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.
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 asteraceouscapitula. 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 1mm wide and long in some species of Wahlenbergia. At the other extreme, it reaches a width of 15cm 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)
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.
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.
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. JSTOR3298542.
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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
↑ 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. S2CID84676862.
↑ "Campanulaceae Juss."Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2022-04-24.
↑ Friis, Else Marie; Crane, Peter R.; Pedersen, Kaj Raunsgaard (August 2011). Early Flowers and Angiosperm Evolution. Cambridge University Press. ISBN9780521592833.
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