Aquilegia

Columbines
flower and fruit of Aquilegia vulgaris (type species)
Scientific classification
Kingdom: Plantae
Division: Magnoliophyta
Class: Magnoliopsida
(unranked): Eudicots
Order: Ranunculales
Family: Ranunculaceae
Subfamily: Thalictroideae
Genus: Aquilegia
L.
Species

60-70, see text

Aquilegia (Columbine; from Latin columba "dove") is a genus of about 60-70 species[1] of perennial plants that are found in meadows, woodlands, and at higher altitudes throughout the Northern Hemisphere, known for the spurred petals [2] of their flowers.

Contents

Etymology

The genus name Aquilegia is derived from the Latin word for eagle (aquila), because the shape of the flower petals are said to resemble an eagle's claw.

"Columbine" is derived from the Latin word for pigeon (columba).

Description

Fruit is a follicle.[3]

Relatives

Columbines are closely related to plants in the genera Actaea (baneberries) and Aconitum (wolfsbanes/monkshoods), which like Aquilegia produce cardiogenic toxins.[4]

Insects

They are used as food plants by some Lepidoptera (butterflies and moths) caterpillars. These are mainly of noctuid moths – noted for feeding on many poisonous plants without harm – like Cabbage Moth (Mamestra brassicae), Dot Moth (Melanchra persicariae) and Mouse Moth (Amphipyra tragopoginis). The Engrailed (Ectropis crepuscularia), a geometer moth, also uses columbine as larval foodplant.

Cultivation

Several species are grown in gardens, including the European Columbine (A. vulgaris), a traditional garden flower in many parts of the world.[5] Numerous cultivars and hybrids have also been developed as well. They are easy to propagate from seed.

Columbine is a perennial, which propagates by seed. It will grow to a height of 15 to 20 inches. It will grow in full sun, however, prefers growing in partial shade and well drained soil, and is able to tolerate average soils and dry soil conditions. Columbine is rated hardiness of Zone 3 so does not require mulching or protection in the winter.[6][7]

Large numbers of hybrids are now available for the garden, since the British A vulgaris was joined by other European and N American varieties. [8] Aquilegia species are very interfertile, and will self sow.[9]

Uses

The flowers of various species of Colombine were consumed in moderation by Native Americans as a condiment with other fresh greens, and are reported to be very sweet, and safe if consumed in small quantities. The plant's seeds and roots are highly poisonous however, and contain cardiogenic toxins which cause both severe gastroenteritis and heart palpitations if consumed as food. Native Americans used very small amounts of Aquilegia root as an effective treatment for ulcers. However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal.[4]

Culture

The Colorado Blue Columbine (A. caerulea) is the official state flower of Colorado (see also Columbine, Colorado).

Evolution

Columbines have been important in the study of evolution. It was found that Sierra Columbine (A. pubescens) and Crimson Columbine (A. formosa) each have specifically adapted pollinators, with hawkmoths that can pollinate one species while usually failing to pollinate the other. Such a "pollination syndrome", being due to flower genetics, ensures reproductive isolation and can be a cause of underlying speciation.[10]

Petal Spur Evolution

Aquilegia petals show an enormous range of petal spur length diversity ranging from a centimeter to the 15 cm spurs of Aquilegia longissima. Interestingly, it was shown that this amazing 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. [11]

Species

Columbine species include:[12]

  • Aquilegia alpina L.
  • Aquilegia atrata W.D.J.Koch
    Dark Columbine
  • Aquilegia atrovinosa
  • Aquilegia aurea Janka
  • Aquilegia barbaricina
    Barbaricina Colombine (doubtfully valid)
  • Aquilegia barnebyi
    Oil Shale Columbine
  • Aquilegia bernardii Gren. & Godr.
  • Aquilegia bertolonii Schott
  • Aquilegia blecicii Podobnik (doubtfully valid)
  • Aquilegia brevistyla
    Smallflower Columbine
  • Aquilegia buergeriana
  • Aquilegia caerulea
    Colorado Blue Columbine
  • Aquilegia canadensis
    Canadian Columbine, Wild Columbine
  • Aquilegia champagnatii Moraldo, E.Nardi & la Valva (doubtfully valid)
  • Aquilegia chrysantha
    Golden Columbine
  • Aquilegia desertorum
    Desert Columbine
  • Aquilegia desolatica
    Desolation Columbine
  • Aquilegia dinarica Beck
  • Aquilegia ecalcarata
  • Aquilegia einseleana F.W.Schultz
  • Aquilegia elegantula
    Western Red Columbine
  • Aquilegia eximia
    Van Houtte's Columbine
  • Aquilegia flabellata
    Fan Columbine, Japanese wodamakinari (including A. akitensis)
  • Aquilegia flavescens
    Yellow Columbine
  • Aquilegia fragrans Benth.
    Fragrant Columbine
  • Aquilegia formosa
    Crimson Columbine, Western Columbine
  • Aquilegia glandulosa
  • Aquilegia grahamii
    Graham's Columbine
  • Aquilegia grata
  • Aquilegia × hybrida
  • Aquilegia incurvata
  • Aquilegia japonica
  • Aquilegia jonesii
    Jones' Columbine
  • Aquilegia karatavica
  • Aquilegia karelini
  • Aquilegia kitaibelii Schott
  • Aquilegia lactiflora
  • Aquilegia laramiensis
    Laramie Columbine
  • Aquilegia litardierei Briq.
  • Aquilegia longissima
    Gray. Longspur Columbine
  • Aquilegia loriae
    Lori's Columbine
  • Aquilegia magellensis F.Conti & Soldano
    Magella Columbine
  • Aquilegia × maruyamana
  • Aquilegia micrantha
    Mancos Columbine
  • Aquilegia moorcroftiana
  • Aquilegia nigricans Baumg.
  • Aquilegia nugorensis Arrigoni & E.Nardi (doubtfully valid)
  • Aquilegia nuragica
  • Aquilegia olympica
  • Aquilegia origami
  • Aquilegia ottonis Orph. ex Boiss.
  • Aquilegia oxysepala
  • Aquilegia pancicii Degen
  • Aquilegia parviflora
  • Aquilegia pubescens
    Sierra Columbine, Coville's Columbine
  • Aquilegia pubiflora
  • Aquilegia pyrenaica DC.
    Pyrenean Columbine
  • Aquilegia rockii
  • Aquilegia saximontana
    Rocky Mountain Columbine
  • Aquilegia scopulorum
    Blue Columbine, Utah Columbine
  • Aquilegia shockleyi
    Desert Columbine
  • Aquilegia sibirica
  • Aquilegia thalictrifolia Schott & Kotschy
  • Aquilegia transsilvanica Schur
  • Aquilegia triternata
    Chiricahua Mountain Columbine
  • Aquilegia truncata Red Columbine
  • Aquilegia turczaninovii
  • Aquilegia viridiflora
  • Aquilegia viscosa Gouan
  • Aquilegia vitalii
  • Aquilegia vulgaris
    Common Columbine, European Columbine, Granny's Nightcap
  • Aquilegia yabeana

See also

Footnotes

  1. ^ Sunset Western Garden Book, 1995:606–607
  2. ^ 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.
  3. ^ Dezhi & Robinson (2001)
  4. ^ a b Tilford (1997)
  5. ^ Nold (2003): p.128
  6. ^ http://www.gardenersnet.com/flower/columbine.htm The Gardener's Network
  7. ^ John Kilmer (1989). The Perennial Encyclopedia ISBN 0886656397
  8. ^ Andrew McIndoe, Kevin Hobbs: Perennials. David & Charles, 2005 ISBN 1558707646, 9781558707641
  9. ^ New England Wild Flower Society Guide to Growing and Propagating Wildflowers of the United States and Canada
  10. ^ Fulton & Hodges (1999), Hodges et al. (2002)
  11. ^ 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.
  12. ^ Dezhi & Robinson (2001), RBGE [2008], USDA [2008]

References

Related Reading