Including Cestraceae (`ineae') Schlechtd., Salpiglossidaceae Hutch., Sclerophylacaceae Miers
Excluding Goetzeaceae
Habit and leaf form. Herbs, shrubs, trees, and lianas (often prickly); non-laticiferous and without coloured juice; resinous, or not resinous. `Normal' plants (usually), or `normal' plants to switch-plants (occasionally). Plants non-succulent. Annual, or biennial, or perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting, or climbing; the climbers stem twiners, or scrambling. Helophytic to xerophytic. Leaves alternate, or alternate to opposite (usually alternate below, but often becoming opposite towards the inflorescence); usually spiral (at least below); `herbaceous', or leathery, or modified into spines; petiolate (mostly), or subsessile, or sessile; non-sheathing; gland-dotted (rarely, e.g. Anthocercis), or not gland-dotted; aromatic, or foetid (assignment of (e.g.) Anthocercis as pleasant/unpleasant being a matter of opinion), or without marked odour (mostly); simple, or compound; epulvinate; when compound, ternate, or pinnate. Lamina dissected, or entire; when simple/dissected, pinnatifid, or spinose; pinnately veined; cross-venulate. Leaves exstipulate; without a persistent basal meristem. Domatia recorded (from 4 genera); represented by hair tufts.
General anatomy. Plants with `crystal sand' (commonly), or without `crystal sand'.
Leaf anatomy. Stomata mainly confined to one surface, or on both surfaces; anomocytic, or anisocytic, or diacytic. Hairs usually present; multicellular. Multicellular hairs branched.
Lamina dorsiventral (usually), or isobilateral; without secretory cavities. The mesophyll containing mucilage cells (rarely), or not containing mucilage cells. Minor leaf veins without phloem transfer cells (5 genera).
Stem anatomy. Cork cambium present; initially deep-seated, or superficial. Nodes unilacunar (with 2 or 3 traces). Primary vascular tissue bicollateral. Internal phloem present. Secondary thickening developing from a conventional cambial ring (usually), or anomalous; from a single cambial ring. `Included' phloem present (occasionally, e.g. Atropa belladonna rhizomes), or absent. Xylem with tracheids, or without tracheids; with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls simple. Wood parenchyma apotracheal, or paratracheal. Sieve-tube plastids S-type.
Reproductive type, pollination. Hermaphrodite (mostly), or monoecious, or andromonoecious, or dioecious (e.g. sometimes in Solanum and Symonanthus). Entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in `inflorescences'. The terminal inflorescence unit apparently cymose. Inflorescences terminal, or axillary, or leaf-opposed (occasionally). Flowers small to medium-sized; fragrant (e.g. Nicotiana), or malodorous (e.g. Anthocercis, if so considered), or odourless (mostly); regular (usually, more or less), or somewhat irregular to very irregular. The floral asymmetry when noticeable) involving the perianth, or involving the androecium, or involving the perianth and involving the androecium. Flowers mostly (4-)5 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk usually present; intrastaminal.
Perianth with distinct calyx and corolla; 10 (nearly always), or 8; 2 whorled; isomerous, or anisomerous. Calyx (4-)5(-6); 1 whorled; gamosepalous; regular (usually), or unequal but not bilabiate; persistent; accrescent, or non-accrescent. Corolla (4-)5; 1 whorled; gamopetalous; contorted and plicate (usually), or imbricate, or valvate, or contorted; rotate, or campanulate, or funnel-shaped, or tubular; regular (usually, more or less), or bilabiate (rarely).
Androecium 5 (usually), or 3-4 (rarely). Androecial members adnate (epipetalous, on the tube); all equal (often), or markedly unequal; free of one another; 1 - whorled. Androecium exclusively of fertile stamens (usually), or including staminodes (Salpiglossideae). Staminodes when present, 1 (Salpiglossis), or 3 (Schizanthus). Stamens 5 (in all but Salpiglossideae), or 2, or 4; didynamous (e.g. Anthocercis), or not didynamous, not tetradynamous (mostly); reduced in number relative to the adjacent perianth (rarely), or isomerous with the perianth; oppositisepalous. Filaments appendiculate (e.g., inconsistently lobed in Anthocercis), or not appendiculate. Anthers connivent (often touching in a ring at their tops), or separate from one another; dehiscing via pores to dehiscing via short slits (terminally), or dehiscing via longitudinal slits; extrorse (e.g. Anthocercis), or introrse (usually, if dehiscence not terminal); tetrasporangiate. Endothecium developing fibrous thickenings, or not developing fibrous thickenings (when the dehiscence is porose). Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer (1 or 2); of the `basic' type (rarely), or of the `dicot' type. Tapetum glandular. Pollen shed in aggregates (rarely), or shed as single grains; in Salpiglossis, in tetrads. Pollen grains aperturate (usually), or nonaperturate; (2-)3-5(-6) - aperturate; colpate, or colporate (or colporoidate), or rugate; 2-celled.
Gynoecium 2. Carpels reduced in number relative to the perianth. Gynoecium syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 2 locular (nearly always), or 3-5 locular (Nicandreae and Datureae, by secondary divisions). Gynoecium oblique (the posterior carpel to the right, as expressed in conventional floral diagams); stylate. Styles 1; attenuate from the ovary; apical. Stigmas 1-2; if regarded as single, 2 - lobed; wet type, or dry type; papillate, or non-papillate; Group II type, or Group III type, or Group IV type. Placentation axile (the placentae usually more or less swollen). Ovules 1-50 per locule (i.e. to `many'); non-arillate; anatropous, or hemianatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type, or Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral (usually), or persistent (e.g. Atropa). Synergids pear-shaped, or hooked (sometimes with filiform apparatus). Endosperm formation cellular, or nuclear, or helobial. Endosperm haustoria usually present; antipodal. Embryogeny solanad (usually), or onagrad (rarely).
Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry, or a drupe. Capsules septicidal (commonly), or loculicidal, or valvular, or circumscissile (Hyoscyamus). Seeds endospermic (usually). Endosperm oily (usually), or not oily (rarely starchy). Seeds with starch (rarely), or without starch. Cotyledons 2. Embryo achlorophyllous (13/21); straight, or straight to curved, or curved (curved through more than a semicircle to annular in Nicandreae, Solaneae and Datureae, but straight to only slightly curved in Cestreae and Salpiglossideae).
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present (mostly), or absent. Iridoids absent. Proanthocyanidins absent. Flavonols present, or absent; kaempferol and quercetin (mostly). Ellagic acid absent (25 species, 14 genera). Arbutin absent. Ursolic acid present. Saponins/sapogenins present, or absent. Aluminium accumulation demonstrated (rarely). Inulin recorded (Solanum, Gibbs 1974). C3. C3 recorded in Datura, Lycium, Lycopersicon, Nicotiana, Petunia, Physalis, Solanum. Anatomy non-C4 type (Cestrum, Datura, Lycium, Nicandra, Physalis, Solanum, Withania).
Geography, cytology. Temperate to tropical. Absent only from cold regions, but with greatest diversity in Central and South America. X = 7-12(+).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Solaniflorae; Solanales. Cronquist's Subclass Asteridae; Solanales. Takhtajan's Subclass Asteridae; Lamianae; Scrophulariales. Species 2000 (or more). Genera about 90; Atropa, Capsicum, Cestrum, Datura, Hyoscyamus, Lycium, Lycopersicon, Mandragora, Nicandra, Nicotiana, Petunia, Physalis, Salpiglossis, Schizanthus, Sclerophylax, Solandra, Solanum, etc.
Family reviewed by D'Arcy in Hawkes, Lester and Skelding (1979).
Economic uses, etc. Products include potato and eggplant (Solanum spp.), and tomato (Lycopersicon). Other edible fruits from Physalis (cape gooseberry, strawberry tomato, jamberberry, sugar cherry, chinese lantern etc., according to the species and variety), Cymphomandra (tamarillo), Capsicum (sweet and chilli peppers), etc. Most produce poisonous alkaloids, and some are commercially important in this connection (Nicotiana, Hyoscyamus, Datura). Many cultivated ornamentals, e.g. Petunia, Lycium, Solanum, Cestrum, Solandra.
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