Including Viburnaceae(`-nideae') Dum. (p.p.)
Excluding Carlemanniaceae, Sambucaceae
Habit and leaf form. Shrubs, or trees (small), or herbs (rarely), or lianas. Self supporting, or climbing; when climbing, stem twiners. Mesophytic. Leaves evergreen, or deciduous; small to medium-sized; opposite (usually), or whorled; petiolate; connate (occasionally), or not connate; simple; epulvinate. Lamina dissected (rarely), or entire; when lobed, pinnatifid; pinnately veined; cross-venulate. Leaves stipulate, or exstipulate; without a persistent basal meristem. Domatia recorded (from two genera); represented by pits, or pockets, or hair tufts.
Leaf anatomy. Minor leaf veins without phloem transfer cells (7 genera).
Stem anatomy. Cork cambium present; initially deep-seated, or superficial. Nodes unilacunar (rarely), or tri-lacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. `Included' phloem absent. Xylem with tracheids; with vessels. Vessels without vestured pits. Wood parenchyma apotracheal, or paratracheal. Pith with diaphragms, or without diaphragms.
Reproductive type, pollination. Hermaphrodite, or polygamomonoecious. Entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in `inflorescences'. The terminal inflorescence unit cymose. Inflorescences axillary. Flowers usually bracteolate; often fragrant; regular to very irregular. The floral asymmetry involving the perianth, or involving the perianth and involving the androecium. Flowers 4-5 merous; cyclic; tetracyclic. Free hypanthium absent.
Perianth with distinct calyx and corolla; (6-)8-10; 2 whorled; isomerous. Calyx (2-)4-5; 1 whorled; gamosepalous (usually), or polysepalous (Weigela); regular; persistent; imbricate, or open in bud; with the odd member posterior. Corolla 4, or 5; 1 whorled; gamopetalous; imbricate; regular, or unequal but not bilabiate, or bilabiate; white, or yellow, or red, or pink, or purple.
Androecium (2-)4, or 5. Androecial members adnate (epipetalous); all equal, or markedly unequal (sometimes didynamous); free of one another; 1 - whorled. Androecium exclusively of fertile stamens. Stamens (2-)4, or 5; isomerous with the perianth; oppositisepalous. Anthers dorsifixed; dehiscing via longitudinal slits; introrse; bilocular; 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, or glandular. Pollen grains aperturate; usually 3 - aperturate; colpate, or porate, or colporate (or colporoidate), or zoniaperturate (sometimes zonorate in Abelia); 3-celled.
Gynoecium 2-5-8; syncarpous; synstylovarious, or eu-syncarpous; inferior. Ovary 2-8 locular. Styles 1; apical. Stigmas 1; capitate; dry type; papillate; Group III type. Placentation axile. Ovules 1-50 per locule (to `many'); pendulous; with dorsal raphe; anatropous; unitegmic; tenuinucellate (usually), or crassinucellate (Viburnum). Embryo-sac development Polygonum-type, or Allium-type, or Adoxa-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped. Endosperm formation cellular. Embryogeny asterad (and other types?).
Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry, or a drupe. Fruits from adjoining flowers not aggregated into compound fruits. Seeds endospermic. Endosperm not ruminate, or ruminate; oily. Cotyledons 2. Embryo achlorophyllous (5/16); straight.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Cynogenic constituents phenylalanine-derived. Alkaloids present, or absent. Iridoids recorded; seco-compounds. Proanthocyanidins present, or absent; when present, cyanidin. Flavonols present (mostly), or absent (Abelia); kaempferol and quercetin (mostly), or quercetin. Ellagic acid absent (11 species, 8 genera). Arbutin absent. Aluminium accumulation not found. C3. C3 recorded in Lonicera, Symphoricarpos, Viburnum. Anatomy non-C4 type (Symphoricarpos).
Geography, cytology. Temperate, or sub-tropical to tropical (at altitude). Widespread, but mostly North temperate and tropical mountains - missing from most of Africa. X = 8 or 9(-12).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Corniflorae; Dipsacales. Cronquist's Subclass Asteridae; Dipsacales. Takhtajan's Subclass Asteridae; Gentiananae; Dipsacales. Species 450. Genera 13; Abelia, Diervilla, Dipelta, Heptacodium, Kolkwitzia, Leycesteria, Linnaea, Lonicera, Symphoricarpos, Triosteum, Viburnum, Weigela, Zabelia.
Economic uses, etc. Cultivated ornamental shrubs and vines from Lonicera, Viburnum, Symphoricarpos, Abelia, Leycesteria, Linnaea, Kolkwitzia; noxious weeds (Lonicera japonica).
Illustrations. capri480.gif capri532.gif capri481.gif capri767.gif
Additional, to be intercalated. Lonicera twining clockwise.
