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The families of flowering plants

L. Watson and M.J. Dallwitz

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 without conspicuous aggregations of leaves. The trees pachycaul. Variously 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. Leaf development not ‘graminaceous’.

General anatomy. Plants with laticifers (these articulated, anastomosing). The laticifers in leaves and in stems (especially in the phloem, often extending elsewhere).

Leaf anatomy. The leaf lamina dorsiventral to centric. Hydathodes very commonly present. Stomata present; mainly confined to one surface (abaxial or sometimes adaxial), or on both surfaces (more often); anomocytic. Hairs present; seemingly always eglandular; unicellular (varying in form and frequency), or multicellular (in some Lobelioideae, otherwise recorded only in Campanumoea). Multicellular hairs when present, nearly always uniseriate. Complex hairs absent. Adaxial hypodermis present (often, in acicular leaves), or absent. Cystoliths commonly present (especially associated with hair bases, and at the leaf margins). Minor leaf veins without phloem transfer cells (Campanula, Jasione).

Axial (stem, wood) anatomy. Young stems with solid internodes, or with spongy internodes, or with hollow internodes. Secretory cavities commonly present; with latex. Cork cambium present (seldom), or absent; when at all active, initially deep-seated, or initially superficial. Nodes unilacunar. Primary vascular tissues seemingly usually in a cylinder, without separate bundles; collateral. Internal phloem present (occasionally, in the form of inverted, collateral medullary bundles), or absent. Cortical bundles present (rarely, e.g. in Campanula), or absent. Medullary bundles present (commonly, sometimes inversely orientated, sometimes consisting only of phloem), or absent. Secondary thickening developing from a conventional cambial ring.

The wood diffuse porous. The vessels small; radially paired, in radial multiples, and clustered. The vessel end-walls simple (usually), or scalariform, or scalariform and simple. The vessels without vestured pits. The axial xylem without tracheids; without fibre tracheids; with libriform fibres; rarely including septate fibres, or without septate fibres. The fibres without spiral thickening. The parenchyma where recorded, in Lobelioideae, scanty paratracheal (elsewhere absent or indistinct). ‘Included’ phloem absent. The wood not storied.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Pollination entomophilous; 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’; when aggregated, in cymes, in racemes, in spikes, and in umbels. The ultimate inflorescence units 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 (sometimes forming an elongate column around the style); 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 (sometimes terminating an androecial column, e.g Centropogon, Burmeistera), 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 (8 genera), or 3-celled (Cephalostigma and Isotoma), or 2-celled and 3-celled (with both conditions in Lobelia).

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, phytochemistry. C3. C3 physiology recorded directly in Campanula. Anatomy non-C4 type (Lobelia). Inulin recorded (very commonly). Cyanogenic (rarely), or not cyanogenic. Cynogenic constituents tyrosine-derived (with triglochinin). Polyacetylenes recorded. Alkaloids present (usually), or absent. Iridoids not detected. Saponins/sapogenins present (rarely), or absent. 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. Aluminium accumulation not found.

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 III core angiosperms; core eudicot; Superorder Asteranae; campanulid. APG IV Order Asterales.

Species 2000. Genera about 90; Adenophora, Apetahia, Astrocodon, Asyneuma, Azorina, Berenice, Brighamia, Burmeistera, Campanula, Canarina, Centropogon, Cephalostigma, 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.


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.)

Illustrations. • Le Maout and Decaisne: Campanula. • Le Maout and Decaisne: Lobelia, Centropogon. • cf. Wahlenbergia subulata, as Lightfootia: Thonner. • Azorina vidalii, as Campanula: Hook. Ic. Pl. 7–8 (1844). • Burmeistera virescens: Standley & Steyermark, Fl. of Guatemala 11 (1963). • Campanula fragilis: Bot. Reg. 1738, 1835. • Campanula glomerata: B. Ent., 1825). • Campanula loeflingii: Bot. Reg. 29 (19), 1843. • Campanula rotundifolia: Eng. Bot. 870 (1866). • Campanula trachelium: B. Ent. 627. • Campanula trachelium: Eng. Bot. 867 (1866). • Centropogon cordifolius: Standley & Steyermark, Fl. of Guatemala 11 (1963). • Codonopsis gracilis, C. inflata and C. javanica: Hooker’s Illustrations of Himalayan plants (1855). • Codonopsis ovata: as Glossocomia ovata, Bot. Reg. 3, 1842. • Downingia pulchella: as Clintonia, Bot. Reg. 1909 (1836). • Isotoma fluviatilis (as Lobelia): Hooker, Fl. Tasmaniae (1860). • Lobelia pedunculata and Lobelia surrepens: Hooker, Fl. Tasmaniae (1860). • Lobelia alsinoides, as trigona: Hook. Ic. Pl. 4 (1841). • Lobelia decurrens: Bot. Reg 1842 (1836). • Lobelia purpurea: Bot. Reg. 1325, 1830. • Lobelia urens: Eng. Bot. 862 (1866). • Michauxia laevigata: Bot. Reg. 1451, 1831. • Nemacladus ramosissimus: Emory, Rep. of U.S. and Mexican Boundary Survey, 2 (1859). • Pratia begonifolia: Bot. Reg. 1373, 1830. • Siphocampylus giganteus: Hook. Ic. Pl. 8 (1844). • Wahlenbergia saxicola: Hooker, Fl. Tasmaniae (1860). • Campanula, Wahlenbergia, Legousia (B. Ent. compilation, 1824–1835). • Lobelia dortmannia, Lobelia urens (Lobelioideae: B. Ent. compilation, 1824–35). • Physoplexis comosa, as Phyteuma: Bot. Mag. 106 (1880).

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 15th April 2018.’.