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Aquilegia

For other uses, seeAquilegia (disambiguation).

Aquilegia(common names:granny's bonnet,[2]columbine) is agenusof about 130species[1]ofperennial plantsthat are found inmeadows,woodlands,and at higher elevations throughout the Northern Hemisphere, known for thespurredpetals[3]of theirflowers.

Columbine
Aquilegia vulgaris
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Ranunculales
Family: Ranunculaceae
Subfamily: Thalictroideae
Genus: Aquilegia
L.
Type species
Aquilegia vulgaris
Species[1]

130, seetext

Synonyms[1]

AquilinaBubani, nom. illeg.

Etymology

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The genus nameAquilegiacomes from theLatin"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

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Perennialherbs,with woody, erect stock,rootsforming thickrhizomes.The basalleavesare compound, 1–3ternate,blades 3-lobed -partite, and lobes lobulate andobtuse.Thecaulineleaves are similar to the basal ones, while the upper ones arebractlike.

Thehermaphrodite(bisexual)flowersare terminal to stem and branches. They are usuallypentamerous(with five spreadingperianthpetaloidsepalsegments). Five tubular honey-leaves[a]are semi erect with a flat limb and spurred orsaccateat the base. The spur is directed backwards and secretesnectar.Stamensare numerous (often more than 50) inwhorlsof 5, the innermost beingscariousstaminodes.There are ten membranaceous intrastaminal scales. There are fivepistilsand thecarpelsare free.[6][7][8]

Thefruithas several (five to 15)follicleswhich are semi erect and slightlyconnatedownwards. 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 allAquilegiaspecies have a ring of staminodia around the base of the stigma, which may help protect against insects.[10][6][7]Chromosome numberis x=7.[8]

Relatives

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Columbines are closely related to plants in the generaActaea(baneberries) andAconitum(wolfsbanes/monkshoods), which likeAquilegiaproduce cardiogenictoxins.[11]

Insects

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They are used as food plants by someLepidoptera(butterfly and moth)caterpillars.These are mainly ofnoctuid moths– noted for feeding on many poisonous plants without harm – such ascabbage moth(Mamestra brassicae),dot moth(Melanchra persicariae) andmouse moth(Amphipyra tragopoginis). Theengrailed(Ectropis crepuscularia), ageometer moth,also uses columbine as a larval food plant. The larvae of thePapaipema leucostigmaalso feed on columbine.[12]

Plants in the genusAquilegiaare a major food source forBombus hortorum,a species of bumblebee. Specifically, they have been found to forage on species ofAquilegia vulgarisin Belgium andAquilegia chrysanthain North America and Belgium. The bees do not show any preference in color of the flowers.[13]

Cultivation

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Columbinecultivar'Magpie'

Columbine is ahardyperennial,whichpropagatesby 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 athardiness zone3 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 EuropeanA. vulgariswas hybridized with other European and North American varieties. [16]Aquilegiaspecies are very interfertile, and will self-sow.[17]Some varieties are short-lived so are better treated asbiennials.

The British National Collection ofAquilegias was held by Mrs Carrie Thomas atKillaynearSwansea.[18]Some time during or before 2014 the collection started to succumb to Aquilegia Downy MildewPeronospora aquilegiicolawhich 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

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Double-floweredAquilegia×hybrida

The flowers of various species of columbine were consumed in moderation byNative Americansas acondimentwith 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 containcardiogenictoxinswhich cause both severegastroenteritisandheart palpitationsif consumed as food. Native Americans used very small amounts ofAquilegiaroot as a treatment forulcers.[21]However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal.[11]

Anacute toxicitytest in mice has demonstrated that ethanol extract mixed withisocytisoside,the main flavonoid compound from the leaves and stems ofAquilegia vulgaris,can be classified as non-toxic, since a dose of 3000 mg/kg did not cause mortality.[22]

Culture

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TheColorado blue columbine(A. coerulea) is the officialstate flowerofColorado(see alsoColumbine, Colorado). It is also used as a symbol of theformer cityofScarboroughin the Canadian province of Ontario.[23]

Wild columbine (Aquilegia canadensis) growing inSleeping Bear Dunes National LakeshorenearGlen Arbor, Michigan

Evolution

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Columbines have been important in the study ofevolution.It was found that theSierra columbine(A. pubescens) andcrimson columbine(A. formosa) each hasadaptedspecifically to apollinator.Bees and hummingbirds are the visitors toA. formosa,whilehawkmothswould only visitA. pubescenswhen given a choice. Such a "pollination syndrome",being due to flower color and orientation controlled by theirgenetics,ensuresreproductive isolationand can be a cause ofspeciation.[24]

Aquilegiapetals show an enormous range of petal spur length diversity ranging from a centimeter to the 15 cm spurs ofAquilegia 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

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

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

See also

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Notes

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  1. ^In theRanunculaceae,a variety of terms are used to describe the whorl of structures between thesepalsand stamens, including honey-leaves, petals, staminodes ornectaries[5]

References

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  1. ^abcdAquilegiaL.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.Retrieved8 April2018.
  3. ^abPuzey, J.R., Gerbode, S.J., Hodges, S.A., Kramer, E.M., Mahadevan, L. (2011) Evolution ofAquilegiaspur 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. ^abCullen et al 1993.
  7. ^abMabberley 1997,p. 50.
  8. ^abTamura 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.ISSN1932-6203.PMC2843724.PMID20352114.
  11. ^abTilford (1997)
  12. ^Roberts, Jason D. (August 19, 2015)."SpeciesPapaipema leucostigma- Columbine Borer Moth - Hodges#9478 ".BugGuide.RetrievedFebruary 5,2019.
  13. ^Macior, Lazarus Walter (1966-03-01). "Foraging Behavior ofBombus(Hymenoptera: Apidae) in Relation toAquilegiaPollination ".American Journal of Botany.53(3): 302–309.doi:10.2307/2439803.JSTOR2439803.
  14. ^"The Gardener's Network".
  15. ^John Kilmer (1989).The Perennial EncyclopediaISBN0-88665-639-7
  16. ^Andrew McIndoe, Kevin Hobbs: Perennials. David & Charles, 2005ISBN1-55870-764-6ISBN978-1-55870-764-1
  17. ^Cullina, William (2000).New England Wild Flower Society Guide to Growing and Propagating Wildflowers of the United States and Canada.Houghton Mifflin Harcourt.ISBN0395966094.
  18. ^"Plant Heritage - National Collections Scheme, UK Garden Plants".Nccpg.Retrieved2014-05-30.
  19. ^"Touchwood Plants".Retrieved2020-06-06.
  20. ^AM 2020.
  21. ^Dermatier, Charmaine."Plant of the Week: Yellow Columbine (Aquilegia flavescens)".US Forest Service.United States Department of Agriculture.Retrieved11 January2020.
  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.PMID12820244.S2CID22144186.Retrieved20 April2023.
  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.PMID17554306.S2CID4412955.
  26. ^Dezhi, Fu; Robinson, Orbélia R. (2001)."Aquilegia Linnaeus".Flora of China.6:278–281.

Bibliography

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Wikimedia Commons has media related toAquilegia.
Wikisourcehas the text of a1911Encyclopædia Britannicaarticle aboutAquilegia.
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