, solitary or paired; sepals 1.5-2 cm; stamens 50-60(-75); leaf blade abaxially with papillae.   5 C. hainanensis
+ Flowers medium size, in inflorescences of 2-6; sepals 0.7-1(-1.3) cm; stamens 8-40; leaf blade abaxially without papillae.   6 C. henryi
10 (7) Ovary densely tomentose, 4-6 mm (including style and stigma); fruit globose, ca. 1 cm, beaked.   7 C. olacifolia
+ Ovary glabrous, 1.5-3 mm; fruit subglobose, 1-1.5 cm, not beaked   (11)
11 (10) Ovary ca. 3 mm, with 4 placentae; fruit 3-7 cm; gynophore 3-5 mm thick; petiole 1-2.2 cm.   8 C. micracantha
+ Ovary ca. 2 mm, with 2 placentae; fruit 0.7-2.5 cm; gynophore 0.5-1 mm thick; petiole 0.5-1 cm   (12)
12 (11) New branches shortly tomentose but eventually glabrescent; leaf blade 2-2.5 × as long as wide; secondary veins 5-8(-10) on each side of midvein   (13)
+ New branches generally glabrous; leaf blade 1.8-4.5(-9) × as long as wide; secondary veins (4 or)5 or 6 on each side of midvein   (14)
13 (12) Leaf blade ovate, base rounded to cuneate but never decurrent on petiole; branchlets with reddish brown to gray trichomes, often with recurved spines.   9 C. bodinieri
+ Leaf blade elliptic, base broadly cuneate and decurrent on petiole; branchlets with rust-colored trichomes, often without spines.   10 C. membranifolia
14 (12) Shrubs; twigs without spines; leaf blade thinly herbaceous, lanceolate, 5-11 cm, base rounded, veins obscure on both surfaces; petals inside densely tomentose.   11 C. sabiifolia
+ Climbers; twigs with spines; leaf blade leathery to thinly leathery, oblong, 9-16 cm, base acute, veins distinct on both surfaces; petals inside glabrous but tomentose at margin   (15)
15 (14) Leaf blade obovate, apex abruptly acuminate.   12 C. sunbisiniana
+ Leaf blade oblong-lanceolate to ovate-lanceolate, apex acute to acuminate.   13 C. acutifolia
16 (1) Inflorescences terminal racemes, with 15-30 flowers.   17 C. assamica
+ Inflorescences terminal corymbs, spikes, umbels, axillary fascicles, or sometimes solitary flowers in axils of leaves   (17)
17 (16) Flowers axillary, solitary; anterior sepal distinctly different from other three; upper petals clawed; leaf blade orbicular to suborbicular   (18)
+ Flowers in terminal corymbs, spikes, axillary fascicles, or umbels (if solitary then sepals ± alike in shape); basal pair of petals without claws or 4 petals ± equal, distinct, and without claws; leaf blade variously ovate, elliptic, or lanceolate   (19)
18 (17) Sepals of inner whorl not saccate or galeate, 3-4.5 mm deep in distal half; leaf blade apex acute, obtuse, or retuse but spine-tipped.   1 C. spinosa
+ Sepals of inner whorl deeply saccate or galeate, 9-15 mm deep in distal half; leaf blade apex shortly mucronate.   2 C. himalayensis
19 (17) Leaf blade abaxially densely covered with persistent brown, rust-colored, or ash-yellow trichomes   (20)
+ Leaf blade abaxially glabrous, glabrescent, or with whitish, grayish, or fulvous trichomes   (24)
20 (19) Leaf blade abaxially with persistent very dense ash-yellow trichomes.   30 C. pubifolia
+ Leaf blade abaxially with persistent brown or rust-colored trichomes   (21)
21 (20) Petiole 1-3(-4) mm; flowering gynophore base glabrous   (22)
+ Petiole longer than 5 mm; flowering gynophore base white pubescent   (23)
22 (21) Gynophore 1.5-2 cm; stamens ca. 2 cm; fruit globose, blackish brown when dry even when immature.   18 C. dasyphylla
+ Gynophore 3-4 cm; stamens ca. 3 cm; fruit ellipsoid, gray when dry.   19 C. wui
23 (21) Ovary (not seen) and fruit surface densely rust-colored tomentose.   20 C. trichocarpa
+ Ovary and young fruit surface smooth and glabrous (mature fruit not known).   21 C. viburnifolia
24 (19) Stamens 28-120, anthers ca. 2 mm; sepals 13 mm or longer, inner pair abaxially puberulent   (25)
+ Stamens 20-80, anthers ca. 1 mm; sepals 3-10 mm, inner pair abaxially glabrous   (29)
25 (24) New branches glabrous; leaf blade leathery, both surfaces glabrous when young   (26)
+ New branches with trichomes; leaf blade subleathery, abaxially with trichomes when young but glabrescent   (27)
26 (25) Leaf blade dark red when dry, base cordate; petiole 1-1.2 cm; fruit ellipsoid, 9-13 cm.   22 C. fohaiensis
+ Leaf blade yellowish green when dry, base broadly cuneate to rounded but not cordate; petiole 1.5-2.5 cm; fruit globose, 4.2-5.5 cm.   23 C. formosana
27 (25) Sepals 1.7-1.8(-2.1) cm.   24 C. yunnanensis
+ Sepals 0.8-1.5 cm   (28)
28 (27) Fruit with 4-8 vertical irregularly crested rib ridges when dry and between ribs irregularly raised; sepals 8-12 mm; stamens 45-50(-65).   25 C. masaikai
+ Fruit smooth when dry; sepals 7-9 mm; stamens 28-36.   26 C. sikkimensis
29 (24) Leaf blade apex shallowly emarginate at least for some leaves, for others obtuse, rounded, or subacute.   27 C. sepiaria
+ Leaf blade apex acute, acuminate, obtuse, or rounded but never emarginate   (30)
30 (29) Inflorescences short axillary racemes or fascicles.   28 C. pubiflora
+ Inflorescences umbellate and sometimes aggregated into panicles   (31)
31 (30) Sepals 2-5(-6) mm; gynophore 0.4-1.2 cm; fruit less than 2(-2.5) cm in diam   (32)
+ Sepals 6-10 mm; gynophore 2-5 cm; fruit often 3-5 cm in diam   (34)
32 (31) Leaf blade base subcordate.   29 C. pachyphylla
+ Leaf blade base cuneate, broadly cuneate, or rounded   (33)
33 (32) Twigs with trichomes but glabrescent; sepals caducous after anthesis; stamens 20-45; placentae 2.   31 C. cantoniensis
+ Twigs glabrous; sepals persistent for a short while after anthesis; stamens 7-9(-12); placentae 4.   32 C. floribunda
34 (31) Inflorescences at least at pedicel base of several flowers with a pair of small yellow spines; spines on flowering twig to 5 mm, recurved to uncinate   (35)
+ Inflorescences without spines; spines on flowering twigs short and straight or absent   (36)
35 (34) Pedicel 0.7-1.7 cm; inflorescences 10-15-flowered (Guangxi and Yunnan).   33 C. chingiana
+ Pedicel 2.5-3.5 cm; inflorescences 6-9-flowered (Hainan and Taiwan).   34 C. lanceolaris
36 (34) Inflorescences a short raceme or corymb, 7-12-flowered; sepals outside shortly pubescent; stamens ca. 25.   35 C. khuamak
+ Inflorescences a subumbel, (1 or)2-5-flowered; sepals outside glabrous; stamens 35 or more   (37)
37 (36) New branches glabrous; leaf blade apex shortly mucronate; stamens ca. 35.   36 C. fengii
+ New branches shortly brown pubescent later glabrescent (but near petiole and node always with residual trichomes); leaf blade apex emarginate (on at least some leaves); stamens 50-70.   37 C. versicolor

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