Aquilegia canadensis, the Canadian columbine, Canada columbine, eastern red columbine, or wild columbine, is a species of flowering plant in the buttercup family. It is a perennial plant that is native to the woodlands and rocky areas in eastern North America in both the United States and southern Canada. It readily hybridizes with other plants in the columbine genus and is a common garden plant.
Description Canadian columbine is an herbaceous perennial growing tall in the wild, occasionally reaching as much as . It produces its stems and leaves from a substantial caudex, a vertically oriented underground stem. Its roots are short-lived and fibrous. The hollow stems are often entirely hairless, but can have very fine, short, and erect glandular hairs on the upper parts or covering the whole stem.
The leaves are two times ternately compound, each leaf divided into three parts with three leaflets, nine in total, though older sources report that they can have three ternate leaves. Leaves attached to the stems are smaller than the basal leaves, those springing directly from the caudex. Basal leaves are much shorter than the flowering stems, reaching lengths of , though more typically including the main petiole.
The leaflets of Canadian columbine are broadly ovate, egg shaped in outline, to almost round, but with wavy teeth or lobes. The surface of the leafletes are always hairless, but the underside can be covered in very fine and short hairs giving a pale appearance. The leaves are green to blue-green, and remains evergreen at temperatures down to and as high as . At these extremes the plant goes dormant and sprouts new leaves when favorable conditions return.
Canadian columbines start blooming in spring, as early as March in their native habitat, or as late as July in early summer. Flowering continues for about one month, dependent on weather conditions. A typical flower has five petals yellow at opening that elongate into red, nectar-bearing spurs and alternate with five red, petal like sepals. However, flowers can have four or six petals and a matching number of sepals. Rarely, flowers in the wild can be entirely salmon, yellow, or white. The flowers are nodding, hanging downwards with the spurs pointing up and the many stamens projecting downwards together. The flowers are showy and measure from the ends of the stamens to the tips of the spurs. Petals measure from end of the spur to the outer edge.
The anthers on the stamens open to release their pollen before the stigmas becomes receptive to aid cross pollination by visitors to the flowers. The flowers produce abundant dilute nectar, and attract hummingbirds, bumblebees, hawkmoths, and other bees. The ruby-throated hummingbird (Archilochus colubris) is the primary pollinator.
After pollination, Canadian columbines produce an upright cluster of five pod-shaped follicles long, which taper into a long, slender beak. When mature, the follicles split along one side to release numerous small shiny, black seeds, 1.5–2 millimeters by 0.5–1 mm. Seeds develop rapidly, reaching maturity around two weeks after flowering.
Plants typically flower in their second year, and natural reseeding allows colonies to persist, despite individual plants having a short lifespan. Plants are shorter lived when growing in rich soils.
Taxonomy
Aquilegia canadensis was given its binomial name by Carl Linnaeus in 1753. It is classified in the genus Aquilegia within the Ranunculaceae family. It has no accepted varieties, but has 11 among its 26 heterotypic synonyms.
Table of Synonyms Name Year Rank Notes Aquilegia australis 1898 species Aquilegia canadensis subsp. americana 1909 subspecies Aquilegia canadensis var. aurea 1889 variety nom. illeg. Aquilegia canadensis var. australis 1946 variety Aquilegia canadensis var. coccinea 1946 variety Aquilegia canadensis var. eminens 1953 variety Aquilegia canadensis var. flaviflora 1888 variety Aquilegia canadensis var. hybrida 1829 variety Aquilegia canadensis var. latiuscula 1946 variety Aquilegia canadensis var. longistyla 1867 variety Aquilegia canadensis var. phippenii 1888 variety Aquilegia canadensis var. vera 1893 variety Aquilegia canadensis var. violacea 1838 variety Aquilegia coccinea 1899 species Aquilegia elegans 1796 species Aquilegia eminens 1914 species Aquilegia flaviflora 1868 species Aquilegia latiuscula 1914 species Aquilegia phoenicantha 1936 species Aquilegia variegata 1794 species
Names The species name, canadensis, is Botanical Latin meaning "belonging to Canada", a reference to where the first scientific specimens were gathered. The common names Canadian columbine or Canada columbine are often used for this species. It is known by the name common columbine for its abundance throughout much of its native range. It is also called red columbine, however this name is used for other species including Aquilegia elegantula. It is additionally known as eastern columbine, eastern red columbine, wild red columbine, wild columbine, American columbine, and rock bells.
Among children the species was called honeysuckle because they would bite off the end of the spurs for the sweet nectar inside, it was most often used in Illinois, Massachusetts, and Wisconsin alongside the name honey horns. Other regional names include meetinghouses from New England, jacket-and-breeches, Jack-in-trousers, and lady's slipper in Massachusetts, red-bells and rock-lily from New Hampshire, and simply bells in Ohio. The names akeley and culverwort in English and or in French are from Quebec while the name cluckies comes from Nova Scotia. Like other species in the genus it is also simply called columbine.
In the Omaha–Ponca language Aquilegia canadensis is called , translating to "black perfume plant", related to one of its uses. In the Pawnee language it is known as or as meaning "black seed".
Distribution and habitat Aquilegia canadensis is a native to eastern and central North America, from New Brunswick to northern Florida and west to Saskatchewan and Texas. It is the sole native columbine from eastern North America, though smallflower columbine (Aquilegia brevistyla) is reported as far east as Clay County, Minnesota and northern parts of Ontario and the non-native European columbine (Aquilegia vulgaris) has escaped from cultivation in Canada, the northeast, and parts of the south.
In Canada it is native, but possibly locally extinct, in the eastern province of New Brunswick. In Quebec it is found in the areas near the St. Lawrence River, but not in the northern or eastern parts of the province. Similarly it is found throughout Southern Ontario, but only in the southern half of Northern Ontario to the west. It also grows in southern Manitoba, but only in the easternmost part of southern Saskatchewan.
In the Midwestern United States Canadian columbine is native to every state, but reaches its western limits in the Dakotas, Nebraska, Kansas, and Oklahoma. Its range extends almost to Wyoming in northern Nebraska, but is otherwise only in the easternmost portions of these states. Though native to Iowa, much of the central part of the state has no record of the species. In Missouri it is found throughout the state except for small areas of the southwest near Kansas and the southeastern lowlands. It is widespread in Minnesota, Wisconsin, and Illinois as well as Indiana, Michigan, and Ohio.
In the Northeastern United States it is recorded in every state from Maine to New Jersey, with records of the species in every county of Pennsylvania. In the Southeastern United States it is native to every state except for Louisiana. However, it is only common in northern parts of the south such as Virginia, West Virginia, and Kentucky. It becomes much less widespread in North Carolina and Tennessee and is only known from a few counties in South Carolina, Georgia, and Alabama. It only grows in the Florida panhandle and is only found in Tishomingo County, Mississippi. Likewise it mostly grows in northern parts of Arkansas. The population in Texas is disjunct from the rest of the range, growing only on the Edwards Plateau.
A. canadensis typically inhabits woodlands, forest edges, rocky slopes, and shaded bluffs. It will prefer well-drained, sandy or rocky soil, and will thrive in an area that experiences light to moderate shade. A. canadensis is best suited for sites that are not waterlogged or overly nutrient-rich. Due to its ability to both self and sexually reproduce, A. canadensis can maintain stable populations across a wide range of habitats.
Ecology
Reproduction Aquilegia canadensis has a mixed mating system, where they are able to both self-pollinate and outcross. The species is fully self-compatible, and has the ability to produce seeds without a pollinator. However, despite being self-compatible, it does not fully compensate for the absence of pollinators, since fruit and seed production is lowered when pollinators are not present.
One major factor that influences the selfing and outcrossing balance is the herkogamy of the anther and stigma. There is genetic variation in the separation of the reproductive structures between different populations, and plants that have greater separation between the anther and stigma have lower rates of autonomous selfing and are more prone to outcrossing. The separation is primarily due to differences in the pistil length, rather than positioning of the structures. The genetic difference in herkogamy between populations suggests that it can evolve through selection to regulate mating strategies.
Pollination is primarily carried out through the Ruby-Throated Hummingbird (Archilochus colubris), which is able to access the nectar stored in the elongated spurs of the flower. The Canadian columbine is one of the eight most important nectar sources for hummingbirds in eastern North America. In addition, bumblebees, hawk moths, and other bees also contribute to outcrossing between flowers. Genetic studies between A. canadensis and its sister species, A. brevistyla, which is primarily bee-pollinated, have shown that the shift to hummingbird pollination is associated with changes to spur curvature, floral pigmentation, and nectar traits.
Reproductive success is also reliant on the size of the population. Plants that grow in larger and more densely populated sites produce significantly more fruits and seeds per plant, in comparison to plants in smaller populations. Spatial isolation by itself does not have a large effect on reproductive output, but smaller population sizes reduces the opportunities for pollinator visits and outcrossing.
The rusty patched bumblebee (Bombus affinis) has been documented cutting holes in the spurs to engage in nectar robbing. Both queens and workers of the rusty patched bumblebee cut holes in the spurs, with some queens collecting pollen before cutting holes to take nectar. Workers either collected pollen or engaged in nectar robbing, not both behaviors at the same flower. Workers and queens both sometimes acquired nectar at the mouth of the flower rather than nectar robbing. Though other species have not been recorded making holes in the nectar spurs, the golden northern bumble bee (Bombus fervidus) do sometimes make use of holes that have already been made as well as collecting nectar from the mouth of the flower. Both honeybees and Lasioglossum sweat bees also use previously created perforations. Many other species of bumblebee collect nectar from Canadian columbine including the two-spotted bumble bee, brown-belted bumblebee, common eastern bumblebee, Nevada bumblebee, and American bumblebee. The half-black bumblebee both collects nectar and pollen from the flowers.
Disease and herbivory