''Aquilegia'' (common names: granny's bonnet, columbine) is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

of about 60–70

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

s that are found in

meadow A meadow ( ) is an open habitat, or field, vegetated by grasses, herbs, and other non-woody plants. Trees or shrubs may sparsely populate meadows, as long as these areas maintain an open character. Meadows may be naturally occurring or artifi ...

woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...

s, and at higher altitudes throughout the Northern Hemisphere, known for the spurred petals Puzey, J.R., Gerbode, S.J., Hodges, S.A., Kramer, E.M., Mahadevan, L. (2011) Evolution of ''Aquilegia'' spur length diversity through changes in cell anisotropy. Proceedings of the Royal Society B. of their

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

Etymology

The genus name Aquilegia comes from the

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

“Aquila”, or “eagle”; this is in obvious reference to the spurred, “hook” shapes within the blooms, that many gardeners say resemble an eagle's talons.

Description

Perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

herbs In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal ...

, with woody, erect stock,

root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...

s forming thick

rhizomes In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

. The basal

leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...

are compound, 1–3

ternate Ternate is a city in the Indonesian province of North Maluku and an island in the Maluku Islands. It was the ''de facto'' provincial capital of North Maluku before Sofifi on the nearby coast of Halmahera became the capital in 2010. It is off the we ...

, blades 3-lobed -partite, and lobes lobulate and obtuse. The

cauline A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...

leaves are similar to the basal ones, while the upper ones are

bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...

like. The

hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...

(bisexual)

flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

are terminal to stem and branches. They are usually pentamerous (with five spreading

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...

petaloid Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usually ...

sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...

segments). Five tubular honey-leaves are semi erect with a flat limb and spurred or saccate at the base. The spur is directed backwards and secretes

nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

Stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

are numerous (often more than 50) in

whorls A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs). Whorls in nature File:Photograph and axial plane floral ...

of 5, the innermost being

scarious This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

staminodes In botany, a staminode is an often rudimentary, sterile or abortive stamen, which means that it does not produce pollen.Jackson, Benjamin, Daydon; ''A Glossary of Botanic Terms with their Derivation and Accent''; Published by Gerald Duckworth & Co. ...

. There are ten membranaceous intrastaminal scales. There are five

pistils Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

and the

Carpels Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

are free. The

fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...

has several (five to 15) follicles which are semi erect and slightly connate downwards. These hold many seeds and are formed at the end of the pistils. The nectar is mainly consumed by long-beaked birds such as hummingbirds. Almost all ''Aquilegia'' species have a ring of staminodia around the base of the stigma, which may help protect against insects.

Chromosome number Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectivel ...

is x=7.

Relatives

Columbines are closely related to plants in the genera '' Actaea'' (baneberries) and ''

Aconitum ''Aconitum'' (), also known as aconite, monkshood, wolf's-bane, leopard's bane, mousebane, women's bane, devil's helmet, queen of poisons, or blue rocket, is a genus of over 250 species of flowering plants belonging to the family Ranunculaceae. ...

'' (wolfsbanes/monkshoods), which like ''Aquilegia'' produce cardiogenic

toxin A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849– ...

s.Tilford (1997)

Insects

They are used as food plants by some

Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...

(butterfly and moth)

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...

s. These are mainly of

noctuid moth The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...

s – noted for feeding on many poisonous plants without harm – such as

cabbage moth The cabbage moth (''Mamestra brassicae'') is primarily known as a pest that is responsible for severe crop damage of a wide variety of plant species. The common name, cabbage moth, is a misnomer as the species feeds on many fruits, vegetables, ...

(''Mamestra brassicae''),

dot moth The dot moth (''Melanchra persicariae'') is a moth of the family Noctuidae. The species was first described by Carl Linnaeus in 1761. It is a very distinctive species with very dark brown, almost black, forewings marked with a large white stigm ...

(''Melanchra persicariae'') and

mouse moth The mouse moth (''Amphipyra tragopoginis'') is a moth of the family Noctuidae. It is a widespread species with a Holarctic distribution. Distribution Europe (except the extreme north, and not occurring in the south of Spain, Sicily, or the Balkan ...

(''Amphipyra tragopoginis''). The engrailed (''Ectropis crepuscularia''), a

geometer moth The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''metr ...

, also uses columbine as a larval food plant. The larvae of the '' Papaipema leucostigma'' also feed on columbine. Plants in the genus ''Aquilegia'' are a major food source for ''

Bombus hortorum ''Bombus hortorum'', the garden bumblebee or small garden bumblebee, is a species of bumblebee found in most of Europe north to 70°N, as well as parts of Asia and New Zealand. It is distinguished from most other bumblebees by its long tongue ...

'', a species of bumblebee. Specifically, they have been found to forage on species of ''

Aquilegia vulgaris ''Aquilegia vulgaris'' is a species of columbine native to Europe with common names that include: European columbine, common columbine, granny's nightcap, and granny's bonnet. It is a flowering herbaceous perennial plant growing to 1.2 m tall ...

'' in Belgium and ''

Aquilegia chrysantha ''Aquilegia chrysantha'', the golden columbine, is a perennial herbaceous plant native to the southwestern United States from extreme southern Utah to Texas and northwestern Mexico. The ferny leaves have three leaflets with three lobes and gr ...

'' in North America and Belgium. The bees do not show any preference in color of the flowers.

Cultivation

Columbine is a hardy

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

, which propagates by seed. It will grow to a height of . It will grow in full sun; however, it prefers growing in partial shade and well drained soil, and is able to tolerate average soils and dry soil conditions. Columbine is rated at

hardiness zone A hardiness zone is a geographic area defined as having a certain average annual minimum temperature, a factor relevant to the survival of many plants. In some systems other statistics are included in the calculations. The original and most wide ...

3 in the United States so does not require mulching or protection in the winter. Large numbers of hybrids are available for the garden, since the European '' A. vulgaris'' was hybridized with other European and North American varieties. ''Aquilegia'' species are very interfertile, and will self-sow. Some varieties are short-lived so are better treated as

biennials A biennial plant is a flowering plant that, generally in a temperate climate, takes two years to complete its biological life cycle. Life cycle In its first year, the biennal plant undergoes primary growth, during which its vegetative structures ...

. The British National Collection of ''Aquilegia''s was held by Mrs Carrie Thomas at Killay near

Swansea Swansea (; cy, Abertawe ) is a coastal city and the second-largest city of Wales. It forms a principal area, officially known as the City and County of Swansea ( cy, links=no, Dinas a Sir Abertawe). The city is the twenty-fifth largest in ...

. Some time during or before 2014 the collection started to succumb to Aquilegia Downy Mildew '' Peronospora aquilegiicola'' which was at the time an emerging disease to which the plants had no resistance. By 2018 the entire collection had been lost. Aquilegia can be grown from seeds or rhizomes.

Uses

Doubled flower of Aquilegia x hybrida 001

The flowers of various species of columbine were consumed in moderation by Native Americans as a

condiment A condiment is a preparation that is added to food, typically after cooking, to impart a specific Flavoring, flavor, to enhance the flavor, or to complement the dish. A table condiment or table sauce is more specifically a condiment that is serv ...

with other fresh greens, and are reported to be very sweet, and safe if consumed in small quantities. The plant's seeds and roots, however, are highly poisonous and contain cardiogenic

s which cause both severe

gastroenteritis Gastroenteritis, also known as infectious diarrhea and gastro, is an inflammation of the gastrointestinal tract including the stomach and intestine. Symptoms may include diarrhea, vomiting, and abdominal pain. Fever, lack of energy, and dehydra ...

and

heart palpitation Palpitations are perceived abnormalities of the heartbeat characterized by awareness of cardiac muscle contractions in the chest, which is further characterized by the hard, fast and/or irregular beatings of the heart. Symptoms include a rap ...

s if consumed as food. Native Americans used very small amounts of ''Aquilegia'' root as a treatment for

ulcer An ulcer is a discontinuity or break in a bodily membrane that impedes normal function of the affected organ. According to Robbins's pathology, "ulcer is the breach of the continuity of skin, epithelium or mucous membrane caused by sloughing o ...

s. However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal. An

acute toxicity Acute toxicity describes the adverse effects of a substance that result either from a single exposure or from multiple exposures in a short period of time (usually less than 24 hours). To be described as ''acute'' toxicity, the adverse effect ...

test in mice has demonstrated that ethanol extract mixed with isocytisoside, the main flavonoid compound from the leaves and stems of ''Aquilegia vulgaris'', can be classified as non-toxic, since a dose of 3000 mg/kg did not cause mortality.

Culture

The

Colorado blue columbine ''Aquilegia coerulea'', the Colorado blue columbine, is a species of flowering plant in the buttercup family Ranunculaceae, native to the Rocky Mountains, USA. ''Aquilegia coerulea'' is the state flower of Colorado. The Latin specific name '' ...

(''A. coerulea'') is the official state flower of

Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...

(see also

Columbine, Colorado Columbine is census-designated place (CDP) in and governed by Jefferson and Arapahoe counties in Colorado, United States. The CDP is a part of the Denver–Aurora–Lakewood, CO Metropolitan Statistical Area. Located primarily in Jefferson Coun ...

). It is also used as a symbol of the former city of

Scarborough Scarborough or Scarboro may refer to: People * Scarborough (surname) * Earl of Scarbrough Places Australia * Scarborough, Western Australia, suburb of Perth * Scarborough, New South Wales, suburb of Wollongong * Scarborough, Queensland, su ...

in the Canadian province of Ontario.

Evolution

Columbines have been important in the study of

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

. It was found that the Sierra columbine (''A. pubescens'') and crimson columbine (''A. formosa'') each has

adapted In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...

specifically to a

pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...

. Bees and hummingbirds are the visitors to ''A. formosa'', while

hawkmoth The Sphingidae are a family of moths ( Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, b ...

s would only visit ''A. pubescens'' when given a choice. Such a "

pollination syndrome Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic (wind and water) or biotic, such as birds, bees, flies, and so forth through a process c ...

", being due to flower color and orientation controlled by their

genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar wor ...

, ensures

reproductive isolation The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring ...

and can be a cause of

speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...

. ''Aquilegia'' petals show an enormous range of petal spur length diversity ranging from a centimeter to the 15 cm spurs of ''Aquilegia longissima''. Selection from pollinator shifts is suggested to have driven these changes in nectar spur length. It was shown that this spur length diversity is achieved solely through changing cell shape, not cell number or cell size. This suggests that a simple microscopic change can result in a dramatic evolutionarily relevant morphological change.

Species

Schwarzviolette Akelei (Aquilegia atrata) 02

Columbine species include:

Notes

References

Bibliography

* ''see also''

Flora Europaea The ''Flora Europaea'' is a 5-volume encyclopedia of plants, published between 1964 and 1993 by Cambridge University Press. The aim was to describe all the national Floras of Europe in a single, authoritative publication to help readers identify ...

* * * ** , in
Allan M. Armitage: Armitage's Native Plants for North American Gardens.Timber Press, 2006
* Dezhi, Fu; Robinson, Orbélia R. (2001): 19. ''Aquilegia. In:'' Wu, Z. Y.; Raven, Peter Hamilton & Hong, D. Y. (eds.): ''Flora of China'' (Vol. 6: Caryophyllaceae through Lardizabalaceae): 278. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
HTML fulltext
* * * Nold, Robert (2003): Columbines: ''Aquilegia'', ''Paraquilegia'', and ''Semiaquilegia''. Timber Press. Preview
at

Google Books Google Books (previously known as Google Book Search, Google Print, and by its code-name Project Ocean) is a service from Google Inc. that searches the full text of books and magazines that Google has scanned, converted to text using optical c ...

* Royal Botanic Garden Edinburgh (RBGE)

008 008, OO8, O08, or 0O8 may refer to: * The Streetwear Brand @008us , inspired by Ian Fleming & Virgil Abloh *"030", the fictional 030 Agent of MI6 * '' 038: Operation Exterminate'', a 1965 Italian action film * '' Explosivo 030'' a 1940 Argentine c ...

Digital

''Aquilegia'' species list
Retrieved 2008-NOV-25. * Tilford, Gregory L. (1997): ''Edible and Medicinal Plants of the West''. Mountain Press Pub., Missoula, Montana. *

United States Department of Agriculture The United States Department of Agriculture (USDA) is the United States federal executive departments, federal executive department responsible for developing and executing federal laws related to farming, forestry, rural economic development, ...

(USDA)

br>USDA Plants Profile: ''Aquilegia''
Retrieved 2008-NOV-25. * * Kramer, E. M. (2009)
Aquilegia: A New Model for Plant Development, Ecology, and Evolution
''Annual Review of Plant Biology,'' Vol. 60. *

External links

{{Taxonbar, from=Q147641 Ranunculaceae genera Poisonous plants