Including Cyclocheilaceae W. Marais, Nyctanth(ac)eae J.G. Agardh, Petreaceae J.G. Agardh, Pyrenaceae Vent., Viticaceae(`-ices') Juss.
Excluding Dicrastylidaceae, Phrymataceae, Stilbaceae, Symphoremataceae
Habit and leaf form. Trees, shrubs, and herbs, or lianas (many); bearing essential oils. `Normal' plants and switch-plants; sometimes with the principal photosynthesizing function transferred to stems. Leaves well developed, or much reduced (occasionally). Self supporting, or climbing; the climbers stem twiners, or scrambling. Mesophytic and xerophytic. Leaves opposite (usually), or whorled, or alternate (rarely); petiolate to sessile; foetid, or without marked odour; simple, or compound; epulvinate; when compound, ternate to pinnate (e.g.Vitex), or palmate. Lamina dissected, or entire; when dissected, pinnatifid; pinnately veined; cross-venulate. Leaves exstipulate; without a persistent basal meristem. Domatia recorded (4 genera); represented by pits, or pockets, or hair tufts.
Leaf anatomy. The mesophyll with sclerencymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (6 genera).
Stem anatomy. Young stems tetragonal (often), or cylindrical, or oval in section. Cork cambium present; initially deep-seated (rarely), or superficial. Nodes unilacunar (1-several traces). Internal phloem absent. Secondary thickening developing from a conventional cambial ring. `Included' phloem absent. Xylem with libriform fibres; with vessels. Vessel end-walls simple. Vessels without vestured pits.
Reproductive type, pollination. Hermaphrodite (usually). Entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers usually aggregated in `inflorescences'; in cymes, in racemes, in spikes, in heads, and in verticils. The terminal inflorescence unit cymose, or racemose. Inflorescences terminal, or axillary; with involucral bracts (often, these commonly coloured), or without involucral bracts; pseudanthial (sometimes), or not pseudanthial. Flowers bracteate; small to medium-sized; very irregular (usually), or regular to somewhat irregular. The floral asymmetry involving the perianth and involving the androecium. Flowers (4-)5(-8) merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk present, or absent; when present, annular.
Perianth with distinct calyx and corolla; (8-)10(-16); 2 whorled; isomerous. Calyx (4-)5(-8); 1 whorled; gamosepalous; unequal but not bilabiate, or regular; persistent; when K5, with the odd member posterior. Corolla (4-)5(-8); 1 whorled; gamopetalous; imbricate; tubular (usually), or campanulate (rarely); unequal but not bilabiate, or bilabiate, or regular (rarely).
Androecium (2-)4(-5). Androecial members adnate (to the corolla tube); markedly unequal (usually), or all equal (rarely); free of one another; 1 - whorled. Androecium exclusively of fertile stamens, or including staminodes. Staminodes when present, 1-3; non-petaloid. Stamens (2-)4(-5) (the posterior member usually, and sometimes the three upper members, reduced or missing); usually didynamous; reduced in number relative to the adjacent perianth (usually), or isomerous with the perianth; oppositisepalous. Anthers connivent (in pairs), or separate from one another; dorsifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer; of the `dicot' type, or of the `monocot' type. Tapetum glandular. Pollen grains aperturate; (2-)3(-5) - aperturate, or 6 - aperturate; colpate, or colporate, or rugate; 2-celled, or 3-celled.
Gynoecium 2 (usually), or 4, or 5. Carpels reduced in number relative to the perianth (usually), or isomerous with the perianth. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 2 locular, or 4 locular, or 8 locular, or 10 locular (the original locules (usually two) early becoming divided by a `false septum' in each - cf. Labiatae). Gynoecium usually median; stylate. Styles 1; attenuate from the ovary, or from a depression at the top of the ovary (but the ovary apex no more than slightly lobed); apical, or lateral. Stigmas 1; 1 - lobed, or 2 - lobed; wet type; papillate; Group III type and Group IV type. Placentation axile. Ovules 2 per locule (in the original locules), or 1 per locule (in the secondary locules); pendulous, or horizontal, or ascending; non-arillate; orthotropous, or hemianatropous, or anatropous; unitegmic; tenuinucellate. Endothelium not differentiated. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; ephemeral (usually), or persistent. Synergids usually hooked (and beaked). Endosperm formation cellular. Endosperm haustoria present; chalazal and micropylar (the latter usually the less well developed). Embryogeny onagrad.
Fruit fleshy (usually), or non-fleshy; dehiscent (rarely), or indehiscent, or a schizocarp; when schizocarpic comprising nutlets, or comprising drupelets; when non-schizocarpic a drupe (usually), or a capsule. Capsules valvular (with 2-4 valves). Seeds non-endospermic. Cotyledons 2 (expanded, flat). Embryo achlorophyllous (5/5); straight.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present, or absent. Iridoids recorded; carbocyclic and seco-compounds. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (5 genera, 6 species). Ursolic acid present. Saponins/sapogenins present, or absent. Aluminium accumulation not found. Sugars transported as oligosaccharides + sucrose. C3. C3 recorded in Verbena. Anatomy non-C4 type (Cleodendron, Lantana, Premna, Stachytarpheta, Verbena, Vitex).
Geography, cytology. Temperate, or sub-tropical to tropical (mainly). Very widespreadtemperate and tropical, but absent from central and Northern Eurasia. X = 5-12.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Lamiiflorae; Lamiales. Cronquist's Subclass Asteridae; Lamiales. Takhtajan's Subclass Asteridae; Lamianae; Lamiales. Species 3000. Genera about 90; Acantholippia, Adelosa, Amasonia, Baillonia, Bouchea, Callicarpa, Caryopteris, Citharexylum, Clerodendrum, Cornutea, Cyclocheilon, Dimetra, Faradaya, Garrettia, Gmelina, Hosea, Karomia, Lantana, Lippia, Neorapinia, Nyctanthes, Oncinocalyx, Oxera, Paravitex, Peronema, Petraea, Premna, Priva, Stachytarpheta, Stylodon, Tectona, Teijsmanniodendron, Teucridium, Urbania, Verbena, Vitex, etc.
Economic uses, etc. Timber from Tectona grandis (teak); some notable ornamentals, e.g. Clerodendrum, Callicarpa, Vitex, Lantana, Verbena; noxious, photosensitizing weeds (Lantana).
Illustrations. verbe470.gif verbe615.gif verbe663.gif
Additional, to be intercalated. Clerodendrum twining clockwise.
