Family Cleomaceae
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. Abaxial epidermis papillose, or not papillose. Stomata present; anomocytic. Hairs present, or absent; commonly glandular (by contrast with Capparidaceae). 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. Plants 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); 2–3 whorled (more or less resolvable into K2+2 C4 with decussate calyx and diagonal petals, cf. Cruciferae, or only theoretically so). 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 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; 2-celled. Gynoecium 2 carpelled. The pistil 2 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1 locular, or 2 locular (with a replum). Locules secondarily divided by ‘false septa’ (the replum), or without ‘false septa’. Gynoecium transverse; stylate, or non-stylate to stylate. Styles 1; apical. Stigmas 1; dry type; papillate; Group II type. Placentation parietal (the two placentas longitudinally divided by a replum, cf. Cruciferae). Ovules in the single cavity 10–100 (to ‘many’); 10–50 per locule (‘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 silicula, or a siliqua. Capsules valvular (the two valves separating from the persistent replum, by contrast with capsular Capparidaceae). Seeds scarcely endospermic, or non-endospermic. Cotyledons 2. Embryo chlorophyllous (1/1); curved, or bent. The radicle lateral, or dorsal. 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 physiology recorded directly in Cleome. C4 physiology recorded directly 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. APG 3 core angiosperms; core eudicot; Superorder Rosanae; malvid; Order Brassicales. 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. Precise comparative data on gynoecium and fruit structure are elusive or non-existent. Illustrations. |
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