Aquilegia typically possess stiff stems and leaves that divide into multipleleaflets. Columbines often have colorful flowers with fivesepals and fivepetals. The petals generally featurenectar spurs which differ in lengths between species. In North America, morphological variations in spurs evolved to suit differentpollinators. Some species and varieties of columbines are naturally spurless. In cultivation, varieties bearing significantly altered physical traits such asdouble flowering are prevalent.
Associated with fertility goddesses inancient Greece andancient Rome, archeological evidence suggestsAquilegia were in cultivation by the 2nd century AD inRoman Britain. Despite often being toxic, columbines have been used by humans as herbal remedies, perfume, and food. Asiantraditional medicine,Indigenous North Americans, andMedieval Europeans have considered portions of the plants to have medicinal uses. Selective breeding and hybridization of columbines has occurred for centuries, with exchanges betweenOld andNew World species creating further diversity.
The 1st-century AD Greek writerDioscorides called columbinesIsopyrum, a name used presently applied toanother genus.[2]: 38 In the 12th century, the abbess and polymathHildegard of Bingen referred to the plants asagleya – from which the genus's name in German,Akelei, derives. The first use ofaquilegia with regards to columbines was in the 13th century byAlbertus Magnus. In the 15th and 16th centuries, the namesColombina,Aquilina, andAquileia came into use. With the Swedish biologistCarl Linnaeus's 1753Species Plantarum, the formal name for the genus becameAquilegia, though limited use ofAquilina persisted in scientific usage until at least 1901.[3]: 23
Several scientific and common names for the genusAquilegia derive from its appearance.[3]: 23 The genus nameAquilegia may come from the Latin word for "eagle",aquila, in reference to the petals' resemblance to eagle talons.[4] Another possible etymology forAquilegia is a derivation from the Latinaquam legere ("to collect water"),[5]aquilegium (a container of water), oraquilex ("dowser" or "water-finder") in reference to the profusion of nectar in the spurs.[6][2]: 37–38 The most common English-language name,columbine, likely originates in thedove-like appearance of thesepals (columba being Latin fordove).[2]: 38 [7]
There are a number of other common names forAquilegia across different languages. In English, these includegranny's bonnet for some plants in the speciesAquilegia vulgaris.[8] The association ofcolumbine with the 1999Columbine High School massacre has led to use of the namesaquilegia andgranny's bonnet used in horticultural circles as a means of avoiding the stigmatized name.[9] In French, the wordancolie is the common name forAquilegia,[8] while individual members of the genus have been calledgants-de-Notre-Dame ("Our Lady's glove"). In Italian,amor-nascoto ("love-born") has been used.[2]: 38–39
Aquilegia is agenus ofherbaceous,perennial flowering plants in the familyRanunculaceae (buttercups). The genus is highly variable in appearance.[2]: 29 [8] Though they are perennials, certain species are short-lived, with some exhibiting lifespans more similar tobiennials and others only flourishing for six to eight years.[2]: 18 [10] Following adormant period in the winter, columbines will grow foliage and have a brief flowering period.[11] Some columbines bloom the first year after sowing, others will bloom in their second.[12] Later,seed heads will emerge and split, sowing new seed. The foliage lives through the summer before wilting and dying going into the fall.[11]
Aquilegia plants grow from slim, woodyrootstocks that comprise the perennial portion of the plants. One or more annualaerial stems rise from the rootstocks each growing season, ultimately dry out following fruiting.[3]: 143–145 Leaves can grow in bothbasal (from the base of the plant) andcauline (from the aerial stem) arrangements. Leaves emanating from closer to the plant's core are generally borne on flexiblepetioles, while leaves further from the core generally lack petioles. Thecompound leaves ofAquilegia are generally ternate (each leaf dividing in threeleaflets), biternate (each leaf dividing into three components that in turn each bear three leaflets, for a total of nine leaflets), or triternate (each leaf divides into three components three times, for a total of 27 leaflets).[8]
The flowering stems emerge from rosettes during the spring and summer.[13] Eachinflorescence appears at the terminus of an aerial stem and can reach 30 cm (12 in) long. Depending the species, an inflorescence will feature one to ten of eithercymes (flower clusters) or solitary flowers.[8] Flower morphology varies across the genus, but all columbine flowers emerge from buds that are initially nodding.[2]: 31 Flowers can be monochromatic or display multiple colors.[13] 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 onlyA. 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.[2]: 33
Theperianth (non-reproductive portion) ofAquilegia flowers generally comprise fivesepals that look likepetals and five petals.[14] Each petal typically comprises two portions: a blade, which are broad and project towards the front of the flower, and anectar spur, anectar-bearing structure which projects backwards.[15]: 376 [2]: 31 The hollow spurs attract pollinators and give columbine flowers a distinctive appearance.[16]: 147 Depending on the species, spurs can have a hooked, horn-like appearance, with straight to coiled spurs also present in the genus.[2]: 31, 38 [7] Some columbines, such asA. ecalcarata, are naturally spurless.[2]: 82 Recessive spurlessness individuals and populations can also be found within typically spurred species.[2]: 31 [8][5]
The reproductive portions of columbine flowers comprise thestamen (male) andgynoecium (female). The stamina, which bear theanthers from whichpollen emerges, formwhorls of five around the gynoecia. The total number of stamen varies between species.[14] There are generally scale-shapedstaminodes between the stamen and femalepistil structures.[8] The flowers undergo three stages ofanthesis: a premale stage, where the flower perianth is open but the anthers are notdehisced (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.[17]
The fanning foliage ofA. flabellata 'Alba'
Aquilegia arebisexual (featuring both male and female organs) and capable ofself-pollination, through either or bothautogamy (does not require assistance from pollinators) andgeitonogamy (requires pollinators).[14] Autogamy has been observed as the primaryfertilization mechanism inA. paui.[17][18]A. formosa andA. eximia may exhibit adichogamy, where male and female organs do not operate simultaneously to prevent self-fertilization. Fertilization viacross pollination also occurs inAquilegia, with pollinators carryingpollen from one flower to thestigma of another.[17]
Aquilegia fruit arefollicles. These follicles have a split on one side and terminate with a curling tip known as abeak.[2]: 32 Columbine seeds are generally obovoid with black, smooth exteriors.[8] Columbine seeds are in adormant state at the point of sowing.[19] In cold storage,A formosa seeds have remained viable for least three years.[20] Seedgermination 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.[19][21]
Thechromosome number for columbines is2n=14.[8][22] Individual plants have been recorded with other anomalous chromosome numbers, ranging up to 2n=32. It is possible thatB chromosomes impact thephenotype and the fertility of individual plants that possess them.[3]: 183–185
A. vulgaris fruit and seeds. Human consumption of columbine seeds and roots can introduce toxins that negatively impact heart health.
Amongcyanophore (organisms that produces a blue color)Aquilegia likeA. vulgaris, thecyanogenic glycosides compoundsdhurrin and triglochinin have been observed. Cyanogenic glycosides generally taste bitter and can be toxic to animals and humans. Ingestion of 20 g (0.71 oz) of freshA. vulgaris leaves by a human was observed as causing convulsions, respiratory distress, and heart failure. A child who consumed 12A. vulgaris flowers experienced weakness of the limbs,cyanosis, drowsiness, andmiosis; all symptoms abated after three hours.[23] Mature seeds and roots contain toxins that, if consumed, are perilous to human heart health.[24]
In flowering plants, the presence ofphenylpropanoids can serve as protection fromultraviolet (UV) light and as a signaling mechanism towards pollinators. A study that examinedA. formosa flowers determined that the petals and sepals had uniform levels of UV-resistant phenylpropanoids.[27]
Leaf miners can leave trails where they have consumed the interior tissue ofAquilegia leaves
Despite its toxicity and in the absence of incentives,[23] some animals consume the fruit and leaves of columbines. Consumption by mammals is not considered a component of theAquilegia reproductive cycle but has been observed. In the case of the endangeredA. paui, one study found that 30% of all fruit was lost to predation by theSoutheastern Spanish ibex.[3]: 27 [18][28]
Also in North America, three species ofPhytomyzaleaf miners lay their eggs onAquilegia:P. aquilegivora in theMidwest,P. aquilegiana in the east, andP. columbinae in the west. Collectively known as the columbine leaf miners, white trails or splotches on leaves indicate where thelarvae consumed the tissue between the leaves' surfaces. The larvae will cut through the leaves,pupating in small puparium on the leaves' undersides. Adults pierce the leaves with theirovipositors to access liquids in the plants, leaving marks.[31] AnotherPhytomyza columbine leaf miner,P. ancholiae, is native to France.[32]
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 incocoons.[31]
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.
Following the evolution of the genus,Aquilegia developed diverse floral features including varied orientation, coloration, and nectar spur morphology to attract differentpollinators, contributing tospeciation.[34] The suite of floral traits that develop to attract a particular set of pollinators are collectively referred to as apollination syndrome.Aquilegia flowers are traditionally divided into three pollination syndromes: bumblebees, hummingbirds, or hawkmoths, each of which are attracted by the plants' nectar. In cases where pollinators are scarce, columbines may adopt autogamy as a primary fertilization method, such as inA. paui.[14]
Eurasian columbines are primarily pollinated by flies, bees, andbumblebees. North American columbines are generally pollinated by bees, bumblebees,hawkmoths, andhummingbirds. Pollination by hummingbirds is more typical to red-flowered North AmericanAquilegia, while pale-flowered columbines may have developed to increase their visibility to hawkmoths in twilight.[2]: 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.[35] Hawkmoths often possess long tongues, permitting them to reach deeper into nectar spurs. The elongated nectar spurs on some columbines prevent hawkmoths from removing nectar from the spurs without also making contact with the reproductive organs of the flower.[36] While hawkmoths are present in Eurasia, there are not Eurasian columbines with the hawkmoth pollination syndrome which includes longer spurs. In North America, the presence of hummingbirds – which are absent in Eurasia and possess tongue lengths that are generally intermediate between other pollinators and hawkmoths – may have functioned as a stepping stone that allowed North AmericanAquilegia to evolve the hawkmoth pollination syndrome.[35]
While a given population ofAquilegia may settle a particular habitat and develop pollination syndromes for certain pollinators, this does not necessarily translate intoecological 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 ofA. micrantha var. mancosana.[3]: 79
Columbines are most commonly assigned to thetribe Isopyreae, though they are sometimes placed within Aquilegieae. The placement of the tribe containingAquilegia has been uncertain, with alternating assignments to twosubfamilies:Thalictroideae and Isopyroideae. Regardless of the placement,Aquilegia forms abasal,paraphyletic group with the generaIsopyrum andThalictrum (meadow-rues) which is characterized by theirplesiomorphy (characteristics shared between clades from theirlast common ancestor) with Berberidaceae.[3]: 31 When placed within themonophyletic Thalictroideae,Aquilegia are the second largest genus in the subfamily in terms oftaxa (described species andsubspecies), behindThalictrum. Columbines are nested in one of the three major clades in the subfamiliy, a clade it shares withSemiaquilegia andUrophysa.Semiaquilegia andAquilegia aresister genera.[38][39]
The broadly acceptedcircumscription ofAquilegia was established by the American botanistPhilip A. Munz in his 1946 monographAquilegia: The Cultivated and Wild Columbines. The only element of Munz's circumscription which has been substantially contended is his inclusion of the spurless Asian speciesA. ecalcarata, which is sometimes instead segregated into the closely related genus of spurless-floweredSemiaquilegia;Semiaquilegia ecalcarata remains the species's common name in cultivation.[3]: 31 [2]: 82 Another spurless columbine,A. micrantha var. mancosana, was also once reassigned toSemiaquilegia. Reassignments toIsopyrum andParaquilegia, such asP. anemonoides in 1920, have been more permanent.[2]: 38, 46–49
Phylogeny of the genusAquilegia, based on Fior et. al. 2013. Some names used by that study are now considered synonyms; the names accepted byPlants of the World Online for those taxa are used here.
There are no good fossils of columbines and other Thalictroideae that indicate how they evolved and radiated.[40][41][38] Genetic evidence suggests that the last common ancestor among Thalictroideae lived in East Asia approximately 36 million years ago, during thelate Eocene.[38]
A 2018 study of genetic evidence indicated thatAquilegia first appeared during theUpper Miocene approximately 6.9 million years ago. The genus split into twoclades 4.8 million years ago, with one clade eventually populating North America and the other radiating across Eurasia.[42] A 2024 study found found that the divergence betweenUrophysa,Semiaquilegia, andAquilegia instead occurred over a relatively short 1 million-year-long period approximately 8 to 9 million years ago.[38] The species is thought to have originated in the mountainous portions of south-central Siberia.[39]
Studies ofAquilegia genetics indicated that North AmericanAquilegia species shared theirlast common ancestor with species from the Asian Far East between 3.84 and 2.99 million years ago. This analysis corresponded with the theory thatAquilegia reached North America via aland bridge over the Bering Strait.[43] 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 preventedAquilegia migration through the region.[39]
Genetic information suggests that thediversification rates of columbines rapidly increased about 3 million years ago,[38] with indications of two independent radiation events occurring around that time: one in North American columbine populations and the other in European populations.[39] Despite the rapid evolution of substantial physical differences across species, genetic divergence remains minimal. This, combined with the presence of relatively few physiological barriers tohybridization, has resulted in columbines displaying exceptionally high degrees of interfertility.[18][44]: 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 ofAquilegia 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.[39]
The nectar spurs present inAquilegia are an unusual evolutionary trait, arising on the ancestor of allAquilegia up to approximately 7 million years ago.[34] In order to determine the gene responsible for the trait, a 2020 paper compared spurredAquilegia taxa against the spurlessA. ecalcarata. This research identified a gene namedPOPOVICH (POP) – named for theSan Antonio Spurs coachGregg Popovich[45] – as responsible for cell proliferation during the early stage of spur development.POP, which encodes aC2H2 zinc-fingertranscription factor, appeared at higher levels in the petals of the spurredAquilegia studied than inA. ecalcarata.[46][34]
The American botanist and gardener Robert Nold attributed the substantial total of named species, subspecies, andvarieties 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.[2]: 41–42 The Italian botanist Enio Nardi stated that authors assessingAquilegia 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.[3]: 33
Thetype species of the genus isA. vulgaris, a European columbine with high levels of physical variability.[2]: 124–125 [3]: 208 Most EuropeanAquilegia are morphologically similar toA. vulgaris, sometimes to the point where visually discerning them fromA. vulgaris is difficult.A. 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".[2]: 35 [3]: 217
Aquilegia ×miniana(pictured) is a naturally occurring hybrid ofA. flavescens andA. formosa.
A lack of genetic and physiological barriers permits columbinehybridization across even distantly related species with high degrees of morphological and ecological differences.[18][44]: 4–6 [8] In natural settings, hybrid columbines may occur wherever the natural ranges of multiple species come into contact.[5]
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 – both natural and cultivated – 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.[3]: 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.[3]: 65 Still, hybridization and subsequentintrogression occurs in North American columbines.[8] Such hybridization across columbines with two different pollination syndromes can be driven by a third pollinator that do not show favoritism towards a particular pollination syndrome.[2]: 33 In the instance of populations of hybrids between the yellow-floweredA. flavescens and red-floweredA. formosa in the northwestern United States, the resultant pink-flowering columbines were initially described as anA. flavescens variety and are now accepted asAquilegia ×miniana.[47]
In China, clades distinguishing eastern and westernA. ecalcarata populations indicategene flow from different species. A study using genetic modeling indicated that the spurlessA. ecalcarata may have developed from two separate mutations from discrete eastern and western populations of the spurredA. kubanica, an instance of a hybridparallel evolution.[48] Further hybridization betweenA. ecalcarata and spurred columbines that share its range is limited by each species's selection for particular pollinators.[3]: 65 However, a short-spurredA. rockii phenotype has developed from hybridization with westernA. ecalcarata.[48]
Aquilegia species have natural ranges which span theNorthern Hemisphere in Eurasia and North America.[1] These ranges encompass theCircumboreal Region, the geographically largestfloristic region in the world.[8] The southern limits of the naturalAquilegia ranges are found in northern African and northern Mexico, with the only native African columbine being theA. ballii of theAtlas Mountains.[49][2]: 127
A. vulgaris, a European columbine which possibly originated in theBalkans,[3]: 208 has spread through both natural radiation and human assistance to become the most widely distributedAquilegia species. Its range has expanded to includeintroduced populations that have sometimes become naturalized in Africa,Macaronesia, the Americas, and Oceania.[2]: 124 [3]: 207–208 The species is also present in Asia, with populations in theRussian Far East andUzbekistan.[50] These introducedA. vulgaris populations typically originated from ornamental cultivation.[3]: 207
Some columbines are narrowlyendemic, with highly restricted ranges.A. paui only has a single population with four subpopulations within a few kilometers of each other in the mountains ofPorts de Tortosa-Beseit, Catalonia.[28][18]A. hinckleyana only natively populates a single location: the basin ofCapote Falls, a waterfall in Texas.[2]: 94 [51] As of 2005[update], the entire population ofA. nuragica – estimated as 10 to 15 individuals – populated an area of approximately 50 m2 (540 sq ft) onSardinia.[52]: 79
CertainAquilegia have been identified as having elevated risks of extinction, with some appearing on theIUCN Red List.[3]: 28–29 Two Sardinian columbines,A. barbaricina andA. nuragica, haveconservation statuses assessed by theIUCN ascritically endangered and the same organization listed the species in their Top 50 Mediterranean Island Plants campaign for conservation.[52]: 76–79 Some columbines, including both rare and common taxa, are the subject governmental regulations. The conflation of taxa has sometimes resulted in uncertainty regarding their conservation statuses.[3]: 28–29, 318
Humans pose a significant threat of impairing columbine population health and driving extinction. Beyond the desirability of the flowers for display, uncommon or rareAquilegia face the risk of destruction bybotanists and others seeking to add them to theirherbariums or privatecollections.[3]: 28–29
Some of the earliest cultivation of columbines may have occurred at Christian cathedrals.
In Europe, cultivation of columbines may have begun over 1700 years ago.Archaeobotanical evidence suggests thatA. vulgaris was cultivated for ornamental purposes in 3rd-century ADRoman Britain.[3]: 25 The discoveries of singularA. vulgaris seeds in burnt waste pits atAlcester andLeicester have been interpreted as evidence of their planting in gardens.[53] Finds of columbines at alate Saxon site nearWinchester Cathedral and three later medieval German sites have also been interpreted as using the plant for gardening.[54] In 12th-century Italy, people may have supportedA. vulgaris orA. atrata populations near religious structures, possibly due to the contemporary treatment of columbines as Christian symbols.[3]: 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.[55] Columbines function well in garden borders and can occupyshade andcottage gardens.[56] Certain columbines are suited for well-drainedrock gardens oralpine gardens. A few, such as the dwarf woodland plantA. flabellata var. pumila, favor moist shaded plantings.[2]: 17 [57]
Lifespans for cultivated columbines are generally short for perennials, with a plant's peak typically occurring in its second year.[58]: 64 Two- to three-year-long lives are typical in cultivatedA. coerulea andA. glandulosa,[44]: 3 withA. vulgaris exhibiting abinnenial-like lifespan.[2]: 18 Conversely,A. chrysantha andA. desertorum are particularly long-lived.[44]: 3 [59] In gardens, columbines will generally live three to four years. This lifespan can be extended bydeadheading, where dead flowers are removed prior to the plant expending the energy needed to produce fruit.[44]: 3 [2]: 18
In cultivation, the seasonal cycle that releases columbine seeds from dormancy can be replicated via astratification where seeds are exposed to two to four weeks of cool temperatures prior to sowing.[21] CultivatedAquilegia typically require well-draining soil. Improperly drained soil can result in the development ofroot rot, caused by either bacteria or fungi. At the end of the growth season, columbines can be protected fromfrost heaving by having their dead foliage removed to near the soil level and mulching once the ground is frozen.[16]: 149 Vernalization – a process by which juvenile plants are exposed to a weeks-long period of cold which mimics seasonal weather – can accelerate the rate at which columbines reach flowering.[60]
If permitted, cultivated columbines drop numerous seeds around themselves, resulting in a rapid proliferation ofseedlings. These seedlings can give the impression that short-lived plants are living longer.[2]: 18 Due to their tendency towards hybridization and – particularly in the case ofF1 hybrids andcultivars (cultivated varieties) – inherent genetic diversity, the seeds of cultivatedAquilegia often do not produce plants true to their type.[58]: 64, 66 [61]: 226 If identical plants are desired,basal cuttings can be performed in early summer.[57]
Several animals are consideredpests of cultivated columbines. Columbine leaf miners of thePhytomyza genus leave white patches or paths on leaves, but the damage is only cosmetic and does not generally require chemical pesticides.[31] The mothsPapaipema lysimachiae andP. nebris (stalk borer) both adversely affect columbines; scraping the ground around impacted plants can destroy the moths' seeds. The larval stage of theErynnis lucilius (columbine duskywing) is known as thecolumbine skipper, and the larvae can chew leaves and bind them together with silk.[16]: 150 Aphid infestation is another frequent issue, requiring rapid intervention to prevent significant destruction.[2]: 21
A. vulgaris 'Nora Barlow' is a double-flowered cultivar known for its coloring.
Columbine cultivars are popular among gardeners, particularly in the Northern Hemisphere.[3]: 25 The cultivars are typically bred in series, with plants sharing traits but varying in color. Some series produce dwarf plants.[57]A. vulgaris has been the basis for many cultivars, distinct in foliage, flower shape, and possession of fragrance.[58]: 64 The single-floweringA. vulgaris cultivar 'Nivea' (also known as 'Munstead White')[62]: 52 received theRoyal Horticultural Society'sAward of Garden Merit.[63][64]: 167
Double-flowered columbines were developed fromA. vulgaris and can be classified into three types. The Flore Pleno group, described in the English herbalistJohn Gerard's 1597 bookHerball, possesses plants where the flowers are elongated and the petals are rounded. The Veraeneana group come in several colors of flower and possess marbled green and gold foliage. The Stellata group, described in the English botanistJohn Parkinson's 1629 bookParadisi in Sole Paradisus Terrestris, has flowers which are star-shaped and have pointed petals. The three-colored, double-flowered cultivar 'Nora Barlow' – first discovered by the botanist and geneticistNora Barlow – is sometimes classified as part of the Stellata group, but displays a greater quantity of particularly narrow sepals than other members of that group.[58]: 64 [64]
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 EurasianAquilegia, the Asian speciesA. oxysepala andA viridiflora resist hybridization with North American columbines. Hybrids withA. vulgaris as one parent frequently look closer toA. vulgaris than any other parent. Many older garden columbines that are suspected hybrids resembleA. vulgaris, making true parentage difficult to determine. The American gardener Robert Nold said that "more has been written about 'Hensol Harebell' [a hybrid ofA. alpina andA. vulgaris which originated in early 20th-century Scotland] than perhaps any other interspecific hybrid", noting its greater affinity toA. vulgaris in appearance.[2]: 132–133, 137
More recent garden hybridization work has primarily utilized North American columbines for their varied greater flower sizes and variety of colors. While columbine flowers often face downwards in a nodding arrangement, hybridizers have introduced theA. coerulea trait of upward-facing flowers. The first such hybrids were the McKana Group (also known as McKana's Giants) – which won the 1935All-America Selections bronze medal for new seed offerings[2]: 138 – 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.[58]: 64–65
In China,A. oxysepala has been used as a dietary supplement and medicine for thousands of years.A. oxysepala has been used there to treatdiseases in women such asirregular menstruation andintermenstrual bleeding. While itsextract's function as an antioxidant is known, with its medicinal use possibly attributable to the extract's good scavenging ofsuperoxide anion radicals, it is inferior to the common dietary supplementascorbic acid.[26] Research has also determinedA. oxysepala to possess antibacterial qualities.[3]: 27
Some Indigenous North American peoples used the roots of columbines to treatulcers.[24] North American peoples have usedA. canadensis andA. chaplinei as anaphrodisiac.[7][66][2]: 38 CrushedA. canadensis seeds were used as a perfume, and the plant was thought to be capable of detecting bewitchment.[2]: 74 TheGoshute people reportedly chewedA. coerulea seeds or utilized the plant's root for medicinal or therapeutic purposes.[67]
Prior to deaths due to overdoses, small quantities of flowers from several columbines species were considered safe for human consumption and were regularly eaten as colorfulgarnishes and parts of salads.[7][24] Several Indigenous North American peoples have been described as eatingA. formosa: theMiwok may have boiled and eaten them with early springgreens, while Hanaksiala and Chehalis children may have sucked nectar from the flowers.[68] Columbine flowers are described as sweet,[24] a flavor attributed to their nectar.[69]
The American botanistVerne Grant repeatedly utilizedAquilegia in research published between the 1950s and the 1990s to explain the role that hybridization,polyploidy, and other processes played in how plant evolution andspeciation occur. Among Grant's works that utilizedAquilegia to illustrate evolutionary patterns and processes was his influential 1971 bookPlant Speciation.[70] The fivespecies groups that Grant proposed in 1952 remains a foundational element for a phylogenetic understanding of columbines.[3]: 97
In 21st-century scientific research ofplant development,ecology, andevolution,Aquilegia has been considered a model system.[38] Utilizing thegenome sequence ofA. coerulea, a study examined polyploidy during the evolution ofeudicots, aclade in which columbines are considered a basal member. This research determined that columbines and all eudicots experienced a sharedtetraploidy, but that only core members of the eudicots clade (which excludes columbines) experienced a sharedhexaploidy.[71]
Aquilegia vulgaris served as a symbol of bodily pleasures within Hieronymus Bosch'sThe Garden of Earthly Delights(detail pictured).
European columbines have been assigned several meanings since the ancient period. Within art,A. vulgaris has been a symbol of both moral and immoral behaviors, as well as an ornamentalmotif.[3]: 19–23 [72] Inancient Greece andancient Rome, the spurs of columbines were interpreted asphallic and the plants were associated with the fertility goddessesAphrodite andVenus.[72] For several centuries, columbines were viewed as symbols ofcuckoldry.[2]: 38 InEnglish literature, columbines have been mentioned with negative connotations.[2]: 38 InWilliam Shakespeare'sElizabethan dramaHamlet, the characterOphelia presentsKing Claudius with flowers that include columbines,[73] where the species is symbolic of deception and serves as an omen of death.[3]: 21
Columbines have several meanings in thelanguage of flowers, a manner of communicating using floral displays. In the 1867 English bookThe Illustrated Language of Flowers by a "Mrs. L. 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".[74] Columbines, due to theirresemblance to doves, have been associated with theHoly Spirit in Christianity since at least the 15th century.[75][72][76]
A. coerulea is thestate flower of Colorado.[77] TheColorado General Assembly passed legislation in 1925 making it illegal to uprootA. coerulea on public lands. The law also limits on how many buds, blossoms, and stems may be picked from the species by a person on public lands.[78] The name of the town ofColumbine Valley, Colorado, ultimately derives from the state flower.[79] Colorado'sA. coerula also served as the inspiration for one of the state's official songs, "Where the Columbines Grow", written in 1909.[80] It was used in the heraldry of theformer city ofScarborough in the Canadian province of Ontario.[81]
The asteroid1063 Aquilegia was named for the genus by the German astronomerKarl 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.[82][83]
^abcdefghijklmnopqrstuvwxyzaaabacadaeafNardi, Enio (2015).Il GenereAquilegia L. (Ranunculaceae) in Italia/The GenusAquilegia (Ranunculaceae) in Italy: Aquilegia Italicarum in Europaearum conspectu descriptio. Translated by Coster-Longman, Christina.Florence: Edizioni Polistampa.ISBN9788859615187.
^Shulkina, Tatyana (2005)."Aquilegia L.".Ornamental Plants From Russia And Adjacent States Of The Former Soviet Union. Rostok.ISBN9785946680325 – via efloras.org.
^abZhang, Keliang; Ji, Yusong; Fu, Guixian; Yao, Linjun; Liu, Huina; Tao, Jun; Walck, Jeffrey L. (July 2021). "Dormancy cycles inAquilegia oxysepala Trautv. et Mey. (Ranunculaceae), a species with non-deep simple morphophysiological dormancy".Plant and Soil.464 (1/2):223–235.Bibcode:2021PlSoi.464..223Z.doi:10.1007/s11104-021-04951-8.ISSN0032-079X.
^"Documented Chromosome Numbers of Plants".Madroño.17 (8):266–268. October 1964.JSTOR41423164.
^abTeuscher, Eberhard; Lindequist, Ulrike (2024).Natural Poisons and Venoms: Plant Toxins: Polyketides, Phenylpropanoids and Further Compounds. De Gruyter STEM. Berlin, Boston:De Gruyter. pp. 300–326.doi:10.1515/9783110728538.ISBN9783110728538.
^Thines, Marco; Denton, Geoffrey J.; Beal, Elizabeth J.; Kilty, Anne; Denton, Jennifer O.; Shin, Hyeon-Dong; Choi, Young-Joon (October 2019). "Peronospora aquilegiicola sp. nov., the downy mildew affecting columbines in the UK is an invasive species from East Asia".European Journal of Plant Pathology.155 (2):515–525.Bibcode:2019EJPP..155..515T.doi:10.1007/s10658-019-01787-y.ISSN1573-8469.S2CID195657188.
^Huang, Lei; Geng, Fang-Dong; Fan, Jing-Jing; Xue, Cheng; Zhang, Xiao-Yan; Kang, Ju-Qing; Zhang, Jian-Qiang; Ren, Yi (2018). "Genetic diversity and evolutionary history of four closely relatedAquilegia species revealed by 10 nuclear gene fragments".Journal of Systematics and Evolution.56 (2):129–138.Bibcode:2018JSyEv..56..129H.doi:10.1111/jse.12298.
_02.jpg)
.jpg)
