Family Aizoaceae
Habit and leaf form. Small shrubs, or herbs. ‘Normal’ plants, or plants of very peculiar vegetative form; commonly more or less neotenic (being often reduced to a single annual pair of leaves, some forms partly subterranean with only a clear window at each leaf tip exposed above-ground). Leaves well developed (usually), or much reduced. Plants succulent. The herbs annual, or perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Xerophytic. Leaves minute to medium-sized; alternate, or opposite (or in false whorls); often terete; fleshy; imbricate to not imbricate; shortly petiolate to sessile; when opposite, connate, or not connate; sheathing, or non-sheathing; mostly simple; epulvinate. Lamina entire; one-veined, or pinnately veined, or palmately veined. Leaves stipulate, or exstipulate. Vegetative buds not scaly. Leaves without a persistent basal meristem. Leaf anatomy. Abaxial epidermis papillose, or not papillose. Stomata anomocytic, or anomocytic and paracytic. Hairs present, or absent; unicellular, or multicellular. Unicellular hairs branched, or unbranched. Urticating hairs absent. Adaxial hypodermis absent. Lamina mostly isobilateral to centric. Main veins embedded. Minor leaf veins without phloem transfer cells (Mesembryanthemum). Stem anatomy. Cork cambium present, or absent; initially deep-seated. Nodes unilacunar. Primary vascular tissue centrifugal. Internal phloem absent. Secondary thickening absent, or developing from a conventional cambial ring, or anomalous; via concentric cambia (in the woodier genera,), or from a single cambial ring. ‘Included’ phloem present, or absent. Xylem with libriform fibres; with vessels. Vessel end-walls usually simple. Primary medullary rays wide. Wood partially storied (VP), or not storied. Sieve-tube plastids P-type; type III (a). Reproductive type, pollination. Plants hermaphrodite (usually), or monoecious (rarely), or dioecious (rarely). Pollination entomophilous (diurnal). Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; (when aggregated) in cymes, or in heads. The ultimate inflorescence unit (when flowers aggregated) cymose. Inflorescences cymes and heads. Flowers small, or medium-sized; regular; not resupinate; cyclic; pentacyclic to polycyclic. Free hypanthium present to absent. Hypogynous disk present, or absent. Perianth sepaline, or with distinct calyx and corolla (generally considered apetalous, but often with colourful, conspicuous staminodal ‘petals’); (1–)4–5(–20), or 20–120 (if the staminodes are interpreted as petals). Calyx (1–)4–5(–8); 1 whorled; gamosepalous (nearly always), or polysepalous (sometimes in Gunniopsis, Galenia); regular, or unequal but not bilabiate (e.g. sometimes with two large members and three smaller); fleshy; persistent; imbricate, or valvate (rarely). Corolla if considered present, 5–120 (of staminodal origin); 1–6 whorled. Androecium (3–)4–5, or 8–10, or 15–200 (i.e. to ‘many’, by branching). Androecial members commonly branched (by dédoublement), or unbranched; when branched/many, maturing centrifugally; free of one another, or coherent; when coherent 3–9 adelphous (?), or 1 adelphous (the filaments basally connate into bundles, or forming a short monadelphous sheath); 1–16 whorled (i.e to ‘many whorls’). Androecium including staminodes (nearly always), or exclusively of fertile stamens (i.e. occasionally no staminodal ‘petals’). Staminodes when present, 3–120 (usually numerous); external to the fertile stamens; petaloid. Stamens (1–)5, or 6–120 (i.e. to ‘many’); reduced in number relative to the adjacent perianth (rarely), or isomerous with the perianth, or diplostemonous to polystemonous; filantherous. Anthers dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer, or initially with more than one middle layer; of the ‘basic’ type, or of the ‘monocot’ type. Tapetum glandular. Pollen monosiphonous; shed as single grains. Pollen grains aperturate; 3 aperturate; colporate (colporoidate), or colpate; spinulose; 3-celled. Gynoecium (1–)2–5(–20) carpelled. The pistil (1–)2–5(–20) celled. Gynoecium syncarpous (rarely pseudomonomerous); synovarious to synstylovarious; superior to inferior. Ovary (1–)2–5(–20) locular. Gynoecium non-stylate to stylate. Styles 1–20 (or absent); apical, or apical to lateral (excentric). Stigmas (1–)2–5(–20); wet type, or dry type; papillate; Group II type, or Group III type. Placentation when unilocular (i.e., rarely) parietal; when plurilocular, axile (typically), or basal (rarely), or axile to apical (e.g. Galenia), or parietal (Mesembryanthemum sensu lato). Ovules (1–)50 per locule; arillate, or non-arillate; anatropous, or campylotropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Endothelium not differentiated. Embryo-sac development Polygonum-type (or unspecified). Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped, or hooked (rarely). Endosperm formation nuclear. Embryogeny caryophyllad, or solanad. Fruit fleshy (rarely), or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry. Capsules loculicidal, or valvular, or septicidal, or circumscissile. Seeds non-endospermic. Perisperm present (mealy). Seeds with starch. Embryo well differentiated. Cotyledons 2. Embryo achlorophyllous (2/2); curved. The radicle dorsal. Seedling. Germination phanerocotylar. Physiology, biochemistry. Not cyanogenic. Alkaloids present (commonly), or absent. Iridoids not detected. Proanthocyanidins present, or absent; when present, cyanidin. Flavonols present, or absent; when present, kaempferol, or kaempferol and quercetin. Ellagic acid absent (4 species, 4 genera). Betalains present. Saponins/sapogenins absent. Aluminium accumulation not found. Plants commonly accumulating free oxalates. C4, or CAM. C4 physiology recorded directly in Lithops, Sesuvium, Trianthema, Zaleya — but Lithops salicola and L. venteri are also listed as CAM. CAM recorded directly in Aizoanthemum, Amoebophyllum, Anisocalyx, Aptenia, Argyroderma, Aridaria, Aspazoma, Astridia, Bergeranthus, Brownanthus, Carpobrotus, Carruanthus, Cephalophyllum, Cheiridopsis, Conophytum, Disphyma, Drosanthemum, Eberlanzia, Faucaria, Fenestraria, Hereroa, Jacobsensis, Lampranthus, Lapidaria, Leipoldtia, Lithops, Malephora, Mesembryanthemum, Mestoklema, Meyerophytum, Mitrophyllum, Monilaria, Nananthus, Opophythum, Platythyra, Prenia, Psilocaulon, Rhombophyllum, Ruschia, Sceletium, Sphalmanthus, Stoeberia, Titanopsis, Trichodiadema, Vanheerdia. Anatomy C4 type (Lithops, Sesuvium, Trianthema, Zaleya), or non-C4 type (Aizoon, Carpobrotus, Corbichonia, Disphyma, Galenia, Mesembryanthemum, Neogunnia, Sesuvium). Geography, cytology. Chiefly southern Africa, but also tropical Africa and Asia, Australia, California, South America. X = 8, 9. Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Caryophylliflorae; Caryophyllales. Cronquist’s Subclass Caryophyllidae; Caryophyllales. APG (1998) Eudicot; core Eudicot; neither Rosid nor Asterid. APG 3 (2009) Order: Caryophyllales. Species 1100. Genera 126; Acrodon, Acrosanthes, Aethephyllum, Aizoanthemum, Aizoön, Aloinopsis, Amphibolia, Antigibbaeum, Antimema, Apatesia, Aptenia, Arenifera, Argyroderma, Aspazoma, Astridia, Bergeranthus, Berrisfordia, Bijlia, Braunsia, Brownanthus, Carpanthea, Carpobrotus, Carruanthus, Caryotophora, Cephalophyllum, Cerochlamys, Chasmatophyllum, Cheiridopsis, Circandra, Cleretum, Conicosia, Conophytum, Corpuscularia, Cylindrophyllum, Cypselea, Dactylopsis, Delosperma, Dicrocaulon, Didymaotus, Dinteranthus, Diplosoma, Disphyma, Dorotheanthus, Dracophilus, Drosanthemopsis, Drosanthemum, Eberlanzia, Ebracteola, Enarganthe, Erepsia, Esterhuysenia, Faucaria, Fenestraria, Frithia, Galenia, Gibbaeum, Glottiphyllum, Gunniopsis, Hallianthus, Hereroa, Herreanthus, Hymenogyne, Imitaria, Jacobsenia, Jensenobotrya, Jordaaniella, Juttadinteria, Khadia, Lampranthus, Lapidaria, Leipoldtia, Lithops, Machairophyllum, Malephora, Mesembryanthemum, Mestoklema, Meyerophytum, Mitrophyllum, Monilaria, Mossia, Muiria, Namaquanthus, Namibia, Nananthus, Nelia, Neohenricia, Octopoma, Odontophorus, Oophytum, Ophthalmophyllum, Orthopterum, Oscularia, Ottosonderia, Phyllobolus, Pleiospilos, Plinthus, Polymita, Psammophora, Pseudobrownanthus, Psilocaulon, Rabiea, Rhinephyllum, Rhombophyllum, Ruschia, Ruschianthemum, Ruschianthus, Saphesia, Schlechteranthus, Schwantesia, Scopelogena, Sesuvium, Skiatophytum, Smicrostigma, Stayneria, Stoeberia, Stomatium, Synaptophyllum, Tanquana, Titanopsis, Trianthema, Trichodiadema, Vanheerdea, Vanzijlia, Wooleya, Zaleya, Zeuktophyllum. Economic uses, etc. Edible fruit from Mesembryanthemum edule (Hottentot fig). |
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