Aquilegia

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Ordo: Ranunculales

Familia: Ranunculaceae
Subfamilia: Thalictroideae
Genus: Aquilegia
Species

A. alpina – A. amaliae – A. apuana – A. aradanica – A. aragonensis – A. atrata – A. atrovinosa – A. atwoodii – A. aurea – A. ballii – A. baltistanica – A. baluchistanica – A. barbaricina – A. barnebyi – A. barykinae – A. bernardii – A. bertolonii – A. blecicii – A. borodinii – A. brevistyla – A. buergeriana – A. canadensis – A. cazorlensis – A. champagnatii – A. chaplinei – A. chitralensis – A. chrysantha – A. coerulea – A. colchica – A. confusa – A. cossoniana – A. cymosa – A. daingolica – A. desertorum – A. desolaticola – A. dichroa – A. dinarica – A. discolor – A. dumeticola – A. ecalcarata – A. einseleana – A. elegantula – A. eximia – A. flabellata – A. flavescens – A. formosa – A. fragrans – A. ganboldii – A. gegica – A. glandulosa – A. gracillima – A. grata – A. grubovii – A. guarensis – A. hebeica – A. hinckleyana – A. hirsutissima – A. hispanica – A. holmgrenii – A. incurvata – A. iulia – A. japonica – A. jonesii – A. kamelinii – A. kansuensis – A. karatavica – A. karelinii – A. kitaibelii – A. kozakii – A. kubanica – A. kurramensis – A. lactiflora – A. laramiensis – A. longissima – A. lucensis – A. magellensis – A. maimanica – A. marcelliana – A. meridionalis – A. micrantha – A. microcentra – A. microphylla – A. montsicciana – A. moorcroftiana – A. nakaoi – A. navajonis – A. nemoralis – A. nevadensis – A. nigricans – A. nikolicii – A. nivalis – A. nivea – A. nugorensis – A. nuragica – A. ochotensis – A. olympica – A. ophiolithica – A. oreophila – A. ottonis – A. oxysepala – A. pancicii – A. parviflora – A. pubescens – A. pubiflora – A. pyrenaica – A. reuteri – A. rockii – A. saxifraga – A. saximontana – A. scopulorum – A. shockleyi – A. sibirica – A. sicula – A. skinneri – A. sternbergii – A. subscaposa – A. tianschanica – A. transsilvanica – A. turczaninowii – A. tuvinica – A. vicaria – A. viridiflora – A. viscosa – A. vitalii – A. vulgaris – A. xinjiangensis – A. synakensis – A. yabeana – A. yangii

Nothospecies

A. × cottia – A. × hybrida – A. × oenipontana
Name

Aquilegia L., Sp. Pl. 1: 533 (1753)

Type species: A. vulgaris L.

Note: There is significant differences in species synonymy across the additional references and secondary sources cited below and WS should try to reflect a broad consensual view where possible.
References
Primary references

Linnaeus, C. 1753. Species Plantarum. Tomus I: 533. Reference page.

Additional references

Barberis, G. & Nardi, E. 2011. A new species of Aquilegia L. from the Ligurian Apennine. Webbia 66(2): 233-234. DOI: 10.1080/00837792.2011.10670898Paywall Reference page.
Bulavkina, A.A. 1937. Aquilegia. Pp. 86–99 in Komarov, V.L. & Schischkin, B.K. (eds.), Flora URSS (Flora Unionis Rerumpublicarum Sovieticarum Socialisticarum) VII. [Nymphaeaceae – Papaveraceae]. xxvi + 615 pp., Academia Scientiarum URSS, Mosqua, Leningrad. DJVU BHL (English translation) Reference page.
Dezhi, F. & Robinson, O.R. 2001. Aquilegia. Pp. 278–281 in Wu, Zh.Y. , Raven, P.H. & Hong, D.Y. (eds.), Flora of China. Volume 6: Caryophyllaceae through Lardizabalaceae. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis, ISBN 1-930723-05-9. efloras Reference page.
Erst, A.S., Shaulo, D.N. & Kuznetsov, A.A. 2013. Aquilegia daingolica (Ranunculaceae), a new species from Mongolia. Sistematicheskie Zametki po Materialam Gerbarii Imeni P. N. Krylova pri Tomskom Gosudarstvennom Universitete 108: 14-22. Reference page.
Erst, A.S., Shaulo, D.N. & Shmakov, A.I. 2013. Aquilegia kamelinii (Ranunculaceae) – a new species from North Asia. [in Russian] Turczaninowia 16(3): 19–24. DOI: 10.14258/turczaninowia.16.3.4 Open access Reference page.
Erst, A.S., Karakulov, A.V. & Luferov, A.N. 2014. Aquilegia barykinae (Ranunculaceae), a new species from the Far East of Russia. Systematic Notes on the materials of P.N. Krylov Herbarium of Tomsk State University 110: 3–8. Reference page.
Erst, A.S., Shaulo, D.N., Luferov, A.N., Kuznetzov, A.A. & Shmakov, A.I. 2014. On taxonomical status Aquilegia kansuensis (Ranunculaceae). Turczaninowia 17(4): 24–25. [in Russian] Reference page.
Erst, A.S., Luferov, A.N., Xiang, K.L. & Wang, W. 2016. A review of the genus Aquilegia L. (Ranunculaceae) in Mongolia. Systematic Notes on the Materials of P.N. Krylov Herbarium of Tomsk State University 114: 37-48. DOI: 10.17223/20764103.114.5 Reference page.
Erst, A.S., Wang, W., Yu, S.X., Xiang, K., Wang, J., Shaulo, D.N., Smirnov, S.V., Kushunina, M., Sukhorukov, A.P. & Nobis, M. 2017. Two new species and four new records of Aquilegia (Ranunculaceae) from China. Phytotaxa 316(2): 121–137. DOI: 10.11646/phytotaxa.316.2.2 Paywall Reference page.
Nardi, E. 2011. Diagnoses of new species of European columbines. Webbia 66:(2): 231-232. DOI: 10.1080/00837792.2011.10670897 Paywall Reference page.
Nardi, E. 2012. Taxonomic and nomenclatural notes on Aquilegia L. (Ranunculaceae). Webbia 67(2): 179-181. DOI: 10.1080/00837792.2012.10670917 Paywall Reference page.
Nardi, E. 2014. Nomenclatural notes on Aquilegia L. (Ranunculaceae) from Europe. Webbia 69(1): 105. DOI: 10.1080/00837792.2014.895892Paywall Reference page.
Nardi, E. 2015. Il genere Aquilegia L. (Ranunculaceae) in Italia: 1-685. Edizioni Polistampa. ISBN 978-8859615187 Reference page.
Niketić, M., Cikovac, P. & Stevanović, V. 2013. Taksonomski i nomenklaturni prilozi o balkanskim kandilkama (Aquilegia L., Ranunculaceae). Bulletin of the Natural History Museum 6: 33-42. DOI: 10.5937/bnhmb1306033N Paywall Reference page.
Shaulo, D.N. & Erst, A.S. 2010. A new species of Columbine (Aquilegia L., Ranunculaceae) from Tuva. Turczaninowia 13(3): 43–45. PDF: (in Russian) Reference page.
Shaulo, D.N. & Erst, A.S. 2011. A new species of Aquilegia (Ranunculaceae) from the West Sayan, North Asia. Turczaninowia 14(3): 28–34. PDF:
Solanas, F.C. & Cabezudo, B. 2018. Aquilegia saxifraga sp. nov. (Ranunculaceae) en la provincia de Málaga (España). Acta Botanica Malacitana 43: 129-136. DOI: 10.24310/abm.v43i0.3512 Open access Reference page.
Whittemore, A.T. 1997. Aquilegia. Pp. - in Flora of North America Editorial Committee (ed.), Flora of North America North of Mexico. Volume 3: Magnoliophyta: Magnoliidae and Hamamelidae. 590 pp. Oxford University Press, New York / Oxford, ISBN 0-19-511246-6. efloras Reference page.

Links

Euro+Med 2006 onwards: Aquilegia in Euro+Med PlantBase – the information resource for Euro-Mediterranean plant diversity. Published on the internet. Accessed: 2021 Mar. 4.
Hassler, M. 2020. Aquilegia. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2020. Species 2000 & ITIS Catalogue of Life. Published on the internet. Accessed: 2020 March 21. Reference page.
Govaerts, R. et al. 2020. Aquilegia in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the internet. Accessed: 2020 March 21. Reference page.
International Plant Names Index. 2020. Aquilegia. Published online. Accessed: 21 March 2020.

Vernacular names
العربية: أنقولية
azərbaycanca: Akvilegiya
Boarisch: Antoniusglocke
беларуская: Ворлікі
català: Corniol
čeština: Orlíček
Cymraeg: Bonet Nain
dansk: Akeleje
Deutsch: Akeleien
dolnoserbski: Akelaja
English: Columbine
Esperanto: Akvilegio
فارسی: تاج‌الملوک
suomi: Akileijat
français: Ancolie
Gaeilge: Colaimbín
hornjoserbsce: Akelaja
magyar: Harangláb, sasfű
հայերեն: Արծվամագիլ
Ido: Aquilegio
日本語: オダマキ属
қазақша: Шөмішгүл
한국어: 매발톱꽃
lietuvių: Sinavadas
македонски: Кандилка
Nederlands: Akelei
norsk nynorsk: Akeleieslekta
ирон: Донæмбырдгæнæг
polski: Orlik
português: Aquilegia
русский: Водосбор
slovenčina: Orlíček
svenska: Aklejsläktet
Türkçe: Hasekiküpesi
українська: Орлики
oʻzbekcha/ўзбекча: Akvilegiya
中文: 耧斗菜属

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

The genus name Aquilegia is derived from the Latin word for eagle (aquila), because of the shape of the flower petals, which are said to resemble an eagle's claw. The common name "columbine" comes from the Latin for "dove", due to the resemblance of the inverted flower to five doves clustered together.[4][5]
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.[7][8][9]

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.[10] Almost all Aquilegia species have a ring of staminodia around the base of the stigma, which may help protect against insects.[11][7][8] Chromosome number is x=7.[9]
Relatives

Columbines are closely related to plants in the genera Actaea (baneberries) and Aconitum (wolfsbanes/monkshoods), which like Aquilegia produce cardiogenic toxins.[12]
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.[13]

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.[14]
Cultivation
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.[15][16]

Large numbers of hybrids are available for the garden, since the European A. vulgaris was hybridized with other European and North American varieties. [17] Aquilegia species are very interfertile, and will self-sow.[18] 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.[19] 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.[20] Aquilegia can be grown from seeds or rhizomes.[21]
Uses
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.[22] However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal.[12]

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.
Culture

The Colorado blue columbine (A. coerulea) is the official state flower of Colorado (see also Columbine, Colorado).
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.[23]

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.[24] 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)
Aquilegia alpina
Fan columbine (Aquilegia flabellata)
Fragrant columbine (Aquilegia fragrans)
Aquilegia × maruyamana
Pyrenean columbine (Aquilegia pyrenaica)

Columbine species include:[25]

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
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

See also

Columbine cup
Nora Barlow

Notes

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[6]

References

"Aquilegia Plants: How To Grow And Take Care Of Aquilegias". Daylilies in Australia. 28 May 2016. Retrieved 8 April 2018.
Sunset Western Garden Book, 1995:606–607
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.
Shorter Oxford English dictionary, 6th ed. United Kingdom: Oxford University Press. 2007. p. 3804. ISBN 978-0199206872.
Bill Neal (1992). Gardener's Latin. London: Robert Hale. p. 10. ISBN 0709051069.
Whittemore & Parfitt 2003.
Cullen et al 1993.
Mabberley 1997, p. 50.
Tamura 1993.
Armstrong, Margaret (1915). Western Wild Flowers. New York and London: C.P. Putnam's Sons, The Knickerbocker Press. p. 134.
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.
Tilford (1997)
Roberts, Jason D. (August 19, 2015). "Species Papaipema leucostigma - Columbine Borer Moth - Hodges#9478". BugGuide. Retrieved February 5, 2019.
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.
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John Kilmer (1989). The Perennial Encyclopedia ISBN 0-88665-639-7
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"Plant Heritage - National Collections Scheme, UK Garden Plants". Nccpg.com. Retrieved 2014-05-30.
"Touchwood Plants". Retrieved 2020-06-06.
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Dermatier, Charmaine. "Plant of the Week: Yellow Columbine (Aquilegia flavescens)". US Forest Service. United States Department of Agriculture. Retrieved 11 January 2020.
Fulton & Hodges (1999), Hodges et al. (2002)
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.

Dezhi, Fu; Robinson, Orbélia R. (2001). "Aquilegia Linnaeus". Flora of China. 6: 278–281.

Bibliography

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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.
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