Myriostoma coliforme | |
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Myriostoma coliforme in the Netherlands | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Class: | Agaricomycetes |
Order: | Geastrales |
Family: | Geastraceae |
Genus: | Myriostoma |
Species: | M. coliforme |
Binomial name | |
Myriostoma coliforme | |
Synonyms [1] | |
Myriostoma coliforme is a fungal species in the family Geastraceae. Basidiocarps resemble earthstars, but the spore sac is supported by multiple columns (instead of a single column) and has multiple ostioles instead of a single, apical ostiole. In the UK, its recommended English name is pepper pot. [2] It has also been called "salt-shaker earthstar". [3] The fungus has a north temperate distribution, but was formerly thought to be more widespread due to confusion with related Myriostoma species. Myriostoma coliforme is an uncommon species and appears on the Red Lists of 12 European countries. In 2004 it was one of 33 species proposed for protection under the Bern Convention by the European Council for Conservation of Fungi.
The species was first mentioned in the scientific literature by Samuel Doody in the second edition of John Ray's Synopsis methodica stirpium Britannicarum in 1696. [4] Doody briefly described the fungus as: "fungus pulverulentus, coli instar perforatus, cum volva stellata" (a powdery fungus, perforated like a colander, with a star-shaped volva), and went on to explain that he found it in 1695 in Kent. [5]
It was first described scientifically as a new species in 1785 from collections made in England by James Dickson, who named it Lycoperdon coliforme. He found it growing in roadside banks and hedgerows among nettles in Suffolk and Norfolk. [6] Nicaise Auguste Desvaux first defined and published the new genus Myriostoma in 1809, with the species renamed Myriostoma anglicum (an illegitimate renaming). [7] Christian Hendrik Persoon had previously placed the species in Geastrum in 1801, [8] while in 1821 Samuel Frederick Gray described the genus Polystoma for it. [9] Myriostoma coliforme received its current and final name when August Carl Joseph Corda moved Dickson's name to Myriostoma in 1842, replacing Desvaux's illegitimate name. [10]
The generic name Myriostoma is derived from the Ancient Greek "Myrios" (ten thousand – as in Myriad) and "stoma" (mouth) and means "thousands of mouths". The specific epithet is derived from the [Latin] words colum, meaning "strainer", and forma, meaning "shape". [11] M.J. Berkeley gave it the English name "Cullenden puff-ball" which also refers to a colander. [12] S.F. Gray called it the "sievelike pill-box". [9]
The fruit bodies start their development underground or buried in leaf debris, linked to a strand of mycelium at the base. As they mature, the exoperidium (the outer tissue layer of the peridium) splits open into 7 to 14 rays which curve backward; this pushes the fruit body above the substrate. Fully opened specimens can reach dimensions of 2–12 cm (0.8–4.7 in) from ray tip to tip. The rays are of unequal size, with tips that often roll back inward. They comprise three distinct layers of tissue. The inner pseudoparenchymatous layer (so named for the resemblance to the tightly packed cells of plant parenchyma) is fleshy and thick when fresh, and initially pale beige but darkening to yellow or brown as it matures, often cracking and peeling off in the process. The exterior mycelial layer, often matted with fine leaf debris or dirt, usually cracks to reveal a middle fibrous layer, which is made of densely packed hyphae 1–2.5 μm wide. The base of the fruit body is concave to vaulted and often covered with adhering dirt. The roughly spherical spore sac (endoperidium) measures 1–5 cm (0.4–2.0 in) in diameter, and is supported by a cluster of short columns shaped like flattened spheres. It is grey-brown and minutely roughened with small (under 0.1 mm high), lightly interconnected warts. There are several to many evenly dispersed mouths, the ostioles, mainly on the upper half of the endoperidium. They are roughly circular with fimbriate edges. [13] The inedible fruit bodies have no distinct taste, although dried specimens develop an odour resembling curry powder or bouillon cubes. [14]
Like earthstars, the fungus uses the force of falling raindrops to help disperse the spores, which are ejected in little bursts when objects (such as rain) strike the outer wall of the spore sac. The gleba is brown to greyish-brown, with a cotton-like texture that, when compressed, allows the endoperidium to flex quickly and create a puff of air that is forced out through the ostioles. This generates a cloud of spores that can then be carried by the wind. [15] There are columellae (sterile structures that start at the base of the gleba and extend through it), which are usually not evident in the mature gleba, but apparent at the base of the spore sac. The columellae are not connected to the ostioles, but rather, terminate within the gleba at some distance from them. The capillitia (sterile strands within the gleba) are long, slender, free, tapering, unbranched, and 3.3–4 μm thick, with thickened walls. The spores are spherical, nonamyloid, and are ornamented with irregularly shaped flaring protuberances up to 1.6 μm high. They measure 6.1–8.0 μm including the ornamentation. [16]
In Europe, Myriostoma coliforme is a distinctive species easily characterized by the multiple openings on its spore sac and the multiple stalks supporting the sac. [15] Historically, it was thought that the holes might have been a result of insects. This was discussed and rejected by Thomas Jenkinson Woodward in 1797:
"It has been doubted whether these mouths might not be accidental, and formed by insects after the expansion of the plant. But this (not to mention their regularity, and that each is furrowed by its border of cilia) is clearly disproved, from the marks of the projections formed by the mouths being seen on the expanded rays, when freshly opened ... I have likewise found an abortive plant, in which the seed did not ripen; but which had numerous projecting papillae on the head, where the mouths should have been formed." [17]
In North America, a second species, Myriostoma calongei , is known from New Mexico. It can be distinguished by its more distinctly verrucose (finely warted) endoperidium, with warts more than 0.1 mm high.
Myriostoma coliforme is saprotrophic, deriving nutrients from decomposing organic matter. Fruit bodies grow grouped in well-drained or sandy soil, often in the partial shade of trees. [13] The species occurs in deciduous forests and mixed forests, gardens, along hedges and grassy road banks, and grazed grasslands. It tends to grow on well-drained south-facing slopes. [18] In Europe, its major habitat is riparian mixed forests dominated by Salix alba and Populus alba along the great rivers. [19] In Hawaii, it has been collected at elevations above 2,000 m (6,600 ft) where it appears to favour the mamane ( Sophora chrysophylla ) forest. [20]
The species is widespread in north temperate regions. [nb 1] It is rare in Europe, where it appears on the Regional Red Lists of 12 countries, and is one of 33 candidate species for listing in Appendix I of the Convention on the Conservation of European Wildlife and Natural Habitats (the "Bern Convention"). [19] Although originally described from England, it was considered extinct in mainland Britain until it was found again in Suffolk in 2006 near Ipswich, one of its original localities; [15] it had been last reported in the country in 1880. [38] The fungus is considered extinct in Switzerland. [39] Its most northerly location is southern Sweden, [40] although it is generally rare in northern Europe. [15] It is similarly widespread but rarely encountered in North America, although there may be isolated localities, like New Mexico, where it is more abundant. [41]
Until 2017, Myriostoma coliforme was thought to be the only species in the genus Myriostoma, with a worldwide distribution. [16] Since then molecular research, based on cladistic analysis of DNA sequences, has revealed at least five additional species in Central and South America, South Africa, and Australia. As a result the distribution of Myriostoma coliforme sensu stricto is uncertain, but it appears to be restricted to the northern hemisphere. [16] [42] [43]
Puffballs are a type of fungus featuring a ball-shaped fruit body that bursts on contact or impact, releasing a cloud of dust-like spores into the surrounding area. Puffballs belong to the division Basidiomycota and encompass several genera, including Calvatia, Calbovista and Lycoperdon. The puffballs were previously treated as a taxonomic group called the Gasteromycetes or Gasteromycetidae, but they are now known to be a polyphyletic assemblage.
Geastrales is an order of gasterocarpic basidiomycetes (fungi) that are related to Cantharellales. The order contains the single family Geastraceae, which includes the "earthstars" formerly placed in Lycoperdales, or Phallales.
Geastrum is a genus of puffball-like mushrooms in the family Geastraceae. Many species are commonly known as earthstars.
Calostoma is a genus of 29 species of gasteroid fungi in the suborder Sclerodermatineae. Like other gasteroid fungi, Calostoma do not have the spore discharge mechanism associated with typical gilled fungi (ballistospory), and instead have enclosed spore-bearing structures. Resembling round puffballs with raised, brightly colored spore openings (ostioles), elevated on a thick, gelatinous stalks, species have been collected in regions of deciduous, temperate, tropical or subtropical forests. Their distribution includes eastern North America, Central America, Asia, and Australasia. The common name given to some species, "prettymouth", alludes to the brightly colored raised openings (ostioles) that may somewhat resemble lips. Other common names include "hotlips" and "puffball in aspic".
Geastrum fornicatum, common name known as the acrobatic earthstar or the arched earthstar, is an inedible species of mushroom in the family Geastraceae. Like other earthstar mushrooms, the thick outer skin splits open at maturity to expose the spore sac to the elements; the specific epithet fornicatum refers to the arched shape of the rays which extend downwards to rest on the mycelial sac and elevate the spore sac.
Clathrus ruber is a species of fungus in the family Phallaceae, and the type species of the genus Clathrus. It is commonly known as the latticed stinkhorn, the basket stinkhorn, or the red cage, alluding to the striking fruit bodies that are shaped somewhat like a round or oval hollow sphere with interlaced or latticed branches. The fungus is saprobic, feeding off decaying woody plant material, and is often found alone or in groups in leaf litter on garden soil, grassy places, or on woodchip garden mulches. Although considered primarily a European species, C. ruber has been introduced to other areas, and now has a wide distribution that includes all continents except Antarctica. The species was illustrated in the scientific literature during the 16th century, but was not officially described until 1729.
Myriostoma is a fungal genus in the family Geastraceae. Basidiocarps resemble earthstars, but the spore sac is supported by multiple columns and has multiple ostioles instead of a single, apical ostiole. Until 2017, the genus was thought to be monotypic with a single, widespread species, Myriostoma coliforme. Recent research has, however, shown that at least six species occur worldwide.
Geastrum pectinatum is an inedible species of mushroom belonging to the earthstar family of fungi. Although young specimens are spherical, fruit body development involves the outer layer of tissue splitting open like a star into 7 to 10 pointed rays that eventually bend back to point downward, revealing a small – 1 to 2.5 cm broad – spore sac. The spore sac is supported by a small radially wrinkled stalk. There is a distinct conical opening (peristome) at the top of the spore sac that is up to 8 mm (0.3 in) long. It is commonly known as the beaked earthstar or the beret earthstar, in reference to the shape of the spore sac and its prominent, protruding peristome. The mass of spores and surrounding cells within the sac, the gleba, is dark-brown, and becomes powdery in mature specimens. Spores are spherical, measuring 4 to 6 micrometers in diameter, with warts on their surfaces. Although uncommon, Geastrum pectinatum has a cosmopolitan distribution, and has been collected in various locations in Europe, North and South America, Asia, Australia and Africa, where it grows on the ground in open woods. Like several other earthstars, crystals of calcium oxalate are found on G. pectinatum, and are thought to be involved in fruit body maturation.
Astraeus is a genus of fungi in the family Diplocystaceae. The genus, which has a cosmopolitan distribution, contains nine species of earthstar mushroom. They are distinguished by the outer layer of flesh (exoperidium) that at maturity splits open in a star-shape manner to reveal a round spore sac. Additionally, they have a strongly hygroscopic character—the rays will open when moist, but when hot and dry will close to protect the spore sac. Species of Astraeus grow on the ground in ectomycorrhizal associations with trees and shrubs. Despite their similar appearance to the Geastrum earthstars Astraeus is not closely related.
Geasteroides is a fungal genus in the family Geastraceae. A monotypic genus, it contains the single species Geasteroides texensis, described by American mycologist William Henry Long in 1917.
Geastrum triplex is a fungus found in the detritus and leaf litter of hardwood forests around the world. It is commonly known as the collared earthstar, the saucered earthstar, or the triple earthstar—and less commonly by the alternative species name Geastrum indicum. It is the largest member of the genus Geastrum and expanded mature specimens can reach a tip-to-tip length of up to 12 centimeters (4.7 in).
The gasteroid fungi are a group of fungi in the Basidiomycota. Species were formerly placed in the obsolete class Gasteromycetes Fr., or the equally obsolete order Gasteromycetales Rea, because they produce spores inside their basidiocarps rather than on an outer surface. However, the class is polyphyletic, as such species—which include puffballs, earthstars, stinkhorns, and false truffles—are not closely related to each other. Because they are often studied as a group, it has been convenient to retain the informal (non-taxonomic) name of "gasteroid fungi".
Geastrum jurei is a species of fungus in the Geastrales, or earthstar fungi. It is known only from Algarrobo, in the Valparaíso province of Chile. It is a fornicate species of earthstar, meaning that the tips of the rays press down so as to raise the spherical spore sac into the air.
Geastrum leptospermum is a species of fungus in the family Geastraceae. It was first described scientifically by American mycologist George F. Atkinson in 1903. The fungus produces small fruit bodies and grows in mosses on tree trunks.
Geastrum quadrifidum, commonly known as the rayed earthstar or four-footed earthstar, is an inedible species of mushroom belonging to the genus Geastrum, or earthstar fungi. First described scientifically by Christian Hendrik Persoon in 1794, G. quadrifidum is a cosmopolitan—but not common—species of Europe, the Americas, Africa, Asia, and Australasia. The fungus is a saprobe, feeding off decomposing organic matter present in the soil and litter of coniferous forests.
Geastrum welwitschii is a species of fungus in the earthstar family. When young and unopened, the fruit bodies resemble small spheres lying in the soil. As the mushroom matures, the thick leathery outer layer of tissue splits star-like to form a number of fleshy arms, which curve downward to reveal the inner spore sac that contains the fertile tissue known as the gleba. The spore sac has a narrow grooved opening at the top where the spores are released. Fully expanded, the fruit bodies are up to 35 mm (1.4 in) wide and 58 mm (2.3 in) tall. First collected from Spain in the mid-19th century, the fungus is distributed in Europe, North America, and Bermuda.
Scleroderma polyrhizum, commonly known as the star earthball or dead man's hand, is a basidiomycete fungus and a member of the genus Scleroderma, or "earthballs". Found in dry, sandy soils, this species begins completely buried before slowly forcing the soil aside as it cracks apart to form a rough, star-shaped body with a diameter of 12–15 cm (4.7–5.9 in). At the center is the dark, brownish spore mass. Widely distributed wherever the soil and climate are favorable, it is known from Asia, Europe, and the Americas.
Calostoma cinnabarinum is a species of gasteroid fungus in the family Sclerodermataceae, and is the type species of the genus Calostoma. It is known by several common names, including stalked puffball-in-aspic and gelatinous stalked-puffball. The fruit body has a distinctive color and overall appearance, featuring a layer of yellowish jelly surrounding a bright red, spherical head approximately 2 centimeters (0.8 in) in diameter atop a red or yellowish brown spongy stipe 1.5 to 4 cm tall. The innermost layer of the head is the gleba, containing clear or slightly yellowish elliptical spores, measuring 14–20 micrometers (µm) long by 6–9 µm across. The spore surface features a pattern of small pits, producing a net-like appearance. A widely distributed species, it grows naturally in eastern North America, Central America, northeastern South America, and East Asia. C. cinnabarinum grows on the ground in deciduous forests, where it forms mycorrhizal associations with oaks.
Myriostoma capillisporum is a fungal species in the family Geastraceae. Basidiocarps resemble earthstars, but the spore sac is supported by multiple columns and has multiple ostioles instead of a single, apical ostiole. The fungus was originally described as a variety of Myriostoma coliforme, based on the distinctive and conspicuous ornamentation of its basidiospores. Recent molecular research, based on cladistic analysis of DNA sequences, has shown that it is a separate species, so far only known from South Africa.
Myriostoma calongei is a fungal species in the family Geastraceae. Basidiocarps resemble earthstars, but the spore sac is supported by multiple columns and has multiple ostioles instead of a single, apical ostiole. The fungus was described from Brazil in 2017 as a result of molecular research, based on cladistic analysis of DNA sequences. Previously, it had been identified as Myriostoma coliforme, now known to be restricted to northern, temperate regions. Myriostoma calongei can be distinguished by the conspicuously verrucose endoperidium. The species has also been confirmed from Argentina, Mexico, and the United States.