The families of flowering plants
~ Caprifoliaceae sensu lato
Habit and leaf form. Herbs. Annual, or biennial, or perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves; sometimes tuberous. Heterophyllous, or not heterophyllous. Leaves opposite (usually), or whorled (rarely); herbaceous, or leathery; petiolate, or subsessile, or sessile; connate, or not connate; simple, or compound; when compound, pinnate. Lamina when simple dissected, or entire; when dissected, pinnatifid; pinnately veined; cross-venulate. Leaves exstipulate.
Leaf anatomy. The leaf lamina in different species variously dorsiventral, or bifacial, or centric (sometimes dorsiventral at the bases of the stems and isobilateral on the upper parts). Stomata present; mainly confined to one surface (sometimes), or on both surfaces (more often); anomocytic (usually), or anisocytic. Hairs present; eglandular and glandular (the former usually lignified, unicellular and sometimes raised on multicellular emergencies, the latter short- to long-stalked with multicellular heads). Lamina without secretory cavities. The mesophyll containing crystals, or without crystals. The crystals when present, usually druses. Main veins not accompanied by sclerenchyma, vertically transcurrent (sometimes, via thin-walled tissue), or embedded. Minor leaf veins with phloem transfer cells (Cephalaria, Dipsacus, Knautia, Pterocephalus, Scabiosa, Succisella).
Axial (stem, wood) anatomy. Young stems cylindrical, or tetragonal; often with hollow internodes. Secretory cavities absent. Cork cambium present; initially deep-seated (usually), or initially superficial. Nodes tri-lacunar, or multilacunar. Primary vascular tissues in a cylinder, without separate bundles (or the initial bundles soon linking); collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring (resulting in early linking of the initial vascular bundles by phloem and prosenchymatous elements).
The vessel end-walls simple (usually), or scalariform and simple. The vessels without vestured pits; with spiral thickening (oftem), or without spiral thickening. The axial xylem with tracheids. Included phloem absent.
Reproductive type, pollination. Plants hermaphrodite. Pollination entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in inflorescences; in heads (these dense, the receptacle usually hairy or scaly). The ultimate inflorescence units cymose. Inflorescences scapiflorous; dense, involucrate heads; with involucral bracts; pseudanthial. Flowers bracteolate (the two bracteoles supposedly joined to form the involucel); small; very irregular; zygomorphic. The floral irregularity involving the perianth and involving the androecium. Flowers cyclic; tetracyclic. Free hypanthium absent.
Perianth with distinct calyx and corolla (the calyx small, variously constituted); 8, or 10; 2 whorled; isomerous. Calyx 4 (by fusion of two members, cf. Veronica), or 5 (but often represented by five or up to ten pappus-like bristles); represented by bristles (often), or not represented by bristles; 1 whorled; polysepalous (of teeth or bristles), or partially gamosepalous (cupular, entire or variously divided); entire, or lobulate, or blunt-lobed, or toothed. Calyx lobes markedly shorter than the tube to markedly longer than the tube. Calyx regular; usually persistent. Epicalyx present (usually, supposedly formed from two fused bracteoles), or absent. Corolla 4 (by fusion of two members), or 5; 1 whorled; gamopetalous; imbricate; funnel-shaped, or tubular; unequal but not bilabiate, or bilabiate; white, or purple, or blue.
Androecium 4, or 2–3 (rarely). Androecial members adnate; all equal, or markedly unequal; free of one another, or coherent; when united, 2 adelphous; 1 whorled. Androecium exclusively of fertile stamens, or including staminodes. Staminodes if present, 1–2 (?); in the same series as the fertile stamens. Stamens 4, or 2–3; inserted in the throat of the corolla tube; didynamous (sometimes), or not didynamous, not tetradynamous; reduced in number relative to the adjacent perianth, or isomerous with the perianth; oppositisepalous; alternating with the corolla members. Anthers dorsifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with one middle layer; of the dicot type. Tapetum amoeboid. Pollen grains aperturate; 3 aperturate, or 4 aperturate; colpate, or porate; 3-celled.
Gynoecium 2 carpelled. Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium syncarpous (but pseudomonomerous); synstylovarious to eu-syncarpous; inferior. Ovary 1 locular. Gynoecium median. Epigynous disk present. Gynoecium non-stylate. Styles 1; apical. Stigmas 1–2; simple, or unequally 2 lobed; dry type; non-papillate; Group II type. Placentation apical. Ovules in the single cavity 1; pendulous; non-arillate; anatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating. Synergids pear-shaped. Hypostase present. Endosperm formation cellular. Embryogeny piperad (where known).
Fruit non-fleshy; indehiscent; achene-like. Dispersal unit the remains of the flower (with the achene enclosed in the epicalyx and the persistent calyx limb). Fruit 1 seeded. Seeds endospermic. Endosperm oily. Embryo well differentiated. Cotyledons 2. Embryo chlorophyllous (4/16); straight.
Seedling. Germination phanerocotylar.
Physiology, phytochemistry. C3. C3 physiology recorded directly in Dipsacus, Scabiosa. Inulin recorded (Cephalaria, Gibbs 1974). Not cyanogenic. Alkaloids present (commonly), or absent. Verbascosides not detected. Iridoids detected; Route I type (normal and seco). Saponins/sapogenins present (rarely), or absent. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (3 species, 3 genera). Aluminium accumulation not found.
Geography, cytology. Holarctic. Temperate and sub-tropical. Old World, chiefly North temperate Eurasia and tropical and southern Africa. X = 5–10. Supposed basic chromosome number of family: 9.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgrens Superorder Corniflorae; Dipsacales. Cronquists Subclass Asteridae; Dipsacales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Dipsacales (as a synonym of Caprifoliaceae).
Species 150. Genera 14; Cephalaria, Dipsacus, Knautia, Lomelosia, Pseudoscabiosa, Pterocephalidium, Pterocephalus, Pycnocomon, Scabiosa, Scabiopsis, Sixalix, Succisa, Succisella, Tremastelma.
General remarks. Comparison of this description with our compilation for Caprifoliaceae sensu stricto (q.v.) offers differences in 14 characters involving leaf anatomy, inflorescence and floral morphology, as well as in embryology and phytochemistry.
Economic uses, etc. Some ornamentals (Scabiosa, Cephalaria); teasels (Dipsacus) are sometimes noxious weeds, and the heads are used for fulling cloth.
Een the dew is
From the teazles jointed cup.
(John Clare 1820, Noon connate leaf bases of Dipsacus
Illustrations. • Technical details: Scabiosa, Dipsacus. • Technical details: Cephalaria (Thonner). • Dipsacus fullonum (B. Ent., 1838). • Dipsacus fullonum: Eng. Bot. 674 (1865). • Knautia arvensis (B. Ent., 1839). • Succisa, Scabiosa (B. Ent. compilation, 1824–35).
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: 13th March 2017. delta-intkey.com/angio’.