Aquilegia

''Aquilegia'', commonly known as columbines, is a genus of perennial flowering plants in the family Ranunculaceae (buttercups). The genus includes between 80 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 been used to create many cultivated varieties.

''Aquilegia'' typically possess stiff stems and leaves divide into multiple leaflets. Columbines often have colorful flowers with five sepals and five petals. The petals generally feature nectar spurs which differ in lengths 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.

Associated with fertility goddesses in ancient Greece and ancient Rome, archeological evidence suggests ''Aquilegia'' were in cultivation by the 2nd century AD in Roman Britain. Despite often being toxic, columbines have been used by humans as herbal remedies, perfume, and food. Asian traditional medicine, 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 exchanges between Old and New World species creating further diversity.

Etymology

The 1st-century AD Greek writer Dioscorides called columbines ''Isopyrum'', a name used presently applied to another genus. In the 12th century, the abbess and polymath Hildegard of Bingen referred to the plants as ''agleya'' from which the genus's name in German, ''Akelei'', derives. The first use of ''aquilegia'' with regards to columbines was in the 13th century by Albertus Magnus. In the 15th and 16th 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.

Several scientific and common names for the genus ''Aquilegia'' derive from its appearance. The genus name ''Aquilegia'' may come from the Latin word for "eagle", '' aquila'', in reference to the petals' resemblance to eagle talons. Another possible etymology for ''Aquilegia'' is a derivation from the Latin ("to collect water"), '' aquilegium'' (a container of water), or ("dowser" or "water-finder") in reference to the profusion of nectar in the spurs. The most common English-language name, '' columbine'', likely originates in the dove-like appearance of the sepals (''columba'' being Latin for ''dove'').

There are a number of other common names for ''Aquilegia'' across different languages. In English, these include ''granny's bonnet'' for some plants in the species ''Aquilegia vulgaris''. In French, the word '' ancolie'' is the common name for ''Aquilegia'', while individual members of the genus have been called ''gants-de-Notre-Dame'' ("Our Lady's glove"), while ''amor-nascoto'' ("love-born") has been used in Italian.

Description

''Aquilegia'' is a genus of herbaceous, perennial flowering plants in the family Ranunculaceae (buttercups). The genus is highly variable in appearance. Though they are perennials, certain species are short-lived, with some exhibiting lifespans more similar to biennials and others only flourishing for six to eight years. Following a dormant period in the winter, columbines will grow foliage and have a brief flowering period. Some columbines bloom the first year after sowing, others will bloom in their second. Later, seed heads will emerge and split, sowing new seed. The foliage lives through the summer before wilting and dying going into the fall.

''Aquilegia'' plants grow from slim, woody rhizomes from which multiple aerial stems rise. The leaves can grow in both basal (from the base of the plant) and cauline (from the 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 (each leaf dividing in three leaflets), 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).

The flowering stems emerge from rosettes during the spring and summer. Each inflorescence appears at the terminus of an aerial stem and can reach long. Depending the species, an inflorescence will feature one to ten of either cymes (flower clusters) or solitary flowers. Flower morphology varies across the genus, but all columbine flowers emerge from buds that are initially nodding. Flowers can be monochromatic or display multiple colors. 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. 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. The American botanist Verne Grant hypothesized that light-colored ''Aquilegia'', especially in North America, might have adopted their shading to increase their visibility to pollinators in twilight.

The perianth (non-reproductive portion) of ''Aquilegia'' flowers generally comprise five sepals that look like petals and five petals. Each petal typically comprises two portions: a blade, which are broad and project towards the front of the flower, and a nectar spur, a nectar-bearing structure which projects backwards. The hollow spurs attract pollinators and give columbine flowers a distinctive appearance. Depending on the species, spurs can have a hooked, horn-like appearance, with straight to coiled spurs also present in the genus. Some columbines, such as '' A. ecalcarata'', are naturally spurless. Recessive spurlessness individuals and populations can also be found within typically spurred species.

The reproductive portions of columbine flowers comprise the stamen (male) and gynoecium (female). The stamen, which bear the anthers from which pollen emerges, form whorls of five around the gynoecium. The total number of stamen varies between species. There are generally scale-shaped staminodes between the stamen and female pistil structures. The flowers undergo three stages of anthesis: a premale stage, where the flower perianth is open but the anthers are not dehisced (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.

''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). Autogamy has been observed as the primary fertilization mechanism in '' A. paui''. '' A. formosa'' and '' A. 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.

''Aquilegia'' fruit are follicles. These follicles have a split on one side and terminate with a curling tip known as a ''beak''. Columbine seeds are generally obovoid with black, smooth exteriors. Columbine seeds are in a dormant state at the point of sowing. 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.

The chromosome number for columbines is ''2n''=14. Individual plants have been recorded with other anomalous chromosome numbers, ranging up to 2''n''=32. It is possible that B chromosomes impact the phenotype and the fertility of individual plants that possess them.

Phytochemisty

Among cyanophore (organisms that produces a blue color) ''Aquilegia'' like ''A. 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 of fresh ''A. vulgaris'' leaves by a human was observed as causing convulsions, respiratory distress, and heart failure. A child who consumed 12 ''A. vulgaris'' flowers experienced weakness of the limbs, cyanosis, drowsiness, and miosis; all symptoms abated after three hours. Mature seeds and roots contain toxins that, if consumed, are perilous to human heart health.

The presence of the antibacterial flavonoid compound isocytisoside has been observed in ''A. vulgaris''. Polyphenols, primarily flavonoids, are the main component of hydroethanolic extract from '' A. oxysepala''. These compounds function as antioxidants. A study of ''A. oxysepala'' extract found it has a good scavenging effect 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.

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. formosa'' flowers determined that the petals and sepals had uniform levels of UV-resistant phenylpropanoids.

Ecology

Despite its toxicity and in the absence of incentives, some animals consume the fruit and leaves of columbines. In the case of the endangered '' A. paui'', one study found that 30% of all fruit was lost to predation by the Southeastern Spanish ibex. Consumption by mammals is not considered a component of the ''Aquilegia'' reproductive cycle.

In the Northeastern United States and Eastern Canada, ''A. 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. In the Western United States, ''Bombus occidentalis from '' A. coerulea'' by opening or using holes cut in the spurs.

Also in North America, three species of ''Phytomyza'' leaf miners lay their eggs on ''Aquilegia'': '' P. aquilegivora'' in the Midwest, '' P. aquilegiana'' in the east, and '' P. 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 puparium on the leaves' undersides. Adults pierce the leaves with their ovipositors to access liquids in the plants, leaving marks. Another ''Phytomyza'' columbine leaf miner, '' P. ancholiae'', is native to France.

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.

Several fungi attack columbine foliage, including '' Ascochyta aquilegiae'', '' Cercospora aquilegiae'', and '' Septoria aquilegiae''. 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.

Pollination

Following the evolution of the genus, ''Aquilegia'' developed diverse floral features including varied nectar spur morphology, orientation, and coloration to attract different pollinators, contributing to speciation. The suite of floral traits that develop to attract a particular set of pollinators are collectively referred to as a pollination 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 in ''A. paui''.

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 to red-flowered North American ''Aquilegia'', while pale-flowered columbines may have developed to increase their visibility to hawkmoths in twilight.

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. 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. 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 American ''Aquilegia'' to evolve the hawkmoth pollination syndrome.

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. micrantha'' var. mancosana.

Taxonomy

Within Linnaean taxonomy, ''Aquilegia'' was first described as a genus in Carl Linnaeus's 1753 ''Species Plantarum''. The genus is typically assigned to the family Ranunculaceae, though a minority of botanists have 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. Another historic assignment, made by the Swedish botanist in 1870, placed ''Aquilegia'' as the sole member of the family Aquilegiaceae.

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'' forms a basal, paraphyletic group with the genera ''Isopyrum'' and ''Thalictrum'' (meadow-rues) which is characterized by their plesiomorphy (characteristics shared between clades from their last common ancestor) with Berberidaceae. When placed within the monophyletic Thalictroideae, ''Aquilegia'' are 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 subfamiliy, a clade it shares with ''Semiaquilegia'' and '' Urophysa''. ''Semiaquilegia'' and ''Aquilegia'' are sister genera.

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 contended is his inclusion of the spurless Asian species '' A. 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. Another spurless columbine, ''A. micrantha'' var. ''mancosana'', was also once reassigned to ''Semiaquilegia''. Reassignments to ''Isopyrum'' and '' Paraquilegia'', such as '' P. anemonoides'' in 1920, have been more permanent.

Evolution

There are no good fossils of columbines and other Thalictroideae that indicate how they evolved and radiated. Genetic evidence suggests that the last common ancestor among Thalictroideae lived in East Asia approximately 36 million years ago, during the late Eocene.

A 2018 study of genetic evidence indicated that ''Aquilegia'' first appeared during the Upper Miocene approximately 6.9 million years ago. The genus split into two clades 4.8 million years ago, with one clade populating North America and the other radiating across Eurasia. A 2024 study found 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. The species is thought to have originated in the mountainous portions of south-central Siberia.

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. 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.

Genetic information suggests that the diversification rates of columbines rapidly increased about 3 million years ago, with indications of two independent radiation events occurring around that time: one in North American columbine populations and the other in European populations. 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.

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.

The nectar spurs present in ''Aquilegia'' are an unusual evolutionary trait, arising on the ancestor of all ''Aquilegia'' up to approximately 7 million years ago. In order to determine the gene responsible for the trait, a 2020 paper compared spurred ''Aquilegia'' taxa against the spurless ''A. ecalcarata''. This research identified a gene named ''POPOVICH'' (''POP'') 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 pedals of the spurred ''Aquilegia'' studied than in ''A. ecalcarata''.

Current species

According to different taxonomic authorities, the genus ''Aquilegia'' comprises between 70 and over 400 taxa. Some totals correspond more closely with Munz's 1946 total of 67, while online Tropicos and the International Plant Names Index have accepted over 200 and 500, respectively. , the Royal Botanic Gardens, Kew's Plants of the World Online accepts 130 species.

The American botanist and 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. 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.

The type species of the genus is '' A. vulgaris'', a European columbine with high levels of physical variability. Most European ''Aquilegia'' are morphologically similar to ''A. vulgaris'', sometimes to the point where visually them discerning from ''A. 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".

Natural hybridization

A lack of genetic and physiological barriers permits columbine hybridization across even distantly related species with high degrees of morphological and ecological differences. In natural settings, hybrid columbines may occur wherever the natural ranges of multiple species come into contact.

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.

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. Still, hybridization and subsequent introgression occurs in North American columbines. 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. In the instance of populations of hybrids between the yellow-flowered '' A. flavescens'' and red-flowered '' A. formosa'' in the northwestern United States, the resultant pink-flowering columbines were initially described as an ''A. flavescens'' variety and are now accepted as ''Aquilegia'' × ''miniana''.

In China, clades distinguishing eastern and western ''A. ecalcarata'' populations indicate gene flow from different species. A study using genetic modeling indicated that the spurless ''A. ecalcarata'' may have developed from two separate mutations from discrete eastern and western populations of the spurred '' A. kubanica'', an instance of a hybrid parallel evolution. Further hybridization between ''A. ecalcarata'' and spurred columbines that share its range is limited by each species's selection for particular pollinators. However, a short-spurred '' A. rockii'' phenotype has developed from hybridization with western ''A. ecalcarata''.

Distribution

''Aquilegia'' species have natural ranges which span the Northern Hemisphere in Eurasia and North America. These ranges encompass the Circumboreal Region, the geographically largest floristic region in the world. The southern limits of the natural ''Aquilegia'' ranges are found in northern African and northern Mexico, with the only native African columbine being the '' A. ballii'' of the Atlas Mountains.

''A. vulgaris'', a European columbine which possibly originated in the Balkans, 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. The species is also present in Asia, with populations in the Russian Far East and Uzbekistan. These introduced ''A. vulgaris'' populations typically originated from ornamental cultivation.

Some columbines are narrowly endemic, with highly restricted ranges. '' A. paui'' only has a single population with four subpopulations within a few kilometers of each other in the mountains of Ports de Tortosa-Beseit, Catalonia. '' A. hinckleyana'' only populates a single location: the basin of Capote Falls, a waterfall in Texas. , the entire population of '' A. nuragica'' estimated as 10 to 15 individuals populated an area of approximately on Sardinia.

Conservation

Certain ''Aquilegia'' have been identified as having elevated risks of extinction, with some appearing on the IUCN Red List. Two Sardinian columbines, '' A. barbaricina'' and ''A. nuragica'', have conservation statuses assessed by the IUCN as critically endangered and the same organization listed the species in their Top 50 Mediterranean Island Plants campaign for conservation. Some columbines, including both rare and common taxa, are the subject governmental regulations.

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.

Cultivation

In Europe, cultivation of columbines may have begun over 1700 years ago. Archaeobotanical evidence suggests that ''A. vulgaris'' was cultivated for ornamental purposes in 3rd-century AD Roman Britain. The discoveries of singular ''A. vulgaris'' seeds in burnt waste pits at Alcester and Leicester have been interpreted as evidence of their planting in gardens. Finds of columbines at a late Saxon site near Winchester Cathedral and three later medieval German sites have also been interpreted as using the plant for gardening. In 12-century Italy, people may have supported ''A. vulgaris'' or '' A. atrata'' populations near religious structures, possibly due to the contemporary treatment of columbines as Christian symbols.

Lifespans for cultivated columbines are generally short for perennials, with a plant's peak typically occurring in its second year. Two- to three-year-long lives are typical in cultivated ''A. coerulea'' and '' A. glandulosa'', with ''A. vulgaris'' exhibiting a binnenial-like lifespan. Conversely, '' A. chrysantha'' and '' A. desertorum'' are particularly long-lived. In gardens, columbines will generally live three to four years. This lifespan can be extended by deadheading, where dead flowers are removed prior to the plant expending the energy needed to produce fruit.

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. Cultivated ''Aquilegia'' typically require well-draining soil. Improperly 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.

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. Due to their tendency towards hybridization and particularly in the case of F1 hybrids and cultivars (cultivated varieties) inherent genetic diversity, the seeds of cultivated ''Aquilegia'' often do not produce plants true to their type.

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. The moths '' Papaipema lysimachiae'' and '' P. nebris'' (stalk borer) both adversely affect columbines; scraping the ground around impacted plants can destroy the moths' seeds. 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. Aphid infestation is another frequent issue, requiring rapid intervention to prevent significant destruction.

Cultivars and cultivated hybrids

Columbine cultivars are popular among gardeners, particularly in the Northern Hemisphere. Artificial hybridization efforts have determined that the degree of interfertility of columbines is not identical across species. While North American columbines easily hybridize with each other and most Eurasian ''Aquilegia'', the Asian species ''A. oxysepala'' and '' A viridiflora'' resist hybridization with North American columbines.

The single-flowering ''A. vulgaris'' cultivar 'Nivea' (also known as 'Munstead White') received the Royal Horticultural Society's Award of Garden Merit.

Double-flowered columbines were developed from ''A. 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 Veraeneana 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' first discovered by the botanist and geneticist Nora 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.

Human uses

Medicinal and herbal

Asian traditional medicine, Indigenous North Americans, and Medieval Europeans have considered columbines plants medicinal herbs. Modern scientific research has determined that columbines can possess antioxidant, antibacterial, and anti-cancer qualities.

In China, ''A. oxysepala'' has been used a dietary supplement and part of traditional Chinese medicine (TCM) for thousands of years. In TCM, ''A. oxysepala'' has been used 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. Research has also determined ''A. oxysepala'' possesses antibacterial qualities.

'' A. sibirica'' has been a significant part of Asian traditional medicine, including traditional Mongolian medicine, and the plant has been used to treat diseases in women, asthma, rheumatism, and cardiovascular diseases. It was also known to inhibit ''Staphylococcus aureus'', one of the bacteria responsible for staphylococcal infections. ''A. sibirica'' also possesses antifungal qualities. Extracts showed the presence of chlorogenic acid and caffeic acid. Extractions performed with heat and methanol extracted more of the medically relevant compounds than those performed at room temperature or with other solvents.

Some Indigenous North American peoples used the roots of columbines to treat ulcers. North American peoples have used '' A. canadensis'' and '' A. chaplinei'' as an aphrodisiac. Crushed ''A. canadensis'' seeds were used as a perfume, and the plant was thought to be capable of detecting bewitchment. The Goshute people reportedly chewed '' A. coerulea'' seeds or utilized the plant's root for medicinal or therapeutic purposes.

Other uses

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 colorful garnishes and parts of salads. Several Indigenous North American peoples have been described as eating ''A. 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. Columbine flowers are described as sweet, a flavor attributed to their nectar.

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''. The five species groups that Grant proposed in 1952 remains a foundational element for a phylogenetic understanding of columbines.

In 21st-century scientific research of plant development, ecology, and evolution, ''Aquilegia'' has been considered a model system. Utilizing the genome sequence of ''A. coerulea'', a 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.

In culture

European columbines shave 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 ornamental motif. 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. For several centuries, columbines were viewed as symbols of cuckoldry. In English literature, columbines have been mentioned with negative connotations. In William Shakespeare's Elizabethan drama ''Hamlet'', the character Ophelia presents King Claudius with flowers that include columbines, where the species is symbolic of deception and serves as an omen of death.

Medieval European artists associated the 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'' (1503–1504) by Hieronymus Bosch, ''A. vulgaris'' serves as a symbol for bodily pleasures. '' Portrait of a Princess'' (1435–1449) by Pisanello depicts multiple '' A. atrata'' at different angles as part of the floral ornamentation that makes that painting characteristic of the international Gothic style.

Columbines have several meanings in the language of flowers, a manner of communicating using floral displays. The 1867 English book ''The 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". Columbines, due to their resemblance to doves, have been associated with the Holy Spirit in Christianity since at least the 15th century.

''A. coerulea'' is the state flower of Colorado. The Colorado General Assembly passed legislation in 1925 making it illegal to uproot '' A. 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. It was used in the heraldry of the former city of Scarborough in the Canadian province of Ontario.

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.

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{{Taxonbar, from=Q147641

Garden plants
Ranunculaceae genera
Taxa described in 1753
Taxa named by Carl Linnaeus