The genus nameAquilegia comes from theLatin "Aquila", or "eagle"; this is in reference to the spurred, "hook" shapes within the blooms, that many gardeners say resemble an eagle's talons.[4]
Perennialherbs, with woody, erect stock,roots forming thickrhizomes. The basalleaves are compound, 1–3ternate, blades 3-lobed -partite, and lobes lobulate andobtuse. Thecauline leaves are similar to the basal ones, while the upper ones arebract like.
Thehermaphrodite (bisexual)flowers are terminal to stem and branches. They are usuallypentamerous (with five spreadingperianthpetaloidsepal segments). Five tubular honey-leaves[a] are semi erect with a flat limb and spurred orsaccate at the base. The spur is directed backwards and secretesnectar.Stamens are numerous (often more than 50) inwhorls of 5, the innermost beingscariousstaminodes. There are ten membranaceous intrastaminal scales. There are fivepistils and thecarpels are free.[6][7][8]
Thefruit has several (five to 15)follicles which are semi erect and slightlyconnate 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 allAquilegia 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]
Columbines are closely related to plants in the generaActaea (baneberries) andAconitum (wolfsbanes/monkshoods), which likeAquilegia produce cardiogenictoxins.[11]
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 leucostigma also feed on columbine.[12]
Plants in the genusAquilegia are a major food source forBombus hortorum, a species of bumblebee. Specifically, they have been found to forage on species ofAquilegia vulgaris in Belgium andAquilegia chrysantha in North America and Belgium. The bees do not show any preference in color of the flowers.[13]
Columbinecultivar 'Magpie' also known as 'William Guiness'[14]
Columbine is ahardyperennial, whichpropagates 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 athardiness 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 EuropeanA. 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 asbiennials.[14]
The British National Collection ofAquilegias was held by Mrs Carrie Thomas atKillay nearSwansea.[19] Some time during or before 2014 the collection started to succumb to Aquilegia Downy MildewPeronospora 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]
The flowers of various species of columbine were consumed in moderation byNative Americans as acondiment 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 containcardiogenictoxins which cause both severegastroenteritis andheart palpitations if consumed as food. Native Americans used very small amounts ofAquilegia root as a treatment forulcers.[22] However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal.[11]
Anacute toxicity test 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.[23]
Aquilegia petals 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.[26] 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]
^In theRanunculaceae, a variety of terms are used to describe the whorl of structures between thesepals and stamens, including honey-leaves, petals, staminodes ornectaries[5]
^abPuzey, J.R., Gerbode, S.J., Hodges, S.A., Kramer, E.M., Mahadevan, L. (2011) Evolution ofAquilegia spur length diversity through changes in cell anisotropy. Proceedings of the Royal Society B.
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.ISBN1-930723-25-3HTML fulltext
Hodges, S. A.; Whittall, J. B.; Fulton, M.; Yang, J. Y. (2002). "Genetics of Floral Traits Influencing Reproductive Isolation betweenAquilegia formosa andAquilegia pubescens".The American Naturalist.159 (S3):S51 –S60.Bibcode:2002ANat..159S..51H.doi:10.1086/338372.PMID18707369.S2CID3399289.
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