Verbascum thapsus
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Verbascum thapsus in Hawaii
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Verbascum thapsus L. |
Verbascum thapsus (Common or Great Mullein) is a species of mullein, native to Europe, northern Africa and Asia, and introduced in the Americas and Australia.
Common mullein is a hairy biennial plant that can grow up to 2 meters. Its small, yellow flowers are densely grouped on the stem, which bolts from a large rosette of leaves. It grows in a wide variety of habitats, but favors well-lit disturbed soils, where it can grow suddenly thanks to its long-lived seeds. Common Mullein is a weedy, but rarely invasive species. While it is not very competitive, and thus not a problem for most cultures, it hosts many insects, such as the tarnished plant bug, that can be harmful to other plants. Although individuals are easy to destroy by hand, populations are difficult to destroy permanently.
V. thapsus is widely used as an herbal remedy with emollient and astringent properties. It is especially recommended for coughs and related problems, but also used in topical applications against a variety of skin problems. The plant was also used to make dyes and torches.
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[edit] Morphology
Common Mullein is a dicotyledonous biennial that produces only a rosette of leaves in its first year of growth.[1][2] The second year it produces a tall 1–2 m stem (stems reaching up to and exceeding 3 m, 10 ft have been reported)[3][4] and ends in a dense spike of flowers,[1] only a few of which flower at the same time.[5] All parts of the plants are covered with star-shaped trichomes.[6][7] The dried stem and fruits usually persist in winter.[5]
The leaves are alternate, thick and decurrent, with a shape varying between oblong and oblanceolate and a size up to 14 cm across and 50 cm long (5 inches wide and 19 inches long).[8][1] Their size diminishes as they get higher on the stem.[2][1] The stem solid stem is nearly an inch (2,5 cm) across and sometimes branched just below the inflorescence,[2] usually when damaged.[9] The plants produces shallow taproots.[1]
Flowers are pentamerous, with a 5-lobed calyx tube and corolla, the latter yellow and an inch or less wide (), and five stamen. They open for a month and a half in mid- to late summer.[7][1] The flowers are almost sessile, with only very short pedicels (2 mm, 0.08 in). The three upper stamens are longer, with their filaments covered with yellow or whitish hair, while the two other stamen have glabrous filaments.[7] The plant produces small ovoid (6 mm, 0.24 in) capsules containing large numbers of minute brown seeds less than a millimeter (0.04 in).[2]
There are three subspecies:[10]
- V. thapsus subsp. thapsus; widespread.
- V. thapsus subsp. crassifolium (Lam.) Murb.; Mediterranean region.
- V. thapsus subsp. giganteum (Willk.) Nyman; Spain, endemic.
The plant is also a parent to several hybrids.[11][12] Of these, the most common is V. × semialbum Chaub. (× V. nigrum).[11] Three—V. × kerneri Fritsch, V. × pterocaulon Franch. and V. × thapsi L. (syn. V. × spurium W.D.J.Koch)—have been reported in North America.[12][13] The others are European.[11]
[edit] Distribution and naturalization
The species has a wide native range including Europe, northern Africa and Asia, from the Azores and Canary Islands east to western China, north to the British Isles, Scandinavia and Siberia, and south to the Himalaya.[10][6][14] It has been introduced throughout the temperate world, and is naturalized in North America (including Hawaii), Australia, Chile, Hispaniola and Argentina (1925).[14][15][16]
In the United States it was imported very early and cultivated for its medicinal property. By the 1630s, it was already found in the wild and had begun spreading so much that Amos Eaton thought it was a native plant, and gives it as such in 1818.[17] In 1839 it is already reported in Michigan and in 1876, in California.[1] It is now found commonly in all the states,[18] and all the Canadian provinces.[19]
[edit] Ecological aspects
Common Mullein most frequently grows as a colonizer of bare and disturbed soil. This is in part due to its intolerance of shade and the very long periods during which the seeds can remain dormant before germination. It is not an agricultural weed, although its presence can be very difficult to completely eradicate, and is especially problematic in overgrazed pastures.[9][1][2] The species is considered a noxious weed in Colorado (Class C),[20] Hawaii[21] and Victoria, Australia (regionally prohibited in the West Gippsland region, and regionally controlled in several others).[22]
V. thapsus favors dry, sandy or gravelly soils, although it can grow in a variety of habitats: meadows, roadsides, forest clearings and pastures. Here it germinates almost solely in bare soil, at temperatures between 10 °C and 40 °C.[9] This ability to grow in a wide range of habitats has been linked to strong phenotype variation rather than adaptation capacities.[23] While it can germinate in total darkness if proper conditions are present (tests give a 35% germination rate under ideal conditions), in the wild, it will only do so if the seeds are exposed, or very close to the soil surface. While it can also appear in areas where some vegetation exist, growth of the rosettes on bare soil is four to seven times more rapid.[9]
Megachilidae bees, notably Anthidium species, use the hair in making their nests.[5] Seeds are generally too small for birds to feed on,[5] although the American Goldfinch has been reported to consume them.[24] Other bird species have been reported to consume the leaves (Hawaiian Goose)[25] or flowers (Palila)[26] of V. thapsus, or to use the plant as a source when foraging for insects (White-headed Woodpecker).[27]
[edit] Life cycle
Common Mullein requires vernalization before it can flowers, which is why it is a biennial.[9] This vernalization is linked to starch degradation in the root activated by low temperatures, and gibberellin application bypasses this requirement.[28] Plants that germinate in autumn overwinter if they are large enough—rosettes less than 15 cm (5.9 in) across die in winter—before flowering the next year. The entire plant usually dies at the end of its second year.[9] Some individuals, especially in the northern parts of the species' range, require a longer growth period and flower in their third year. Under better conditions, some individuals flower in the first year.[29] Triennial individuals were found to produce less seeds than biennial and annual ones. While year of flowering and size are linked to the environment, most other characteristics appear to be genetic.[30]
A given flower is open only for a single day. Flowers are autogamous and will self-pollinate if they have not been pollinated by insects during the day. While many insects visit the flowers, only some bees actually accomplish pollination.[9][1][2] Visitors include halictid bees and hoverflies.[5]
The seeds of V. thapsus maintain their germinative powers for decades, up to a hundred years, according to some studies.[31] Because of this, and because the plant is an extremely prolific seed bearer (up to 180,000[1][9] or 240,000[2]), it remains in the soil seed bank for extended periods of time, and can sprout from apparently bare ground,[9] or shortly after forest fires long after previous plants have died.[2] Common Mullein rarely establishes on new grounds without human intervention because its seeds are not dispersed very far. Seed dispersion requires the stem to be moved by wind or animal movement; 75% of the seeds fall within a meter of the parent plant, and 93% fall within five meters.[9]
[edit] Agricultural impacts and control
Common Mullein is not considered an agricultural weed because it cannot compete with established plants, and is easily crowded out by them, except in areas where vegetation is sparse to begin with, such as Californian semi-desertic areas of the Eastern Sierra Nevada. In such ecological contexts, it crowds out native herbs and grasses. Its tendency to appear after forest fires also disturbs the normal ecological succession.[9][2]
Despite not being an agricultural weed in itself, it hosts a number of insects and diseases, including both pests and beneficial insects.[32] It is also a potential reservoir of the Cucumber mosaic virus.[33] A study found V. thapsus hosts insects from 29 different families. Most of the pests found were western flower thrips (Frankliniella occidentalis), Lygus species such as the tarnished plant bug (L. lineolaris), and various spider mites from the family Tetranychidae. These make the plant a potential reservoir for overwintering pests.[32]
Other insects commonly found on Common Mullein feeds on Verbascum species in general. They include mullein thrips (Haplothrips verbasci),[32] Gymnaetron tetrum and the Mullein Moth (Cucullia verbasci).[1] Useful insects are also hosted by Common Mullein, including predatory mites of the Galendromus, Typhlodromus and Amblyseius genera (Phytoseiidae), the minute pirate bug (Orius tristicolor)[32] and the mullein plant bug (Campylomma verbasci).[34] The plant's ability to host both pests and beneficials makes it potentially useful to help stabilize populations of insects used for biological control in other cultures, like Campylomma verbasci and Dicyphus hesperus (Miridae), a predator of whiteflies.[35] A number of Lepidoptera species, including the Stalk borer (Papaipema nebris) and Gray Hairstreak (Strymon melinus), also use V. thapsus as a host plant.[36]
Control of the plant, when desired, is best managed via mechanical means, such as hand pulling and hoeing, preferably followed by sowing of native plants. Animals rarely graze it because of its irritating hairs, and liquid herbicides require surfactants to be effective, as the hair causes water to roll off the plant, much like the lotus effect. Burning is ineffective, as it only creates new bare areas for seedlings to colonize.[9][1][2] G. tetrum and Cucullia verbasci usually have little effect on V. thapsus populations.[1] Goats and chickens have been proposed to control Mullein.[9] Effective (when used with a surfactant) contact herbicides include glyphosate,[1][2] triclopyr[1] and sulfurometuron-methyl.[2] Ground herbicides, like tebuthiuron, are also effective, but recreate bare ground and require repeated application to prevent regrowth.[9]
[edit] Names
V. thapsus is known by a variety of names. "Common mullein" is the usual name in North America,[37][38] but "Great Mullein" is the one used in the UK.[39][40] Vernacular names include innumerable references to the plant's hairiness: "Woolly," "Velvet" or "Blanket Mullein,"[41][40] "Beggar's," "Moses'," "Poor Man's," "Our Lady's" or "Old Man's Blanket,"[38][39][42] and so on ("Flannel" is another generic name).
Some names refer to the plant's size and shape: "Shepherd's Club(s)" or "Staff," "Aaron's Rod" (a name it shares with a number of other plants with tall, yellow inflorescences), and a plethora of other "X's Staff" and "X's Rod."[39][38][43] The plant is still called "Velvet" or "Mullein Dock"—"Dock" is a British name applied to any broad-leaved plant.[44]
The specific epithet thapsus was first used by Theophrastus (as θάψος, "thapsos")[45] for an unspecified herb from the Ancient Greek settlement of Thapsos, near modern Syracuse, Sicily,[46][45] though it is often assimilated to the ancient Tunisian city of Thapsus.[47]
[edit] Uses
Common Mullein has been used since ancient times as a remedy for skin, throat and breathing ailments. It has long had a medicinal reputation, especially as an astringent and emollient. It contains mucilage, several saponins, coumarin and glycosides. Dioscorides recommended it for diseases of the lung and it is nowadays widely available in health and herbal stores. Non-medical uses have included dyeing and making torches.
[edit] Medical uses
Dioscorides first recommended the plant 2000 years ago, against pulmonary diseases,[48] and this has remained one of its primary uses, especially against cough. Leaf decoctions or herbal teas were used for expectoration, consumption, dry cough, bronchitis, sore throat and hemorrhoids. Leaves were also smoked against pulmonary ailments, a tradition that was rapidly transmitted to Native American peoples.[39][49] Natives also used the plants to make syrups against croup. The combination of expectorant saponins and emollient mucilage makes the plant particularly effective for cough. All preparations meant to be drunk have to be finely filtered to eliminate the irritating hairs.[28]
Oil from the flowers was used against catarrhs, colics and, in Germany, earaches, frostbite, eczema and other external conditions.[39] Topical application of various V. thapsus-based preparations was recommended for the treatment of warts,[50] boils, carbuncles, hemorrhoids, and chilblains, amongst others.[39][49] Recent studies have found that Common Mullein (especially the flowers) contains glycyrrhizin compounds with bactericide and potential anti-tumoral action.[51] Different extracts have varying levels of efficiency against bacterias.[28] In Germany, a governmental commission sanctioned medicinal use of the plant for catarrhs.[52] It was also part of the National Formulary in the United States[49] and United Kingdom.[39] The plant's leaves, in addition to the seeds, have been reported to contain rotenone, although quantities are unknown.[53]
In Spanish, Great Mullein is called Gordolobo, a name transferred to Gnaphalium conoideum. G. conoideum is used similarly by the Mexican Aztecs, which caused confusion in naming, and both are sold under the name "Gordolobo," which has lead to at least one case of poisoning due to confusion of G. conoideum with Senecio longilobus.[54]
[edit] Other uses
Like many ancient plants (Pliny the Elder describes it in his Naturalis Historia),[55] Common Mullein was linked to witches,[39] although the relationship remained generally ambiguous, and the plant was also widely held to ward off curses and evil spirits.[39][49][48][28] The seeds contain several compounds (saponins, glycosides, coumarin, rotenone) that cause breathing problems in fish, and have been widely used for fishing.[1][56]
The flowers provide dyes of bright yellow or green, and have been used for hair coloring.[39][53] The dried leaves and hair were made into candle wicks, or put into shoes to help with insulating them. The dried stems were also dipped into suet or wax to make torches.[39][49]
[edit] References
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- ^ Coutlee, Ellen L. (December 1963). "Maintenance Behavior of the American Goldfinch". Wilson Bulletin 75 (4): 342-357. ISSN 0043-5643.
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- ^ Watts, Elsevier's dictionary of plant names, pp.302, 634.
- ^ a b (French)Carnoy, A. (1959). Dictionnaire étymologique des noms grecs de plantes. Louvain: Publications Universitaires. OCLC 3284108.
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- ^ In book 25, Pliny describes "two principal kinds [of verbascum]" (Latin, English) thought to be V. thapsus and V. sinuatum. The precise attribution of a third kind is unclear
- ^ Wilhelm, Gene (April 1974). "The mullein: Plant piscicide of the mountain folk culture". Geographic Review 64 (2): 235-252. DOI:10.2307/213812.