The families of flowering plants.                                                                                                                                                                

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~Caprifoliaceae, Adoxaceae sensu lato

ExcludingViburnaceae Dum.

Habit and leaf form. Small trees, or shrubs, or herbs (few); leptocaul. Mesophytic. Leaves deciduous; opposite; petiolate; foetid; compound; pinnate. Lamina pinnately veined; cross-venulate. Leaves stipulate (sometimes with four or more stipules side by side at a node), or exstipulate. Stipules when present, often scaly. Lamina margins serrate. Vegetative buds scaly. Leaves without a persistent basal meristem.

General anatomy. Plantswith ‘crystal sand’ (abundant).

Leaf anatomy. Stomata present; mostly anomocytic.

Stem anatomy. Cork cambium present; initially superficial. Nodes multilacunar (at least in S. niger with one lateral trace on each side of the node bifurcating, to be shared by the opposing leaves of the pair: a very peculiar phenomenon). Secondary thickening developing from a conventional cambial ring. Xylem with libriform fibres. Vessels without vestured pits. Wood partially storied (VP); parenchyma scanty paratracheal.

Reproductive type, pollination. Fertile flowershermaphrodite. Plants hermaphrodite. Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in cymes, in corymbs, and in panicles. The ultimate inflorescence unit cymose. Inflorescences terminal; large, repeatedly branched, compound, flat topped, umbelliform corymbs and thyrses. Flowers bracteolate (the pedicels articulate); small, or medium-sized; fragrant, or malodorous (according to opinion); regular; 3–5 merous; cyclic; tetracyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; 6–10; 2 whorled; isomerous. Calyx 3–5; 1 whorled; gamosepalous; regular; persistent; open in bud; with the median member posterior. Corolla 3–5; 1 whorled; gamopetalous; imbricate, or valvate; regular (rotate); white.

Androecium 3–5. Androecial members adnate (to the corolla); free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens 3–5; isomerous with the perianth; oppositisepalous; alternating with the corolla members. Anthers dehiscing via longitudinal slits; extrorse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate; 3 aperturate; colporate; 3-celled.

Gynoecium 3–5 carpelled. Carpels reduced in number relative to the perianth to isomerous with the perianth. The pistil 3–5 celled. Gynoecium syncarpous; synstylovarious; inferior. Ovary 3–5 locular. Epigynous disk present, or absent. Gynoecium more or less non-stylate. Stigmas 3–5; dry type; papillate; Group II type. Placentation axile, or apical. Ovules 1 per locule; pendulous; epitropous; with ventral raphe; anatropous; unitegmic; tenuinucellate. Embryo-sac development Adoxa-type (Adoxa type). Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped. Endosperm formation cellular.

Fruit fleshy; indehiscent; a drupe (berrylike). The drupes with separable pyrenes (with 3–5 one-seeded pyrenes). Fruit 3–5 seeded. Seeds endospermic. Endosperm oily. Embryo chlorophyllous (1/1); straight.

Physiology, biochemistry. Cyanogenic, or not cyanogenic (in different species). Iridoids detected; ‘Route I’ type (normal and seco). Proanthocyanidins absent. Flavonols present; kaempferol and quercetin. Ellagic acid absent. Arbutin absent. Ursolic acid present. C3. C3 physiology recorded directly in Sambucus.

Geography, cytology. Temperate to tropical. Widespread, but absent from Amazonia, Arabia, India, Western Australia and Pacific, and represented in Africa by only one species in the Eastern mountains.

Taxonomy.Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Corniflorae; Cornales. Cronquist’s Subclass Asteridae; Dipsacales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Dipsacales (as a synonym of Adoxaceae).

Species 40. Genera 1; only genus, Sambucus.

General remarks. RbcL sequence analyses by Backlund and Bremer (1997) imply close relationship between Adoxa, Sambucus and Viburnum; strongly supporting the taxonomic integrity of Bentham and Hooker’s tribe Caprifoliaceae-Sambuceae (= Adoxaceae sensu lato, e.g. Judd et al. 1984) but suggesting that these genera are relatively distant from Caprifoliaceae. Perhaps their true affinities (a bone of contention since the nineteenth century) have now been resolved.

Economic uses, etc. Edible ‘elderberry’.


  • Technical details: Sambucus. 
  • Sambucus nigra: Eng. Bot. 637 (1865). 
  • Sambucus nigra (Lindley).
  • Sambucus ebulus: Eng. Bot. 638 (1865).
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