Linaria cymbalaria var. minor Goiran ex Fiori & Bég.
Linaria hederifolia Steud.
Linaria hederifolia St.-Lag.
Tursitis cymbalaria(L.) Raf.
Cymbalaria muralis, commonly called ivy-leaved toadflax or Kenilworth ivy, is a low, spreading, viney plant with small purple flowers, native to southern Europe. It belongs to the plantain family (Plantaginaceae), and is introduced to many other temperate locations. The flower stalk is unusual for seeking light until it is fertilized, after which it grows away from the light. Other names include coliseum ivy, Oxford ivy, mother of thousands, and wandering sailor.
Cymbalaria muralis spreads quickly, growing 10–80cm (3.9–31.5in) tall.[3][4][5] It is a biennial or short lived perennial plant.[6] Its roots are thin and fiberous for reaching into cracks.[7] Its rounded, heart-shaped, or kidney shaped leaves are 5–40millimeters long, 6–60millimeters wide, and are supported by nearly all alternating, thin stems, 10–22millimeters long, and usually purple in color.[5][3][4] The leaves are either smooth (glaberous) or with widely scattered hairs in the subspecies muralis[3] or quite fuzzy (villous) in the subspecies visianii.[8] They may have from three to seven lobes,[5] but most often has five lobes.[4] The leaves are relatively thick and often blushed with purple on their undersides.[4]
Cymbalaria muralis flower
Cymbalaria muralis has small flowers that strongly resemble those of a snapdragon.[5] They have sepals at the base of the flower that have lobes 1.5–3millimeters long and are sharply pointed. The petals form a closed tube 3–5millimeters long. The lower lip of the flower expanded upwards to block the tube of the flower similarly to the well known snapdragon flowers (palate inflated). The two cushion shaped parts of the lip under the mouth have yellow spots.[4] The lower lobes of the flower are spreading while the upper ones stand up, with rounded ends and 2–3millimeters long.[3] The majority of the flower is lilac in color with a yellow throat[5] with darker lines on the upper lobes.[4] Rarely the flowers may be all or nearly all white. At the rear of the flower there is a short spur that is about ⅓ the total length of the flower.[4]
Flowering is dependent on local conditions. In Britain it may flower from May until the end of November,[9] while in Eastern Europe it does so from June through July with seeds in September.[10] In North America it may bloom from May to October.[3] In New Zealand it blooms all year long, from January to December.[11]
Cymbalaria muralis producing fruits and seeds
The flowers are pollinated by bees, but are also self-compatible.[4] Once a flower is fertilized it forms a globular capsule 3–5millimeters in diameter.[3] Each of the capsules has two openings, each with three valves.[12] The seeds are quite small, just 0.5–1millimeter with a crescent shape covered in minute warty bumps.[3] This plant has an unusual method for planting its own seeds. The flower stalk is initially positively phototropic and moves towards the light. After fertilisation, it becomes negatively phototropic ("scototropic") and moves away from the light. This results in seed capsules being pushed into dark crevices of rock walls, where it is more likely to germinate.[13]
Taxonomy
Cymbalaria muralis botanical illustration from Deutschlands Flora in Abbildungen, 1796
As with many plants, Cymbalaria muralis was given its first scientific name and description by the early taxonomist Carl Linnaeus. He placed it in the genus Antirrhinum with plants commonly called snapdragons in 1753 as Antirrhinum cymbalaria. Its taxonomic history since that date is quite complex with twenty-two species and five subspecies that are considered to be taxonomic synonyms as of 2024.[1]
The very first is a reclassification in genus Linaria, the toadflaxes, as Linaria cymbalaria by the botanist Philip Miller in 1768. Four other names were published after this in Linaria that had acceptance for a time and Linaria cymbalaria continued to be used alongside other names as late as 1902.[1]
A spelling variation (see Orthographical variant) created by the botanist Noël Martin Joseph de Necker 1773 when he spelled it Antirrhinum cimbalaria. The species was described and given a taxonomic superfluous name six years later in 1779 by the naturalist Jean-Baptiste Lamarck as Antirrhinum hederaceum. Likewise the botanist Richard Anthony Salisbury gave it a similar superfluous name, Antirrhinum hederifolium, in 1796. It was described four more times under different names in Antirrhinum in the 1800s.[1]
As of 2024 there are two widely accepted subspecies. The autonymCymbalaria muralis subsp. muralis and Cymbalaria muralis subsp. visianii.[1][20] In some sources Cymbalaria muralis subsp. pubescens continues to be listed as a valid subspecies, but it is not listed as valid by most sources.[21]
Cymbalaria muralis subsp. muralis
This subspecies is hairless or nearly so in most parts.[3] It is widely spread and commonly encountered across much of the temperate world, see distribution for details.[22]
Cymbalaria muralis subsp. visianii
This subspecies was first described in 1925 as Cymbalaria muralis f. visianii by Sándor Jávorka using an incomplete description by Jenő Béla Kümmerle.[23] In 1972 the Irish botanist D. A. Webb described it with its current name making the distinction from Cymbalaria pilosa, which as of 2024 is regarded as a synonym of Sibthorpia europaea by POWO.[24] All parts of the plants are covered in fine hairs (villous) except for the seed pods, which are hairless.[25]
Names
Cymbalaria muralis flower side view
The species name, "muralis", comes from the Latin "mūrālis" for walls and relating to them, for its habit of growing from them.[26][27]
One of its most frequent common names in English, "ivy-leaved toadflax", is a compound name that describes the appearance of the plant. The leaves of the species are similar to those of ivy. Its flowers are similar to those of the related Linaria genus, which is called toadflax because it is a frequent weed of flax grown as a crop and the flowers reminded people of a toad's mouth.[28] The common name "Kenilworth ivy" is speculated to come from the English town of Kenilworth or the castle of the same name which is near where the plant first appeared in England in the 1600s.[6]
Related to its habit of growing on ruins it is also called "coliseum ivy", a name that has been in use since 1864.[6][29] From its first location of introduction in England it is also called "Oxford ivy" and was previously called "Oxford weed".[30] The common name "mother of thousands" is applied to this species and also to Saxifraga stolonifera, Tolmiea menziesii, and Soleirolia soleirolii.[31][32][33] Other less common names related to its ivy like appearance include "ivy-weed", "ivy-wort", and "Kentucky-ivy".[34] It is also called "pennywort", but this name is shared with many other vaguely round leafed herbaceous plants.[30][34] Less common names shared with other plants include "cancer root" (with Conopholis americana and Epifagus virginiana),[34] and "wandering sailor"[30] (with genus Lysimachia[35]), and "Wandering Jew" (with Saxifraga stolonifera and Tradescantia fluminensis).[34] Two other nautical names include "climbing sailor" and "roving sailor".[34] It has also been called "pedlar's basket", "rabbits", "roving Jenny", and "Devil's ribbon".[7][34]
Distribution
Cymbalaria muralis growing on a wall
POWO regards the native range for this species to be limited to five areas of southern Europe; Austria, France, Italy, Switzerland, and the former Yugoslavia, identical to their recorded native range for Cymbalaria muralis subsp. muralis.[1][22] Similarly the World Plants database records it as native to the same areas, but more specifically to Slovenia, Croatia, Bosnia & Hercegovina, Serbia, and Kosovo in the former Yugoslavia and to the small state of San Marino on the Italian peninsula. It also shows it as native to additional areas north east of the Alps such as Czechia, Slovakia, and Liechtenstein, and many Atlantic Ocean islands such as the Azores, Madeira, the Canary Islands, and the Cape Verde Islands.[36]
Cymbalaria muralis subsp. visianii recorded by POWO as growing in just Italy and the former Yugoslavia.[23] The World Plants database largely agrees listing it as native to the central and southern parts of Italy and to Croatia, but also listing it as introduced in Germany.[36]
From this original range it spread to much of Europe and to the rest of the world as either an accidental introduction or because of its use as a ornamental plant.[37] Its first record outside of cultivation in Great Britain is from 1640.[4] A frequently repeated story is that the plants were introduced accidentally as part of a shipment of statuary to Oxford.[30] However, this is unlikely since it was recorded growing in an English garden in 1617 in Droxford.[38] Thereafter it became a popular ornamental plant that was widely planted in the United Kingdom through the 19th century.[39] Regardless of its status it is recorded from Ireland and Portugal in the west in every country as far east as Ukraine, Poland, and Sweden, and according to POWO it may also be found in the Baltic States, northwestern Russia, and the North Caucasus[1] with World Plants recording it in Belarus and Georgia.[36]
In Africa it grows in Morocco, Algeria, and Tunisia in the north and in South Africa it grows in the Cape Provinces and the Northern Provinces floristic areas. In Asia it is now found in Turkey, Jordan, North Korea, South Korea, and the eastern Himalayas.[1]
In North America Cymbalaria muralis has been recorded growing outside cultivation in Bermuda, Canada, Costa Rica, Guatemala, Honduras, Jamaica, Mexico, and the United States.[1][36] In the US it is generally agreed that it grows in much of the Midwest, north-east, and New England as far south as South Carolina, Tennessee, and Arkansas and as far west as Wisconsin, Illinois, and Missouri with the exception of Maine and New Hampshire. It is also found on the west coast in California, Oregon, and Washington State. It is more scattered in the interior states, listed as growing in Colorado, Nebraska, South Dakota, and Utah, with the last of these recorded by USDA Natural Resources Conservation Service PLANTS database (PLANTS) and World Plants, but not POWO.[40][1][36] In Canada it agreed that it grows in five provinces, British Columbia, Manitoba, New Brunswick, Ontario, and Québec. However PLANTS and World Plants also record it in Nova Scotia[40][36] while POWO alone records it on the island of Newfoundland.[1]
In South America it has been observed in Colombia, Ecuador, and Peru in the north.[36] It has also been recorded in the south and southeast of Brazil, Uruguay, Bolivia, and the north of Argentina.[1] In mainland Chile it has been reported from Valparaíso, in Santiago, the Maule Region, and the Biobío Region.[36]
On oceanic islands it has been introduced to Maui in Hawaii,[40][36] both the North and South Island of New Zealand,[41] Bermuda, Mauritius, St. Helena,[1] and the Juan Fernández Islands.[36] It is also found in six Australian states, Western Australia, South Australia, Victoria, New South Wales, the south of Queensland, and the island state of Tasmania.[41]
Habitat
The original habitat for Cymbalaria muralis was the narrow niches of rock faces and cliffs.[42] In cooler climates like Lower Silesia in Poland it grows in warm microclimates such as on south or west facing walls of structures or the embankments of rivers.[37] It is considered very characteristic of the vegetation that grows on walls.[43]
In hot climates it becomes a high altitude specialist, for example growing from 1100–1300meters in elevation in Costa Rica.[44]
Ecology
Cymbalaria muralis is a generalist, attracting a wide range of pollinators, including bees, flies, and butterflies.[45] Their roots are associated with arbuscular mycorrhiza, a group of funguses that partner with plants.[46]
At least three aphid species Myzus ornatus (violet aphid), Myzus persicae (green peach aphid), and Myzus cymbalariae are commonly found on Cymbalaria muralis. The last of these was first observed feeding on ivy-leaved toadflax and is named for the species.[47] The plant is also a host for tomato ringspot virus.[48]
Uses
Edibility
Ivy-leaved toadflax is sometimes used as a salad green in Southern Europe.[9] Its taste is described as like raw garden peas[49] or bitter with a sharp numbing sensation,[9] however it has also been noted as slightly toxic by researchers Marion Cooper and Anthony Johnson.[50]
Cultivation
In gardens ivy-leaved toadflax is planted as a groundcover, particularly on rocky slopes, and planted in wall cracks. Once established it will reproduce on its own, both from seeding and stems rooting themselves.[6] It is winter hardy in USDA zones 5 to 9.[5] According to the garden author Carolyn Singer it is deer resistant.[51]
Related Research Articles
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Linaria vulgaris, the common toadflax, yellow toadflax or butter-and-eggs, is a species of flowering plant in the family Plantaginaceae, native to Europe, Siberia and Central Asia. It has also been introduced and is now common in North America.
Antirrhinum majus, the common snapdragon, is a species of flowering plant belonging to the genus Antirrhinum. The plant was placed in the family Plantaginaceae following a revision of its prior classical family, Scrophulariaceae.
Nuttallanthus canadensis, the blue toadflax, Canada toadflax, or old-field toadflax, is a species of Nuttallanthus in the family Plantaginaceae, native to eastern North America from Ontario east to Nova Scotia and south to Texas and Florida.
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Anchusa officinalis, also knowns as common bugloss or common alkanet, is a species of flowering plant in the borage family. It is native to Europe and small parts of western Asia, but has been escaped from cultivation to grow in additional locations in Europe and the Americas. The flowers are noted for their popularity with bumblebees due to a large nectar flow. The plants have been used in traditional medicines, but were falling out of favor by the early 1800s. They are still planted in gardens for their popularity with bees and their blue flowers.
Lactuca muralis, the wall lettuce, is a perennial flowering plant in the tribe Cichorieae within the family Asteraceae, also referred to as Mycelis muralis.
Linaria bipartita is a species of flowering plant in the plantain family known by the common name clovenlip toadflax. It is native to Morocco, but it can be found elsewhere as an introduced species and it is cultivated as an ornamental plant. It is an annual herb growing 10 to 30 centimeters tall with linear leaves 3 to 5 centimeters in length. The inflorescence is a raceme of flowers occupying the top of the stem. The flower is about 2 centimeters long with five lobes arranged into two lips with a spur at the end. The flower is often reddish purple in color, but flowers of many different colors are bred for the garden. The fruit is a spherical capsule about 2 millimeters wide.
Linaria maroccana is a species of flowering plant in the plantain family known by the common names Moroccan toadflax and annual toadflax. It is native to Morocco, but it can be found elsewhere growing wild as an introduced species, such as California. It is a readily available ornamental plant for the flower garden. This is an annual herb growing erect to approach a maximum height of 50 cm (20 in), its stem with linear leaves 2–4 cm (0.8–1.6 in) long. The inflorescence is a raceme of flowers occupying the top of the stem. At the base of each flower is a calyx with five narrow, pointed lobes. The flower is 2–4 cm (0.8–1.6 in) long with five lobes arranged into two lips with a spur at the end. The flower is often purple in color with white near the throat, but flowers of many different colors are bred for the garden. Dwarf cultivars are also available.
Linaria dalmatica is a herbaceous, short-lived perennial plant native to western Asia and southeastern Europe that has become a weed in other areas. The family this plant now belongs to is the Plantaginaceae Family. Previously, it belonged to the Scrophulariaceae (Figwort) family. Its common names include Balkan toadflax, broadleaf toadflax, and Dalmatian toadflax. Linaria dalmatica has unique yellow flowers with an orange center that draw individuals to purchase them to display in their gardens. The distribution of L. dalmatica to North America can be attributed to use as a fabric dye, folk remedies and as an ornamental plant. However, it is now classified as a weed in both Canada and the U.S.A.
Symphyotrichum lanceolatum is a species of flowering plant in the family Asteraceae native to North America. Common names include panicled aster, lance-leaved aster, and white panicled aster. It is a perennial, herbaceous plant that may reach 1.5 meters tall or more, sometimes approaching 2 m. The lance-shaped leaves are generally hairless but may feel slightly rough to the touch on the top because of tiny bristles. The flowers grow in clusters and branch in panicles. They have 16–50 white ray florets that are up to 14 millimeters long and sometimes tinged pink or purple. The flower centers consist of disk florets that begin as yellow and become purple as they mature.
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Townsendia grandiflora, commonly called largeflower ground-daisy, largeflower townsendia, or showy townsendia is an inconspcious plant of the foothills of the Rocky Mountains and nearby shortgrass prairies. They are usually found in somewhat eroded or rocky areas such as the sides of hills, banks, and mesas. They are part of the Townsendia genus, which can be difficult for both amateur and expert botanists to correctly identify. A short-lived plant, they are nonetheless grown by rock garden enthusiasts for their relatively large daisy-like flowers.
↑ Kuprianova, L.A. (1968). "Genus 1326. CYMBALARIA Medic". In Komaroav, V.L; Schischkin, B.K.; Bobrov, E.G.; Shetler, Stanwyn G.; Fet, Galina N. (eds.). Flora of the U.S.S.R. Vol.XXII: Solanaceae and Scrophulariacea. Jerusalem, Israel: Israel Program for Scientific Translations. p.156. Retrieved 9 January 2024.
↑ Steudel, Ernst Gottlieb (1821). Nomenclator Botanicus (vol. 1) (in Latin). Stuttgardtiae et Tubingae, sumtibus I.G. Cottae. p.482. Retrieved 1 January 2024.
1 2 3 4 5 6 7 8 9 10 Hassler, Michael (3 January 2024). "Cymbalaria". World Plants. Synonymic Checklist and Distribution of the World Flora. Version 18.3. Retrieved 5 January 2024.
1 2 Szczęśniak, Ewa; Świerkosz, Krzysztof (2003). "Cymbalaria muralis P.Gaertn., B.Mey. & Schreb. and Cymbalarietum muralis Görs 1966 in Lower Silesia – expansion or regression?". In Zając, Adam; Zając, Maria; Zemanek, Bogdan (eds.). Phytogeographical Problems of Synanthropic Plants. Cracow, Poland: Jagiellonian University. pp.185–193. doi:10.6084/M9.FIGSHARE.105653. ISBN8391516148. Retrieved 5 January 2024.
↑ Guzmán, B.; Gómez, J. M.; Vargas, P. (July 2017). "Is floral morphology a good predictor of floral visitors to Antirrhineae (snapdragons and relatives)?". Plant Biology. 19 (4): 515–524. Bibcode:2017PlBio..19..515G. doi:10.1111/plb.12567. PMID28316136.
↑ Harley, J. L.; Harley, E. L. (February 1987). "A Check-list of Mycorrhiza in the British Flora". New Phytologist. 105 (s1): 69. doi:10.1111/j.1469-8137.1987.tb00674.x.
↑ Stroyan, H. L. G. (December 1967). "A Replacement Name in Myzus Passerini (Homoptera: Aphidoidea)". Proceedings of the Royal Entomological Society of London. Series B, Taxonomy. 36 (11–12): 186. doi:10.1111/j.1365-3113.1967.tb00536.x.
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