Aquilegia

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Columbine
Wald-Akelei.JPG
Aquilegia vulgaris
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Ranunculales
Family: Ranunculaceae
Subfamily: Thalictroideae
Genus: Aquilegia
L.
Type species
Aquilegia vulgaris
L.
Species [1]

130, see text

Synonyms [1]

AquilinaBubani, nom. illeg.

Aquilegia (common names: granny's bonnet, [2] columbine) is a genus of about 130 species [1] of perennial plants that are found in meadows, woodlands, and at higher elevations throughout the Northern Hemisphere, known for the spurred petals [3] of their flowers.

Contents

Etymology

The genus name Aquilegia comes from the Latin "Aquila", or "eagle"; this is in obvious reference to the spurred, "hook" shapes within the blooms, that many gardeners say resemble an eagle's talons. [4]

Description

Perennial herbs, with woody, erect stock, roots forming thick rhizomes. The basal leaves are compound, 1–3 ternate, blades 3-lobed -partite, and lobes lobulate and obtuse. The cauline leaves are similar to the basal ones, while the upper ones are bract like.

The hermaphrodite (bisexual) flowers are terminal to stem and branches. They are usually pentamerous (with five spreading perianth petaloid sepal segments). Five tubular honey-leaves [a] are semi erect with a flat limb and spurred or saccate at the base. The spur is directed backwards and secretes nectar. Stamens are numerous (often more than 50) in whorls of 5, the innermost being scarious staminodes. There are ten membranaceous intrastaminal scales. There are five pistils and the carpels are free. [6] [7] [8]

The fruit has several (five to 15) follicles which are semi erect and slightly connate downwards. These hold many seeds and are formed at the end of the pistils. The nectar is mainly consumed by long-beaked birds such as hummingbirds. [9] Almost all Aquilegia species have a ring of staminodia around the base of the stigma, which may help protect against insects. [10] [6] [7] Chromosome number is x=7. [8]

Relatives

Columbines are closely related to plants in the genera Actaea (baneberries) and Aconitum (wolfsbanes/monkshoods), which like Aquilegia produce cardiogenic toxins. [11]

Insects

They are used as food plants by some Lepidoptera (butterfly and moth) caterpillars. These are mainly of noctuid moths – noted for feeding on many poisonous plants without harm – such as cabbage moth (Mamestra brassicae), dot moth (Melanchra persicariae) and mouse moth (Amphipyra tragopoginis). The engrailed (Ectropis crepuscularia), a geometer moth, also uses columbine as a larval food plant. The larvae of the Papaipema leucostigma also feed on columbine. [12]

Plants in the genus Aquilegia are a major food source for Bombus hortorum , a species of bumblebee. Specifically, they have been found to forage on species of Aquilegia vulgaris in Belgium and Aquilegia chrysantha in North America and Belgium. The bees do not show any preference in color of the flowers. [13]

Cultivation

Columbine cultivar 'Magpie' Aquilegia columbine magpie cultivar 2.jpg
Columbine cultivar 'Magpie'

Columbine is a hardy perennial, which propagates by seed. It will grow to a height of 40–50 cm (15–20 in). It will grow in full sun; however, it prefers growing in partial shade and well drained soil, and is able to tolerate average soils and dry soil conditions. Columbine is rated at hardiness zone 3 in the United States so does not require mulching or protection in the winter. [14] [15]

Large numbers of hybrids are available for the garden, since the European A. vulgaris was hybridized with other European and North American varieties. [16] Aquilegia species are very interfertile, and will self-sow. [17] Some varieties are short-lived so are better treated as biennials.

The British National Collection of Aquilegias was held by Mrs Carrie Thomas at Killay near Swansea. [18] Some time during or before 2014 the collection started to succumb to Aquilegia Downy Mildew Peronospora aquilegiicola which was at the time an emerging disease to which the plants had no resistance. By 2018 the entire collection had been lost. [19] Aquilegia can be grown from seeds or rhizomes. [20]

Uses

Double-flowered Aquilegia x hybrida Doubled flower of Aquilegia x hybrida 001.JPG
Double-flowered Aquilegia × hybrida

The flowers of various species of columbine were consumed in moderation by Native Americans as a condiment with other fresh greens, and are reported to be very sweet, and safe if consumed in small quantities. The plant's seeds and roots, however, are highly poisonous and contain cardiogenic toxins which cause both severe gastroenteritis and heart palpitations if consumed as food. Native Americans used very small amounts of Aquilegia root as a treatment for ulcers. [21] However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal. [11]

An acute toxicity test in mice has demonstrated that ethanol extract mixed with isocytisoside, the main flavonoid compound from the leaves and stems of Aquilegia vulgaris, can be classified as non-toxic, since a dose of 3000 mg/kg did not cause mortality. [22]

Culture

The Colorado blue columbine (A. coerulea) is the official state flower of Colorado (see also Columbine, Colorado). It is also used as a symbol of the former city of Scarborough in the Canadian province of Ontario. [23]

Wild columbine (Aquilegia canadensis) growing in Sleeping Bear Dunes National Lakeshore near Glen Arbor, Michigan Wild Columbine Michigan.jpg
Wild columbine (Aquilegia canadensis) growing in Sleeping Bear Dunes National Lakeshore near Glen Arbor, Michigan

Evolution

Columbines have been important in the study of evolution. It was found that the Sierra columbine (A. pubescens) and crimson columbine (A. formosa) each has adapted specifically to a pollinator. Bees and hummingbirds are the visitors to A. formosa, while hawkmoths would only visit A. pubescens when given a choice. Such a "pollination syndrome", being due to flower color and orientation controlled by their genetics, ensures reproductive isolation and can be a cause of speciation. [24]

Aquilegia petals show an enormous range of petal spur length diversity ranging from a centimeter to the 15 cm spurs of Aquilegia longissima. Selection from pollinator shifts is suggested to have driven these changes in nectar spur length. [25] It was shown that this spur length diversity is achieved solely through changing cell shape, not cell number or cell size. This suggests that a simple microscopic change can result in a dramatic evolutionarily relevant morphological change. [3]

Species

Dark columbine (Aquilegia atrata) Schwarzviolette Akelei (Aquilegia atrata) 02.jpg
Dark columbine (Aquilegia atrata)
Aquilegia alpina Aquilegia alpina1JUSA.jpg
Aquilegia alpina
Fan columbine (Aquilegia flabellata) Aquilegia flabellata1.jpg
Fan columbine (Aquilegia flabellata)
Fragrant columbine (Aquilegia fragrans) Aquilegia fragrans 2.jpg
Fragrant columbine (Aquilegia fragrans)
Aquilegia x maruyamana Aquilegia x maruyamana1.jpg
Aquilegia × maruyamana
Pyrenean columbine (Aquilegia pyrenaica) Aquilegia pyrenaica1JUSA.jpg
Pyrenean columbine (Aquilegia pyrenaica)
Aquilegia longissima Aquilegia longissima.jpg
Aquilegia longissima

130 columbine species are accepted. [1] [26]

See also

Notes

  1. ↑ In the Ranunculaceae, a variety of terms are used to describe the whorl of structures between the sepals and stamens, including honey-leaves, petals, staminodes or nectaries [5]

Related Research Articles

<i>Nepeta</i> Genus of flowering plants, known for effect on cats (catnip) in the mint family (Lamiaceae)

Nepeta is a genus of flowering plants in the family Lamiaceae. The genus name, from Latin nepeta (“catnip”), is reportedly in reference to Nepete, an ancient Etruscan city. There are 295 accepted species.

<i>Aquilegia formosa</i> Western North American species of columbine

Aquilegia formosa, the crimson columbine, western columbine, or (ambiguously) "red columbine", is a common wildflower native to western North America, from Alaska to Baja California, and eastward to Montana and Wyoming.

<i>Aquilegia coerulea</i> North American species of columbine

Aquilegia coerulea, the Colorado columbine, Rocky Mountain columbine, or blue columbine, is a species of flowering plant in the buttercup family Ranunculaceae, native to the Rocky Mountains and some of the surrounding states of the western United States. It is the state flower of Colorado. The Latin specific name coerulea means "sky blue".

<i>Aquilegia pubescens</i> Californian endemic species of columbine

Aquilegia pubescens is a high-altitude species of columbine known by the common names Sierra columbine, alpine columbine and Coville's columbine. Its flowers are large and usually a creamy white.

<i>Aquilegia chrysantha</i> North American species of columbine

Aquilegia chrysantha, the golden columbine, is a perennial herbaceous flowering plant native to the southwestern United States and northwestern Mexico.

<i>Aquilegia eximia</i> Californian species of columbine

Aquilegia eximia, the serpentine columbine or Van Houtte's columbine, is a perennial species of flowering plant in the family Ranunculaceae, endemic to California.

<i>Aquilegia flavescens</i> North American species of columbine

Aquilegia flavescens, the yellow columbine, is a perennial species of flowering plant in the family Ranunculaceae, native to the Rocky Mountains of the United States and Canada.

<i>Aquilegia elegantula</i> North American species of columbine

Aquilegia elegantula, the western red columbine, is a perennial species of flowering plant in the family Ranunculaceae, native to the Southwestern United States and northern Mexico.

<i>Aquilegia micrantha <span style="font-style:normal;">var.</span> grahamii</i> Variety of flowering plant in the family Ranunculaceae

Aquilegia micrantha var. grahamii, common name Graham's columbine, is a variety of perennial flowering plant in the family Ranunculaceae, endemic to Utah in the United States.

<i>Aquilegia longissima</i> North American species of columbine

Aquilegia longissima, the long-spur columbine or long-spurred columbine, is a rare perennial flowering plant in the family Ranunculaceae that is native to northern Mexico, Texas, and Arizona.

<span class="mw-page-title-main">Nectar spur</span> Nectar spur, secrets

A nectar spur is a hollow extension of a part of a flower. The spur may arise from various parts of the flower: the sepals, petals, or hypanthium, and often contain tissues that secrete nectar (nectaries). Nectar spurs are present in many clades across the angiosperms, and are often cited as an example of convergent evolution.

<i>Aquilegia alpina</i> Alpine European species of columbine

Aquilegia alpina, the alpine columbine or breath of God, is a perennial species of flowering plant in the family Ranunculaceae, native to the western and central Alps.

<i>Aquilegia olympica</i> Species of flowering plant in the family Ranunculaceae

Aquilegia olympica is a perennial flowering plant in the family Ranunculaceae, native to Turkey, Iran, and the Caucasus.

<i>Aquilegia ottonis</i> European species of columbine

Aquilegia ottonis is a species of flowering plant in the buttercup family. It has a broad distribution in Europe; it is native to Greece, Italy, and Albania. Plants produce blue-purple flowers which, based on pollination syndromes, are thought to be pollinated by bumblebees. A. ottonis is named after King Otto of Greece; the specific epithet 'ottonis' is the third declension of 'otto'.

<i>Aquilegia ecalcarata</i> Chinese endemic species of columbine

Aquilegia ecalcarata, the spurless columbine or false columbine, is a perennial species of flowering plant in the family Ranunculaceae, native to central China.

<i>Aquilegia sibirica</i> Species of flowering plant

Aquilegia sibirica, the Siberian columbine, is a species of flowering plant in the family Ranunculaceae native to the north-central Asian regions of Siberia, northern Mongolia, Kazakhstan, and Xinjiang. A hardy perennial plant, it prefers temperate environments. The Siberian columbine can be between 1 foot (0.30 m) and 2 feet (0.61 m) tall with flowers that are lilac-blue and white in color.

<i>Aquilegia buergeriana</i> Japanese species of columbine

Aquilegia buergeriana is a perennial species of flowering plant in the family Ranunculaceae, native to Japan.

<i>Aquilegia desertorum</i> North American species of columbine

Aquilegia desertorum, the desert columbine, is a perennial species of flowering plant in the family Ranunculaceae, native to the southwestern United States.

<i>Aquilegia glandulosa</i> North Asian species of columbine

Aquilegia glandulosa, the Altai columbine or Siberian columbine, is a perennial species of flowering plant in the family Ranunculaceae, native to northern and central Asia.

<i>Aquilegia micrantha</i> Species of flowering plants

Aquilegia micrantha, the Mancos columbine or Bluff City columbine, is a perennial species of plant in the family Ranunculaceae, native to Utah, Colorado, and Arizona.

References

  1. 1 2 3 4 Aquilegia L. Plants of the World Online . Retrieved 17 May 2024.
  2. ↑ "Aquilegia Plants: How To Grow And Take Care Of Aquilegias". Daylilies in Australia. 28 May 2016. Retrieved 8 April 2018.
  3. 1 2 Puzey, J.R., Gerbode, S.J., Hodges, S.A., Kramer, E.M., Mahadevan, L. (2011) Evolution of Aquilegia spur length diversity through changes in cell anisotropy. Proceedings of the Royal Society B.
  4. ↑ "Columbine: A Flower of Meanings". San Francisco, CA: Sutro Stewards. 25 July 2017.
  5. ↑ Whittemore & Parfitt 2003.
  6. 1 2 Cullen et al 1993.
  7. 1 2 Mabberley 1997, p. 50.
  8. 1 2 Tamura 1993.
  9. ↑ Armstrong, Margaret (1915). Western Wild Flowers. New York and London: C.P. Putnam's Sons, The Knickerbocker Press. p. 134.
  10. ↑ Voelckel, Claudia; Borevitz, Justin O.; Kramer, Elena M.; Hodges, Scott A. (2010-03-23). "Within and between Whorls: Comparative Transcriptional Profiling of Aquilegia and Arabidopsis". PLOS ONE. 5 (3): e9735. Bibcode:2010PLoSO...5.9735V. doi: 10.1371/journal.pone.0009735 . ISSN   1932-6203. PMC   2843724 . PMID   20352114.
  11. 1 2 Tilford (1997)
  12. ↑ Roberts, Jason D. (August 19, 2015). "Species Papaipema leucostigma - Columbine Borer Moth - Hodges#9478". BugGuide. Retrieved February 5, 2019.
  13. ↑ Macior, Lazarus Walter (1966-03-01). "Foraging Behavior of Bombus (Hymenoptera: Apidae) in Relation to Aquilegia Pollination". American Journal of Botany. 53 (3): 302–309. doi:10.2307/2439803. JSTOR   2439803.
  14. ↑ "The Gardener's Network".
  15. ↑ John Kilmer (1989). The Perennial Encyclopedia ISBN   0-88665-639-7
  16. ↑ Andrew McIndoe, Kevin Hobbs: Perennials. David & Charles, 2005 ISBN   1-55870-764-6 ISBN   978-1-55870-764-1
  17. ↑ Cullina, William (2000). New England Wild Flower Society Guide to Growing and Propagating Wildflowers of the United States and Canada. Houghton Mifflin Harcourt. ISBN   0395966094.
  18. ↑ "Plant Heritage - National Collections Scheme, UK Garden Plants". Nccpg.com. Retrieved 2014-05-30.
  19. ↑ "Touchwood Plants" . Retrieved 2020-06-06.
  20. ↑ AM 2020.
  21. ↑ Dermatier, Charmaine. "Plant of the Week: Yellow Columbine (Aquilegia flavescens)". US Forest Service. United States Department of Agriculture. Retrieved 11 January 2020.
  22. ↑ Adamska, T (June 2003). "Hepatoprotective effect of the extract and isocytisoside from Aquilegia vulgaris". Phytother Res. 17 (6): 691–696. doi:10.1002/ptr.1233. PMID   12820244. S2CID   22144186 . Retrieved 20 April 2023.
  23. ↑ "City of Scarborough".
  24. ↑ Fulton & Hodges (1999), Hodges et al. (2002)
  25. ↑ Whittall, Justen B.; Hodges, Scott A. (7 June 2007). "Pollinator shifts drive increasingly long nectar spurs in columbine flowers". Nature. 447 (7145): 706–709. Bibcode:2007Natur.447..706W. doi:10.1038/nature05857. PMID   17554306. S2CID   4412955.
  26. ↑ Dezhi, Fu; Robinson, Orbélia R. (2001). "Aquilegia Linnaeus". Flora of China. 6: 278–281.

Bibliography

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