Campanulaceae

Campanulaceae
Campanula cespitosa
Scientific classification
Kingdom:Plantae
Clade:Angiosperms
Clade:Eudicots
Clade:Asterids
Order:Asterales
Family:Campanulaceae
Juss.[1]
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) is an almost forgotten vegetable.

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, such as Hawaii (110 species), Juan Fernandez Islands (4 spp.), Madeira (4 spp.), the Mascarenes (5 spp.), Saint Helena (5 spp.) and Tahiti (5 spp.) Continental areas with high diversity are Greece and the Balkans, Caucasia, eastern Himalaya, the northern Andes, South Africa and California - northern Mexico.

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.

Most current classifications include the segregate family Lobeliaceae in Campanulaceae as subfamily Lobelioideae. The genera Cyphia, Cyphocarpus, Nemacladus, Parishella and Pseudonemacladus are treated in one to three separate subfamilies.

Description

Although most Campanulaceae are perennial herbs, there is also a large number of annuals such as Legousia, comprising mostly field weeds, and Cyphocarpus, ephemerals of the Atacama desert. Isotoma hypocrateriformis is a succulent annual from Australia's dry interior. There are also biennials, e.g. Campanula medium. Insular and tropical montane species in particular are often more or less woody and/or have the leaves in a dense rosette. Both traits occur in Cyanea leptostegia from the rain-forests of Kauai, the tallest of Campanulaceae. Lysipomia comprises 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. Brighamia are large stem succulents with ephemeral leaves found on dry coastal cliffs in Hawaii. Far wetter is the habitat of the submerged aquatics Lobelia dortmanna, an isoetid common in oligotrophic lakes in the taiga zone, and Howellia aquatilis, an elodeid growing in ponds in SW North America.

Leaves are often alternate, more rarely opposite or whorled. They are also simple (Petromarula one of very few exceptions), and without stipules.

Flowers are bisexual (dioecious in Dialypetalum) and protandrous. Petals are fused into a corolla that may be bell- or star-shaped in subfamily Campanuloideae, while tubular and bilaterally symmetric in most Lobelioideae. There are most commonly 5-8 lobes. Blue of various shades is the most common petal colour, but purple, red, pink, orange, yellow, white, and green also occur.

Stamens, equal in number to the petals, may be fused into a tube, as in all species of Lobelioideae. The style is in various ways involved in the "presentation" of the pollen, as in several other families of the order Asterales.

Carpel number is usually 2, 3 or 5 (8 in Ostrowskia), and corresponds to the number of stigmatic lobes. The ovary is inferior or, very rarely, superior (e.g. Cyananthus).

Fruits are berries or capsules.

Subfamilies and genera

Campanuloideae
Lobelioideae
Cyphioideae

Fossil record

The earliest known occurrence of Campanulaceae pollen is from Oligocene strata.[3] 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.[4][5]

Chemical compounds

Members of subfamily Lobelioideae contain the alkaloid lobeline. The principal storage carbohydrate of Campanulaceae is inulin, a polyfructan 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. 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.
  • Cosner, M. E.; Raubeson, L. A.; Jansen, R. K. (2007). "Chloroplast DNA rearrangements in Campanulaceae: phylogenetic utility of highly rearranged genomes" (PDF). 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.

References

  1. ↑ 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, retrieved 2010-12-10
  2. ↑ 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.
  3. ↑ Friis, Else Marie; Crane, Peter R.; Pedersen, Kaj Raunsgaard (August 2011). Early Flowers and Angiosperm Evolution. Cambridge University Press. ISBN 9780521592833.
  4. ↑ http://www.amjbot.org/content/early/2016/02/09/ajb.1500450.full.pdf
  5. ↑ 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)". New Phytologist. 210 (4): 1430–1442. doi:10.1111/nph.13920. PMC 4950005. PMID 26990796.
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