The families of flowering plants
Including Cestrineae (Cestraceae) Schlechtd., Salpiglossidaceae Hutch., Sclerophylacaceae Miers; excluding Duckeodendraceae, Goetzeaceae.
Habit and leaf form. Herbs, shrubs, trees, and lianas (often prickly); resinous, or not resinous. Normal plants (usually), or normal plants to switch-plants (occasionally). Annual, or biennial, or perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves, or with terminal aggregations of leaves (e.g. in mature Anthocercis viscosa). 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 (rarely very small and ericoid, e.g. in Fabiana imbricata); 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; leaf development not graminaceous. Domatia occurring in the family (known from 4 genera); manifested as hair tufts.
General anatomy. Plants with crystal sand (commonly), or without crystal sand.
Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial (occasionally isobilateral). Stomata mainly confined to one surface, or on both surfaces; anomocytic, or anisocytic, or diacytic. Hairs usually present (representing diverse forms, including complexly branched, tufted, stellate and peltate types: see illustration); eglandular and glandular; unicellular and multicellular. Multicellular hairs uniseriate and multiseriate; commonly branched. Complex hairs present. The mesophyll containing mucilage cells (rarely), or not containing mucilage cells. Minor leaf veins without phloem transfer cells (5 genera).
Axial (stem, wood) anatomy. Cork cambium present; initially deep-seated, or initially superficial. Nodes unilacunar (with 2 or 3 traces). Primary vascular tissues in a cylinder, without separate bundles; bicollateral. Internal phloem universally present (as strands, or a continuous ring). Secondary thickening developing from a conventional cambial ring (usually), or anomalous. The anomalous secondary thickening when present, via concentric cambia, or from a single cambial ring (?).
The wood ring porous to semi-ring porous, or diffuse porous. The vessel end-walls simple. The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem with tracheids, or without tracheids; with fibre tracheids (rarely), or without fibre tracheids; with libriform fibres, or without libriform fibres; including septate fibres (rarely), or without septate fibres. The parenchyma apotracheal, or paratracheal. Included phloem present (occasionally, e.g. Atropa belladonna rhizomes), or absent.
Reproductive type, pollination. Plants hermaphrodite (mostly), or monoecious, or andromonoecious, or dioecious (e.g., sometimes in Solanum and Symonanthus). Pollination entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in inflorescences. The ultimate inflorescence units when present, 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 irregularity (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, or 11–14; 2 whorled; isomerous, or anisomerous. Calyx (4–)5(–7); 1 whorled; gamosepalous. Calyx lobes markedly shorter than the tube to markedly longer than the tube. Calyx regular (usually), or unequal but not bilabiate; persistent; accrescent, or non-accrescent. Corolla (4–)5(–7); 1 whorled; gamopetalous. Corolla lobes markedly shorter than the tube to markedly longer than the tube. Corolla 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), or unequal but not bilabiate (sometimes).
Androecium 5 (usually), or 3–4 (rarely), or 6–7 (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); in the same series as the fertile stamens; representing when present, the posterior median member (Salpiglossis), or the posterior median member and the posterior-lateral pair (Schizanthus). Fertile stamens representing the posterior median member, the posterior-lateral pair, and the anterior-lateral pair (mostly), or the posterior-lateral pair and the anterior-lateral pair (Salpiglossis), or the anterior-lateral pair (Schizanthus). Stamens 5 (in all but Salpiglossideae), or 2, or 4; inserted near the base of the corolla tube, or midway down the corolla tube, or in the throat of the corolla tube; 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; alternating with the corolla members. 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; dorsifixed, or basifixed; 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 (recorded in 15 genera).
Gynoecium 2 carpelled. Carpels reduced in number relative to the perianth. The pistil 2 celled (usually), or 3–5 celled (Nicandreae and Datureae). Gynoecium syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 2 locular (but sometimes complicated by secondary divisions). Locules secondarily divided by false septa (Nicandreae and Datureae), or without false septa. 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; semi-cylindric. 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, phytochemistry. C3. C3 physiology recorded directly in Datura, Lycium, Lycopersicon, Nicotiana, Petunia, Physalis, Solanum. Anatomy non-C4 type (Cestrum, Datura, Lycium, Nicandra, Physalis, Solanum, Withania). Sugars transported as sucrose (in Datura, Solanum). Inulin recorded (Solanum, Gibbs 1974). Cyanogenic, or not cyanogenic. Alkaloids present (mostly), or absent. Anthraquinones detected (Fabiana); polyacetate derived. Arbutin absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins absent. Flavonols present, or absent; kaempferol and quercetin (mostly). Ellagic acid absent (25 species, 14 genera). Ursolic acid present. Aluminium accumulation demonstrated (rarely). Sieve-tube plastids S-type.
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. Dahlgrens Superorder Solaniflorae; Solanales. Cronquists Subclass Asteridae; Solanales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; lamiid; Order Solanales.
Species 2000 (or more). Genera about 95; Acnistus, Anisodus, Anthocercis, Anthotroche, Archiphysalis, Althenaea, Atropa, Atropanthe, Benthamiella, Bouchetia, Brachistus, Browallia, Brugmansia, Brunfelsia, Calibrachoa, Capsicum, Cestrum, Chamaesaracha, Combera, Crenidium, Cuatresia, Cyphanthera, Cyphomandra, Datura, Deprea, Discopodium, Duboisia, Dunalia, Dyssochroma, Ectozma, Exodeconus, Fabiana, Grabowskia, Grammosolen, Hawkesiophyton, Heteranthia, Hunzikeria, Hyoscyamus, Iochroma, Jaborosa, Jaltomata, Juanulloa, Latua, Leptoglossis, Leucophysalis, Lycianthes, Lycium, Lycopersicon, Mandragora, Margaranthus, Markea, Melananthus, Mellissia, Merinthopodium, Metternichia, Nectouxia, Nicandra, Nicotiana, Nierembergia, Nothocestrum, Oryctes, Pantacantha, Parabouchetia, Pauia, Petunia, Phrodus, Physalis, Physochlaina, Plowmania, Protoschwenckia, Przewalskia, Quincula, Rahowardiana, Reyesia, Salpichroa, Salpiglossis, Saracha, Schizanthus, Schultesianthus, Schwenckia, Sclerophylax , Scopolia, Sessea, Sesseopsis, Solandra, Solanum, Streptosolen, Symonanthus, Trianaea, Triguera, Tubocapsicum, Vassobia, Vestia, Withania, Witheringia.
General remarks. Family reviewed by DArcy 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.
And shrieks, like
mandrakes, torn out of the earth,
That living mortals, hearing them, go mad
(Romeo and Juliet, iv.,3)
Atropa, too, that, as the
Shows her black fruit to tempt and to betray
(Charlotte Smith, quoted by Ann Pratt, Wild Flowers (1857))
The pipe, with
solemn interposing puff,
Makes half a sentence at a time enough
(William Cowper, Tobacco)
When potatoes, leaves, or haulms are green,
To livestock must they neer be gien
(ancient anon, re. solanine poisoning.)
Illustrations. • Technical details: Atropa, Mandragora, Solanum. • Technical details: Cestrum. • Technical details: Datura. • Technical details: Discopodium. • Technical details: Hyoscyamus. • Technical details: Jaborosa. • Technical details: Lycium. • Technical details: Nicotiana. • Anthocercis viscosa: habit (photo). • Anthocercis viscosa: flower (photo). • Anthocercis littorea (photo). • Anthocercis littorea (close-up photo). • Anthocercis littorea: Bot. Reg. 212, 1817. • Atropa belladonna (B. Ent.). • Atropa belladonna: Eng. Bot. 934 (1866). • Atropa belladonna (J. E. Sowerby, 1861). • Browallia americana: Bot. Mag. 34, 1788. • Browallia grandiflora: Bot. Reg. 1384, 1830. • Cestrum purpureum: as Habrothamnus purpureus, Bot. Reg. 1844, 43. • Cyphanthera tasmanica (as Anthocercis): Hooker, Fl. Tasmaniae (1860). • Datura (Brugmansia) sanguinea: as Brugmansia bicolor, Bot. Reg. 1739, 1835. • Datura stramonium (J. E. Sowerby, 1861). • Fabiana imbricata: Bot. Reg. 1839, 59. • Grabowskia boerhaaviaefolia: Bot. Reg. 1985, 1837. • Hyoscyamus niger (B. Ent.). • Hyocyamus niger (J. E. Sowerby, 1861). • Lycium barbarum: Eng. Bot. 933 (1866). • Solanum dulcamara (B. Ent.). • Solanum dulcamara (J. E. Sowerby, 1861). • Solanum nigrum (B. Ent.). • Solanum nigrum (J. E. Sowerby, 1861). • Solanum uncinellum Lindl.: Bot. Reg. xxvi, 15 (1840). • Leaf hairs of Duboisia, Lycium, Melananthus, Sessea, Solanum,: Solereder, 1908.
We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.
Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 19th October 2016. delta-intkey.com’.