| Aquilegia vulgaris | |
|---|---|
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Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Order: | Ranunculales |
| Family: | Ranunculaceae |
| Genus: | Aquilegia |
| Species: | A. vulgaris |
| Binomial name | |
| Aquilegia vulgaris | |
| Synonyms[1][2]: 200–202 | |
List
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Aquilegia vulgaris is a species ofperennial flowering plant of the genusAquilegia (columbine) in the familyRanunculaceae. Commonly called thecommon columbine,European crowfoot, andgranny's bonnet, it presently possesses the most expansive range and greatest morphological variability in its genus. The current wild range ofA. vulgaris includes itsnative range in Europe as well asintroduced populations in Asia, Oceania, North America (where it has becomenaturalized), and South America.
The wild form ofA. vulgaris can grow flowering stems that 90 cm (35 in) tall from and often form a bushy clump at its base. In their native range, the species blooms from May to June. The flowers, with diameters measuring up to 60 mm (2.4 in) across, are typically blue or purple and possesspetals with structures known asnectar spurs.
Associated with fertility goddesses inancient Greece andancient Rome, archeological evidence suggestsA. vulgaris was in cultivation by the 2nd century AD inRoman Britain. The species represented virtuous behaviour, theHoly Spirit, and theTrinity withinChristian art; in other contexts, such asWilliam Shakespeare'sHamlet,A. vulgaris was a malevolent symbol. While it has been treated as an herbal remedy since theMiddle Ages, some chemicals within the plant are toxic to humans. The species was given itsscientific binomial name byCarl Linnaeus in his 1753Species Plantarum.
Some horticultural varieties, known ascultivars, that were developed by the 16th century have remained popular with gardeners. Cultivars ofA. vulgaris have continued to be developed, as havehybrids crossing it with other columbines. The resulted plants produce an array of colours anddouble-flowered examples.
Aquilegia vulgaris is aperennialherbaceousflowering plant of the genusAquilegia in the familyRanunculaceae. AnA. vulgaris plant possesses a thickrootstock that can be either simple or branched, with one or two floweringstems.[2]: 203 A. vulgaris plants often form bushy clumps from which their thin stems project upward.[3] The aerial stems grow to between 50 cm (20 in) and 90 cm (35 in) tall and can be between 3 mm (0.12 in) and 5 mm (0.20 in) wide at their bases.[2]: 203 [4]: 46 These stems are leafy and are pubescent (covered in hairs). The hairs are whitish and are absent on some plants of the species. The branches from the stems are glabrous (covered in glands) and pubescent, with the top portions showing more pubescence than below. The hairs are small and whitish.[2]: 203
The basal leaves (leaves attached to the base of the plant) are present in large quantities and can reach between 22 cm (8.7 in) and 40 cm (16 in) long when including theirpetioles. The petioles themselves can reach between 14 cm (5.5 in) and 27 cm (11 in) long. These leaves are biternate, with eachleaflet itself subdivided in three. The thin leaf blades of basal leaves are glabrous on their top sides. They areglaucous (pale blue-grey) and pubescent beneath. There are also cauline leaves (leaves attached to an aerial stem) which are borne on petioles between 0.1 cm (0.039 in) and 11 cm (4.3 in) in length. The cauline leaves, which appear singularly or in pairs, are similar to the basal leaves but become smaller and simplified the further up a stem they appear.[2]: 203–204
WildA. vulgaris plants flower between May and June in their native range.[2]: 204 In North America, whereA. vulgaris isnaturalized,[5]: 125 plants flower with a spring to summer bloom from May to July.[6] Theinflorescences of the species can feature between three and eighteen flowers. Thebracts are a bit glabrous or scarcely pubescent and are downy beneath. The bracts split into three segments that are each lanceolate in shape. Thepeduncles are particularly downy and are densely covered by small hair-like structures calledtrichomes.[2]: 204
WildA. vulgaris flowers, particularly those on the form that is sometimes calledAquilegia vulgaris subsp. vulgaris, are typically blue or purple, with rare examples of white and reddish flowers.[2]: 204 [6][7]: 379 The flowers are in a nodding orientation (facing downwards) and have diameters of between 30 mm (1.2 in) and 60 mm (2.4 in). The primary flowers of an inflorescence are proportionally larger – possessing flowers considered medium- to small-sized within the genus – than the secondary flowers of that inflorescence.[2]: 204
The five petaloidsepals of the flower can be oriented divergent or perpendicular to thefloral axis.[8]: 49 [6][7]: 376 These sepals are ovate to ovate-lanceolate in shape, coming to acute tips. The sepals of a flower are shorter than itspetals, measuring between 15 mm (0.59 in) and 30 mm (1.2 in) long and 6 mm (0.24 in) to 16 mm (0.63 in) wide. The inner structure of the flowers form a cup-like appearance.[2]: 204
WildAquilegia vulgaris, as with otherAquilegia, have five petals which possessnectar spurs, a form ofnectar-bearing structure.[7]: 376 [5]: 31 The petals are medium-sized within the genus and are isotropic, with lengths and widths of between 22 mm (0.87 in) and 34 mm (1.3 in). The broad portion (thelimb) of the petal is shorter than its nectar spur. The limbs have broad, rounded ends and measure 10 mm (0.39 in) to 14 mm (0.55 in) long and 8 mm (0.31 in) to 13 mm (0.51 in) wide. The obconical spurs can be hooked or, more rarely, curved. The spurs range in length from 12 mm (0.47 in) to 20 mm (0.79 in) long and are between 4 mm (0.16 in) and 8 mm (0.31 in) at their opening (thethroat).[2]: 204 The spurs have an even tapper as they narrow towards their ends.[6]
Each flower features multiplestamens,[5]: 32 which measure between 9 mm (0.35 in) and 13 mm (0.51 in) long. The fruit of the plant arefollicles which are between 15 mm (0.59 in) and 25 mm (0.98 in) long.[6] Members ofAquilegia produce largely quantities of seeds, which are black.[5]: 32 A. vulgaris seeds have shiny surfaces and lackelaiosome (a fleshy mass present on some seeds).[4]: 46 [9] The species's seeds are between 2.2 mm (0.087 in) and 2.5 mm (0.098 in) long and have been recorded as weighing 1.53 mg (0.0236 gr).[4]: 46 [9] The plant'schromosome number is 2n=14.[6]
Aphids, a type of insect, are known to attack columbines. This is particularly the case forA. vulgaris andhybrids closely related to it. Aphid infestations can result in stunted growth, stickyhoneydew accumulations on the basal leaves, and flowers that are deformed or fail to open.[5]: 20–21
Pollination can be achieved throughpollinators visiting flowers to access thenectar stored in the spurs. Studies across EuropeanA. vulgaris populations have long identified visitation bybumblebees (Bombus).Hummingbirds have also been observed to visit cultivated specimens of the species.[10]
AmongAquilegia withcyanophores (cells that produce a blue colour) 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.[11]
TheRoyal Botanic Gardens, Kew's Useful Plants and Fungi of Colombia project has identifiedA. vulgaris as poisonous.[1] An acute toxicity test in mice showed that ethanol extract and the mainflavonoid compoundisocytisoside from the leaves and stems of the species can be classified as nontoxic since a dose of 3000 mg/kg did not cause mortality in mice.[12]Nicholas Culpeper recommended the seeds taken in wine to speed the process of childbirth. In modern herbal medicine it is used as anastringent anddiuretic.[13]
The first reliable descriptions ofAquilegia vulgaris were written bymedieval European mystics and scientists including the 12th-century abbessHildegard of Bingen – who considered the plant's herbal functions – and the 13th-century friarAlbertus Magnus.[2]: 23 Thebinomial nameAquilegia vulgaris was assigned by the Swedish biologistCarl Linnaeus in his 1753 bookSpecies Plantarum.[5]: 124
A vulgaris is the most morphologically variable species ofAquilegia.[5]: 124 It is also at the head of aspecies complex that comprises the majority of EuropeanAquilegia, with a minority morphologically aligning with theAquilegia alpina complex.[5]: 35 [2]: 23 TheA. vulgaris complex is typified as possessing longer spurs than other Eurasian species, an adaptation suited to pollination by bumblebees with longer tongues. This difference in pollination adaptations corresponds with the distinction between the North AmericanAquilegia complexes:Aquilegia canadensis is adapted tohummingbird pollination and theAquilegia coerulea complex is adapted to moth pollination.[14]: 355
The genus nameAquilegia may come from the Latin word for "eagle",aquila, in reference to the petals' resemblance to eagle talons.[15] A more likely etymology forAquilegia is a derivation from the Latinaquam legere "to collect water",aquilegium (a container of water), oraquilex ("dowser" or "water-finder") in reference to the profusion of nectar in the spurs.[16][5]: 37–38 The specific namevulgaris is Latin for "common".[17][5]: 124
Common names for the species include the common columbine, European crowfoot, and granny's bonnet.[3][18] The wordcolumbine, the common name for species in the genus, derives from the Latin wordcolumba, meaning "dove", a reference to the flowers' appearance being similar to a group of doves.[5]: 37–38 The namegranny's bonnet is derived from the petals' resemblance tobonnets.[19]
Originally a European species,Aquilegia vulgaris possibly originated in theBalkans.[2]: 208 It has since spread to become the most widely distributedAquilegia species.[5]: 124 Its range has expanded – both through natural radiation and human assistance – to includeintroduced populations that have sometimes become naturalized in Africa,Macaronesia, the Americas, and Oceania.[2]: 207–208 The species is also present in Asia, with populations in theRussian Far East andUzbekistan.[1] These introduced populations typically originated from ornamental cultivation.[2]: 207
In Europe, the species ranges northward into southernScandinavia and England.[4]: 46 The boundaries of species's distribution innorthern Europe, simplified byA. vulgaris being the exclusive member of the genus in this region, has been understood since the 19th century.[2]: 208 Thesouthern European distribution of the species is less defined, as its radiation through theIberian,Italian, and Balkan peninsulas has brought it into contact with other columbines in that region withintrogressive effects.[2]: 208 The species also ranges east into western Russia.[4]: 46 Aquilegia ballii, sometimes considered a variety ofA. vulgaris, inhabits theAtlas Mountains and is the onlyAquilegia in Africa.[5]: 127
The American botanist and gardener Robert Nold, saying that there was little evidence for the previous species's absence in any part of Europe and that expansion was a natural process, viewed categorizations of any European populations ofA. vulgaris as naturalized rather than as native were "tenuous at best".[5]: 125 TheRoyal Botanic Gardens, Kew'sPlants of the World Online (POWO) records the species as native to Albania, Austria, the Balkans, theBaltic states, Belarus, Belgium, the British Isles, Bulgaria,Corsica, continental France, Germany, the Iberians and Italian peninsulas, theLow Countries, Poland,Sicily, Switzerland, and parts of European Russia. POWO considers the populations in Denmark, theCaucasus, Scandinavia, Ukraine, and portions of European Russian as introduced.[1]
The Italian botanist Enio Nardi considered several hypotheses for howA. vulgaris reached theBritish Isles. The hypothesis suggesting the earliest arrival time suggests the plant spread over a plateau that connected the British Isles toContinental Europe during theLate Miocene (prior to 5.333 million years ago). Other hypotheses suggest a later arrival, including duringQuaternary glaciations or as recently as within the period ofrecorded history.[2]: 131
Introduced populations ofA. vulgaris live in Macaronesia, a series ofarchipelagos in the Atlantic Ocean off theNorth African and European coasts. The species has been repeatedly recorded as present on theAzores,Canary Islands, andMadeira since at least 1932. OnTenerife, one of the Canary Islands, the plant has been reported as present since at least 1974.[2]: 208
The species is naturalized in North America,[5]: 125 where it escaped from cultivation as an ornamental plant. It is established within cooler environments on the continent.[20] While most of the naturalized populations are the wild form with blue or purple flowers, others descend from horticultural forms with white, red, or pink flowers.[6][20] Some forms of the species present on the continent are likely the descendants of hybrids betweenA. vulgaris and otherAquilegia. Populations ofA. vulgaris exist on both the Pacific and Atlantic coasts of Canada and the United States.[6]A. vulgaris was among several foreign species proposed as thenational flowers of the United States during the early 20th century.[21]
In Oceania, the species has been introduced toNew South Wales,Tasmania, andVictoria in Australia and both theNorth andSouth Islands of New Zealand. Introduced populations have been recorded in South America since at least 1845, when it was recorded as present in Chilean cultivation.A. vulgaris populations in Chilea and Argentina have both been recorded in the 21st century.[2]: 208
Archaeobotanical evidence suggests thatAquilegia vulgaris was cultivated for ornamental purposes in 3rd-century ADRoman Britain.[2]: 25 The discoveries of singularA. vulgaris seeds in burnt waste pits atAlcester andLeicester have been interpreted as evidence of their planting in gardens.[22] 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.[23] In 12-century Italy, people may have supportedA. vulgaris orAquilegia atrata populations near religious structures, possibly due to the contemporary treatment of columbines as Christian symbols.[2]: 25
By the 16th century,selectively bred horticulturalA. vulgaris were being recorded in Europe. Linnaeus made mention of some horticulturalA. vulgaris when describing the species in 1753 in what was a rare inclusion of cultivated plants within his work. Presently,A. vulgaris is a frequently cultivated garden plant, though the wild form of the species is rare in such settings. It is most commonly grown in its many cultivated forms, known ascultivars.[24]: 167 Some of these cultivars were developed centuries ago and remain popular among gardeners in the northern parts of the globe. Some cultivars produce particular colours, while others were bred asdouble-flowered plants where stamens produce petals or have sepal-likeperianths.[2]: 25
The species is suited to sunny and partially shaded settings withinUSDA hardiness zones 3 through 10.[24]: 167 A. vulgaris plants generally only live three or four years in garden settings and can be interpreted as havingbiennial-like lifespans. Within their lifetime, each plant can produce dozens ofseedlings.Deadheading (removing dead flowers) before a plant expends the energy necessary to produce seeds can extend the lifespan of any columbine.[5]: 18
The single-flowering cultivar 'Nivea' (also known as 'Munstead White')[8] received theRoyal Horticultural Society'sAward of Garden Merit.[25][24]: 167 'Nivea' plants grow to 90 cm (35 in) tall, have grey-green foliage, and possess white flowers. Other single-flowering cultivars include 'Heidi' and 'Hensol Harebell', both of which can reach can reach 60 cm (24 in) tall and have nodding flowers. 'Heidi' plants have purple-red stems and pink flowers, while 'Hensol Harebell' flowers are short-spurred andmauve-blue.[24]: 167
The double-flowered cultivars ofA. vulgaris are divided into groups. Within the Flore Pleno group, which includes the white-flowered 'Graeme Iddon' and mauve-blue-flowered 'Rougham Star', the flowers are elongated and the petals are rounded. The cultivars of the Veraeneana group come in several colours of flower and possess green and gold marbled foliage.[24]: 167
The Stellata group's double flowers are star-shaped and have pointed petals. This group includes the dark purple-flowered 'Black Barlow', the blue-flowered 'Blue Barlow', the multicoloured 'Nora Barlow', and the light-pink flower 'Rose Barlow'.[24]: 167 Named forNora Barlow, who was a British botanist andCharles Darwin's granddaughter, 'Nora Barlow' originated in herCambridgeshire garden during the early 1980s. Since the introduction of 'Nora Barlow' into cultivation by the horticulturistAlan Bloom, it has become widely sold. Hybridization has altered the cultivar's appearance from nodding flowers that were equally green, white, and pink to forward-facing flowers with little green.[26]: 67
European columbines such asAquilegia vulgaris have been assigned several meanings since the ancient period. Within art,A. vulgaris has been a symbol of both moral and immoral behaviours, as well as an ornamentalmotif.[2]: 19–23 [27] Inancient Greece andancient Rome, the spurs of columbines were interpreted asphallic and the plants were associated with the fertility goddessesAphrodite andVenus.[27] InWilliam Shakespeare'sElizabethan dramaHamlet, the characterOphelia presentsKing Claudius with flowers that includeA. vulgaris,[28] where the species is symbolic of deception and serves as an omen of death.[2]: 21 InThe Garden of Earthly Delights (1503–1504) byHieronymus Bosch,A. vulgaris serves as a symbol for bodily pleasures.[2]: 21
Medieval European artistsassociated the plant with Christian sacredness and sublimity, withFlemish painters of the 15th century frequently depicting the plant in prominent locations within theirChristian artworks.[2]: 19 Among the Flemish works of this period featuringA. vulgaris areRogier van der Weyden'sLamentation of Christ (c. 1450),Hans Memling'sThe Last Judgment (1470–1472) andFlowers in a Jug (c. 1485), andHugo van der Goes'sPortinari Altarpiece (1475–1476).[2]: 19
With itsdove-like flowers, the species came to symbolize theHoly Spirit, such as in a 1497 Parisianbook of hours by German printer Thielman Kerver.[29][27] The interpretation of the five spurs as five doves saw columbines called "Five birds together" in Austria. A set of seven columbine flowers was associated with theSeven gifts of the Holy Spirit. The numeric symbolism of columbines was extended to its leaves, with their three-lobed form drawing associations with theTrinity.[27]
| International | |
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| Other | |
