Capparis L.

Shrubs or rarely trees, erect, scandent or sprawling; shoots generally armed with stipular spines, infrequently unarmed; base of shoots sometimes surrounded by cataphylls; indumentum simple or with stellate hairs. Leaves petiolate, alternate, simple, entire, rarely reduced and apparently leafless. Flowers bisexual, arranged in panicles, racemes, corymbs or umbels, sometimes solitary, axillary or supraaxillary in vertical series or 3 - 4 conferted on condensed lateral shoots; bracts early caducous. Sepals 4, mostly caducous after anthesis, in two unequal whorls of two each; the outer concave and overlapping inner ones, imbricate or valvate, free or almost so, rarely outer pair connate in bud; inner pair always free and flat. Petals 4, caducous after anthesis, mostly unequal, oblong or obovate, not clawed; the upper pair cohering around disc, but not connate; lower pair free. Receptacle flat with small adaxial disc. Stamens 8 to many, inserted on torus; androphore absent. Ovary ovoid or spherical on a gynophore not nmuch elongating but often incrassate in fruit, unilocular, 4-many-ovuled; placentae 2 - 6; style short; stigma obscure to capitate. Berry on a slender or thickened stipe, globose, elongate or ovoid; pericarp leathery, corky or smooth, ribbed or sculptured, indehiscent or tardily dehiscent; seeds 1 - many, reniform, embedded in pulp; embryo convolute.

Pantropic in America, Africa, Asia, Australia, sometimes in temperate regions of Europe, China, Australia, S. Africa and S. America; ca 240 species, 29 species in India.

Literature. JACOBS, M. (1965) The genus Capparis (Capparaceae) from Indus to the Pacific. Blumea 12: 385 - 541. JAFRI, S. M. H. (1956) The genus Capparis in West Pakistan, Afghanistan and N.W. Himalaya. Pakistan J. Forestry 6: 191 - 192. NICOLSON, D. H. (1975) The reinstatement of Capparis rheedii DC. (Capparaceae). Bull. Bot. Surv. India 17: 160 - 161. ST.JOHN, H. (1965). Revision of Capparis spinosa and its African, Asian and Pacific relatives. Micronesia 2: 25 - 44. SUNDARA RAGHAVAN, R. & R. S. RAO (1965) Critical notes on three species of Capparis Linn. from peninsular India. J. Bombay Nat. Hist. Soc. 62: 412 - 424.

Notes. Plants usually noctiflorous, with flowers opening at dusk and pollinated by butterflies, moths or other nocturnal insects. Seed dispersal is effected by mammals or birds. Sterility is prevalent in both sexes. Pollen longiaxis, oval,3-colporate. Ectoaperture colpi long extending to pole. Ectoaperture more or less circular, with granular teeth. Exine ornamentation psillate, reticulate, rugose or perforate. Pollen quite homogenous.

KEY TO THE SPECIES

1a. Leaves small, only on young twigs, early caducous; plants apparently leafless

7. Capparis decidua

b. Leaves large, persistent, plants prominently leafy

2

2a. Flowers solitary, axillary (rarely 2 - 4 conferted together on lateral axis as in C. brevispina or C. nilgiriensis)

3

b. Flowers generally not solitary and axillary, but variously arranged, (rarely a few flowers axillary if inflorescence is a corymb or in supra-axillary series)

9

3a. Upper petals with a yellow or purplish blotch; ovary densely hairy

4

b. Upper petals not blotched as above; ovary glabrous

7

4a. Leaves with basal two pairs of nerves conferted towards base

5

b. Leaves with arching nerves, never conferted towards base

6

5a. Leaves with prominent reticulation and acute apex; flowers 8 - 10 cm across; gynophore glabrous during anthesis

21. Capparis rheedei

b. Leaves with obscure reticulation and obtuse apex; flowers 1.5 - 1.8 cm across, gynophore hairy during anthesis

10. Capparis flavicans

6a. Leaves stiff, recurved at margins, mucronate at tip; fruits beaked, with smooth pericarp

3. Capparis brevispina

b. Leaves flexuous, not recurved at margins, not mucronate at tip; fruits not beaked, with warted or corrugated pericarp

18. Capparis nilgiriensis

7a. Outer sepals connate in bud with free tip; inner sepals petaloid; fruits with sculptured pericarp

8. Capparis divaricata

b. All sepals free, none petaloid; fruits with ribbed pericarp

8

8a. Twigs with brownish stellate indumentum; petioles 2 - 3 mm long; pedicels 1 - 1.5 cm long

13. Capparis grandiflora

b. Twigs with cobweb-like tomentum, at length glabrescent; petioles more than 5 mm long; pedicels 3 - 8 cm long

27. Capparis spinosa

9a. Flowers in supra-axillaly vertical series of 2-10 or even more (rarely solitaty, axillary)

10

b. Flowers otherwise, never in supra-axillaty vertical series

15

10a. Flowers appearing before leaves, hence flowering shoots apparently leafless

11

b. Flowers not developing before leaves, hence flowering shoots always leafy

12

11a. Flowers 2- 4 -stichous, many, extending for 10 - 15 cm or more, small, hardly 4 - 6 mm across; sepals less than 5 mm long; fruits 8 - 10 mm across

17. Capparis multiflora

b. Flowers not 2 - 4-stichous, 2 - 6 (10) together, large, 2.5 - 4 cm across; sepals more than 7 mm long; fruits 4 - 6 mm across

29. Capparis zeylanica

12a. Cataphylls present, at least at base

13

b. Cataphylls absent

14

13a. Leaves drying glaucous, prominently reticulate; overy 3 mm or less long, glabrous; fruits large, exceeding 3 cm across, many-seeded

15. Capparis micracantha

b. Leaves drying brown, indistinctly reticulate; overy 4 - 6 mm long, densely tomentose; fruits small, under 1 cm across, 1 (-3)-seeded

19. Capparis olacifolia

14a. Shoots with recurved spines; stamens 16 - 20

28. Capparis tenera

b. Shoots generally unarmed, if armed, spines straight; stamens exceeding 20

1. Capparis acutifolia

15a. Inflorescence racemose or paniculate, if umbellate, the umbels arranged to form a panicle

16

b. Inflorescence neither racemose, nor paniculate; flowers either in solitaly umbels, subumbels or in corymbs

20

16a. Inflorescence a terminal or subterminal raceme

2. Capparis assamica

b. Inflorescence variously paniculate

17

17a. Cataphylls prominent; flowers in axillary racemose bundles, arranged in the form of a panicle

20. Capparis pachyphylla

b. Cataphylis absent; flowers in axillary sabumbels or corymbs, arranged in the form of a panicle

18

18a. Gynophore exceeding 3 cm in lenath; fruits exceeding 2.5 cm across

26. Capparis sikkimensis

b. Gynophore under 1.5 cm in length; fruits under 2.5 cm across

19

19a. Midrib of leaves flattish or raised at basal part; panicles dense-flowered; stamens 8 - 12; placentae 2; fruits 1.8 - 2 cm across with a leathery pericarp

11. Capparis floribunda

b. Midrib of leaves sunken at basal part; panicles lax-flowered; stamens 18 - 22 placentae 4; fruits under 1.8 cm with thin pericarp

4. Capparis cantoniensis

20a. Flowers in corymbs (with a few subterminal solitaty flowers intermixed at base)

21

b. Flowers umbellate or subumbellate

24

21a. Leaves with olive green velvety indumentum; veins close, more or less parallel, mostly without intermediary nerves

14. Capparis grandis

b. Leaves otherwise, neither velvety pubescent, nor with close parallel nerves

22

22a. Flowers large 10 - 12 cm across; stamens 80 - 120; fruits 10 cm or more across when fully mature

16. Capparis moonii

b. Flowers medium-sized, 3 - 6 cm across; stamens under 60; fruits up to 6 cm across when fully mature

23

23a. Sepals and petals glabrous; gynophore 3.5 - 5.5 cm long; fruits orange-yellow, globose, 5 - 6 cm across, many-seeded

23. Capparis roxburghii

b. Sepals and petals pubescent; gynophore 2.5 - 3.5 cm long; fruits deep purple, umbonate, 2.5 - 3 cm across, 1 - 4-seeded

6. Capparis cleghornii

24a. Cataphylls conspicuous, at least at base

25

b. Cataphylls absent

27

25a. Gynophore 20 - 25 mm long

5. Capparis cinerea

b. Gynophore under 15 mm long

26

26a. Leaves linear to oblong-lanceolate; flowers purple or violet; fruits ellipsoid, 6 or more seeded

9. Capparis diversifolia

b. Leaves elliptic to oblong; flowers greenish white; fruits spindle-shaped, 1 - 2-seeded

12. Capparis fusifera

27a. Mostly unarmed, if armed spines few, straight and short

25. Capparis shevaroyensis

b. Densely armed, spines otherwise

28

28a. Erect Shrubs; leaves with cordate base; spines dimorphic, either long and acicular or sharply recurved or both in the same plant

22. Capparis rotundifolia

b. Scandent shrubs; leaves cuneate, rounded or rarely subcordate at base; spines uniform, recurved

24. Capparis sepiaria

List of Lower Taxa