Cleomaceae (Pax) Airy Shaw

~ Capparidaceae

Including Cleomaceae Horan. (p.p.)

Habit and leaf form. Herbs (mostly), or shrubs (a few), or trees (a few), or lianas (a few); glandular, resinous. Annual (mostly), or perennial. Self supporting, or climbing (a few). Leaves alternate; spiral; petiolate; non-sheathing; aromatic, or without marked odour (?), or foetid; compound; unifoliolate (rarely, the leaves `simple'), or ternate, or palmate ((1-)3-7 foliolate); stipulate. Stipules scaly, or leafy, or spiny.

Leaf anatomy. Stomata present; anomocytic.

Lamina dorsiventral to centric. Minor leaf veins without phloem transfer cells (Cleome, Isomeris).

Stem anatomy. Secondary thickening developing from a conventional cambial ring. Vessel end-walls simple. Wood parenchyma paratracheal.

Reproductive type, pollination. Hermaphrodite, or andromonoecious.

Inflorescence, floral, fruit and seed morphology. Flowers somewhat irregular to very irregular. Floral receptacle developing a gynophore, or developing an androphore and developing a gynophore. Hypogynous disk present.

Perianth with distinct calyx and corolla; (4-)8(-12); 2-4 whorled (more or less resolvable into K2+2 C2+2, or only theoretically so); isomerous. Calyx (2-)4(-6); 1 whorled, or 2 whorled; polysepalous, or gamosepalous; persistent, or not persistent; imbricate, or open in bud. Corolla (2-)4(-6); 1-2 whorled; polypetalous, or gamopetalous (connate or not). Petals clawed.

Androecium 4-100 (the four primordia often branching to give many stamens). Androecial members branched (from 4 primordia), or unbranched; when many/branched, maturing centrifugally; free of the perianth; united with the gynoecium (fused to the gynophore), or free of the gynoecium; all equal, or markedly unequal. Androecium exclusively of fertile stamens, or including staminodes. Stamens 4-50; tetradynamous (rarely), or not didynamous, not tetradynamous; basically oppositisepalous (i.e. considering the primordia). Anthers dehiscing via longitudinal slits; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Anther wall initially with more than one middle layer. Tapetum glandular. Pollen grains aperturate; (2-)3(-4) - aperturate; colporate.

Gynoecium 2; syncarpous; eu-syncarpous; superior. Ovary 1 locular. Gynoecium transverse. Styles 1; apical. Stigmas 1; dry type; papillate; Group II type. Placentation parietal. Ovules in the single cavity 10-100 (`many'); campylotropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped (with diverging tips), or hooked. Endosperm formation nuclear. Embryogeny onagrad.

Fruit non-fleshy; dehiscent; a siliqua. Seeds scarcely endospermic, or non-endospermic. Cotyledons 2. Embryo chlorophyllous (1/1); curved, or bent.

Physiology, biochemistry. Mustard-oils present. Cyanogenic, or not cyanogenic. Alkaloids absent. Proanthocyanidins absent. Flavonols present; kaempferol and quercetin, or quercetin. Ellagic acid present (Cleome gigantea), or absent (C. spinosa, Polanisia). Aluminium accumulation not found. C3 and C4. C3 recorded in Cleome. C4 recorded in Gynandropsis. Anatomy C4 type (Gynandropsis), or non-C4 type (Cleome).

Geography, cytology. Sub-tropical and tropical. Widespread tropical and subtropical.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren's Superorder Violiflorae; Capparales. Cronquist's Subclass Dilleniidae; Capparales. Takhtajan's Subclass Dilleniidae; Dillenianae; Capparales. Species 275. Genera about 8; Buhsia, Cleome, Cleomella, Gynandropsis, Haptocarpum, Physostemon, Polanisia, Wislizenia.

Separation from Capparidaceae may be unsustainable, since difficulties are encountered in assigning the genera.

Illustrations. cleom215.gif cleom233.gif cleom358.gif