Aquilegia

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Columbine
Wald-Akelei.JPG
Aquilegia vulgaris
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Ranunculales
Family: Ranunculaceae
Subfamily: Thalictroideae
Genus: Aquilegia
L. (1753)
Type species
Aquilegia vulgaris
L.
Species [1]

130, see text

Synonyms [1]

AquilinaBubani (1901), nom. illeg.

Aquilegia (common names: granny's bonnet, [2] columbine) is a genus of about 130 species [1] of perennial plants that are found in meadows, woodlands, and at higher elevations throughout the Northern Hemisphere, known for the spurred petals [3] of their flowers.

Contents

Etymology

The genus name Aquilegia comes from the Latin "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. [4]

Description

Perennial herbs, with woody, erect stock, roots forming thick rhizomes. The basal leaves are compound, 1–3 ternate, blades 3-lobed -partite, and lobes lobulate and obtuse. The cauline leaves are similar to the basal ones, while the upper ones are bract like.

The hermaphrodite (bisexual) flowers are terminal to stem and branches. They are usually pentamerous (with five spreading perianth petaloid sepal segments). Five tubular honey-leaves [lower-alpha 1] are semi erect with a flat limb and spurred or saccate at the base. The spur is directed backwards and secretes nectar. Stamens are numerous (often more than 50) in whorls of 5, the innermost being scarious staminodes. There are ten membranaceous intrastaminal scales. There are five pistils and the carpels are free. [6] [7] [8]

The fruit 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. [9] Almost all Aquilegia species have a ring of staminodia around the base of the stigma, which may help protect against insects. [10] [6] [7] Chromosome number is x=7. [8]

Relatives

Columbines are closely related to plants in the genera Actaea (baneberries) and Aconitum (wolfsbanes/monkshoods), which like Aquilegia produce cardiogenic toxins. [11]

Insects

They are used as food plants by some Lepidoptera (butterfly and moth) caterpillars. These are mainly of noctuid moths noted for feeding on many poisonous plants without harm such as cabbage moth (Mamestra brassicae), dot moth (Melanchra persicariae) and mouse moth (Amphipyra tragopoginis). The engrailed (Ectropis crepuscularia), a geometer moth, also uses columbine as a larval food plant. The larvae of the Papaipema leucostigma also feed on columbine. [12]

Plants in the genus Aquilegia are a major food source for Bombus hortorum , a species of bumblebee. Specifically, they have been found to forage on species of Aquilegia vulgaris in Belgium and Aquilegia chrysantha in North America and Belgium. The bees do not show any preference in color of the flowers. [13]

Cultivation

Columbine cultivar 'Magpie' Aquilegia columbine magpie cultivar 2.jpg
Columbine cultivar 'Magpie'

Columbine is a hardy perennial, which propagates by seed. It will grow to a height of 40–50 cm (15–20 in). 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 3 in the United States so does not require mulching or protection in the winter. [14] [15]

Large numbers of hybrids are available for the garden, since the European A. vulgaris was hybridized with other European and North American varieties. [16] Aquilegia species are very interfertile, and will self-sow. [17] Some varieties are short-lived so are better treated as biennials.

The British National Collection of Aquilegias was held by Mrs Carrie Thomas at Killay near Swansea. [18] 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. [19] Aquilegia can be grown from seeds or rhizomes. [20]

Uses

Double-flowered Aquilegia x hybrida Doubled flower of Aquilegia x hybrida 001.JPG
Double-flowered Aquilegia × hybrida

The flowers of various species of columbine 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, however, are highly poisonous 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 a treatment for ulcers. [21] However, the medical use of this plant is better avoided due to its high toxicity; columbine poisonings may be fatal. [11]

An acute toxicity 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. [22]

Culture

The Colorado blue columbine (A. coerulea) is the official state flower of Colorado (see also Columbine, Colorado). It is also used as a symbol of the former city of Scarborough in the Canadian province of Ontario. [23]

Wild columbine (Aquilegia canadensis) growing in Sleeping Bear Dunes National Lakeshore near Glen Arbor, Michigan Wild Columbine Michigan.jpg
Wild columbine (Aquilegia canadensis) growing in Sleeping Bear Dunes National Lakeshore near Glen Arbor, Michigan

Evolution

Columbines have been important in the study of evolution. It was found that the Sierra columbine (A. pubescens) and crimson columbine (A. formosa) each has adapted specifically to a pollinator. Bees and hummingbirds are the visitors to A. formosa, while hawkmoths would only visit A. pubescens when given a choice. Such a "pollination syndrome", being due to flower color and orientation controlled by their genetics, ensures reproductive isolation and can be a cause of speciation. [24]

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. [25] 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. [3]

Species

Dark columbine (Aquilegia atrata) Schwarzviolette Akelei (Aquilegia atrata) 02.jpg
Dark columbine (Aquilegia atrata)
Aquilegia alpina Aquilegia alpina1JUSA.jpg
Aquilegia alpina
Fan columbine (Aquilegia flabellata) Aquilegia flabellata1.jpg
Fan columbine (Aquilegia flabellata)
Fragrant columbine (Aquilegia fragrans) Aquilegia fragrans 2.jpg
Fragrant columbine (Aquilegia fragrans)
Aquilegia x maruyamana Aquilegia x maruyamana1.jpg
Aquilegia × maruyamana
Pyrenean columbine (Aquilegia pyrenaica) Aquilegia pyrenaica1JUSA.jpg
Pyrenean columbine (Aquilegia pyrenaica)

130 columbine species are accepted. [1] [26]

See also

Notes

  1. In the Ranunculaceae, a variety of terms are used to describe the whorl of structures between the sepals and stamens, including honey-leaves, petals, staminodes or nectaries [5]

Related Research Articles

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<i>Linaria</i> Genus of flowering plants in the family Plantaginaceae

Linaria is a genus of almost 200 species of flowering plants, one of several related groups commonly called toadflax. They are annuals and herbaceous perennials, and the largest genus in the Antirrhineae tribe of the plantain family Plantaginaceae.

<i>Pulsatilla</i> Genus of flowering plants in the buttercup family

The genus Pulsatilla contains about 40 species of herbaceous perennial plants native to meadows and prairies of North America, Europe, and Asia. Derived from the Hebrew word for Passover, "pasakh", the common name pasque flower refers to the Easter (Passover) flowering period, in the spring. Common names include pasque flower, wind flower, prairie crocus, Easter flower, and meadow anemone. Several species are valued ornamentals because of their finely-dissected leaves, solitary bell-shaped flowers, and plumed seed heads. The showy part of the flower consists of sepals, not petals.

<i>Aquilegia formosa</i> Western North American species of columbine

Aquilegia formosa, the crimson columbine, western columbine, or (ambiguously) "red columbine", is a common wildflower native to western North America, from Alaska to Baja California, and eastward to Montana and Wyoming.

<i>Saponaria</i> Genus of flowering plants

Saponaria is a genus of flowering plants in the family Caryophyllaceae, native to Asia and Europe, and are commonly known as soapworts. They are herbaceous perennials and annuals, some with woody bases. The flowers are abundant, five-petalled and usually in shades of pink or white. The genus is closely related to the genus Silene, being distinguished from these by having only two styles in the flower. It is also related to Gypsophila, but its calyx is cylindrical rather than bell-shaped.

<i>Aquilegia canadensis</i> Common North American species of columbine

Aquilegia canadensis, the Canadian or Canada columbine, eastern red columbine, or wild columbine, is a species of flowering plant in the buttercup family Ranunculaceae. It is an herbaceous perennial native to woodland and rocky slopes in eastern North America, prized for its red and yellow flowers. It readily hybridizes with other species in the genus Aquilegia.

<i>Aquilegia coerulea</i> North American species of 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.

<i>Angelonia</i> Genus of flowering plants

Angelonia is a genus of about 30 species which occur from Mexico to Argentina and is classified in the Plantaginaceae. They are herbaceous plants occurring mainly in arid and semi-arid habitats. Most Angelonia species can be found in Northeastern Brazil in the seasonally-dry tropical forest namely Caatinga. The flowers of Angelonia are highly specialized for pollination because they have hairs in the inner corolla, which produces oils collected by oil bee pollinators, especially of the genus Centris.

<span class="mw-page-title-main">Pollination syndrome</span> Flower traits that attract pollinators

Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic or biotic, such as birds, bees, flies, and so forth through a process called pollinator-mediated selection. These traits include flower shape, size, colour, odour, reward type and amount, nectar composition, timing of flowering, etc. For example, tubular red flowers with copious nectar often attract birds; foul smelling flowers attract carrion flies or beetles, etc.

<i>Aquilegia pubescens</i> Californian endemic species of columbine

Aquilegia pubescens is a high-altitude species of columbine known by the common names Sierra columbine, alpine columbine and Coville's columbine. Its flowers are large and usually a creamy white.

<i>Aquilegia elegantula</i> North American species of columbine

Aquilegia elegantula is a species of columbine known by the common name western red columbine. It is native to the southwestern United States, particularly the Four Corners states, and northern Mexico, where it grows in moist areas in mountain coniferous forests. It is a rhizomatous perennial herb growing 10 t 60 centimeters tall. The green leaf blades are borne on long, slender petioles and divided into three leaflets which each have rounded lobes along the front edges. The flower has five long petals up to 3 centimeters in length including their elongated, knob-tipped spurs. The petals are bright red in the spurs and lighten to yellow-green or orange at the tips. Between the petals are the oval-shaped sepals, which are reddish to yellowish in color and are held parallel to the petals. Flowers often droop such that the mouth is toward the ground and the spurs point up; the flowers are pollinated by the broad-tailed hummingbird.

<i>Aquilegia micrantha <span style="font-style:normal;">var.</span> grahamii</i> Species of columbine endemic to Utah

Aquilegia micrantha var. grahamii is a variety of flowering plant in the buttercup family known by the common name Graham's columbine. It is endemic to Utah in the United States, where it is known only from Uintah County. It occurs in three canyons along the Uinta Mountains. There are an estimated 5,000 to 10,000 individuals.

<i>Aquilegia pyrenaica</i> Pyrenean endemic species of columbine

Aquilegia pyrenaica, common name Pyrenean columbine, is a species of flowering plant in the family Ranunculaceae. It is endemic to the Pyrenees where it grows on grassland and in rocky places. It was first described in 1805 by Augustin Pyramus de Candolle who gave it the name Aquilegia pyrenaica.

<i>Aquilegia longissima</i> North American species of columbine

Aquilegia longissima, the long-spur columbine, is a rare perennial herb that is native to northern Mexico, Trans-Pecos Texas and southern Arizona. In the original species description by Asa Gray in 1883, Aquilegia longissima is distinguished from the shorter-spurred golden columbine Aquilegia chrysantha based on the extremely long spur over 10 cm long, narrow spatulate petals and very slender spurs that usually hang straight down. The petal spurs have been known to reach 16 cm in length, the longest nectary spurs of any Eudicot. The plants typically flower in the fall after monsoon rains and are found in oak-pine-juniper woodlands in shaded canyons near intermittent streams or on talus slopes.

<span class="mw-page-title-main">Nectar spur</span>

A nectar spur is a hollow extension of a part of a flower. The spur may arise from various parts of the flower: the sepals, petals, or hypanthium, and often contain tissues that secrete nectar (nectaries). Nectar spurs are present in many clades across the angiosperms, and are often cited as an example of convergent evolution.

<i>Aquilegia ottonis</i> European species of columbine

Aquilegia ottonis is a species of flowering plant in the buttercup family. It has a broad distribution in Europe; it is native to Greece, Italy, and Albania. Plants produce blue-purple flowers which, based on pollination syndromes, are thought to be pollinated by bumblebees. A. ottonis is named after King Otto of Greece; the specific epithet 'ottonis' is the third declension of 'otto'.

<i>Aquilegia sibirica</i> Species of flowering plant

Aquilegia sibirica, the Siberian columbine, is a species of flowering plant in the family Ranunculaceae native to the north-central Asian regions of Siberia, northern Mongolia, Kazakhstan, and Xinjiang. A hardy perennial plant, it prefers temperate environments. The Siberian columbine can be between 1 foot (0.30 m) and 2 feet (0.61 m) tall with flowers that are lilac-blue and white in color.

<i>Aquilegia glandulosa</i> North Asian species of columbine

Aquilegia glandulosa, the Siberian columbine, is a perennial species of plant in the family Ranunculaceae, native to northern and central Asia.

<i>Aquilegia micrantha</i> Species of flowering plants

Aquilegia micrantha, the Mancos columbine or Bluff City columbine, is a perennial species of plant in the family Ranunculaceae, native to Utah, Colorado, and Arizona.

<i>Aquilegia <span style="font-style:normal;">×</span> hybrida</i> Species of flowering plants

Aquilegia × hybrida is a hybrid species of columbine, the result of a cross between Aquilegia vulgaris and Aquilegia canadensis.

References

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  11. 1 2 Tilford (1997)
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Bibliography