Caprifoliaceae Juss.
Including Diervillaceae (Raf.) Pyck., Linnaeaceae (Raf.) Backlund, Loniceraceae von Vest, Viburnideae (Viburnaceae) Dum. (p.p.)
Excluding Carlemanniaceae, Sambucaceae, 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; 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 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. Plants hermaphrodite, or polygamomonoecious. 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 terminal 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.
Perianth with 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.
Seedling. Germination phanerocotylar.
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 (1998) Eudicot; core Eudicot; Asterid; Euasterid II; Dipsacales. Species about 330. Genera 12; Abelia, Diervilla, Dipelta, Heptacodium, Kolkwitzia, Leycesteria, Linnaea, Lonicera, Symphoricarpos, Triosteum, Weigela, Zabelia.
Analyses of rbcL sequences by Backlund and Bremer (1997) indicate 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) propose 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).
Illustrations. • Lonicera caprifolium. • Technical details (Lonicera, Symphoricarpos). • Technical details (Linnaea, Lonicera).
Quotations
Sleep thou, and I will wind thee in my arms,
So doth
the woodbine - the sweet honeysuckle -
Gently entwist
(‘Midsummer
Night’s Dream’, iv., 1)
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
