Smilacaceae Vent.

Excluding Ripogonaceae

Habit and leaf form. Scandent shrubs and herbs, or lianas. Perennial; with neither basal nor terminal aggregations of leaves; rhizomatous, or tuberous. Climbing; tendril climbers (the tendrils from the petiole bases), or petiole twiners (the tendrils sometimes reduced to points). Mesophytic and xerophytic. Leaves evergreen; alternate (mostly), or opposite; usually leathery; petiolate to subsessile; sheathing (rarely), or non-sheathing. Leaf sheaths when present, with free margins. Leaves foetid, or without marked odour; simple (though the tendrils, sometimes called `stipules', have also been seen as representing the midveins of lateral leaflets). Lamina entire; lanceolate, or ovate; one-veined, or palmately veined (curved-convergent); cross-venulate; cordate, or hastate, or attenuate at the base. Leaves stipulate (ostensibly, if the tendril bases be so interpreted), or exstipulate. Lamina margins entire.

Leaf anatomy. Stomata present; anomocytic. Hairs present, or absent.

Minor leaf veins without phloem transfer cells (Smilax). Vessels present, or absent; end-walls scalariform.

Stem anatomy. Secretory cavities present. Cork cambium present. Secondary thickening absent. Xylem with vessels. Vessel end-walls scalariform.

Root anatomy. Root xylem with vessels. Vessel end-walls scalariform.

Reproductive type, pollination. Dioecious. Entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in `inflorescences'; in cymes, or in racemes, or in spikes, or in umbels, or in verticils. Inflorescences not scapiflorous; terminal, or axillary; often of one or more superimposed verticils appearing as racemes, spikes or umbels. Flowers regular; 3 merous; cyclic. Perigone tube present (short or long), or absent.

Perianth of `tepals'; 6; free, or joined; 2 whorled; isomerous; petaloid (but fairly inconspicuous); similar in the two whorls to different in the two whorls (the inner members more or less reduced, sometimes fringed).

Androecium 3 (rarely), or 6 (usually), or 9-18 (rarely). Androecial members free of the perianth; free of one another, or coherent; sometimes 1 - adelphous (filaments fused into a tube or column); (1-)2(-6) - whorled. Androecium exclusively of fertile stamens. Stamens 6 (usually), or 3, or 9, or 12, or 15, or 18; isomerous with the perianth to polystemonous. Anthers basifixed; dehiscing via longitudinal slits; introrse, or latrorse; unilocular; tetrasporangiate. The thickenings spiral. Microsporogenesis successive. The initial microspore tetrads isobilateral. Tapetum glandular. Pollen shed as single grains. Pollen grains nonaperturate (or indistinctly so); 2-celled.

Gynoecium 1 (rarely), or 3. Carpels isomerous with the perianth, or reduced in number relative to the perianth (rarely). Gynoecium monomerous (rarely), or syncarpous; of one carpel (rarely), or synovarious to synstylovarious; superior. Carpel stylate; apically stigmatic; when monocarpellary 1 ovuled, or 2 ovuled. Ovary when syncarpous 3 locular. Gynoecium stylate. Styles 3; free (usually), or partially joined; apical. Stigmas dry type; papillate; Group II type. Placentation axile. Ovules 1 per locule, or 2 per locule; pendulous; non-arillate; orthotropous (usually), or hemianatropous, or campylotropous; bitegmic; crassinucellate. Embryo-sac development Polygonum-type. Endosperm formation nuclear.

Fruit fleshy. The fruiting carpel when monocarpellary, baccate. Fruit when syncarpous (i.e. usually) indehiscent; a berry; usually 3 seeded. Seeds endospermic. Endosperm oily. Embryo well differentiated (mostly small). Embryo straight. Testa without phytomelan.

Physiology, biochemistry. Proanthocyanidins absent. Flavonols present, or absent; when present, quercetin, or kaempferol and quercetin. Ellagic acid absent. Saponins/sapogenins present. C3. C3 recorded in Smilax. Anatomy non-C4 type (Smilax).

Geography, cytology. Holarctic, Paleotropical, Neotropical, Cape, and Australian. Temperate (warm), or sub-tropical to tropical. Widespread, tropical, subtropical and warm temperate. X = 10, 13-16.

Taxonomy. Subclass Monocotyledonae. Superorder Liliiflorae; Dioscoreales. Species 370. Genera 3; Heterosmilax, Pseudosmilax, Smilax.

Illustrations. smila215.gif smila855.gif