~ Caprifoliaceae s. l.
Habit and leaf form. Herbs. Annual, or biennial, or perennial; with a basal aggregation of leaves, or with neither basal nor terminal 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. Stomata present; anomocytic (usually), or anisocytic.
Minor leaf veins with phloem transfer cells (Cephalaria, Dipsacus, Knautia, Pterocephalus, Scabiosa, Succisella).
Stem anatomy. Young stems cylindrical, or tetragonal. Cork cambium present; initially deep-seated (usually), or superficial. Nodes tri-lacunar, or multilacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent. Xylem with tracheids. Vessel end-walls simple, or scalariform and simple.
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 unit 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, biochemistry. Not cyanogenic. Alkaloids present (commonly), or absent. Iridoids detected; ‘Route I’ type (normal and seco). Verbascosides not detected. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (3 species, 3 genera). Saponins/sapogenins present (rarely), or absent. Aluminium accumulation not found. Inulin recorded (Cephalaria, Gibbs 1974). C3. C3 physiology recorded directly in Dipsacus, Scabiosa.
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. Dahlgren’s Superorder Corniflorae; Dipsacales. Cronquist’s 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.
Economic uses, etc. Some ornamentals (Scabiosa, Cephalaria); teasels (Dipsacus) are sometimes noxious weeds, and the heads are used for fulling cloth.
• Technical details: Scabiosa, Dipsacus.
• Technical details: Cephalaria (Thonner).
• Dipsacus fullonum (B. Ent.).
• Dipsacus fullonum: Eng. Bot. 674 (1865).
• Knautia arvensis (B. Ent.).
• Succisa, Scabiosa (B. Ent. compilation).
E’en the dew is parched up
From the teazle’s jointed cup.
(John Clare 1820, ‘Noon’ — connate leaf bases of Dipsacus