Aquilegia

Genus of perennial plants (columbine)

For other uses, see Aquilegia (disambiguation).

Columbine

Aquilegia vulgaris
Scientific classification
Kingdom:  | Plantae
Clade:  | Tracheophytes
Clade:  | Angiosperms
Clade:  | Eudicots
Order:  | Ranunculales
Family:  | Ranunculaceae
Subfamily:  | Thalictroideae
Genus:  | Aquilegia

Type species
Aquilegia vulgaris

Species

60-70, see text

Aquilegia (common names: granny's bonnet,[1] columbine) is a genus of about 60–70 species[2] of perennial plants that are found in meadows, woodlands, and at higher altitudes throughout the Northern Hemisphere, known for the spurred petals [3] of their flowers.

## Etymology[edit]

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[edit]

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[edit]

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

## Insects[edit]

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[edit]

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[edit]

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[edit]

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

## Evolution[edit]

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[edit]

Dark columbine (Aquilegia atrata) Aquilegia alpina Fan columbine (Aquilegia flabellata) Fragrant columbine (Aquilegia fragrans) Aquilegia × maruyamana Pyrenean columbine (Aquilegia pyrenaica)

Columbine species include:[26]

* Aquilegia alpina L. – alpine columbine
* Aquilegia atrata W.D.J.Koch – dark columbine
* Aquilegia atrovinosa
* Aquilegia aurea Janka
* Aquilegia barbaricina – Barbaricina columbine (doubtfully valid)
* Aquilegia barnebyi – oil shale columbine
* Aquilegia bernardii Gren. & Godr.
* Aquilegia bertolonii Schott – Bertoloni columbine
* Aquilegia blecicii Podobnik (doubtfully valid)
* Aquilegia brevistyla – smallflower columbine
* Aquilegia buergeriana
* Aquilegia canadensis – Canadian columbine, wild columbine
* Aquilegia champagnatii Moraldo, E.Nardi & la Valva (doubtfully valid)
* Aquilegia chrysantha – golden columbine
* Aquilegia coerulea – Colorado blue columbine
* Aquilegia desertorum – desert columbine
* Aquilegia desolatica – desolation columbine
* Aquilegia dinarica Beck
* Aquilegia ecalcarata
* Aquilegia einseleana F.W.Schultz
* Aquilegia elegantula – western red columbine
* Aquilegia eximia – Van Houtte's columbine
* Aquilegia flabellata – fan columbine, Japanese wodamakinari (including A. akitensis)
* Aquilegia flavescens – yellow columbine
* Aquilegia fragrans Benth. – fragrant columbine
* Aquilegia formosa – crimson columbine, western columbine
* Aquilegia glandulosa
* Aquilegia grahamii – Graham's columbine
* Aquilegia grata
* Aquilegia × hybrida
* Aquilegia incurvata
* Aquilegia japonica
* Aquilegia jonesii – Jones' columbine
* Aquilegia karatavica
* Aquilegia karelini
* Aquilegia kitaibelii Schott
* Aquilegia lactiflora

* Aquilegia laramiensis – Laramie columbine
* Aquilegia litardierei Briq.
* Aquilegia longissima – Gray. – longspur columbine
* Aquilegia loriae – Lori's columbine
* Aquilegia magellensis F.Conti & Soldano – Magella columbine
* Aquilegia × maruyamana
* Aquilegia micrantha – Mancos columbine
* Aquilegia moorcroftiana
* Aquilegia nigricans Baumg. – Bulgarian columbine
* Aquilegia nugorensis Arrigoni & E.Nardi (doubtfully valid)
* Aquilegia nuragica – Nuragica columbine
* Aquilegia olympica Boiss.
* Aquilegia origami
* Aquilegia ottonis Orph. ex Boiss.
* Aquilegia oxysepala
* Aquilegia pancicii Degen
* Aquilegia parviflora
* Aquilegia pubescens – Sierra columbine, Coville's columbine
* Aquilegia pubiflora
* Aquilegia pyrenaica DC. – Pyrenean columbine
* Aquilegia rockii
* Aquilegia saximontana – Rocky Mountain columbine
* Aquilegia scopulorum – blue columbine, Utah columbine
* Aquilegia shockleyi – desert columbine
* Aquilegia sibirica – Siberian columbine
* Aquilegia thalictrifolia Schott & Kotschy
* Aquilegia transsilvanica Schur
* Aquilegia triternata – Chiricahua Mountain columbine
* Aquilegia truncata – red columbine
* Aquilegia turczaninovii
* Aquilegia viridiflora Pall. – green columbine, green-flowered columbine
* Aquilegia viscosa Gouan
* Aquilegia vitalii
* Aquilegia vulgaris – common columbine, European columbine, granny's nightcap
* Aquilegia yabeana

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## See also[edit]

* Columbine cup
* Nora Barlow

## Notes[edit]

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]

## References[edit]

1. ^ "Aquilegia Plants: How To Grow And Take Care Of Aquilegias". Daylilies in Australia. 28 May 2016. Retrieved 8 April 2018.
2. ^ Sunset Western Garden Book, 1995:606–607
3. ^ a b 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. ^ a b Cullen et al 1993.
7. ^ a b Mabberley 1997, p. 50.
8. ^ a b 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. ^ a b 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[edit]

* Cullen, J; et al. (1993) [1964]. "Aquilegia L.". In Tutin, T. G.; Burges, N. A.; Edmondson, J. R. (eds.). Flora Europaea. Volume 1, Psilotaceae to Platanaceae (2 ed.). Cambridge University Press. pp. 287–290. ISBN 978-0-521-41007-6. see also Flora Europaea
* Mabberley, D. J. (1997) [1987]. The Plant-Book: A Portable Dictionary of the Vascular Plants (2nd ed.). Cambridge University Press. ISBN 978-0-521-41421-0.
* Whittemore, Alan T.; Parfitt, Bruce D. (2003). "Ranunculaceae Jussieu". Flora of North America. New York: Oxford University Press. Retrieved 2 February 2021.
* Kubitzki, Klaus; Rohwer, Jens G.; Bittrich, Volker, eds. (1993). The Families and Genera of Vascular Plants. II Flowering plants - Dicotyledons. Magnoliid, Hamamelid and Caryophyllid families. Berlin: Springer. ISBN 978-3540555094.
* Tamura, M (1993-07-28). Aquilegia. Springer. p. 580\. ISBN 9783540555094., in Kubitzki et al (1993)
* Allan M. Armitage: Armitage's Native Plants for North American Gardens.Timber Press, 2006 ISBN 0-88192-760-0 ISBN 978-0-88192-760-3
* Dezhi, Fu; Robinson, Orbélia R. (2001): 19. Aquilegia. In: Wu, Z. Y.; Raven, Peter Hamilton & Hong, D. Y. (eds.): Flora of China (Vol. 6: Caryophyllaceae through Lardizabalaceae): 278. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis. ISBN 1-930723-25-3 HTML fulltext
* Fulton, M.; Hodges, S. A. (1999). "Floral isolation between Aquilegia formosa and Aquilegia pubescens". Proceedings of the Royal Society B: Biological Sciences. 266 (1435): 2247–2252. doi:10.1098/rspb.1999.0915. PMC 1690454.
* Hodges, S. A.; Whittall, J. B.; Fulton, M.; Yang, J. Y. (2002). "Genetics of Floral Traits Influencing Reproductive Isolation between Aquilegia formosa and Aquilegia pubescens". The American Naturalist. 159: S51–S60. doi:10.1086/338372. PMID 18707369. S2CID 3399289.
* Nold, Robert (2003): Columbines: Aquilegia, Paraquilegia, and Semiaquilegia. Timber Press. ISBN 0-88192-588-8 Preview at Google Books
* Royal Botanic Garden Edinburgh (RBGE) [2008]: Digital Flora Europaea: Aquilegia species list. Retrieved 2008-NOV-25.
* Tilford, Gregory L. (1997): Edible and Medicinal Plants of the West. Mountain Press Pub., Missoula, Montana. ISBN 0-87842-359-1
* United States Department of Agriculture (USDA) [2008]: USDA Plants Profile: Aquilegia. Retrieved 2008-NOV-25.
* Puzey, J. R.; Gerbode, S. J.; Hodges, S. A.; Kramer, E. M.; Mahadevan, L. (2011). "Evolution of spur-length diversity in Aquilegia petals is achieved solely through cell-shape anisotropy" (PDF). Proceedings of the Royal Society B: Biological Sciences. 279 (1733): 1640–1645. doi:10.1098/rspb.2011.1873. PMC 3282339. PMID 22090381.
* Kramer, E. M. (2009). Aquilegia: A New Model for Plant Development, Ecology, and Evolution Annual Review of Plant Biology, Vol. 60.
* "How to Grow Columbine". American Meadows. 2020. Retrieved 3 February 2021.

## External links[edit]

Wikimedia Commons has media related to Aquilegia.

Wikisource has the text of a 1911 Encyclopædia Britannica article about Aquilegia.

Taxon identifiers|

* Wikidata: Q147641
* Wikispecies: Aquilegia
* APDB: 188248
* APNI: 130744
* BOLD: 121221
* CoL: ZHP
* EoL: 37461
* EPPO: 1AQIG
* FNA: 102388
* FoAO2: Aquilegia
* FoC: 102388
* GBIF: 3033167
* GRIN: 884
* iNaturalist: 48245
* IPNI: 33064-1
* IRMNG: 1025084
* ITIS: 18728
* NBN: NHMSYS0000456001
* NCBI: 3450
* NZOR: e6ef1470-c015-49a9-9098-1c71e3b224c7
* Open Tree of Life: 964055
* PLANTS: AQUIL
* POWO: urn:lsid:ipni.org:names:325954-2
* Tropicos: 40018252
* uBio: 5865999
* VASCAN: 838
* VicFlora: 6e3fc015-bf90-4d24-807c-ff74b0561c9c
* WFO: wfo-4000002807

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