The families of flowering plants.
IncludingDiervillaceae (Raf.) Pyck., Linnaeaceae (Raf.) Backlund, Loniceraceae von Vest, Viburnideae (Viburnaceae) Dum. (p.p.)Excluding Carlemanniaceae,Dipsacaceae, Morinaceae, Sambucaceae, Valerianaceae,Viburnaceae
Habit and leaf form. Shrubs, or trees (small), or herbs (rarely), or lianas. Self supporting, or climbing; when climbing, stem twiners; Lonicera twining clockwise. Mesophytic. Leaves evergreen, or deciduous; small to medium-sized; opposite (usually), or whorled; ‘herbaceous’ (mostly), or leathery; petiolate; connate (occasionally), or not connate; simple; epulvinate. Lamina dissected (exemplified in Leycesteria), or entire; when lobed, pinnatifid; pinnately veined; cross-venulate. Leaves stipulate (sometimes large, e.g. Pentapyxis, = Leycesteria), or exstipulate; without a persistent basal meristem. Domatia occurring in the family (from two genera); manifested as 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. Plants hermaphrodite, or polygamomonoecious. Pollination entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely, Triostem), or aggregated in ‘inflorescences’; if solitary, axillary; in cymes, or in racemes, or in spikes, or in heads, or in verticils. The ultimate inflorescence unit cymose (basically, but the flowers are commonly in axillary pairs, and in Lonicera the members of each pair are sometimes basally congenitally fused). Inflorescences terminal, or axillary. Flowers usually bracteolate; often fragrant; nearly always somewhat irregular to very irregular (except in Pentapyxis = Leycesteria p.p. and a few species of Lonicera); nearly always at least slightly zygomorphic. The floral irregularity involving the perianth (often only the corolla), or involving the perianth and involving the androecium. Flowers 4–5 merous; cyclic; tetracyclic. Free hypanthium absent.
Perianthwith distinct calyx and corolla; 8–10; 2 whorled; isomerous. Calyx (2–)4, or 5; 1 whorled; gamosepalous (usually), or polysepalous (Weigela); unequal but not bilabiate, or regular; non-fleshy; persistent, or not persistent; non-accrescent; imbricate, or open in bud; with the median member posterior. Corolla 4, or 5; 1 whorled; gamopetalous; imbricate; campanulate, or funnel-shaped, or tubular; unequal but not bilabiate, or bilabiate (very rarely equal); white, or yellow, or red, or pink, or purple (or various combinations).
Androecium (2–)4, or 5. Androecial members adnate (epipetalous); all equal, or markedly unequal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens (2–)4, or 5; inserted near the base of the corolla tube, or midway down the corolla tube, or in the throat of the corolla tube; didynamous, or not didynamous, not tetradynamous; isomerous with the perianth; oppositisepalous. Anthers separate from one another; 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 carpelled. Carpels reduced in number relative to the perianth, or isomerous with the perianth, or increased in number relative to the perianth. The pistil 2–5(–8) celled. Gynoecium syncarpous; eu-syncarpous;inferior. Ovary 2–8 locular. Epigynous disk present, or absent. Gynoecium stylate (elongate). Styles 1; apical. Stigmas 1; 1–5 lobed; capitate (usually), or truncate; wet type; papillate; Group III type. Placentation axile to apical. Ovules 1–50 per locule (to ‘many’); pendulous;apotropous; with dorsal raphe; often biseriate; non-arillate; anatropous; unitegmic; tenuinucellate. 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 achene-like, or a berry (the fused pairs of flowers of some Lonicera species resulting in compound berries), or a drupe (e.g. Symphoricarpos, Triosteum).Capsules valvular (Weigela). The drupes with separable pyrenes. Fruit 5–50 seeded (to ‘many’). Seeds endospermic. Endosperm not ruminate; oily. Cotyledons 2. Embryo achlorophyllous (5/16); straight.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Cynogenic constituents phenylalanine-derived. Alkaloids present, or absent. Iridoids detected; ‘Route I’ type (normal and seco). Proanthocyanidins present, or absent; when present, cyanidin. Flavonols present (mostly), or absent (Abelia); kaempferol and quercetin (mostly), or quercetin. Ellagic acid absent (11 species, 7 genera). Arbutin absent. Aluminium accumulation not found. C3. C3 physiology recorded directly in Lonicera, Symphoricarpos. 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. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Dipsacales.
Species about 330. Genera 12; Abelia, Diervilla, Dipelta, Heptacodium,Kolkwitzia, Leycesteria, Linnaea, Lonicera, Symphoricarpos,Triosteum, Weigela, Zabelia.
General remarks. Analyses of rbcL sequences by Backlund and Bremer (1997) indicated that Viburnum should be removed from Caprifoliaceae; that it is closely related to Adoxa and Sambucus; and that these three genera (i.e., the tribe Sambuceae of the Bentham and Hooker Caprifoliaceae, 1876) are related to Apiales or Cornales rather than to Dipsacales. Backlund and Pyck (1998) proposed Diervillaceae (Diervilla, Weigela and Linnaeaceae (Abelia, Dipelta,Kolkwitzia, Linnaea, Zabelia) as families separated from but closely related to Caprifoliaceae sensu stricto, without adequately revising the family descriptions.
Economic uses, etc. Cultivated ornamental shrubs and vines from Lonicera, Symphoricarpos, Abelia,Leycesteria, Linnaea, Kolkwitzia; noxious weeds (Lonicera japonica).
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