Campanulaceae Juss.
Including Ciliovallaceae Dulac, Cyananthaceae J.G. Agardh, Cyphiaceae DC., Cyphocarpaceae Miers, Jasionideae (Jasionaceae) Dum., Lobeliaceae R.Br., Nemacladaceae Nutt.
Excluding Pentaphragmataceae, Sphenocleaceae
Habit and leaf form. Herbs (mostly), or trees to shrubs (a few); laticiferous. The herbs mostly perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. The trees pachycaul. Hydrophytic to xerophytic; when hydrophytic, rooted. Leaves of hydrophytes submerged and emergent. Leaves usually alternate, or opposite (sometimes), or whorled (sometimes); petiolate, or subsessile; sheathing, or non-sheathing. Leaf sheaths with free margins. Leaves not gland-dotted; simple (usually), or compound (occasionally); epulvinate; when compound, pinnate. Lamina when simple, dissected, or entire; linear to orbicular; when simple-dissected, pinnatifid, or palmatifid. Leaves exstipulate. Lamina margins commonly crenate, or serrate, or dentate. Leaves without a persistent basal meristem.
General anatomy. Plants with laticifers (articulated, anastomosing). The laticifers in leaves and in stems (especially in the phloem, often extending elsewhere).
Leaf anatomy. Hydathodes very commonly present. Stomata present; mainly confined to one surface, or on both surfaces (commonly); anomocytic.
Lamina dorsiventral to centric. Cystoliths commonly present. Minor leaf veins without phloem transfer cells (Campanula, Jasione).
Stem anatomy. Secretory cavities commonly present; with latex. Cork cambium present, or absent (?); initially deep-seated, or superficial. Nodes unilacunar. Primary vascular tissue seemingly usually in a cylinder, without separate bundles. Cortical bundles present (rarely, e.g. in Campanula), or absent. Medullary bundles present (commonly), or absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent. Xylem without tracheids; without fibre tracheids; with libriform fibres. Vessel end-walls simple (usually), or scalariform, or scalariform and simple. Wood parenchyma absent or indistinct.
Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Entomophilous. Pollination mechanism conspicuously specialized (via modifications of the style, with sterile tissue covering the stigmas at anthesis. Active in most Lobelioideae, usually passive).
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes, in racemes, in spikes, and in umbels. The terminal inflorescence unit cymose, or racemose. Inflorescences scapiflorous, or not scapiflorous; terminal, or axillary; with involucral bracts, or without involucral bracts; pseudanthial, or not pseudanthial. Flowers medium-sized to large; regular to somewhat irregular; 5 merous; cyclic; tetracyclic.
Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous. Calyx 5; 1 whorled; polysepalous, or gamosepalous (depending on interpretation — the ‘tube’ nearly always being united with the ovary); basally appendaged (e.g. in Campanula, with adjoining pairs of sepals contributing to each appendage), or neither appendaged nor spurred; imbricate, or valvate. Epicalyx present (sometimes), or absent. Corolla 5; 1 whorled; gamopetalous (commonly), or polypetalous (e.g. Jasione); valvate; often campanulate; bilabiate, or regular; blue (predominantly), or white, or yellow, or red, or pink, or purple.
Androecium 5. Androecial members free of the perianth, or adnate (then low down on the corolla); free of one another, or coherent; 1 whorled. Androecium exclusively of fertile stamens. Stamens 5; inserted when epipetalous, near the base of the corolla tube; isomerous with the perianth; oppositisepalous; alternating with the corolla members. Anthers cohering, or separate from one another; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate; (2–)3–12 aperturate; colpate, or porate, or colporate (or colporoidate), or foraminate, or rugate (rarely); 2-celled, or 3-celled.
Gynoecium 2 carpelled, or 3 carpelled, or 5 carpelled. The pistil 2 celled, or 3 celled, or 5(–10) celled. Gynoecium syncarpous; synstylovarious; inferior (usually), or superior (rarely). Ovary 2 locular, or 3 locular, or 5(–10) locular. Locules secondarily divided by ‘false septa’ (occasionally), or without ‘false septa’. Styles 1. Stigmas 2, or 3, or 5 (as many as the carpels); wet type, or dry type; papillate, or non-papillate; Group II type and Group IV type. Placentation axile. Ovules 10–50 per locule (i.e. ‘many’); horizontal; non-arillate; anatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped, or hooked (and occasionally with filiform apparatus). Endosperm formation cellular. Endosperm haustoria present; chalazal and micropylar. Embryogeny solanad.
Fruit fleshy (rarely), or non-fleshy (nearly always); dehiscent, or indehiscent; a capsule (usually), or a berry. Capsules septicidal, or loculicidal, or valvular, or splitting irregularly (i.e. variously dehiscent). Seeds endospermic. Endosperm oily (rarely starchy). Seeds small; winged, or wingless. Seeds with starch (rarely), or without starch. Cotyledons 2. Embryo achlorophyllous (4/6); straight.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic (rarely), or not cyanogenic. Cynogenic constituents tyrosine-derived (with triglochinin). Polyacetylenes recorded. Alkaloids present (usually), or absent. Iridoids not detected. Proanthocyanidins present (rarely), or absent; when present, cyanidin (Centropogon). Flavonols present (rarely), or absent; when present, kaempferol and quercetin, or quercetin. Ellagic acid absent (11 species, 7 genera). Ursolic acid present, or absent. Saponins/sapogenins present (rarely), or absent. Aluminium accumulation not found. Inulin recorded (very commonly). C3. C3 physiology recorded directly in Campanula. Anatomy non-C4 type (Lobelia).
Geography, cytology. Holarctic, Paleotropical, Neotropical, Cape, Australian, and Antarctic. Frigid zone to tropical. Cosmopolitan, except tropical Africa. X = 6–17.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Asteriflorae; Campanulales. Cronquist’s Subclass Asteridae; Campanulales. APG (1998) Eudicot; core Eudicot; Asterid; Euasterid II; Asterales. Species 2000. Genera about 90; Adenophora, Apetahia, Astrocodon, Asyneuma, Azorina, Berenice, Brighamia, Burmeistera, Campanula, Canarina, Centropogon, Clermontia, Codonopsis, Craterocapsa, Cryptocodon, Cyananthus, Cyanea, Cylindrocarpa, Cyphia, Cyphocarpus, Delissea, Diastatea, Dielsantha, Downingia, Echinocodon, Edraianthus, Feeria, Githopsis, Grammatotheca, Gunillaea, Hanabusaya, Heterochaenia, Heterocodon, Heterotoma, Hippobroma, Homocodon, Howellia, Hypsela, Isotoma, Jasione, Legenere, Legousia, Leptocodon, Lightfootia, Lobelia, Lysipomia, Merciera, Michauxia, Microcodon, Monopsis, Musschia, Namacodon, Nemacladus, Nesocodon, Numaeacampa, Ostrowskia, Palmerella, Parishella, Peracarpa, Petromarula, Physoplexis, Phyteuma, Platycodon, Popoviocodonia, Porterella, Pratia, Prismatocarpus, Peudonemacladus, Rhigiophyllum, Roella, Rollandia, Ruthiella, Sclerotheca, Sergia, Siphocampylus, Siphocodon, Solenopsis, Symphyandra, Theilera, Trachelium, Treichelia, Trematolobelia, Trimeris, Triodanis, Unigenes, Wahlenbergia, Zeugandra.
Economic uses, etc. Numerous ornamentals from Lobelia, Wahlenbergia, Codonopsis, Jasione, etc., and more than 120 species of Campanula.
Illustrations. • Campanula, Wahlenbergia, Legousia. • Lobelia (Lobelioideae - Lobelia). • Campanula glomerata. • Technical details (Campanula). • Technical details (Lobelia, Centropogon). • Technical details Lightfootia).
Quotations
And there with hispid leaves and blooms
Of
darken’d sapphire, richly swinging,
The Bell-flower nettle-leaved
illumes
With azure light the woods; while bringing
Around it troops of
insect things,
With merry song and dancing wings
(Quoted by Ann Pratt,
‘Wild Flowers’ (1857), unattributed - Campanula trachelium)
When glowworm found in lanes remote,
Is murder’d for its shining
coat,
And put in flowers that Nature weaves
With hollow shapes and
silken leaves,
Such as the Canterbury Bell,
Serving for lamp or lantern
well
(John Clare, quoted by Ann Pratt, ‘Wild Flowers’ (1857) -
Campanula spp.)
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).
