Capparis in Flora of China @ efloras.org

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

List of lower taxa

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Capparis (PDF)

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