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

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

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

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


Description
Although most Campanulaceae are herbs (sometimes climbing, as in ), there is also a large number of e.g. species of . Isotoma hypocrateriformis is a annual from Australia's dry interior. There are also , e.g. the commonly cultivated ornamental ( bells). Many perennial campanuloids grow in rock-crevices, such as () and Petromarula pinnata (). Some lobelioids also grow on rocks, e.g. the peculiar perennial stem succulent in . 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). are minute cushion plants of the high , while giant rosette-forming lobelias (e.g., ) 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 common in lakes in the zone of North America and Europe, and Howellia aquatilis, an 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 .

Tubers occur in several genera, e.g. .

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

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

are ( in ) and . 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 . 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 and some Campanuloideae (e.g. )

Within the family pollen grains are often , less commonly , , or .

Carpel number is usually 2, 3 or 5 (8 in ), 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 , pollen is, already in the bud stage, released into the tube formed by the . 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 . Brighamia and have pale or white flowers with a long-tubed corolla, and are pollinated by . Pollination by lizards has been reported for and mauritianus.

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

Berries are a common fruit-type in Lobelioideae ( , , , Cyanea etc.), whilst rare in Campanuloideae ( 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. The Angiosperm Phylogeny Website divides the family into five subfamilies.Stevens, P.F. (2001 onwards). "Campanulaceae". Angiosperm Phylogeny Website. Retrieved 2022-04-23.


Fossil record
The earliest known occurrence of Campanulaceae is from .
(2011). 9780521592833, Cambridge University Press.
Earliest Campanulaceae dated, are seeds of † Campanula paleopyramidalis from 17-16 million years old deposits in the , , . It is a close relative of the extant Campanula pyramidalis.[2]


Chemical compounds
Members of subfamily Lobelioideae contain the alkaloid . The principal storage carbohydrate of Campanulaceae is , a also occurring in the related .


Literature

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