{{short description|Genus of flowering plants}} {{Featured article}} {{Use American English|date=May 2025}} {{Other uses}} {{Automatic taxobox |name = Columbine |image = Wald-Akelei.JPG |image_alt = Photograph of a flowering ''Aquilegia vulgaris'' plant |image_caption = ''Aquilegia vulgaris'' |taxon = Aquilegia |authority = L. |subdivision_ranks = Species |subdivision = Approx. 130, see list |subdivision_ref = <ref name = POWO>{{Cite powo | id=325954-2 |title=''Aquilegia'' L.|access-date=23 February 2026}}</ref> |type_species = ''Aquilegia vulgaris'' |type_species_authority = L. |synonyms = ''Aquilina'' {{small|Bubani, nom. illeg.}} |synonyms_ref = <ref name =POWO/> }} '''''Aquilegia''''', commonly known as '''columbines''', is a genus of perennial flowering plants in the family Ranunculaceae (buttercups). The genus includes between 70 and 400 taxa (described species and subspecies) with natural ranges across the Northern Hemisphere. Natural and introduced populations of ''Aquilegia'' exist on all continents but Antarctica. Known for their high physical variability and ease of hybridization, columbines are popular garden plants and have created many cultivated varieties.

''Aquilegia'' plants typically possess stiff stems and leaves that divide into multiple leaflets. Columbines often have colorful flowers with five sepals and five petals. The petals generally feature nectar spurs which differ in length between species. In North America, morphological variations in spurs evolved to suit different pollinators. Some species and varieties of columbines are naturally spurless. In cultivation, varieties bearing significantly altered physical traits such as double flowering are prevalent.

Columbines were associated with fertility goddesses in ancient Greek and ancient Roman religion, and connected to Christian religious concepts. Archeological evidence suggests ''Aquilegia'' plants were in cultivation by the 3rd century AD in Roman Britain, and they remain popular in gardens worldwide. Despite often being toxic, columbines have been in use by humans as herbal remedies, perfume, and food. Asian traditional medicine practitioners, Indigenous North Americans, and Medieval Europeans have considered portions of the plants to have medicinal uses. Selective breeding and hybridization of columbines has occurred for centuries, with genetic exchanges between Old and New World species creating further diversity.

==Etymology== The 1st-century AD Greek writer Dioscorides referred to columbine plants as ''Isopyrum'', a name that is now applied to another genus, ''Isopyrum''.<ref name=Nold/>{{rp|38}} In the 12th century, the abbess and polymath Hildegard of Bingen referred to the plants as ''agleya''{{snd}} from which the genus's name in German, ''Akelei'', derives. The first use of ''aquilegia'' with regard to columbines was in the 13th century by Albertus Magnus. In the 16th and 17th centuries, the names ''Colombina'', ''Aquilina'', and ''Aquileia'' came into use. With the Swedish biologist Carl Linnaeus's 1753 ''Species Plantarum'', the formal name for the genus became ''Aquilegia'', though limited use of ''Aquilina'' persisted in scientific usage until at least 1901.<ref name=Nardi/>{{rp|23}}

Several scientific and common names for the genus ''Aquilegia'' derive from its appearance.<ref name=Nardi/>{{rp|23}} The genus name ''Aquilegia'' may come from the Latin word for "eagle", ''aquila'', in reference to the petals' resemblance to eagle talons.<ref>{{cite web|url=https://www.wildflower.org/plants/result.php?id_plant=AQCHC|title=''Aquilegia chrysantha'' var. ''chaplinei''|website=wildflower.org|publisher=Lady Bird Johnson Wildflower Center|access-date=11 May 2025|archive-url= https://web.archive.org/web/20240613052630/https://www.wildflower.org/plants/result.php?id_plant=AQCHC|archive-date=13 June 2024|url-status=live}}</ref> Another possible etymology for ''Aquilegia'' is a derivation from the Latin {{lang|la|aquam legere}} ("to collect water"),<ref name=Weber/> ''aquilegium'' (a container of water), or {{lang|la|aquilex}} ("dowser" or "water-finder") in reference to the profusion of nectar in the spurs.<ref>{{cite web|url=https://dryades.units.it/Grappa/index.php?procedure=taxon_page&id=10000042&num=30877|title=''Aquilegia confusa'' Rota|work=Portale alla Flora del Monte Grappa|publisher=University of Trieste|access-date=11 May 2025|archive-url= https://web.archive.org/web/20241210035437/https://dryades.units.it/Grappa/index.php?procedure=taxon_page&id=10000042&num=30877|archive-date=10 December 2024|url-status=live|lang=it}}</ref><ref name=Nold/>{{rp|37–38}} The most common English-language name, ''columbine'', is founded in the dove-like appearance of the sepals and derived from the Italian ''colombino'' and French ''colombin'', which were in turn derived from ''columba'', Latin for ''dove''.<ref name=Nold/>{{rp|38}}<ref name=CE/><ref>{{cite journal|title=The Etymology of Columbine|first=E. J.|last=Hill|author-link=Ellsworth Jerome Hill|journal=The Plant World|date=September 1902|volume=2|number=9|pp=175–177|jstor=43477174|jstor-access=free}}</ref>

There are a number of other common names for ''Aquilegia'' across different languages. In English, these include ''granny's bonnet'' for various species, but particularly ''Aquilegia vulgaris''.<ref>{{Cite OED|Granny's Bonnet, N. |5952472021 }}</ref> An older common name for columbines is ''lion's herb'', from the folk belief that lions fed on it for courage.<ref>{{Cite book |date=1949 |editor-last1=Leach |editor-first1=Maria |editor-link1=Maria Leach |editor-last2=Fried |editor-first2=Jerome |title=Standard Dictionary of Folklore, Mythology, and Legend |url=https://archive.org/details/standarddictiona0000mari |url-access=registration |volume=One, A–I |location=New York |publisher=Funk & Wagnalls |page=242 |oclc=23850031}}</ref> In French, the word ''ancolie'' is the common name for ''Aquilegia'',<ref name=FNA/> while individual members of the genus have been called ''gants-de-Notre-Dame'' ("Our Lady's glove"). In Italian, ''amor-nascoto'' ("love-born") has been used.<ref name=Nold/>{{rp|38–39}}

==Description== thumb|right|alt=Photograph of flowers of Aquilegia atrata|''A.&nbsp;atrata'' flowers ''Aquilegia'' is a genus of herbaceous, perennial flowering plants in the family Ranunculaceae (buttercups). The genus is highly variable in appearance.<ref name=Nold>{{cite book|title=Columbines: ''Aquilegia'', ''Paraquilegia'', and ''Semiaquilegia''|first=Robert|last=Nold|publisher=Timber Press|date=2003|location=Portland, OR|isbn=0881925888|url=https://archive.org/details/columbines00robe|url-access=registration|via=Internet Archive}}</ref>{{rp|29}}<ref name=FNA>{{cite book|chapter-url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=102388|chapter=''Aquilegia''|title=Flora of North America: North of Mexico|volume=3|isbn=9780195112467|publisher=Oxford University Press|location=New York|date=1997|via=eFloras.org}}</ref> Though they are perennials, certain species are short-lived, with some exhibiting lifespans more similar to biennials and others only flourishing for three to six years.<ref name=Nold/>{{rp|18}}<ref name=Ornamental>{{cite book|chapter-url=http://www.efloras.org/florataxon.aspx?flora_id=120&taxon_id=102388|title=Ornamental Plants From Russia And Adjacent States of the Former Soviet Union|first=Tatyana|last=Shulkina|date=2005|chapter=''Aquilegia'' L.|publisher=Rostok|isbn=9785946680325|location=Saint Petersburg|via=efloras.org}}</ref> Following a dormant period in the winter, columbines will grow foliage and have a brief flowering period.<ref name=SouthernLiving/><ref name=Nold/>{{rp|15–16}} Some columbines bloom the first year after sowing, others will first bloom in their second year.<ref name=Nold/>{{rp|13}} Later, seed heads will emerge and split, sowing new seed. The foliage lives through the summer before wilting and dying going into the fall.<ref name=SouthernLiving>{{cite web|url=https://www.southernliving.com/columbine-plants-7570572|title=How To Grow And Care For Columbine Flowers|first=Peggy|last=Riccio|editor-first=Khara|editor-last=Scheppmann|work=Southern Living|date=25 February 2025|access-date=28 May 2025}}</ref><ref name=PollinationBiology/>

''Aquilegia'' plants grow from slim, woody rootstocks that comprise the perennial portion of the plants. One or more annual aerial stems rise from the rootstocks each growing season, ultimately drying out following fruiting.<ref name=Nardi/>{{rp|143–145}} Leaves can grow in both basal (from where the plant meets the soil) and cauline (from an aerial stem) arrangements. Leaves emanating from closer to the plant's core are generally borne on flexible petioles, while leaves further from the core generally lack petioles. The compound leaves of ''Aquilegia'' are generally ternate, biternate, or triternate. Ternate leaves each divide into three leaflets, biternate leaves divide into three components that each in turn bear three leaflets for a total of nine leaflets, and triternate leaves divide into three components three times for a total of 27 leaflets.<ref name=FNA/>

The flowering stems emerge from rosettes during the spring and summer.<ref name=RHSA>{{cite web|url=https://www.rhs.org.uk/plants/aquilegia|title=''Aquilegia''|website=rhs.org.uk|publisher=Royal Horticultural Society|access-date=16 May 2025}}</ref> Each inflorescence appears at the terminus of an aerial stem and can reach {{cvt|30|cm}} long. Depending on the species, an inflorescence will feature one to ten of either cymes (flower clusters) or solitary flowers.<ref name=FNA/> Flower morphology varies across the genus, but all columbine flowers emerge from buds that are initially nodding (facing downwards).<ref name=Nold/>{{rp|31}} Flowers can be monochromatic or display multiple colors.<ref name=RHSA/> The typical flower color for columbines is blue in shades ranging into purple and nearly black shades. Blue flowering is especially the norm in European columbines, where only ''A.&nbsp;aurea'' possesses yellow flowers. In North America, yellow and red flowers are typical, with blue and blue-purple flowers almost exclusive to high-altitude species.<ref name=Nold/>{{rp|33}}

The perianth (non-reproductive portion) on an ''Aquilegia'' flower generally comprises five sepals that look like petals and five petals.<ref name=PollinationSystems/> Each petal typically comprises two portions: a blade, which is broad and projects towards the front of the flower, and a nectar spur, which is a nectar-bearing structure which projects backwards.<ref name=FI>{{cite book|chapter-url=https://bibdigital.rjb.csic.es/medias/71/a5/be/6f/71a5be6f-b4bc-432f-9a6a-baa17a5d8ccb/files/Fl_iber1.pdf|chapter=''Aquilegia'' L.|title=Flora Iberica: Plantas Vasculares de la Península Ibérica e Islas Baleares|trans-title=Flora Iberica: Vascular Plants of the Iberian Peninsula and Balearic Islands|language=es|date=1986|volume=1: Lycopodiaceae–Papaveraceae|location=Madrid|publisher=Real Jardín Botánico de Madrid|isbn=8400062221|editor-first1=S.|editor-last1=Castroviejo|editor-first2=M.|editor-last2=Laínz|editor-first3=G.|editor-last3=López González|editor-first4=P.|editor-last4=Montserrat|editor-first5=F.|editor-last5=Muñoz Garmendia|editor-first6=J.|editor-last6=Paiva|editor-first7=L.|editor-last7=Villar}}</ref>{{rp|376}}<ref name=Nold/>{{rp|31}} The hollow spurs attract pollinators and give columbine flowers a distinctive appearance.<ref name=Problem>{{cite book|title=Rodale's Flower Garden Problem Solver|first1=Jeff|last1=Ball|first2=Liz|last2=Ball|date=1990|isbn=0878578684|publisher=Rodale Press|location=Emmaus, PA}}</ref>{{rp|147}} Some species possess spurs with hooked, horn-like appearances, while other species can have spurs that are either straight or coiled.<ref name=Nold/>{{rp|31, 38}}<ref name=CE/> Some columbines, such as ''A.&nbsp;ecalcarata'', are naturally spurless.<ref name=Nold/>{{rp|82}} Individuals and populations can also be found with recessive spurlessness within typically spurred species.<ref name=Nold/>{{rp|31}}<ref name=FNA/><ref name=Weber/>

''Aquilegia'' are bisexual (featuring both male and female organs) and capable of self-pollination, through either or both autogamy (does not require assistance from pollinators) and geitonogamy (requires pollinators).<ref name=PollinationSystems/> Autogamy has been observed as the primary fertilization mechanism in ''A.&nbsp;paui''.<ref name=Shift/> ''A.&nbsp;formosa'' and ''A.&nbsp;eximia'' may exhibit adichogamy, where male and female organs do not operate simultaneously to prevent self-fertilization. Fertilization via cross-pollination also occurs in ''Aquilegia'', with pollinators carrying pollen from one flower to the stigma of another.<ref name=PollinationBiology>{{cite journal|title=Pollination Biology and Secondary Pollinators in Seven North American ''Aquilegia'' Species|first1=Marion|last1=Chartier|first2=Constantin|last2=Kopper|first3=Michael|last3=Münch|first4=Olivia Messinger|last4=Carril|first5=Sergio|last5=Díaz-Infante|first6=Margarita|last6=Lachmayer|first7=Silvia|last7=Ulrich|first8=Joseph&nbsp;S.|last8=Wilson|first9=Jürg|last9=Schönenberger|first10=Maria|last10=Von Balthazar|journal=International Journal of Plant Sciences|volume=186|number=1|date=January 2025|pages=20–37|doi=10.1086/733068|bibcode=2025IJPlS.186...20C |issn=1058-5893|eissn=1537-5315}}</ref>

thumb|right|alt=Photograph of fanning foliage of Aquilegia flabellata|The fanning foliage of ''A.&nbsp;flabellata'' 'Alba' The reproductive portions of a columbine flower comprise the stamen (male) and gynoecium (female). The stamens, which bear the anthers from which pollen emerges, form a whorl of five around the gynoecium. The total number of stamens varies between species.<ref name=PollinationSystems/> ''Aquilegia'' is the only Ranunculaceae genus where each species has its stamens in a whorled arrangement and has syncarpy (fused carpels) in its gynoecium, with this consistency resulting from the presence of nectar spurs.<ref>{{cite book|title=Floral Diagrams: An Aid to Understanding Flower Morphology and Evolution|first=Louis P.|last=Ronse De Craene|date=2022|orig-date=2010|edition=2nd|publisher=Cambridge University Press|location=Cambridge|doi=10.1017/9781108919074|isbn=9781108825733|lccn=2021050296|page=160}}</ref> Generally, there are scale-shaped staminodes between the stamen and female pistil structures.<ref name=FNA/> ''Aquilegia'' are protandrous, meaning that a flower's anthers release its pollen prior to the flower's stigmas being receptive to pollen.<ref name=Nold/>{{rp|27}} The flowers undergo three stages of anthesis: a premale stage, where the flower perianth is open but the anthers are not dehiscenced (split to expose pollen); the male stage where with the perianth present and the anthers dehiscenced, and a postmale stage where the anthers have withered but the perianth remains.<ref name=PollinationBiology/>

''Aquilegia'' fruit comprise follicles. These follicles have a split on one side and terminate with a curling tip known as a ''beak''.<ref name=Nold/>{{rp|32}} Columbine seeds are generally obovoid with black, smooth exteriors.<ref name=FNA/> Columbine seeds are in a dormant state at the point of sowing.<ref name=Dormacy/> ''Aquilegia'' seeds remain viable longer than many Ranunculaceae, though germination is faster in newer seeds;<ref name=Nold/>{{rp|23}} in cold storage, ''A.&nbsp;formosa'' seeds have remained viable for at least three years.<ref>{{cite web|url=https://www.nrcs.usda.gov/plantmaterials/orpmcmt9921.pdf|title=Protocol Information: ''Aquilegia formosa'' Fisch. ex DC.|publisher=Corvallis Plant Materials Center, United States Department of Agriculture|location=Corvallis, OR|date=2009|access-date=11 August 2025}}</ref> Seed germination is primarily dependent on temperature, with seeds typically requiring a multi-month period of summer temperatures followed by a multi-week to multi-month exposure to winter temperatures (vernalization) prior to germinating once temperatures warm with the arrival of spring. This prevents seedlings from emerging until there are survivable environmental conditions.<ref name=Dormacy>{{cite journal|title=Dormancy Cycles in ''Aquilegia oxysepala'' Trautv. et Mey. (Ranunculaceae), A Species with Non-Deep Simple Morphophysiological Dormancy|first1=Keliang|last1=Zhang|first2=Yusong|last2=Ji|first3=Guixian|last3=Fu|first4=Linjun|last4=Yao|first5=Huina|last5=Liu|first6=Jun|last6=Tao|first7=Jeffrey&nbsp;L.|last7=Walck|journal=Plant and Soil|date=July 2021|volume=464|number=1/2|pages=223–235|doi=10.1007/s11104-021-04951-8|bibcode=2021PlSoi.464..223Z |issn=0032-079X}}</ref><ref name=Sowing>{{cite journal|title=Dissecting Seed Dormancy and Germination in ''Aquilegia barbaricina'', Through Thermal Kinetics of Embryo Growth|first1=M.|last1=Porceddu|first2=E.|last2=Mattana|first3=H.&nbsp;W.|last3=Pritchard|first4=G.|last4=Bacchetta|journal=Plant Biology|doi=10.1111/plb.12610|pmid=28762612|date=2017|pages=983–993|volume=19|number=6|bibcode=2017PlBio..19..983P |hdl=11584/222396|hdl-access=free}}</ref>

The chromosome number for columbines is ''2n''=14 (seven pairs of chromosomes).<ref name=FNA/><ref>{{cite journal|title=Documented Chromosome Numbers of Plants|journal=Madroño|volume=17|number=8|date=October 1964|pages=266–268 |jstor=41423164}}</ref> Individual plants have been recorded with other anomalous chromosome numbers. It is possible that B chromosomes impact the phenotype and the fertility of the individual plants that possess them.<ref name=Nardi/>{{rp|183–185}}

===Phytochemistry=== thumb|alt=Photograph of fruit and seeds of Aquilegia vulgaris|''A.&nbsp;vulgaris'' fruit and seeds. Human consumption of columbine seeds and roots can introduce toxins that negatively impact heart health. Among ''Aquilegia'' that have cyanophores (cells that produce a blue color) like ''A.&nbsp;vulgaris'', the cyanogenic glycosides compounds dhurrin and triglochinin have been observed. Cyanogenic glycosides generally taste bitter and can be toxic to animals and humans. Ingestion of {{cvt|20|g}} of fresh ''A.&nbsp;vulgaris'' leaves by a human was observed to cause convulsions, respiratory distress, and heart failure. A child who consumed 12 ''A.&nbsp;vulgaris'' flowers experienced weakness of the limbs, cyanosis, drowsiness, and miosis; all symptoms abated after three hours.<ref name=Toxicity>{{cite book|title=Natural Poisons and Venoms: Plant Toxins: Polyketides, Phenylpropanoids and Further Compounds|first1=Eberhard|last1=Teuscher|first2=Ulrike|last2=Lindequist|series=De Gruyter STEM|publisher=De Gruyter|date=2024|isbn=9783110728538|doi=10.1515/9783110728538|location=Berlin; Boston|pages=300–326}}</ref> Mature seeds and roots contain toxins that, if consumed, are perilous to human heart health.<ref name=FSAF/>

The presence of the antibacterial flavonoid compound isocytisoside has been observed in ''A.&nbsp;vulgaris''.<ref name=Antibacterial>{{cite journal|title=Antibacterial phenolic compounds from the flowering plants of Asia and the Pacific: coming to the light|journal=Pharmaceutical Biology|first1=Mazdida|last1=Sulaiman|first2=Layane|last2=Ebehairy|first3=Veeranoot|last3=Nissapatorn|first4=Mohammed|last4=Rahmatullah|first5=Jhonnel|last5=Villegas|first6=Helina Jean|last6=Dupa|first7=Ricksterlie&nbsp;C.|last7=Verzosa|first8=Karma&nbsp;G.|last8=Dolma|first9=Muhamad|last9=Shabaz|first10=Scholastica|last10=Lanting|first11=Nor Azizun|last11=Rusdi|first12=Nor Hayati|last12=Abdullah|first13=Mohammed Khaled|last13=Bin Break|first14=Teng Jin|last14=Khoo|first15=Wei|last15=Wang|first16=Christophe|last16=Wiart|doi=10.1080/13880209.2024.2407530|volume=62|number=1|date=11 October 2024|doi-access=free|pmid=39392281|pages=713–766|pmc=11486068 }}</ref> Polyphenols, primarily flavonoids, are the main component of hydroethanolic extract (produced with a solvent made from a mixture of water and ethanol) from ''A.&nbsp;oxysepala''. These compounds function as antioxidants. A study of ''A.&nbsp;oxysepala'' extract found it has a good scavenging effect (a capacity to remove or inactivate undesired substances) on DPPH, superoxide anion, and hydroxyl radicals, but a poor scavenging capacity towards hydrogen peroxide. For all these, ascorbic acid has a superior scavenging effect to the extract.<ref name=AOExtract>{{cite journal|title=Evaluation of Antioxidant Activities of ''Aquilegia oxysepala'' Hydroethanolic Extract|first1=Yan|last1=Yu|first2=Zhi-Biao|last2=Yi|first3=Yi-Zeng|last3=Liang|journal=Journal of Food Biochemistry|date=August 2009|volume=33|number=4|pages=500–505|doi=10.1111/j.1745-4514.2009.00233.x|issn=0145-8884}}</ref>

In flowering plants, the presence of phenylpropanoids can serve as protection from ultraviolet (UV) light and as a signaling mechanism towards pollinators. A study that examined ''A.&nbsp;formosa'' flowers determined that the petals and sepals had uniform levels of UV-resistant phenylpropanoids.<ref>{{cite journal|title=Transcontinental patterns in floral pigment abundance among animal-pollinated species|first1=Eduardo|last1=Narbona|first2=Jose&nbsp;C.|last2=Del&nbsp;Valle|first3=Justen&nbsp;B.|last3=Whittall|first14=Melissa|last14=León-Osper|first4=M.&nbsp;Luisa|last4=Buide|first5=Iñigo|last5=Pulgar|first6=Maria Gabriela|last6=Gutierrez Camargo|first7=Leonor Patricia|last7=Cerdeira Morellato|first8=Nancy|last8=Rodríguez-Castañeda|first9=Victor|last9=Rossi|first10=Katie|last10=Conrad|first11=Joey|last11=Hernandez-Mena|first12=Pedro&nbsp;L.|last12=Ortiz|first13=Montserrat|last13=Arista|journal=Scientific Reports|volume=15|number=1|date=7 May 2025|page=15927 |doi=10.1038/s41598-025-94709-4|doi-access=free|pmid=40335586 |pmc=12059069 |bibcode=2025NatSR..1515927N }}</ref>

==Ecology== thumb|right|alt=Photograph of leaf miner trails on Aquilegia canadensis leaf|Leaf miners can leave trails where they have consumed the interior tissue of ''Aquilegia'' leaves. Despite the plants' toxicity and in the absence of incentives for human consumption,<ref name=Toxicity/> some animals consume the fruit and leaves of columbines. Consumption by mammals is not considered a component of the ''Aquilegia'' reproductive cycle but has been observed. In the case of the endangered ''A.&nbsp;paui'', one study cited a personal observation that 30% of all fruit was lost to goat predation.<ref name=Shift/><ref name=Conservation>{{cite journal|url=https://www.cambridge.org/core/journals/oryx/article/conservation-assessment-of-aquilegia-paui-ranunculaceae-a-case-study-of-an-extremely-narrow-endemic/9E66060BEEABA24D662CDB23D2DEE239|title=Conservation Assessment of ''Aquilegia paui'' (Ranunculaceae): A Case Study of an Extremely Narrow Endemic|last1=Martinell|first1=M. Carmen|first2=Jordi|last2=López-Pujol|first3=Cèsar|last3=Blanché|first4=Julián|last4=Molero|first5=Llorenç|last5=Sàez|journal=Oryx|date=2011|volume=45|number=2|pages=187–190|doi=10.1017/S0030605310001754|via=Cambridge University Press|archive-date=27 April 2025|access-date=16 May 2025|archive-url=https://web.archive.org/web/20250427225231/https://www.cambridge.org/core/journals/oryx/article/conservation-assessment-of-aquilegia-paui-ranunculaceae-a-case-study-of-an-extremely-narrow-endemic/9E66060BEEABA24D662CDB23D2DEE239|url-status=live|hdl=10261/50366|hdl-access=free}}</ref>

In the Northeastern United States and Eastern Canada, ''A.&nbsp;canadensis'' serves as the host plant for the butterfly ''Erynnis lucilius'' (columbine duskywing). In two periods, the first from April to June and the second from July to September, the butterflies lay their eggs on the underside of the columbine leaves. The latterly laid brood overwinters as caterpillars in the plant litter around the columbine.<ref>{{cite web|url=https://www.fs.usda.gov/wildflowers/pollinators/pollinator-of-the-month/columbine-duskywing.shtml|title=Pollinator of the Month: Columbine Duskywing (''Erynnis lucilius'')|first=Melissa|last=Simpson|publisher=United States Forest Service|website=fs.usda.gov|access-date=28 May 2025}}</ref> In the Western United States, ''Bombus occidentalis'' (western bumblebee) has been observed nectar robbing from ''A.&nbsp;coerulea'' by opening or using holes cut in the spurs.<ref>{{cite journal|last1=Pyke|first1=Graham H.|title=Local Geographic Distributions of Bumblebees Near Crested Butte, Colorado: Competition and Community Structure|journal=Ecology|date=1982|volume=63|issue=2|pages=555–573|doi=10.2307/1938970 |jstor=1938970|bibcode=1982Ecol...63..555P|issn=0012-9658}}</ref>

Also in North America, three leaf miner species in the genus ''Phytomyza'' lay their eggs on ''Aquilegia'': ''P.&nbsp;aquilegiana'' in the east, ''P.&nbsp;aquilegivora'' in the Midwestern United States, and ''P.&nbsp;columbinae'' in the west. Collectively known as the columbine leaf miners, white trails or splotches on leaves indicate where the larvae consumed the tissue between the leaves' surfaces. The larvae will cut through the leaves, pupating in small puparia on the leaves' undersides. Adults pierce the leaves with their ovipositors to access liquids in the plants, leaving marks.<ref name=NAPests>{{cite web|url=https://hort.extension.wisc.edu/articles/common-columbine-pests-columbine-leafminer-and-columbine-sawfly/|title=Common Columbine Pests: Columbine Leafminer and Columbine Sawfly|first=Susan|last=Mahr|publisher=Wisconsin Horticulture, University of Wisconsin–Madison|website=hort.extension.wisc.edu|access-date=28 May 2025}}</ref> Another ''Phytomyza'' columbine leaf miner, ''P.&nbsp;ancholiae'', is native to France.<ref>{{cite journal|url=https://www.biodiversitylibrary.org/page/10626219|title=Mémoire pour servir a l'histoire des Diptères dont les larves minent les feuilles des plantes|first=Claude Charles|last=Goureau|author-link=Claude Charles Goureau|lang=fr|pages=153–155|journal=Annales de la Société entomologique de France|volume=2|number=9|date=1851|via=Biodiversity Heritage Library}}</ref>

Originally from Europe, ''Pristiphora rufipes'' (columbine sawfly) is now also found in Canada and the United States. After developing from eggs laid on columbine leaves in late spring, the green larvae will eat the leaves from the outside in during their active period from April to June. In cases where many larvae are on the same plant, only the stem and flowers may go uneaten. The larvae mature within a few weeks, after which they drop from the plants and pupate in cocoons.<ref name=NAPests/>

Several fungi attack columbine foliage, including ''Ascochyta aquilegiae'', ''Cercospora aquilegiae'', and ''Septoria aquilegiae''.<ref name=Problem/> The fungus-like oomycete species ''Peronospora aquilegiicola'', a type of downy mildew, originated in East Asian ''Aquilegia'' and ''Semiaquilegia'' populations. It was first reported on columbines in the United Kingdom in 2013, resulting in discussion about quarantining measures to prevent its spread to Continental Europe.<ref>{{Cite journal|last1=Thines|first1=Marco|last2=Denton|first2=Geoffrey&nbsp;J.|last3=Beal|first3=Elizabeth&nbsp;J.|last4=Kilty|first4=Anne|last5=Denton|first5=Jennifer&nbsp;O.|last6=Shin|first6=Hyeon-Dong|last7=Choi|first7=Young-Joon|date=October 2019|title=''Peronospora aquilegiicola'' sp. nov., the Downy Mildew Affecting Columbines in the UK is an Invasive Species from East Asia|journal=European Journal of Plant Pathology|volume=155|issue=2|pages=515–525|doi=10.1007/s10658-019-01787-y|bibcode=2019EJPP..155..515T|s2cid=195657188|issn=1573-8469}}</ref>

===Pollination=== thumb|upright=1.2|right|alt=Photograph of a hybrid columbine being pollinated by a hawkmoth|Some North American columbines adapted to pollination by hawkmoths ''(pictured)'' and hummingbirds. Longer nectar spurs require nectar-seeking hawkmoths to make contact with the columbine's reproductive organs. ''Aquilegia'' developed diverse floral features including varied orientation, coloration, and nectar spur morphology to attract different pollinators, contributing to speciation.<ref name=POPOVICH/> The suite of floral traits that develop to attract a particular set of pollinators are collectively referred to as a flower's pollination syndrome (also called a floral syndrome). ''Aquilegia'' flowers are traditionally divided into three pollination syndromes: bumblebees, hummingbirds, or hawkmoths, each of which is attracted by the plants' nectar.<ref name=PollinationSystems/> In addition to these primary pollinators, pollen-collecting insects frequently function as secondary pollinators among columbines. A 2026-published study determined that pollination syndromes in ''Aquilegia'' species were primarily driven by the three sets of nectar-collecting pollinators but found no evidence that secondary, pollen-collecting pollinators imposed any evolutionary pressure on floral traits.<ref>{{cite journal|title=Floral Syndromes in ''Aquilegia'' (Ranunculaceae) are Associated with Nectar- But Not Pollen-Collecting Pollinators|first1=Anna-Sophie|last1=Hawranek|first2=Maria|last2=Von Balthazar|first3=Marion|last3=Chartier|first4=Jürg|last4=Schönenberger|date=16 January 2026|journal=Annals of Botany|doi=10.1093/aob/mcaf333|doi-access=free}}</ref> In cases where pollinators are scarce, columbines may adopt autogamy as a primary fertilization method, such as in ''A.&nbsp;paui''.<ref name=PollinationSystems>{{cite journal|title=Pollination and Reproductive Systems in Columbines (''Aquilegia'', Ranunculaceae): Review and Insights|first1=Maria|last1=Von Balthazar|first2=Margarita|last2=Lachmayer|first3=Anna-Sophie|last3=Hawranek|first4=Constantin|last4=Kopper|first5=Jürg|last5=Schönenberger|first6=Marion|last6=Chartier|journal=International Journal of Plant Sciences|volume=186|number=1|date=January 2025|pages=38–52|doi=10.1086/733070|issn=1058-5893|eissn=1537-5315|bibcode=2025IJPlS.186...38V}}</ref>

Eurasian columbines are primarily pollinated by flies, bees, and bumblebees. North American columbines are generally pollinated by bees, bumblebees, hawkmoths, and hummingbirds. Pollination by hummingbirds is more typical for red-flowered North American ''Aquilegia'', while pale-flowered columbines may have developed to increase their visibility to hawkmoths in twilight.<ref name=Nold/>{{rp|32–33}}

Nectar spur length on particular columbines is often correlated to their associated pollinators. While nectar spur length in Eurasia varies little, there is substantial variation in North American spur length.<ref name=Increasingly/> There, hawkmoths often possess long tongues, which permit them to reach deep into nectar spurs. In areas of hawkmoth pollination, the elongated nectar spurs of ''A.&nbsp;coerulea'' prevent them from removing nectar without making contact with the pollen.<ref>{{Cite journal|last1=Brunet|first1=Johanne|date=June 2009|title=Pollinators of the Rocky Mountain Columbine: Temporal Variation, Functional Groups and Associations With Floral Traits|url=https://www.researchgate.net/publication/24400848 |journal=Annals of Botany|volume=103|issue=9|pages=1567–1578|doi=10.1093/aob/mcp096|issn=1095-8290|pmc=2701757|pmid=19414518|via=ResearchGate}}</ref> While hawkmoths are present in Eurasia, there are no Eurasian columbines with the hawkmoth pollination syndrome which includes longer spurs. In North America, the presence of hummingbirds{{snd}} which are absent in Eurasia and possess tongue lengths that are generally intermediate between other pollinators and hawkmoths{{snd}} may have functioned as a stepping stone that allowed North American ''Aquilegia'' to evolve the hawkmoth pollination syndrome.<ref name=Increasingly>{{cite journal|title=Pollinator Shifts Drive Increasingly Long Nectar Spurs in Columbine Flowers|first1=Justen B.|last1=Whittal|first2=Scott A.|last2=Hodges|journal=Nature|volume=447|date=June 2007|issue=7145 |pages=706–709 |doi=10.1038/nature05857|pmid=17554306 |bibcode=2007Natur.447..706W }}</ref>

While a given population of ''Aquilegia'' may settle a particular habitat and develop pollination syndromes for certain pollinators, this does not necessarily translate into ecological speciation with genetic barriers between species. The likelihood of such speciation increases when floral mutations and pollinator behavioral changes coincide with isolated, small populations, as in the case of ''A.&nbsp;micrantha'' var.&nbsp;mancosana.<ref name=Nardi/>{{rp|79}}

==Taxonomy== Within Linnaean taxonomy, ''Aquilegia'' was first described as a genus in Carl Linnaeus's 1753 ''Species Plantarum''.<ref name=Nardi/>{{rp|23}} The genus is typically assigned to the family Ranunculaceae, though a minority of botanists formerly considered it a member of the family Helleboraceae. The latter placement, first made by the French botanist Jean-Louis-Auguste Loiseleur-Deslongchamps in 1819, was premised on Helleboraceae fruiting almost universally occurring with a follicle.<ref name=Nold/>{{rp|29}}<ref>{{cite web|url=https://www.fs.usda.gov/wildflowers/plant-of-the-week/aquilegia_caerulea.shtml|title=Plant of the Week: Colorado Columbine (''Aquilegia caerulea'' James)|first=Steve|last=Popovich|publisher=United States Forest Service|website=fs.usda.gov|access-date=12 May 2025|archive-url=https://web.archive.org/web/20230524172921/https://www.fs.usda.gov/wildflowers/plant-of-the-week/aquilegia_caerulea.shtml|archive-date=24 May 2023}}</ref> Another historic assignment, made by the Swedish botanist {{ill|Nils Lilja|sv|Nils Lilja}} in 1870, placed ''Aquilegia'' as the sole member of the family Aquilegiaceae.<ref name=Nardi/>{{rp|31}}

Columbines are most commonly assigned to the tribe Isopyreae, though they are sometimes placed within Aquilegieae. The placement of the tribe containing ''Aquilegia'' has been uncertain, with alternating assignments to two subfamilies: Thalictroideae and Isopyroideae. Regardless of the placement, ''Aquilegia'' and the genera ''Isopyrum'' and ''Thalictrum'' (meadow-rues) form a basal, paraphyletic group which is characterized by their plesiomorphy (characteristics shared between clades from their last common ancestor) with Berberidaceae.<ref name=Nardi/>{{rp|31}} When placed within the monophyletic Thalictroideae, ''Aquilegia'' is the second largest genus in the subfamily in terms of taxa (described species and subspecies), behind ''Thalictrum''. Columbines are nested in one of the three major clades in the subfamily, a clade it shares with ''Semiaquilegia'' and ''Urophysa''. ''Semiaquilegia'' and ''Aquilegia'' are sister genera.<ref name=East>{{cite journal|url=https://academic.oup.com/aob/article/134/7/1251/7743276|title=Out of and In East Asia: Phylogeny, Biogeography and Diversification of Thalictroideae (Ranunculaceae) in the Northern Hemisphere|first1=Yuan-Yuan|last1=Ling|first2=Huan-Wen|last2=Peng|first3=Lian|last3=Lian|first4=Andrey&nbsp;S.|last4=Erst|author-link4=Andrey Erst|first5=Kun-Li|last5=Xiang|first6=Wei|last6=Wang|journal=Annals of Botany|volume=134|issue=7|date=16 December 2024|pages=1251–1262|pmid=39196797|doi=10.1093/aob/mcae148|doi-access=free|pmc=11688531 }}</ref><ref name=Extended>{{cite journal|title=Extended Phylogeny of ''Aquilegia'': The Biogeographical and Ecological Patterns of Two Simultaneous But Contrasting Radiations|first1=Jesús M.|last1=Bastida|first2=Julio M.|last2=Alcántara|first3=Pedro&nbsp;J.|last3=Rey|first4=Pablo|last4=Vargas|first5=Carlos&nbsp;M.|last5=Herrera|journal=Plant Systematics and Evolution|volume=284|number=3/4|date=February 2010|pages=171–185|jstor=23673233|url=https://digital.csic.es/handle/10261/36746|doi=10.1007/s00606-009-0243-z|bibcode=2010PSyEv.284..171B |hdl=10261/36746|hdl-access=free}}</ref>

The broadly accepted circumscription of ''Aquilegia'' was established by the American botanist Philip A. Munz in his 1946 monograph ''Aquilegia: The Cultivated and Wild Columbines''. The only element of Munz's circumscription which has been substantially contested is his inclusion of the spurless Asian species ''A.&nbsp;ecalcarata'', which is sometimes instead segregated into the closely related genus of spurless-flowered ''Semiaquilegia''; ''Semiaquilegia ecalcarata'' remains the species's common name in cultivation.<ref name=Nardi/>{{rp|31}}<ref name=Nold/>{{rp|82}} Another spurless columbine, ''A.&nbsp;micrantha'' var.&nbsp;''mancosana'', was also once reassigned to ''Semiaquilegia''. Reassignments to ''Isopyrum'' and ''Paraquilegia'', such as ''P.&nbsp;anemonoides'' in 1920, have been more permanent.<ref name=Nold/>{{rp|38, 46–49}}

===Evolution=== {{Cladogram of Aquilegia species}} There are no good fossils of columbines and other Thalictroideae that indicate how they evolved and radiated.<ref name=Fior2013>{{cite journal|title=Spatiotemporal Reconstruction of the ''Aquilegia'' Rapid Radiation Through Next-Generation Sequencing of Rapidly Evolving cpDNA Regions|first1=Simone|last1=Fior|first2=Mingai|last2=Li|first3=Bengt|last3=Oxelman|first4=Roberto|last4=Viola|first5=Scott&nbsp;A.|last5=Hodges|first6=Lino|last6=Ometto|first7=Claudio|last7=Varotto|journal=New Phytologist|date=April 2013|volume=198|issue=2|doi=10.1111/nph.12163|url=https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.12163|pages=579–592|pmid=23379348 |bibcode=2013NewPh.198..579F }}</ref><ref name=Hodges1994>{{cite journal|title=Columbines: A Geographically Widespread Species Flock|first1=Scott A.|last1=Hodges|first2=Michael L.|last2=Arnold|journal=Proceedings of the National Academy of Sciences of the United States of America|date=24 May 1994|volume=91|number=11|pages=5129–5132|doi=10.1073/pnas.91.11.5129 |doi-access=free|jstor=2364926|pmid=8197196 |pmc=43945|bibcode=1994PNAS...91.5129H }}</ref><ref name=East/> Genetic evidence published in a 2024 study suggests that the last common ancestor among Thalictroideae lived in East Asia approximately 36 million years ago, during the late Eocene.<ref name=East/>

A 2018 study of genetic evidence indicated that ''Aquilegia'' first appeared during the Upper Miocene approximately 6.9 million years ago. According to this study, the genus split into two clades 4.8 million years ago, with one clade eventually populating North America and the other radiating across Eurasia.<ref>{{cite journal|title=Genetic Diversity and Evolutionary History of Four Closely Related ''Aquilegia'' Species Revealed By 10 Nuclear Gene Fragments|doi=10.1111/jse.12298|journal=Journal of Systematics and Evolution|volume=56|number=2|date=2018|pages=129–138|first1=Lei|last1=Huang|first2=Fang-Dong|last2=Geng|first3=Jing-Jing|last3=Fan|first4=Cheng|last4=Xue|first5=Xiao-Yan|last5=Zhang|first6=Ju-Qing|last6=Kang|first7=Jian-Qiang|last7=Zhang|first8=Yi|last8=Ren|bibcode=2018JSyEv..56..129H }}</ref> The 2024 study found that the divergence between ''Urophysa'', ''Semiaquilegia'', and ''Aquilegia'' instead occurred over a relatively short 1 million-year-long period approximately 8 to 9 million years ago.<ref name=East/> The genus ''Aquilegia'' likely originated in the mountainous portions of south-central Siberia.<ref name=Extended/>

Studies of ''Aquilegia'' genetics indicated that North American ''Aquilegia'' species shared their last common ancestor with species from the Asian Far East between 3.84 and 2.99 million years ago. This analysis corresponded with the theory that ''Aquilegia'' reached North America via a land bridge over the Bering Strait.<ref name=Fior2013/> While there were several periods after this date range where the Beringian land bridge connected Asia and North America, these occurred when climatic conditions would have prevented ''Aquilegia'' migration through the region.<ref name=Extended/>

Genetic information suggests that the diversification rates of columbines rapidly increased about 3 million years ago,<ref name=East/> with indications of two independent radiation events occurring around that time: one in North American columbine populations and the other in European populations.<ref name=Extended/> Despite the rapid evolution of substantial physical differences across species, genetic divergence remains minimal. This, combined with the presence of relatively few physiological barriers to hybridization, has resulted in columbines displaying exceptionally high degrees of interfertility.<ref name=Shift>{{cite journal|title=Shift Towards Autogamy in the Extremely Marrow Endemic ''Aquilegia paui'' and Comparison with Its Widespread Close Relative ''A. vulgaris'' (Ranunculaceae)|first1=M. Carmen|last1=Martinell|first2=Ana|last2=Rovira|first3=Cèsar|last3=Blanché|first4=Maria|last4=Bosch|journal=Plant Systematics and Evolution|date=August 2011|volume=295|number=1/4|pages=72–83|doi=10.1007/s00606-011-0463-x |jstor=43558220|bibcode=2011PSyEv.295...73M }}</ref><ref name=Munz>{{cite book|title=Aquilegia: The Cultivated and Wild Columbines|series=Gentes Herbarum|volume=VII|location=Ithaca, NY|author-link=Philip A. Munz|first=Philip A.|last=Munz|date=25 March 1946|publisher=The Bailey Hortorium of the New York State College of Agriculture at Cornell University|url=https://archive.org/details/sim_gentes-herbarum_1946-03-25_7_1|via=Internet Archive}}</ref>{{rp|4–5}}

Among Asian and European columbines, differences in floral morphology and pollinators are lower between species than between North American species. However, there are approximately the same number of ''Aquilegia'' species across the three continents. This suggests that pollinator specialization played a dominant role in North American columbine speciation while habitat specialization was the primary driver of Asian and European columbine speciation.<ref name=Extended/>

The nectar spurs present in ''Aquilegia'' are an unusual evolutionary trait, arising on the ancestor of all ''Aquilegia'' approximately 5–7 million years ago.<ref name=POPOVICH/> In order to determine the gene responsible for the trait, a 2020 paper compared spurred ''Aquilegia'' taxa against the spurless ''A.&nbsp;ecalcarata''. This research identified a gene named ''POPOVICH'' (''POP''){{snd}} for the then-San Antonio Spurs coach Gregg Popovich due to the gene's regulatory role resembling a coach's management of a team<ref>{{cite news|url=https://www.eurekalert.org/news-releases/547270|title=The Popovich of Floral Nectar Spurs|work=EurekAlert!|publisher=American Association for the Advancement of Science|date=13 November 2020|access-date=1 October 2025}}</ref>{{snd}} as responsible for cell proliferation during the early stage of spur development. ''POP'', which encodes a C2H2 zinc-finger transcription factor, appeared at higher levels in the petals of the spurred ''Aquilegia'' studied than in ''A.&nbsp;ecalcarata''.<ref name=POPOVICH>{{cite journal |title=''POPOVICH'', Encoding a C2H2 Zinc-Finger Transcription Factor, Plays a Centeal Role in the Development of a Key Innovation, Floral Nectar Spurs, in ''Aquilegia''|journal=Proceedings of the National Academy of Sciences of the United States of America|publisher=National Academy of Sciences |volume=117|number=36|date=8 September 2020|jstor=26969162|last1=Ballerini|first1=Evangeline S.|last2=Min |first2=Ya|last3=Edwards|first3=Molly&nbsp;B.|last4=Kramer|first4=Elena&nbsp;M.|last5=Hodges|first5=Scott&nbsp;A.|pages=22552–22560|doi=10.1073/pnas.2006912117|pmid=32848061|pmc=7486772|bibcode=2020PNAS..11722552B|doi-access=free}}</ref>

===Current species=== {{Main|List of Aquilegia species}} According to different taxonomic authorities, the genus ''Aquilegia'' comprises between 70 and over 400 taxa (including species and subspecies).<ref>{{cite journal|url=https://pmc.ncbi.nlm.nih.gov/articles/PMC3091760/|title=Genomic Tools Development for ''Aquilegia'': Construction of a BAC-based Physical Map|first1=Guang-Chen|last1=Fang|first2=Barbara&nbsp;P.|last2=Blackmon|first3=David&nbsp;C.|last3=Henry|first4=Margaret&nbsp;E.|last4=Staton|first5=Christopher&nbsp;A.|last5=Saski|first6=Scott&nbsp;A.|last6=Hodges|first7=Jeff&nbsp;P.|last7=Tomkins|first8=Hong|last8=Luo|journal=BMC Genomics|date=8 November 2010|page=1|volume=11|number=621|pmid=21059242|doi=10.1186/1471-2164-11-621|doi-access=free}}</ref><ref name=Nardi/>{{rp|33}} Some totals correspond more closely with Munz's 1946 total of 67, while Tropicos records over 200 names{{snd}} both legitimate and illegitimate{{snd}} for taxa within the genus.<ref name=Nardi/>{{rp|33}} {{As of|2026|}}, the Royal Botanic Gardens, Kew's ''Plants of the World Online'' accepts 130 species.<ref name=POWO/>

The American gardener Robert Nold attributed the substantial total of named species, subspecies, and varieties to the 19th-century practice of assigning names to even minutely distinct specimens. However, Nold also held that overly broad species could increase the number of varietal names.<ref name=Nold/>{{rp|41–42}} The Italian botanist Enio Nardi stated that authors assessing ''Aquilegia'' as containing fewer than 100 species "either mask or underestimate their splitting into subspecies, many of which were originally described at the species level" and remain accepted as species in taxonomic indices.<ref name=Nardi/>{{rp|33}}

The type species of the genus is ''A.&nbsp;vulgaris'', a European columbine with high levels of physical variability.<ref name=Nold/>{{rp|124–125}}<ref name=Nardi>{{cite book|title=Il Genere ''Aquilegia'' L. (Ranunculaceae) in Italia/The Genus ''Aquilegia'' (Ranunculaceae) in Italy: Aquilegia Italicarum in Europaearum Conspectu Descriptio|first=Enio|last=Nardi|publisher=Edizioni Polistampa|date=2015|location=Florence|translator-last1=Coster-Longman|translator-first1=Christina|isbn=9788859615187}}</ref>{{rp|208}} Most European ''Aquilegia'' are morphologically similar to ''A.&nbsp;vulgaris'', sometimes to the point where visually discerning them from ''A.&nbsp;vulgaris'' is difficult. ''A.&nbsp;vulgaris'' is sometimes considered to encompass Iberian and North African columbines that are not accepted as separate species for reasons that Nardi said were founded in "tradition, more cultural than scientific".<ref name=Nold/>{{rp|35}}<ref name=Nardi/>{{rp|217}}

===Natural hybridization=== thumb|right|alt=Photo of an Aquilegia x miniana plant|''Aquilegia'' × ''miniana'' ''(pictured)'' is a naturally occurring hybrid of ''A.&nbsp;flavescens'' and ''A.&nbsp;formosa''. A lack of genetic and physiological barriers permits columbine hybridization across even distantly related species that possess substantial morphological and ecological differences.<ref name=Shift/><ref name=Munz/>{{rp|4–6}}<ref name=FNA/> In natural settings, hybrid columbines may occur wherever the natural ranges of multiple species come into contact.<ref name=Weber>{{cite book|title=Colorado Flora: Western Slope|first=William A.|last=Weber|author-link=William Alfred Weber|date=1987|publisher=Colorado Associated University Press|location=Boulder, CO|isbn=0870811673|lccn=87-70012|page=295}}</ref>

While artificial pollination has determined the extent of the genus's interfertility, breeding between plants within the same species is generally more common even in settings{{snd}} both natural and cultivated{{snd}} where multiple columbine species are present. A significant barrier to hybridization occurring naturally is the proclivity of pollinators to preferentially support infraspecific crossbreeding due to the pollinators' recognition of familiar flower typology.<ref name=Nardi/>{{rp|65}}

In North America, species with flowers adapted to hummingbird and hawkmoth pollination have far reduced natural hybridization with species that do not share these adaptations.<ref name=Nardi/>{{rp|65}} Still, hybridization and subsequent introgression occurs in North American columbines.<ref name=FNA/> Such hybridization across columbines with two different pollination syndromes can be driven by a third pollinator that does not show favoritism towards a particular pollination syndrome.<ref name=Nold/>{{rp|33}} In the instance of populations of hybrids between the yellow-flowered ''A.&nbsp;flavescens'' and red-flowered ''A.&nbsp;formosa'' in the northwestern United States, the resultant pink-flowering columbines were initially described as an ''A.&nbsp;flavescens'' variety and are now accepted as ''Aquilegia'' × ''miniana''.<ref>{{cite journal|url=https://phytokeys.pensoft.net/article/99170/|title=The Correct Name for an ''Aquilegia'' (Ranunculaceae) Hybrid of the Parentage ''Aquilegia flavescens'' × ''A.&nbsp;formosa''|first=Quentin C. B.|last=Cronk|journal=PhytoKeys|date=2023 |issue=220|doi=10.3897/phytokeys.220.99170|doi-access=free|issn=1314-2011|pages=31–38|pmid=37251608 |pmc=10209508 |bibcode=2023PhytK.220...31C}}</ref>

In China, clades distinguishing eastern and western ''A.&nbsp;ecalcarata'' populations indicate gene flow from different species. A study using genetic modeling indicated that the spurless ''A.&nbsp;ecalcarata'' may have developed from two separate mutations from discrete eastern and western populations of the spurred ''A.&nbsp;kubanica'', an instance of a hybrid parallel evolution.<ref name=Parallel/> Further hybridization between ''A.&nbsp;ecalcarata'' and the spurred columbines that share its range is limited by each species's selection for particular pollinators.<ref name=Nardi/>{{rp|65}} However, a short-spurred ''A.&nbsp;rockii'' phenotype has developed from hybridization with Western ''A.&nbsp;ecalcarata''.<ref name=Parallel>{{cite journal|title=Genomics of Hybrid Parallel Origin in ''Aquilegia ecalcarata''|journal=BMC Ecology and Evolution|first1=Fang-Dong|last1=Geng|first2=Miao-Qing|last2=Liu|first3=Xue-Dong|last3=Zhang|first4=Lu-Zhen|last4=Wang|first5=Meng-Fan|last5=Lei|date=2024 |volume=24|number=75|page=75 |doi=10.1186/s12862-024-02266-7|doi-access=free|pmid=38844857 |bibcode=2024BMCEE..24...75G }}</ref>

==Distribution== ''Aquilegia'' species have natural ranges spanning the Northern Hemisphere in Eurasia and North America.<ref name=POWO/> These ranges encompass the Circumboreal Region, the geographically largest floristic region in the world.<ref name=FNA/> The southern limits of the natural ''Aquilegia'' ranges are found in northern Africa and northern Mexico, with the only native African columbine being ''A.&nbsp;ballii'' of the Atlas Mountains.<ref name=FOP>{{cite web|url=http://legacy.tropicos.org/Name/40018252?projectid=32|title=''Aquilegia''|work=Flora of Pakistan|publisher=Missouri Botanical Garden|access-date=16 May 2025|via=Tropicos}}</ref><ref name=Nold/>{{rp|127}}

''A.&nbsp;vulgaris'', a European columbine which possibly originated in the Balkans,<ref name=Nardi/>{{rp|208}} has spread through both natural radiation and human assistance to become the most widely distributed ''Aquilegia'' species. Its range has expanded to include introduced populations that have sometimes become naturalized in Africa, Macaronesia, the Americas, and Oceania.<ref name=Nold/>{{rp|124}}<ref name=Nardi/>{{rp|207–208}} Naturalized populations of ''A.&nbsp;vulgaris'' that originated from gardens are the only columbines observed growing in the wild in Australia, with recorded observations in New South Wales, Victoria, and Tasmania.<ref>{{cite book|url=https://www.dcceew.gov.au/sites/default/files/env/pages/bf97195f-fe6d-4017-ac00-e16cd0249814/files/flora-australia-02-winteraceae-platanaceae.pdf|chapter=''Aquilegia'' L.|first=J. A.|last=Jeanes|title=Flora of Australia: Winteraceae to Platanaceae|series=Flora of Australia|volume=2|page=296|date=2007|editor-first=Annette&nbsp;J.&nbsp;G.|editor-last=Wilson|isbn=9780643059672|publisher=Australian Biological Resources Study and CSIRO Publishing|location=Melbourne}}</ref> The species is also present in Asia, with populations in the Russian Far East and Uzbekistan<ref>{{cite POWO|title=''Aquilegia vulgaris'' L.|id=306409-2|access-date=16 May 2025}}</ref> and where they also typically originated from ornamental cultivation.<ref name=Nardi/>{{rp|207}}

Some columbines are narrowly endemic, with highly restricted ranges. ''A.&nbsp;paui'' only has a single population with four subpopulations within a few kilometers of each other in the Ports de Tortosa-Beseit mountains of Catalonia.<ref name=Conservation/><ref name=Shift/> ''A.&nbsp;hinckleyana'' only natively populates a single location: the basin of Capote Falls, a waterfall in Texas.<ref name=Nold/>{{rp|94}}<ref>{{cite web|url=https://www.wildflower.org/plants/result.php?id_plant=aqchh|title=''Aquilegia chrysantha'' var. ''hinckleyana''|website=wildflower.org|publisher=Lady Bird Johnson Wildflower Center|access-date=15 May 2025}}</ref> {{As of|2005}}, the entire population of ''A.&nbsp;nuragica''{{snd}} estimated as 10 to 15 individuals{{snd}} occupied an area of approximately {{cvt|50|m2}} on Sardinia.<ref name=Top50>{{cite book|url=https://top50.iucn-mpsg.org/uploads/files/TOP50%20Med%20Island%20Plants%20English.pdf|archive-url=https://web.archive.org/web/20120405135852/http://data.iucn.org/dbtw-wpd/edocs/2005-025.pdf|archive-date=5 April 2012|title=The Top 50 Mediterranean Plants|editor-first1=Bertrand|editor-last1=de&nbsp;Montmollin|editor-first2=Wendy|editor-last2=Strahm|location=Gland, Switzerland|date=2005|publisher=International Union for Conservation of Nature|via=Archive.org|isbn=9782831708324|url-status=live}}</ref>{{rp|79}}

===Conservation=== Certain ''Aquilegia'' have been identified as having elevated risks of extinction, with some appearing on the IUCN Red List.<ref name=Nardi/>{{rp|29}} Two Sardinian columbines, ''A.&nbsp;barbaricina'' and ''A.&nbsp;nuragica'', have conservation statuses assessed by the IUCN as critically endangered and the same organization listed the species in their 2005 Top 50 Mediterranean Island Plants campaign for conservation.<ref name=Top50/>{{rp|76–79}} Some columbines, including both rare and common taxa, are subject to governmental regulations intended to protect individuals and populations. The conflation of taxa has sometimes resulted in uncertainty regarding their conservation statuses.<ref name=Nardi/>{{rp|29; 318}}

Humans pose a significant threat of impairing columbine population health and driving extinction. Beyond the desirability of the flowers for display, uncommon or rare ''Aquilegia'' face the risk of destruction by botanists and others seeking to add them to their herbariums or private collections.<ref name=Nardi/>{{rp|29}}

==Cultivation== thumb|alt=Photograph of purple columbines growing in a garden near the Washington National Cathedral|Some of the earliest cultivation of columbines may have occurred at Christian cathedrals. In Europe, cultivation of columbines may have begun over 1,700 years ago. Archaeobotanical evidence suggests that ''A.&nbsp;vulgaris'' was cultivated for ornamental purposes in 3rd-century&nbsp;AD Roman Britain.<ref name=Nardi/>{{rp|25}} Two ''A.&nbsp;vulgaris'' seeds in burnt waste pits, one at Alcester and another at Leicester, have been interpreted as evidence of their planting in Roman gardens.<ref>{{cite journal|url=https://img1.wsimg.com/blobby/go/c09070e7-cc09-40af-ae91-d4366196a62c/downloads/Circaea_12-2.pdf?ver=1740510670957|title=Evidence for Food and Fodder from Plant Remains at Causeway Lane, Leicester, U.K.|first=Angela|last=Mockton|journal=Circaea|date=1996|volume=12|number=2|pages=254–255}}</ref><ref name=Moffet>{{cite journal|url=https://img1.wsimg.com/blobby/go/c09070e7-cc09-40af-ae91-d4366196a62c/downloads/Circaea_5-2.pdf?ver=1775147326029|title=Gardening in Roman Alcester|first=Lisa|last=Moffett|date=1988|journal=Circaea|volume=5|number=2|pages=74–76}}</ref> Finds of columbines at a late Saxon site near Winchester Cathedral and three later medieval German sites may indicate the plants were used for gardening.<ref name=Moffet/> In 12th-century Italy, people may have supported ''A.&nbsp;vulgaris'' or ''A.&nbsp;atrata'' populations near religious structures, possibly due to the then-contemporary treatment of columbines as Christian symbols.<ref name=Nardi/>{{rp|25}}

''Aquilegia'' are grown for a variety of purposes. Cut flowers can be collected once the flower opens. The dried fruit can also be used in ornamental arrangements.<ref>{{cite web|url=http://www.gardening.cornell.edu/homegardening/scene37a5.html|title=Columbine, Hybrid|website=gardening.cornell.edu|publisher=Cornell University|access-date=17 October 2025}}</ref> Columbines function well in garden borders and can occupy shade and cottage gardens.<ref>{{cite web|url=https://plants.ces.ncsu.edu/plants/aquilegia-x-hybrida/|title=''Aquilegia x hybrida''|publisher=North Carolina State University|website=plants.ces.ncsu.edu|access-date=17 October 2025}}</ref> Certain columbines are suited for well-drained rock gardens or alpine gardens. A few, such as the dwarf woodland plant A.&nbsp;flabellata var.&nbsp;''pumila'', favor moist shaded plantings.<ref name=Nold/>{{rp|17}}<ref name=RHSHow>{{cite web|url=https://www.rhs.org.uk/plants/aquilegia/growing-guide|title=How to Grow Aquilegias|publisher=Royal Horticultural Society|access-date=17 October 2025}}</ref>

Lifespans for cultivated columbines are generally relatively short for perennials, with a plant's peak typically occurring in its second year.<ref name=AHS>{{cite book|title=Encyclopedia of Perennials|editor-last1=Rice|editor-first1=Graham|editor-last2=Bluemel|editor-first2=Kurt|publisher=American Horticultural Society, DK Publishing|location=New York|date=2006|isbn=9780756613433}}</ref>{{rp|64}} Two- to three-year-long lives are typical in cultivated ''A.&nbsp;coerulea'' and ''A.&nbsp;glandulosa'',<ref name=Munz/>{{rp|3}} with ''A.&nbsp;vulgaris'' exhibiting a biennial-like lifespan.<ref name=Nold/>{{rp|18}} Conversely, ''A.&nbsp;chrysantha'' and ''A.&nbsp;desertorum'' are particularly long-lived.<ref name=Munz/>{{rp|3}}<ref>{{cite web|url=https://cwelwnp.usu.edu/westernnativeplants/plantlist_view.php?id=58&name=aquilegiadesertorum|title=Desert Columbine in the Landscape|first=Stephen|last=Love|work=Native Plants for the Intermountain West|publisher=University of Wyoming Extension|via=Utah State University|access-date=15 May 2025}}</ref> In gardens, columbines will generally live three to four years. This lifespan can be extended by deadheading, where dead flowers are removed prior to a plant expending the energy needed to produce fruit.<ref name=Munz/>{{rp|3}}<ref name=Nold/>{{rp|18}}

In cultivation, the seasonal cycle that releases columbine seeds from dormancy can be replicated via a stratification where seeds are exposed to two to four weeks of cool temperatures prior to sowing.<ref name=Sowing/> Cultivated ''Aquilegia'' typically require well-draining soil. Poorly drained soil can result in the development of root rot, caused by either bacteria or fungi. At the end of the growth season, columbines can be protected from frost heaving by having their dead foliage removed to near the soil level and mulching once the ground is frozen.<ref name=Problem/>{{rp|149}} Vernalization{{snd}} a process by which juvenile plants are exposed to a weeks-long period of cold which mimics seasonal weather{{snd}} can accelerate the rate at which columbines reach flowering.<ref>{{cite magazine|url= https://www.canr.msu.edu/resources/aquilegia|title=Production Tips for Top Performers: ''Aquilegia vulgaris''|magazine=Greenhouse Grower|first1=Cathy|last1=Whitman|first2=Sonali|last2=Padhye|date=August 2008|pages=44–46|via=Michigan State University}}</ref>

If permitted, cultivated columbines drop numerous seeds around themselves, resulting in a rapid proliferation of seedlings. These seedlings can give the impression that short-lived plants are living longer.<ref name=Nold/>{{rp|18}} Due to their tendency towards hybridization and{{snd}} particularly in the case of F1 hybrids and cultivars (cultivated varieties){{snd}}inherent genetic diversity, the seeds of cultivated ''Aquilegia'' often do not produce plants true to their type.<ref name=AHS/>{{rp|64, 66}}<ref name=DK>{{cite book|title=Encyclopedia of Plants & Flowers|publisher=American Horticultural Society, DK|editor-first=Christopher|editor-last=Brickell|orig-date=1989|date=2011|isbn=978-0-7566-6857-0|location=New York City}}</ref>{{rp|226}} If identical plants are desired, basal cuttings can be performed in early summer.<ref name=RHSHow/> Propagation by division can be achieved by pulling plants apart in the late winter prior to new growth occurring. Artificial self-pollination can be performed by placing mesh cages around developing flowers to block access by pollinators. Pollen from the anther of a later-opening flower can be brushed onto the stigma of an earlier-opening flower. The cages can then be removed once the fruit begins developing.<ref name=Nold/>{{rp|27}}

Several animals are considered pests of cultivated columbines. Columbine leaf miners of the ''Phytomyza'' genus leave white patches or paths on leaves, but the damage is only cosmetic and does not generally require chemical pesticides.<ref name=NAPests/> The moths ''Papaipema lysimachiae'' and ''P.&nbsp;nebris'' (stalk borer) both adversely affect columbines; scraping the ground around impacted plants can destroy the moths' eggs. The larval stage of the ''Erynnis lucilius'' (columbine duskywing) is known as the ''columbine skipper'', and the larvae can chew leaves and bind them together with silk.<ref name=Problem/>{{rp|150}} Aphid infestation is another frequent issue, requiring rapid intervention to prevent significant destruction.<ref name=Nold/>{{rp|21}}

===Cultivars and cultivated hybrids=== thumb|right|alt=Aquilegia vulgaris Nora Barlow flowers|''A. vulgaris'' 'Nora Barlow' is a double-flowered cultivar known for its coloring. Columbine cultivars are popular among gardeners, particularly in the Northern Hemisphere.<ref name=Nardi/>{{rp|25}} The cultivars are frequently bred in series, with plants sharing traits but varying in color. Some series produce dwarf plants.<ref name=RHSHow/> ''A.&nbsp;vulgaris'' has been the basis for many cultivars, distinct in foliage, flower shape, and possession of fragrance.<ref name=AHS/>{{rp|64}} The single-flowering ''A.&nbsp;vulgaris'' cultivar 'Nivea' (also known as 'Munstead White')<ref name="Perennials">{{cite book|title=Perennials for American Gardens|first1=Ruth Rogers|last1=Clausen|first2=Nicolas H.|last2=Ekstrom|publisher=Random House|date=1989|location=New York|isbn=0394557409|lccn=88-43209}}</ref>{{rp|52}} received the Royal Horticultural Society's Award of Garden Merit.<ref>{{cite web|url=https://www.rhs.org.uk/Plants/112596/i-Aquilegia-vulgaris-i-Nivea/Details|title=RHS Plant Selector - Aquilegia vulgaris 'Nivea' |publisher=Royal Horticultural Society|access-date=13 January 2018}}</ref><ref name="Hogan" />{{rp|167}}

Double-flowered columbines were developed from ''A.&nbsp;vulgaris'' and can be classified into three types. The Flore Pleno group, described in the English herbalist John Gerard's 1597 book ''Herball'', possesses plants where the flowers are elongated and the petals are rounded. The Vervaeneana group come in several colors of flower and possess marbled green and gold foliage. The Stellata group, described in the English botanist John Parkinson's 1629 book ''Paradisi in Sole Paradisus Terrestris'', has flowers which are star-shaped and have pointed petals. The three-colored, double-flowered cultivar 'Nora Barlow'{{snd}} first discovered by the botanist and geneticist Nora Barlow{{snd}} is sometimes classified as part of the Stellata group, but displays a greater quantity of particularly narrow sepals than other members of that group.<ref name=AHS/>{{rp|64}}<ref name=Hogan>{{cite book|title=Flora: A Gardener's Encyclopedia|editor-first=Sean|editor-last=Hogan|publisher=Timber Press|location=Portland, OR|date=2003|isbn=0881925381|page=167}}</ref>

Experimental hybridization attempts have determined that the degree of interfertility of columbines is not identical across the genus. While North American columbines easily hybridize with each other and most Eurasian ''Aquilegia'', the Asian species ''A.&nbsp;oxysepala'' and ''A.&nbsp;viridiflora'' resist hybridization with North American columbines. Hybrids with ''A.&nbsp;vulgaris'' as one parent frequently look closer to ''A.&nbsp;vulgaris'' than any other parent. Many older garden columbines that are suspected hybrids resemble ''A.&nbsp;vulgaris'', making true parentage difficult to determine. According to the American gardener Robert Nold, "more has been written about 'Hensol Harebell' [a hybrid of ''A.&nbsp;alpina'' and ''A.&nbsp;vulgaris'' which originated in early 20th-century Scotland] than perhaps any other interspecific hybrid". Nold noted the variety's great affinity to ''A.&nbsp;vulgaris'' in appearance.<ref name=Nold/>{{rp|132–133, 137}}

More recent garden hybridization work has primarily utilized North American columbines for their greater flower sizes and variety of colors. While columbine flowers often face downwards in a nodding arrangement, hybridizers have introduced the ''A.&nbsp;coerulea'' trait of upward-facing flowers. The first such hybrids were the McKana Group (also known as McKana's Giants){{snd}} which won the 1935 All-America Selections bronze medal for new seed offerings<ref name=Nold/>{{rp|138}}{{snd}} and Mrs. Scott-Elliot hybrids. However, these large-flowered columbines produced unpredictably colored flowers. Newer F1 hybrids series have larger blooms with greater uniformity in color.<ref name=AHS/>{{rp|64–65}}

==Human uses== ===Medicinal and herbal=== thumb|right|alt=Photograph of a Aquilegia oxysepala flower|''A.&nbsp;oxysepala'' has been a part of traditional Chinese medicine for thousands of years. Asian traditional medicine practitioners, Indigenous North Americans, and medieval Europeans have treated columbine plants as medicinal herbs. Modern scientific research has determined that columbines can possess antioxidant, antibacterial, and anti-cancer qualities.<ref name=Nardi/>{{rp|27}}

In China, ''A.&nbsp;oxysepala'' has been used as a dietary supplement and medicine for thousands of years. ''A.&nbsp;oxysepala'' has been used there to treat diseases in women such as irregular menstruation and intermenstrual bleeding. While its extract's function as an antioxidant is known, with its medicinal use possibly attributable to the extract's good scavenging of superoxide anion radicals, it is inferior to the common dietary supplement ascorbic acid.<ref name=AOExtract/> Research has also determined ''A.&nbsp;oxysepala'' to possess antibacterial qualities.<ref name=Nardi/>{{rp|27}}

''A.&nbsp;sibirica'' has been a significant part of Asian traditional medicine, including traditional Mongolian medicine, and this plant has been used to treat diseases in women, asthma, rheumatism, and cardiovascular diseases. Scientific research has demonstrated the species can inhibit ''Staphylococcus aureus'', one of the bacteria responsible for staphylococcal infections. ''A.&nbsp;sibirica'' also possesses antifungal qualities. Extracts showed the presence of chlorogenic acid and caffeic acid, with the former known to inhibit the pathogenic yeast ''Candida albicans''.<ref name=Medicinal>{{cite journal|title=Antifungal Activity of Mongolian Medicinal Plant Extracts|journal=Natural Product Research|volume=34 |number=4|date=2020|doi=10.1080/14786419.2019.1610960|url=https://www.tandfonline.com/doi/abs/10.1080/14786419.2019.1610960 |last1=Giordani |first1=Cristiano|last2=Simonetti|first2=Giovanna|last3=Natsagdorj|first3=Damdinsuren|last4=Choijamts|first4=Gotov|last5=Ghirga|first5=Francesca|last6=Calcaterra|first6=Andrea|last7=Quaglio|first7=Deborah|last8=De Angelis |first8=Giulia|last9=Toniolo|first9=Chiara|last10=Pasqua|first10=Gabriella|pages=449–455|pmid=31135192|s2cid=167220925|access-date=15 May 2025|archive-date=13 August 2023 |archive-url=https://web.archive.org/web/20230813212628/https://www.tandfonline.com/doi/abs/10.1080/14786419.2019.1610960|url-status=live}}</ref>

Some Indigenous North American peoples used the roots of columbines to treat ulcers.<ref name=FSAF/> North American peoples have used ''A.&nbsp;canadensis'' and ''A.&nbsp;chaplinei'' as an aphrodisiac.<ref name=CE>{{cite encyclopedia|url=https://www.thecanadianencyclopedia.ca/en/article/columbine|title=Columbine|encyclopedia=The Canadian Encyclopedia|first=Roger|last=Vick|publisher=Historica Canada|location=Toronto|date=6 February 2006|access-date=15 May 2025}}</ref><ref>{{cite book|url=https://www.degruyter.com/document/doi/10.7560/727205-007/html|chapter=5|title=The Guadalupe Mountains of Texas|first1=Michael|last1=Allender|first2=Alan|last2=Tennant|publisher=University of Texas Press|location=Austin, TX|date=1997|orig-date=1980|page=78|isbn=029270481X|lccn=80-16821|doi=10.7560/727205-007|url-access=subscription}}</ref><ref name=Nold/>{{rp|38}} Crushed ''A.&nbsp;canadensis'' seeds were used as a perfume, and the plant was thought to be capable of detecting bewitchment.<ref name=Nold/>{{rp|74}} The Goshute people reportedly chewed ''A.&nbsp;coerulea'' seeds or utilized the plant's root for medicinal or therapeutic purposes.<ref>{{cite web|last1=Whittemore|first1=Alan T.|title=''Aquilegia coerulea''|url=http://floranorthamerica.org/Aquilegia_coerulea |website=Flora of North America|access-date=26 May 2025|date=6 November 2020}}</ref>

===Other uses=== Before deaths from overdoses were reported, small quantities of flowers from several columbines species were considered safe for human consumption and were regularly eaten as colorful garnishes and parts of salads.<ref name=CE/><ref name=FSAF>{{cite web|url=https://www.fs.usda.gov/wildflowers/plant-of-the-week/Aquilegia-flavescens.shtml|title=Plant of the Week: Yellow Columbine (''Aquilegia flavescens'')|first=Charmaine|last=Delmatier|publisher=United States Forest Service|website=fs.usda.gov|date=2014|access-date=18 May 2025}}</ref> Several Indigenous North American peoples have been described as eating ''A.&nbsp;formosa'': the Miwok may have boiled and eaten them with early spring greens, while Hanaksiala and Chehalis children may have sucked nectar from the flowers.<ref>{{cite web|url=https://depts.washington.edu/hortlib/pal/on-the-edibility-of-columbine-flowers/|title=On the Edibility of Columbine Flowers|publisher=Elisabeth C. Miller Library, University of Washington|website=depts.washington.edu/hortlib|access-date=18 May 2025}}</ref> Columbine flowers are described as sweet,<ref name=FSAF/> a flavor attributed to their nectar.<ref>{{cite web|url=https://pfaf.org/user/Plant.aspx?LatinName=Aquilegia+shockleyi|title=Aquilegia shockleyi - Eastw.|website=pfaf.org|publisher=Plants for a Future|access-date=18 May 2025}}</ref>

The American botanist Verne Grant repeatedly utilized ''Aquilegia'' in research published between the 1950s and the 1990s to explain the role that hybridization, polyploidy, and other processes played in how plant evolution and speciation occur. Among Grant's works that utilized ''Aquilegia'' to illustrate evolutionary patterns and processes was his influential 1971 book ''Plant Speciation''.<ref name=Studies>{{cite journal|title=Verne Grant and Evolutionary Studies of ''Aquilegia''|first1=Scott A.|last1=Hodges|first2=Michelle|last2=Fulton|first3=Ji&nbsp;Y.|last3=Yang|first4=Justen&nbsp;B.|last4=Whittall|journal=New Phytologist|volume=161|issue=1|date=January 2004|pages=113–120|doi=10.1046/j.1469-8137.2003.00950.x|doi-access=free|bibcode=2004NewPh.161..113H }}</ref> The five species groups that Grant proposed in 1952 remain foundational elements for understanding the phylogeny of columbines.<ref name=Nardi/>{{rp|97}}

In 21st-century scientific research of plant development, ecology, and evolution, ''Aquilegia'' has been considered a model system for studying the relevant mechanisms.<ref name=East/> Utilizing the genome sequence of ''A.&nbsp;coerulea'', a 2019-published study examined polyploidy during the evolution of eudicots, a clade in which columbines are considered a basal member. This research determined that columbines and all eudicots experienced a shared tetraploidy, but that only core members of the eudicots clade (which excludes columbines) experienced a shared hexaploidy.<ref name=Eudicots>{{cite journal|title=The ''Aquilegia'' Genome Reveals a Hybrid Origin of Core Eudicots|journal=Genome Biology|first1=Gökçe|last1=Aköz|first2=Magnus|last2=Nordborg|date=28 November 2019|volume=20|number=256|page=256 |pmid=31779695|doi=10.1186/s13059-019-1888-8|doi-access=free|pmc=6883705 }}</ref>

==In culture== thumb|right|alt=Detail of the painting The Garden of Earthly Delights containing a columbine flower|''A. vulgaris'' serves as a symbol of bodily pleasures within Hieronymus Bosch's ''The Garden of Earthly Delights'' ''(detail pictured, with blue columbines in the center)''. European columbines have been assigned several meanings since the ancient period. Within art, ''A.&nbsp;vulgaris'' has been a symbol of both moral and immoral behaviors, as well as an ornamental motif.<ref name=Nardi/>{{rp|19–23}}<ref name=Symbolism>{{cite journal|url=https://academic.oup.com/jxb/article/60/6/1535/519453|title=Symbolism of Plants: Examples from European-Mediterranean Culture Presented with Biology and History of Art: May: Columbine|journal=Journal of Experimental Botany|volume=60|issue=6|date=April 2009|first1=Riklef|last1=Kandeler|first2=Wolfram R.|last2=Ullrich|pages=1535–1536|doi=10.1093/jxb/erp087|pmid=19363205 |via=academic.oup.com}}</ref> In ancient Greece and ancient Rome, the spurs of columbines were interpreted as phallic and the plants were associated with the fertility goddesses Aphrodite and Venus.<ref name=Symbolism/> By the early 17th century, columbines were viewed as symbols of cuckoldry due to their horn-shaped nectar spurs.<ref name=Nold/>{{rp|38}}<ref name=NQ/> In English literature, columbines have been mentioned with negative connotations.<ref name=Nold/>{{rp|38}} In ''Hamlet'', an Elizabethan drama by William Shakespeare, Ophelia presents King Claudius with flowers that include columbines,<ref name=Dwyer>{{cite journal|title=Garden Plants and Wildflowers in Hamlet|first=John|last=Dwyer|volume=24|number=2|journal=Australian Garden History|date=2012 |pages=5–8, 34|jstor=24918848}}</ref> where columbines are symbolic of cuckoldry, deception, and impending death.<ref name=Nardi/>{{rp|21}}<ref name=NQ>{{cite journal|url=https://go.gale.com/ps/i.do?id=GALE%7CA15103323&sid=googleScholar&v=2.1&it=r&linkaccess=abs&issn=00293970&sw=w&u=nu_main&p=AONE&aty=ip|title=Ophelia's Flowers Again|journal=Notes and Queries|volume=41|number=1|page=42|first1=Robert|last1=Painter|first2=Brian|last2=Parker|date=March 1994}}</ref>

Medieval European artists associated columbines with Christian sacredness and sublimity, with Flemish painters of the 15th century frequently depicting them in prominent locations within their Christian artworks. In ''The Garden of Earthly Delights'' ({{circa|1503–1504}}) by Hieronymus Bosch, ''A.&nbsp;vulgaris'' serves as a symbol for bodily pleasures. ''Portrait of a Princess'' (1435–1449) by Pisanello depicts multiple ''A.&nbsp;atrata'' at different angles as part of the floral ornamentation that makes that painting characteristic of the international Gothic style.<ref name=Nardi/>{{rp|19–21}}

Columbines have several meanings in the language of flowers, a manner of communicating using floral displays. In the 1867 English book ''The Illustrated Language of Flowers'' by a "Mrs.&nbsp;L.&nbsp;Burke", columbines are generally described as communicating "folly". The same book identifies purple columbines with "resolve to win" and red columbines with "anxious and trembling".<ref>{{cite book|url=https://archive.org/details/illustratedlang00burka/page/16/mode/2up|title=The Illustrated Language of Flowers|first=L.|last=Burke|location=London|publisher=George Routledge & Sons|date=1867|page=16|via=Archive.org}}</ref> Columbines, due to their resemblance to doves, have been associated with the Holy Spirit in Christianity since at least the 15th century.<ref>{{cite journal|title=The Cover Design|journal=The Library Quarterly|first=Evelyn A.|last=Walker|date=October 2003|volume=73 |issue=4 |pages=466–468|doi=10.1086/603442 |jstor=4309687}}</ref><ref name=Symbolism/><ref>{{cite web|url=https://libguides.nybg.org/languageflowers|title=The Language of Flowers: Home|website=libguides.nybg.org|publisher=LuEsther T. Mertz Library, New York Botanical Garden|location=New York City|access-date=18 May 2025}}</ref>

thumb|right|upright=.75|alt=Photograph of an Aquilegia coerulea flower|''A. coerulea'' is the state flower of Colorado. ''A.&nbsp;coerulea'' is the state flower of Colorado.<ref name=American>{{cite magazine|title=All-American Columbines|first=Robert|last=Nold|magazine=Horticulture|volume=101|issue=2|date=April 2004|pages=52–57}}</ref> The Colorado General Assembly passed legislation in 1925 making it illegal to uproot ''A.&nbsp;coerulea'' and puts limits on how many buds, blossoms, and stems may be picked from the species by a person; these restrictions apply only on public lands.<ref name=Colorado>{{cite encyclopedia|url= https://coloradoencyclopedia.org/article/state-flower|title=State Flower|encyclopedia=Colorado Encyclopedia|location=Denver|publisher=History Colorado|access-date=18 May 2025}}</ref> The name of the town of Columbine Valley, Colorado, ultimately derives from the state flower.<ref>{{cite work|url=https://columbinevalley.org/wp-content/uploads/2020/01/Town-of-Columbine-Valley-2020-Master-Plan.pdf|title=Town of Columbine Valley Master Plan 2020|publisher=Town of Columbine Valley|date=2020|page=9}}</ref> Colorado's ''A.&nbsp;coerulea'' also served as the inspiration for one of the state's official songs, "Where the Columbines Grow", written in 1909.<ref>{{cite work|url=https://i2i.org/wp-content/uploads/2015/10/IP-3-2015b-Columbines-1.pdf|title=Reclaiming the Centennial State’s Centennial Song: The Facts About "Where the Columbines Grow"|first=Robert G.|last=Natelson|date=September 2015|pages=8, 10|publisher=Independence Institute}}</ref> Following the 1999 Columbine High School massacre in Columbine, Colorado, the columbine became a symbol of remembrance in Colorado.<ref>{{cite book|title=Comprehending Columbine|first=Ralph W.|last=Larkin|author-link=Ralph Larkin|date=2007|publisher=Temple University Press|location=Philadelphia|isbn=9781592134922|lccn=2006020365|pages=115–116}}</ref> {{As of|2023}}, a specialty license plate depicting a columbine in honor of the victims and survivors of the massacre was the most popular specialty plate in Colorado.<ref>{{cite news|url=https://www.cpr.org/2023/01/30/colorado-dmv-specialty-license-plates/|title=Colorado Has More Than 200 Specialty License Plates. Here's What They Mean and How to Get One|first=May|last=Ortega|date=30 January 2023|access-date=26 February 2026|website=cpr.org|publisher=Colorado Public Radio}}</ref> The columbine was also used in the heraldry of the former city of Scarborough in the Canadian province of Ontario.<ref>{{Cite web|url=https://www.gg.ca/en/heraldry/public-register/project/606|title=City of Scarborough|publisher=Governor General of Canada|website=gg.ca|access-date=31 May 2025}}</ref>

The asteroid 1063 Aquilegia was named for the genus by the German astronomer Karl Reinmuth. He submitted a list of 66 newly named asteroids in the early 1930s, including a sequence of 28 asteroids that were all named after plants, in particular flowering plants.<ref>{{cite book|title=Dictionary of Minor Planet Names|last=Schmadel|first=Lutz D.|publisher=Springer Berlin Heidelberg|location=Berlin|page=91|date=2007|isbn=978-3-540-00238-3|doi=10.1007/978-3-540-29925-7_1064|chapter=(1063) Aquilegia}}</ref><ref>{{cite book|title=Dictionary of Minor Planet Names|last=Schmadel|first=Lutz D.|publisher=Springer Berlin Heidelberg|page=90|date=2007|isbn=978-3-540-00238-3|location=Berlin|doi=10.1007/978-3-540-29925-7_1055|chapter=(1054) Forsytia}}</ref>

==References== {{reflist}}

==External links== *{{Commons category-inline|Aquilegia}}

{{Portal bar|Biology|Plants|Medicine}} {{Taxonbar|from=Q147641}}

Category:Aquilegia Category:Garden plants Category:Medicinal plants Category:Ranunculaceae genera Category:Taxa described in 1753 Category:Botanical taxa named by Carl Linnaeus