Trees or shrubs, evergreen or deciduous, monoecious or dioecious; indumentum of stellate
and sometimes simple hairs, usually mixed with sessile granular yellow, orange or red gland
dots. Leaves alternate, subopposite or opposite, decussate, often unequal in size and shape at
each node, entire or sometimes palmately lobed or toothed, penninerved or palmately 3 - 9-
nerved and sometimes peltate at base, sometimes with hairy domatia, usually with discoid
almost immersed glands near the base on upper surface; stipules minute, often caducous.
Inflorescences terminal or axillary, simple or branched racemes or spikes. Male flowers: 1 – 9 per
node; sepals 3 or 4, valvate; petals and disc absent; stamens 20 - 50 (- 150), free, crowded on a
central flat or convex receptacle; anthers globose to oblong, basi- to dorsifixed; thecae parallel,
often separated by wide thickened and widened connective; disc rarely present; pistillode
minute or absent. Female flowers: solitary or rarely paired at nodes along the rachis; calyx
spathaceous, deeply 3 - 6-lobed or minutely toothed at length; ovary (2 -) 3 (- 4)-loculed, rarely
up to 5-loculed, usually echinate, muricate or tubercled; styles 2 or 3, simple, free or shortly
connate at base, erecto-patent, spreading or recurved, papillose or laciniate. Fruits capsular, (2
-) 3 (- 4)-loculed, dehiscing into bivalved parts, tubercled, muricate or echinate; seeds usually
3, rarely 2, ovoid to oblong or globose, black to red, ecarunculate; testa crustaceous; endosperm
fleshy; cotyledons broader than radicles.
Paleotropical, ca 135 species, of which only two are in Africa, the rest in Asia; 17 species
in India.
KEY TO THE SPECIES
1a . Leaves penninerved, not palmately nerved at base
2
b. Leaves palmately 3 - 9-nerved at base, penninerved above
7
2 a. Leaves strictly opposite
3
b. Leaves alternate or opposite in young shoots
4
3 a. Branchlets terete; leaves of each pair at each node unequal in size, one being of normal size and
the other stipuliform
b. Leaves alternate, tomentose beneath; male receptacle not central; fruits densely fulvous
tomentose; seeds black
11
11 a. Leaves narrowly oblong-ovate, mostly twice as long as broad, with red granules beneath;
stamens 15 - 30; styles 2; fruits 3 or 4-loculed, densely clothed with crimson-red resinous
powder
b. Leaves broadly ovate-deltoid, mostly as long as broad, yellow tomentose and glandular beneath;
stamens 40 - 80; styles 2; fruits 2-loculed, densely fulvous tomentose, not with red resinous
powdery glands
16 a. Leaves nearly oblong to elliptic or ovate-elliptic, 10 – 35 cm long; lateral nerves 5 – 10 pairs;
racemes 6 – 22 cm long, much shorter than the leaves
