Marasmius rotula

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***Marasmius rotula*** is a common species of agaric fungus in the family Marasmiaceae. Widespread in the Northern Hemisphere, it is commonly known variously as the **pinwheel mushroom**, the **pinwheel marasmius**, the **little wheel**, the **collared parachute**, or the **horse hair fungus**. The type species of the genus *Marasmius*, *M. rotula* was first described scientifically in 1772 by mycologist Giovanni Antonio Scopoli and assigned its current name in 1838 by Elias Fries.

The fruit bodies, or mushrooms, of *M. rotula* are characterized by their whitish, thin, and membranous caps up to Template:Convert wide that are sunken in the center, and pleated with scalloped margins. The slender and wiry black hollow stems measure up to Template:Convert long by Template:Convert thick. On the underside of the caps are widely spaced white gills that are attached to a collar encircling the stem. The mushrooms grow in groups or clusters on decaying wood such as fallen twigs and sticks, moss-covered logs, and stumps.

Unlike other mushrooms known to release spores in response to a circadian rhythm, spore release in *M. rotula* is dependent upon sufficient moisture. Dried mushrooms may revive after rehydrating and continue to release spores for up to three weeks—a sustained spore production of markedly longer duration than other typical agarics. There are several species of *Marasmius* with which *M. rotula* might be confused due to somewhat similar overall appearances, but differences in size, gill arrangement, and substrate are usually sufficient field characteristics to distinguish them. Although *M. rotula* mushrooms are not generally considered edible, they produce a unique peroxidase enzyme that is attracting research interest for possible use in bioengineering applications.

## Taxonomy

The species was first described by Italian mycologist Giovanni Antonio Scopoli as *Agaricus rotula* in 1772. In 1821 Elias Magnus Fries redescribed the mushroom in *Systema Mycologicum*, and later transferred it to *Marasmius* in his 1838 *Epicrisis Systematis Mycologici*. Synonyms include names derived from generic transfers to *Androsaceus* by Narcisse Théophile Patouillard in 1887, and to *Chamaeceras* by Otto Kuntze in 1898; both of these genera are now obsolete and have since been sunk back into *Marasmius*.

In his 1821 *A Natural Arrangement of British Plants*, Samuel Frederick Gray introduced the generic name *Micromphale*, including the species *Micromphale collariatum*, which was based on William Withering's 1796 *Merulius collariatus*. In 1946 Alexander H. Smith and Rolf Singer proposed to conserve the name *Marasmius* over *Micromphale*; the latter had nomenclatorial priority as it was published first. The generic name *Marasmius*, with *M. rotula* as the lectotype species, was later conserved at the 1954 Paris Congress on Botanical Nomenclature. *M. rotula* is also the type species of section *Marasmius* within the genus. This grouping of species is characterized by inamyloid flesh, a cap cuticle with broom cells (finger-like projections common to *Marasmius* species) ornamented with numerous warts, gills usually attached to a collar surrounding the stem, and the presence of black rhizomorphs on the stem.

Several varieties of *M. rotula* have been described. Miles Berkeley and Moses Ashley Curtis named var. *fuscus* in 1869 for its brown cap. In 1887 Pier Andrea Saccardo described var. *microcephalus* from Italy, with caps half the normal size. It is now understood, however, that fruit body morphology is variable and dependent upon environmental conditions. Joseph Schröter described var. *phyllophyla* in 1889, but that taxon is now treated as *Marasmius bulliardii*.

*Marasmius rotula* is commonly known as the "pinwheel mushroom", the "pinwheel Marasmius", the "collared parachute", or the "horse hair fungus". This latter name is shared with other *Marasmius* species, including *M. androsaceus* and *M. crinis-equi*. Gray called it the "collared dimple-stool". The name "little wheel fungus" is suggestive of the collar to which the gills are attached like the spokes of a wheel, like the specific epithet, which is a diminutive of *rota*, the Latin word for "wheel".

## Description

 File:Marasmius rotula 224073.jpg
The cap of the fruit body is thin and membranous, measuring Template:Convert in diameter. It has a convex shape slightly depressed in the center, conspicuous furrows in an outline of the gills, and scalloped edges. Young, unexpanded caps are yellowish brown; as the cap expands, the color lightens to whitish or light pinkish-white, often with a darker, sometimes brown center. The variety *fusca* has brown caps. The white or slightly yellowish flesh is very thin, reaching about 0.25–1.5 mm thick in the central part of the cap, and even thinner at the margin.

Gills are attached to a collar, never to the stem, although some specimens have the collar pressed close enough to it that this characteristic may be less obvious. Widely spaced, they have the same whitish to pale yellow color as the flesh, and typically number between 16 and 22. They are initially narrow, but thicken downward to about 1–3 mm at the exposed edge. The stem is Template:Convert long and up to Template:Convert thick, with a smooth, sometimes shiny surface. It is tough, hollow, and either straight or with some curving. The color is blackish-brown up to a lighter, almost translucent apex. The base of the stem may be connected to dark brown or black root-like rhizomorphs 0.1–0.3 mm thick. Mature specimens display no veil.
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Details of the fruit bodies' appearance, color in particular, are somewhat variable and dependent on growing conditions. For example, specimens growing on logs in oak and hickory forests in the spring tend to have more yellowish-white, depressed caps than those found in the same location in autumn, which are light yellow brown and more convex in shape. The fruit body development of *M. rotula* is hemiangiocarpous, with an hymenium that is only partially enclosed by basidiocarp tissues. Robert Kühner showed that a cortina-like tissue covers the young gills before the expanding cap breaks away from the stem. In unfavorable weather conditions, however, the mushrooms may fail to develop normally and instead produce semi-gasteroid basidiocarps.

### Microscopic characteristics

Viewed in deposit, such as with a spore print, the spores of *Marasmius rotula* appear white or pale yellow. Under an optical microscope, they are hyaline (translucent), teardrop- or pip-shaped, and have dimensions of 7–10 by 3–5 µm. The basidia (spore-producing cells) are four-spored, club-shaped or nearly so, and 21–21 by 4–17 µm. Along the edge of the gill, interspersed among the basidia, are non-reproductive cells, the cheilocystidia; these are club-shaped with rough wart-like protuberances on the surface. The gill edges further feature broom cells, which are variably shaped, thin-walled, and measure 7–32 by 2.5–20 µm. Their apical surfaces are covered with yellowish, blunt, and conical warts or incrustations 0.2–1.5 by 0.1–1 µm.

## Similar species

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There are several less-common species of *Marasmius* with which *M. rotula* might be confused due to somewhat similar overall appearances, but differences in size, gill arrangement, and substrate are usually sufficient field characteristics to distinguish between them. For example, *Marasmius capillaris* has a pale tan cap with a white center, and grows on oak leaves without forming clusters. Furthermore, its cap is evenly rounded, unlike the pleated and furrowed cap of *M. rotula*, and its stem is somewhat thinner (usually less than 0.3 mm) and slightly darker in color.

*M. rotula* is distinguished from *M. bulliardii* by its larger size, and greater number of gills. *M. limosus* is found in marshes, where it fruits on the dead stems of reeds and rushes. *Tetrapyrgos nigripes* (formerly treated in *Marasmius*) has white caps that are Template:Convert in diameter, attached gills that are sometimes slightly decurrent, a dark stem covered with tiny white hairs that give it a powdered appearance, and triangular to star-shaped spores. *M. neorotula*, described from Brazil, was considered by its discoverer Rolf Singer to be closely related to *M. rotula*. In addition to its tropical distribution, it can distinguished from *M. rotula* by its smaller size and more widely spaced gills. *M. rotuloides*, known only from montane forests of Trinidad, can only be reliably distinguished from *M. rotula* by microscopic characteristics: it has smaller, ovoid spores measuring 5 by 2.5 µm.

Other *Marasmius* species with a pinwheel arrangement of gills are readily distinguished from *M. rotula* by differences in color, including the orange *M. siccus*, the pink *M. pulcherripes*, and the rust *M. fulvoferrugineus*. *Mycena corticola* is smaller than *Marasmius rotula*, has a pale pink-brown cap, and is usually found growing singly or in small groups on bark near the base of living trees.

## Ecology and distribution

*Marasmius rotula* is a saprobic species and as such obtains nutrients by decomposing dead organic matter. It grows in deciduous forests and fruits in groups or clusters on dead wood (especially beech), woody debris such as twigs or sticks, and occasionally on rotting leaves. The fruit bodies, which are easily overlooked because of their diminutive size, are often present in abundance after rains. The species is relatively intolerant of low water potentials, and will grow poorly or not at all under water stress conditions. It is unable to degrade leaf litter until it becomes more fragmented and more compacted so that the water-holding capacity increases in the deeper layers of the soil. The magnolia warbler has been noted to line its nests with the fruit bodies' stems.

 File:Marasmius rotula 89484.jpg
In 1975 American mycologist Martina S. Gilliam investigated the periodicity of spore release in *M. rotula* and concluded that spore discharge did not follow a regular circadian rhythm, as is typical of agaric and bolete mushrooms, but rather was dependent on rain. A threshold of rainfall is required to elicit a spore discharge response and the duration of peak spore discharge correlates with the amount of rainfall, rather than its duration. Furthermore, Gilliam noted that spore prints were more readily obtained if the stem ends were placed in water, suggesting that water must enter through the fruit body for discharge to occur.

Like those of many other species of *Marasmius*, the fruit bodies of *M. rotula* can desiccate and shrivel in dry periods, then revive when sufficient moisture is available again in the form of rain or high humidity. Gilliam's study demonstrated that revived fruit bodies were capable of discharging spores over a period of at least three weeks, whereas previous studies using similar methods with other Agaricomycetes showed spore discharge occurred over a shorter period of up to six days after revival. The potential for sustained spore production and discharge may be due to the growth of new basidioles (immature basidia) during periods of growth, which then complete maturation when the mushroom revives. This may also explain why the gills become thicker as the mushroom matures.

The fungus is widespread and common in its preferred habitats in North America, Europe, and northern Asia. Although far less common in southerly locations, isolated collections have been reported from Africa (Congo, Nigeria, Sierra Leone, and Tanzania) and South Asia (India). In North America *M. rotula* is most common in the eastern part of the continent.

## Uses

*Marasmius rotula* is generally considered inedible, but is not poisonous. The mushroom has no distinguishable odor, and its flavor varies from mild or bitter. Louis Krieger, writing in *National Geographic* in the 1920s, noted that the mushroom was used as an addition to gravies and, when used to garnish venison, "adds the appropriate touch of the wild woodlands." The fruit bodies will bioaccumulate cadmium: a study of the metal concentration of 15 wild mushroom species of India showed that *M. rotula* accumulated the highest concentration of that metal.

A peroxidase enzyme known as *Mro*APO (*Marasmius rotula* aromatic peroxygenase) is attracting research interest for possible applications in biocatalysis. In general, enzymes that catalyze oxygen-transfer reactions are of great utility in chemical synthesis since they work selectively and under ambient conditions. Fungal peroxidases can catalyze oxidations that are difficult for the organic chemist, including those involving aromatic substrates such as aniline, 4-aminophenol, hydroquinone, resorcinol, catechol, and paracetamol. The *M. rotula* enzyme is the first fungal peroxygenase that can be produced in high yields. It is highly stable over a wide pH range, and in a variety of organic solvents. The enzyme has other potential for use as a biosensor for aromatic substances in environmental analysis and drug monitoring.

## References

1. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Scopoli 1772`
2. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Fries 1821`
3. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Fries 1838`
4. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Patouillard 1887`
5. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Kuntze 1898`
6. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Kirk 2008`
7. ↑ 7.0 7.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gray 1821`
8. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Withering 1796`
9. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Singer 1946`
10. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Taxon 1953`
11. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Taxon 1954`
12. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gilliam 1976 p112`
13. ↑ 13.0 13.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Berkeley 1869`
14. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Saccardo 1887`
15. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Schröter 1885`
16. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlMycoBank: Marasmius rotula var. phyllophila`
17. ↑ 17.0 17.1 17.2 17.3 17.4 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Roody 2003`
18. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `BMS`
19. ↑ 19.0 19.1 19.2 19.3 19.4 19.5 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Miller 2006`
20. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Arora 1986`
21. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Smith 2005`
22. ↑ 22.0 22.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Orr 1979`
23. ↑ 23.0 23.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Bessette 1997`
24. ↑ 24.0 24.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `McKnight 1987`
25. ↑ 25.0 25.1 25.2 25.3 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Jordan 2004`
26. ↑ 26.0 26.1 26.2 26.3 26.4 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gilliam 1976 pp122–7`
27. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gilliam 1976 p3`
28. ↑ 28.0 28.1 28.2 Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlMarasmius rotula (MushroomExpert.Com)`
29. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gilliam 1976 p121`
30. ↑ 30.0 30.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Roberts 2011`
31. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Healy 2008`
32. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Singer 1989`
33. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Dennis 1951`
34. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Kauffman 1918`
35. ↑ 35.0 35.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Dix 1984`
36. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Koske 1986`
37. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Schorger 1946`
38. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Roenneberg 2001`
39. ↑ 39.0 39.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gilliam 1975`
40. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Beeli 1928`
41. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Idu 2008`
42. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Beeli 1938`
43. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Vězda 1975`
44. ↑ 44.0 44.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Das 2005`
45. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Krieger 1920`
46. ↑ 46.0 46.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Yarman 2012`
47. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gröbe 2011`

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