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Campanulaceae
	Campanula cespitosa
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Asterales
Family:	Campanulaceae; Juss.
Genera
	See text

The family Campanulaceae (also bellflower family), of the order Asterales, contains nearly 2400 species in 84 genera of herbaceous plants, shrubs, and rarely small trees, often with milky sap.^[2] Among them are several familiar garden plants belonging to the genera Campanula (bellflower), Lobelia, and Platycodon (balloonflower). Campanula rapunculus (rampion or r. bellflower) and Codonopsis lanceolata are eaten as vegetables. Lobelia inflata (indian tobacco), L. siphilitica and L. tupa (devil's tobacco) and others have been used as medicinal plants. Campanula rapunculoides (creeping bellflower) may be a troublesome weed, particularly in gardens, while Legousia spp. may occur in arable fields.

Most current classifications include the segregate family Lobeliaceae in Campanulaceae as subfamily Lobelioideae. A third subfamily, Cyphioideae, includes the genus Cyphia, and sometimes also the genera Cyphocarpus, Nemacladus, Parishella and Pseudonemacladus. Alternatively, the last three genera are placed in Nemacladoideae, while Cyphocarpus is placed in its own subfamily, Cyphocarpoideae.

This family is almost cosmopolitan, occurring on all continents except Antarctica. In addition, species of the family are native to many remote oceanic islands and archipelagos. Hawaii is particularly rich, with well over 100 endemic species of Hawaiian lobelioids. Continental areas with high diversity are South Africa, California and the northern Andes.

Habitats range from extreme deserts to rainforests and lakes, from the tropics to the high Arctic (Campanula uniflora), and from sea cliffs to high alpine habitats.

Description

Although most Campanulaceae are perennial herbs (sometimes climbing, as in Codonopsis), there is also a large number of annuals e.g. species of Legousia. Isotoma hypocrateriformis is a succulent annual from Australia's dry interior. There are also biennials, e.g. the commonly cultivated ornamental Campanula medium (Canterbury bells). Many perennial campanuloids grow in rock-crevices, such as Musschia aurea (Madeira) and Petromarula pinnata (Crete). Some lobelioids also grow on rocks, e.g. the peculiar perennial stem succulent Brighamia rockii in Hawaii. Insular and tropical montane species in particular are often more or less woody and may bear the leaves in a dense rosette. When, in addition, the plant is unbranched, the result may be a palm- or treefern-like habit, as in species of the hawaiian genus Cyanea, which comprises the tallest of Campanulaceae, C. leptostegia (up to 14 m). Lysipomia are minute cushion plants of the high Andes, while gigant rosette-forming lobelias (e.g., Lobelia deckenii) are a characteristic component of the vegetation in the alpine zone on the tropical African volcanoes. In the Himalaya Campanula modesta and Cyananthus microphyllus reach even higher, probably setting the altitudinal record for the family at 4800 m. Several species are associated with freshwater, such as Lobelia dortmanna, an isoetid common in oligotrophic lakes in the boreal zone of North America and Europe, and Howellia aquatilis, an elodeid growing in ponds in SW North America.

There is usually abundant, white latex, but occasionally the exudate is clear and/or very sparse, as in Jasione.

Tubers occur in several genera, e.g. Cyphia.

Leaves are often alternate, more rarely opposite (e. g. Codonopsis) or whorled (Ostrowskia). They are simple (Petromarula one of very few exceptions) entire (repeatedly divided in spp. of Cyanea), but often with dentate margin. Stipules are absent.

Inflorescences are quite diverse, including both cymose and racemose types. In Jasione they are strongly condensed and resemble asteraceous capitula. In a few species, e. g. Cyananthus lobatus, flowers are solitary.

Flowers are bisexual (dioecious in Dialypetalum) and protandrous. Petals are fused into a corolla with 3 to 8 lobes. It may be bell- or star-shaped in subfamily Campanuloideae, while tubular and bilaterally symmetric in most Lobelioideae. Blue of various shades is the most common petal colour, but purple, red, pink, orange, yellow, white, and green also occur. The corolla may be down to 1 mm wide and long in some species of Wahlenbergia. At the other extreme, it reaches a width of 15 cm in Ostrowskia.

Stamens are equal in number to, and alternating with the petals. Anthers may be fused into a tube, as in all species of Lobelioideae and some Campanuloideae (e.g. Symphyandra)

Within the family pollen grains are often tricolporate, less commonly triporate, tricolpate, or pantoporate.

Carpel number is usually 2, 3 or 5 (8 in Ostrowskia), and corresponds to the number of stigmatic lobes.

The style is in various ways involved in the "presentation" of the pollen, as in several other families of the order Asterales. In Lobelioideae, pollen is, already in the bud stage, released into the tube formed by the anthers. During flowering, it is pushed up by the elongating style and "presented" to visiting pollinators at the apex of the tube, a mechanism described as a pollen pump. The style eventually protrudes through the anther tube, and becomes receptive to pollen. In Campanuloideae, the pollen is instead packed between hairs on the style, gradually being released as the hairs invaginate. Subsequently, the stigmatic lobes unfold, and become receptive.

Bees and birds (particularly hummingbirds and hawaiian honeycreepers) are probably the most common pollinators of Campanulaceae. A few confirmed and many probable cases of bat-pollination are known, particularly in the genus Burmeistera. Brighamia and Hippobroma have pale or white flowers with a long-tubed corolla, and are pollinated by hawkmoths. Pollination by lizards has been reported for Musschia aurea and Nesocodon mauritianus.

The ovary is usually inferior or, in some species, semi-inferior. Very rarely is it completely superior (e.g. Cyananthus). In Campanumoea javanica, calyx and corolla diverge from the ovary at different levels.

Berries are a common fruit-type in Lobelioideae (Burmeistera, Clermontia, Centropogon, Cyanea etc.), whilst rare in Campanuloideae (Canarina being one of few examples). Capsules, with very varying modes of dehiscence, are otherwise the predominating fruit type in the family.

Seeds are mostly small (<2 mm) and numerous.

Subfamilies and genera

95 genera are accepted.^[3] The Angiosperm Phylogeny Website divides the family into five subfamilies.^[4]

Campanuloideae

Adenophora Fisch – Europe and Asia
Asyneuma Griseb. & Schenk – southern Europe and Asia
Berenice Tul. – Réunion
Campanula L. (synonyms Azorina Feer and Theodorovia Kolak.) – mostly northern hemisphere
Campanulastrum Small
Canarina L. – Canary Islands and East Africa
Codonopsis Wall. – eastern Asia
Craterocapsa Hilliard & B.L.Burtt – South Africa
Cryptocodon Fed. – C Asia
Cyananthus Wall. ex Benth. – E Asia
Cyclocodon Griff. ex Hook.f. & Thomson – tropical and subtropical Asia and New Guinea
Cylindrocarpa Regel – Central Asia
Eastwoodiella Morin – California
Echinocodon D.Y.Hong – China
Edraianthus A.DC. – SE Europe and W Asia
Favratia Feer – Alps
Feeria Buser – Morocco
× Fockeanthus H.R.Wehrh. – central Europe
Githopsis Nutt. – W N America
Gunillaea Thulin – Tropical Africa and Madagascar
Hanabusaya Nakai – Korea
Heterochaenia A.DC. – Réunion
Heterocodon Nutt. – SW N America
Himalacodon D.Y.Hong & Qiang Wang – Himalayas
Homocodon D.Y.Hong – China
Jasione L. – Europe and SW Asia
Kericodon Cupido – Cape Provinces
Legousia Durande – Europe and N Africa
Melanocalyx (Fed.) Morin – subarctic Eurasia and North America and Rocky Mountains
Merciera A.DC. – South Africa
Michauxia L'Hér. – Middle East
Microcodon A.DC. – South Africa
Muehlbergella Feer
Musschia Dumort. – Madeira
Namacodon Thulin – southwestern Africa
Nesocodon Thulin – Mauritius
Ostrowskia Regel – Central Asia
Palustricodon Morin – central and eastern North America
Pankycodon D.Y.Hong & X.T.Ma – Himalayas
Peracarpa Hook.f. & Thomson – Southeast Asia
Petromarula Vent. ex R.Hedw. – Crete
Physoplexis Schur – Alps
Phyteuma L. – Europe and Asia
Platycodon A.DC. – eastern Asia
Poolea Morin – Texas
Prismatocarpus L'Hér. – Southern Africa
Protocodon Morin – Florida
Pseudocodon D.Y.Hong & H.Sun
Ravenella Morin – California
Rhigiophyllum Hochst. – South Africa
Roella L. – South Africa
Rotanthella Morin – Florida
Sergia Fed. – C Asia
Siphocodon Turcz.- South Africa
Smithiastrum Morin – California and Oregon
Theilera E.Phillips – South Africa
Trachelium Tourn. ex L. – SE Europe, Middle East and C Asia
Treichelia Vatke – South Africa
Triodanis Raf. – Americas and southern Europe
Wahlenbergia Serra, M.B.Crespo & Lammers (synonym Hesperocodon Eddie & Cupido) – mostly Southern Hemisphere
Zeugandra P.H.Davis – Iran

Lobelioideae

Brighamia A.Gray – Hawaii
Burmeistera H.Karst. & Triana – N Andes and C America
Centropogon C.Presl. – Neotropics
Clermontia Gaudich. – Hawaii
Cyanea Gaudich. – Hawaii
Delissea Gaudich. – Hawaii
Dialypetalum Benth. – Madagascar
Diastatea Scheidw. – Neotropics
Dielsantha E.Wimm. – Tropical Africa
Downingia Torr. – W N America and S S America
Grammatotheca C.Presl. – South Africa
Heterotoma Zucc. – Mexico
Hippobroma G.Don – W Indies
Howellia A.Gray – SW N America
Isotoma (R.Br.) Lindl. – Australia
Legenere McVaugh – California
Lithotoma E.B.Knox – Australia
Lobelia Plum. ex L. (synonyms include Hypsela, Laurentia, Pratia) – cosmopolitan
Lysipomia Kunth – Andes
Monopsis Salisb. – tropical and southern Africa and Comoros
Palmerella A.Gray – California
Porterella Torr. – SW N America
Ruthiella Steenis – New Guinea
Sclerotheca A.DC. (synonym Apetahia Baill.) – Society Islands
Siphocampylus Pohl – Neotropics
Solenopsis C.Presl – S Europe and N Africa
Trematolobelia Zahlbr. ex Rock – Hawaii
Trimeris C.Presl – St. Helena
Unigenes E.Wimm. – South Africa
Wimmeranthus Rzed. – southwestern Mexico
Wimmerella Serra, M.B.Crespo & Lammers – South Africa

Cyphioideae

Cyphia P.J.Bergius – Africa

Cyphocarpoideae

Cyphocarpus Miers – northern Chile

Nemacladoideae

Nemacladus Nutt. (syn. Parishella) – SW N America
Pseudonemacladus McVaugh – Mexico

Fossil record

The earliest known occurrence of Campanulaceae pollen is from Oligocene strata.^[5] Earliest Campanulaceae macrofossils dated, are seeds of †Campanula paleopyramidalis from 17-16 million years old Miocene deposits in the Nowy Sacz, Carpathians, Poland. It is a close relative of the extant Campanula pyramidalis.^[6]^[7]

Chemical compounds

Members of subfamily Lobelioideae contain the alkaloid lobeline. The principal storage carbohydrate of Campanulaceae is inulin, a fructan also occurring in the related Asteraceae.

Literature

Lammers, T.G. (2007). World Checklist and Bibliography of Campanulaceae. Richmond, Surrey, United Kingdom: Royal Botanic Gardens, Kew.
Fedorov, A.; Kovanda, M. (1976). "Campanulaceae". In T.G. Tutin; V.H. Heywood; N.A. Burges; D.M. Moore; D.H. Valentine; S.M. Walters; D.A. Webb (eds.). Flora Europaea. Cambridge University Press. pp. 74–93.
Borsch, T.; Korotkova, N.; Raus, T.; Lobin, W.; Loehne, C. (2009). "The petD group II intron as a genus and species level marker: Utility for tree inference and species identification in the diverse genus Campanula (Campanulaceae)". Willdenowia. 39: 7–33. doi:10.3372/wi.39.39101.
Roquet, C.; Sáez, L.; Aldasoro, J. J.; Alfonso, S.; Alarcón, M. L.; Garcia-Jacas, N. (2008). "Natural delineation, molecular phylogeny and floral evolution in Campanula". Systematic Botany. 33: 203–217. doi:10.1600/036364408783887465. S2CID 86780466.
Cosner, M. E.; Raubeson, L. A.; Jansen, R. K. (2007). "Chloroplast DNA rearrangements in Campanulaceae: phylogenetic utility of highly rearranged genomes". BMC Evolutionary Biology. 4 (27): 1–17. doi:10.1186/1471-2148-4-27. PMC 516026. PMID 15324459.
Eddie, W. M. M.; Shulkina, T.; Gaskin, J.; Haberle, R. C.; Jansen, R. K. (2003). "Phylogeny of Campanulaceae s. str. inferred from ITS sequences of nuclear ribosomal DNA". Annals of the Missouri Botanical Garden. 90 (4): 554–575. doi:10.2307/3298542. JSTOR 3298542.

References

^ Angiosperm Phylogeny Group (2009), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III", Botanical Journal of the Linnean Society, 161 (2): 105–121, doi:10.1111/j.1095-8339.2009.00996.x, hdl:10654/18083
^ Lammers, Thomas (2011). "Revision of the Infrageneric Classification of Lobelia L. (Campanulaceae: Lobelioideae)". Annals of the Missouri Botanical Garden. 98: 37–62. doi:10.3417/2007150. S2CID 84676862.
^ "Campanulaceae Juss". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 23 August 2024.
^ Stevens, P.F. (2001 onwards). "Campanulaceae". Angiosperm Phylogeny Website. Retrieved 2022-04-23.
^ Friis, Else Marie; Crane, Peter R.; Pedersen, Kaj Raunsgaard (August 2011). Early Flowers and Angiosperm Evolution. Cambridge University Press. ISBN 9780521592833.
^ [1]^{[dead link‍]}
^ Lagomarsino, L. P.; Condamine, F. L.; Antonelli, A; Mulch, A; Davis, C. C. (2016). "The abiotic and biotic drivers of rapid diversification in Andean bellflowers (Campanulaceae)" (PDF). New Phytologist. 210 (4): 1430–1442. doi:10.1111/nph.13920. PMC 4950005. PMID 26990796.