Campanulaceae | |
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Campanula cespitosa | |
Scientific classification ![]() | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
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 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.
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.
95 genera are accepted. [3] The Angiosperm Phylogeny Website divides the family into five subfamilies. [4]
As of August 2024 [update] , Plants of the World Online includes some genera in Campanulaceae that are not included by the Angiosperm Phylogeny Website: [3] [4]
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]
Members of subfamily Lobelioideae contain the alkaloid lobeline. The principal storage carbohydrate of Campanulaceae is inulin, a fructan also occurring in the related Asteraceae.
Asterales is an order of dicotyledonous flowering plants that includes the large family Asteraceae known for composite flowers made of florets, and ten families related to the Asteraceae. While asterids in general are characterized by fused petals, composite flowers consisting of many florets create the false appearance of separate petals.
Lobelia is a genus of flowering plants in the family Campanulaceae comprising 415 species, with a subcosmopolitan distribution primarily in tropical to warm temperate regions of the world, a few species extending into cooler temperate regions. They are known generally as lobelias.
Hamamelidaceae, commonly referred to as the witch-hazel family, is a family of flowering plants in the order Saxifragales. The clade consists of shrubs and small trees positioned within the woody clade of the core Saxifragales. An earlier system, the Cronquist system, recognized Hamamelidaceae in the Hamamelidales order.
Campanula is the type genus of the Campanulaceae family of flowering plants. Campanula are commonly known as bellflowers and take both their common and scientific names from the bell-shaped flowers—campanula is Latin for "little bell".
The Marantaceae are a family, the arrowroot family, of flowering plants consisting of 31 genera and around 530 species, defining it as one of the most species-rich families in its order. Species of this family are found in lowland tropical forests of Africa, Asia, and the Americas. The majority (80%) of the species are found in the American tropics, followed by Asian (11%) and African (9%) tropics. They are commonly called the prayer-plant family and are also known for their unique secondary pollination presentation.
Lobelioideae is a subfamily of the plant family Campanulaceae. It contains 32 genera, totalling about 1200 species. Some of the larger genera are Lobelia, Siphocampylus, Centropogon, Burmeistera and Cyanea.
The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the largest on any island archipelago, with over 125 species. The six genera involved can be broadly separated based on growth habit: Clermontia are typically branched shrubs or small trees, up to 7 metres (23 ft) tall, with fleshy fruits; Cyanea and Delissea are typically unbranched or branching only at the base, with a cluster of relatively broad leaves at the apex and fleshy fruits; Lobelia and Trematolobelia have long thin leaves down a single, non-woody stem and capsular fruits with wind-dispersed seeds; and the peculiar Brighamia have a short, thick stem with a dense cluster of broad leaves, elongate white flowers, and capsular fruits. The relationships among the genera and sections remains unsettled as of April 2022.
Codonopsis is a genus of flowering plant in the family Campanulaceae. As currently recognized, Codonopsis includes two other groups sometimes separated as distinct genera, i.e. Campanumoea and Leptocodon. The enlarged genus Codonopsis is widespread across eastern, southern, central, and southeastern Asia, including China, Japan, the Russian Far East, Kazakhstan, the Indian Subcontinent, Iran, Indochina, Indonesia, etc.
Loasaceae is a family of 15–20 genera and about 200–260 species of flowering plants in the order Cornales, native to the Americas and Africa. Members of the family include annual, biennial and perennial herbaceous plants, and a few shrubs and small trees. Members of the subfamily Loasoideae are known to exhibit rapid thigmonastic stamen movement when pollinators are present.
Burmeistera is a genus of flowering plants in the bellflower family, Campanulaceae. There are around 130 species distributed in Central and South America. This genus represents a rapid evolutionary radiation with species having diverged within only the last 2.6 million years.
Campanula rapunculus, common name rampion bellflower, rampion, rover bellflower, or rapunzel, is a species of bellflower (Campanula) in the family Campanulaceae.
Giant lobelia is a common name for several plants in the genus Lobelia; many of these species are restricted to high altitude, alpine ecosystems. In East African highlands as many as 20 species were listed in 1934 (Bruce). A selection of species names is below.
Cyanea is a genus of flowering plants in the family Campanulaceae that are endemic to Hawaii. The name Cyanea in Hawaiian is hāhā.
Cyanea dolichopoda was a species of shrub in the bellflower family that was endemic to Kauai. It was discovered in 1990 and has not been located in the wild since 1992. Like other Cyanea it is known as haha in Hawaiian.
Cyanea kuhihewa is a rare species of flowering plant in the bellflower family known by the common name Limahuli Valley cyanea. It is endemic to Kauai, where only two mature plants are known from a single wild population. Like other Cyanea it is known as haha in Hawaiian.
Cyanea remyi is a rare species of flowering plant in the bellflower family known by the common name Remy's cyanea. It is endemic to Hawaii, where it is known only from the island of Kauai. It is a federally listed endangered species of the United States. Like other Cyanea, it is known as haha in Hawaiian.
Legousia is a genus of flowering plants in the family Campanulaceae, native to Europe. Species in the genus used to be placed under genus Specularia along with plants in genera Triodanis and Heterocodon as well as some species in genus Campanula. However, the division has been confirmed evolutionarily comprehensive by a Campanulaceae phylogeny based on DNA molecular evidence.
Cyphia is a genus of flowering plants in the bellflower family, native to Africa; and particularly South Africa. It has been placed in its own subfamily, Cyphioideae. It is closely related to the genus Lobelia.Cyphia comes in shades of white to mauve, whereas lobelias have strong colours of blue to purple. Species in this genus have a bilabiate corolla; with 3 lobes on top and 2 below.
Smithiastrum is a genus of plants in the Campanulaceae or bellflower family found in the western United States. Its members are commonly known as harebells and were previously included in the family's type genus, Campanula, and were segregated when it was found that the Campanula was polyphyletic. As of 2023, Smithiastrum comprises two species.