Myriostoma
Encyclopedia
Myriostoma is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Geastraceae
Geastraceae
The earthstars are the family Geastraceae of gasterocarpic basidiomycetes . It includes the genera Geastrum and Myriostoma. About sixty-four species are classified in this family, divided among eight genera....

. The genus is monotypic
Monotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...

, containing the single species Myriostoma coliforme. It is an earthstar
Geastrales
Geastrales is an order of gasterocarpic basidiomycetes that relates to Cantharellales. It includes the genera Geastrum and Myriostoma.The common name is "earthstars."...

, so named because outer wall of the spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

-bearing body splits open into the shape of a star. The inedible fungus has a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...

, and has been found in Africa, Asia, North and South America, and Europe, where it grows on humus-rich forest or woodland floor, especially on well-drained and sandy soils. This somewhat rare fungus was included in 2004 in a list of 33 species proposed for protection under the Bern Convention
Convention on the Conservation of European Wildlife and Natural Habitats
The Bern Convention on the Conservation of European Wildlife and Natural Habitats 1979, also known as the Bern Convention , came into force on June 1, 1982....

 by the European Council for Conservation of Fungi, and included on the Red Lists
Regional Red List
A Regional Red List is a report of the threatened status of species within a certain country or region. It is based on the IUCN Red List of Threatened Species, an inventory of the conservation status of species on a global scale...

 of 12 European countries.

The fruit body
Sporocarp (fungi)
In fungi, the sporocarp is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne...

, initially shaped like a puffball
Puffball
A puffball is a member of any of several groups of fungus in the division Basidiomycota. The puffballs were previously treated as a taxonomic group called the Gasteromycetes or Gasteromycetidae, but they are now known to be a polyphyletic assemblage. The distinguishing feature of all puffballs is...

, is encased within an outer covering that splits open from the top to form rays. These rays curve down to expose an inner papery spore case, the endoperidium, which contains the fertile gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

. The fungus is unique among the earthstars in having a spore case that is supported by multiple stalks, as is perforated by several small holes suggestive of its common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

s salt-shaker earthstar and pepperpot. It is the largest of the earthstar fungi, and reaches diameters of up to 12 cm (4.7 in). Its spores have elongated warts that creates a ridge-like pattern on the surface described as being reticulate.

Taxonomy, phylogeny, and naming

The species was first mentioned in the scientific literature by Samuel Doody
Samuel Doody
-Life:The eldest of the second family of his father, John Doody, an apothecary in Staffordshire who later moved to London where he had a shop in The Strand, he was born in Staffordshire 28 May 1656...

 in the second edition of John Ray
John Ray
John Ray was an English naturalist, sometimes referred to as the father of English natural history. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after "having ascertained that such had been the practice of his family before him".He published important works on botany,...

's Synopsis methodica Stirpium Brittanicorum in 1696. Ray described the mushroom like so: "fungus pulverulentus coli instar perforatus, cum volva stellata", and went on to explain that he has found it in 1695 in Kent
Kent
Kent is a county in southeast England, and is one of the home counties. It borders East Sussex, Surrey and Greater London and has a defined boundary with Essex in the middle of the Thames Estuary. The ceremonial county boundaries of Kent include the shire county of Kent and the unitary borough of...

. It was first described scientifically
Species description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...

 as a new species from England in 1776 by James Dickson
James Dickson
Sir James Robert Dickson, KCMG was an Australian politician and businessman, the 13th Premier of Queensland and a member of the first federal ministry....

, who named it Lycoperdon coliforme. He found it growing in roadside banks and hedgerows among nettles in Suffolk
Suffolk
Suffolk is a non-metropolitan county of historic origin in East Anglia, England. It has borders with Norfolk to the north, Cambridgeshire to the west and Essex to the south. The North Sea lies to the east...

 and Norfolk
Norfolk
Norfolk is a low-lying county in the East of England. It has borders with Lincolnshire to the west, Cambridgeshire to the west and southwest and Suffolk to the south. Its northern and eastern boundaries are the North Sea coast and to the north-west the county is bordered by The Wash. The county...

. Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 called it Geastrum coliforme in 1801, and Gray called it Polystoma coliforme in 1821. In North America, the fungus was reported from Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...

 by Charles Horton Peck
Charles Horton Peck
Charles Horton Peck, born March 30, 1833 in Sand Lake, New York, died 1917 in Albany, New York, was an American mycologist of the 19th and early 20th centuries...

, collected in Florida by Lucien Underwood in 1891, and had notes published about it by Andrew Price Morgan
Andrew Price Morgan
Andrew Price Morgan was an American botanist. He investigated the flora of the Miami Valley in Ohio. While his interest included flowering plants, as noted by his Flora of the Miami Valley, Ohio, his special interest was in fungi. Morgan worked as a teacher in Dayton...

 in American Naturalist, April 1892. Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...

 named it Bovistoides simplex in 1919, but in 1942, William Henry Long examined the specimen and concluded that it was a weathered spore sac of M. coliforme that had become detached from the outer star-shaped exoperidium. This conclusion was confirmed in a later study of the type material. The genus Myriostoma was named and described by Nicaise Auguste Desvaux
Nicaise Auguste Desvaux
-Works:*Journal de Botanique, appliquée à l'Agriculture, à la Pharmacie, à la Médecine et aux Arts .*Observations sur les plantes des environs d'Angers ....

 published in the Journal de Botanique, Rédigé par une Société di Botanistes in 1809.

Myriostoma had been classified in the Geastraceae
Geastraceae
The earthstars are the family Geastraceae of gasterocarpic basidiomycetes . It includes the genera Geastrum and Myriostoma. About sixty-four species are classified in this family, divided among eight genera....

 family until British mycologist Donald Dring (1973) placed it in the Astraceae (Sclerodermatales) based on the presence of the trabeculae (stout columns which extend from the peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...

 to the central core of the fruit body) in the gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

, and the absence of a true hymenium
Hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some cells develop into sterile cells called cystidia or...

. In his 1989 monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...

, Sunhede placed in again in the Geastraceae. Phylogenetic analysis has shown that Myriostoma and Geastrum
Geastrum
Geastrum is a genus of mushroom in the family Geastraceae. Many species are known commonly as earthstars.The name comes from geo meaning earth and aster meaning star....

form a sister group to the Phallales
Phallales
The Phallales are an order of fungi that is more or less synonymous with the gomphoid-phalloid clade. The order contains two families, the Claustulaceae and the Phallaceae, which collectively contain 26 genera and 88 species.-External links:...

, but are not closely related to the Lycoperdales
Lycoperdales
The Lycoperdales are a now outdated order of fungi. The order included some well-known types such as the giant puffball, the earthstars, and other tuberous fungi...

.

Other taxa formerly classified as Myriostoma include Myriostoma anglicum Desv. (1809) (the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

, and the variety Myriostoma coliforme var. coliforme (Dicks.) Corda (1842) and Myriostoma coliforme var. capillisporum V.J. Staněk (1958). These are all considered synonymous
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

 with Myriostoma coliforme, given its current name by August Carl Joseph Corda
August Carl Joseph Corda
August Carl Joseph Corda was a Czech physician and mycologist.-Early life and education:Corda was born in Reichenberg , Bohemia on November 15, 1809. Corda's father was a textile seller...

 in 1842. M. coliforme is the sole species in Myriostoma, making the genus monotypic
Monotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...

. Because the original type material has been lost, in 1989 Sunhede suggested that Dickson's illustration in his 1776 publication (tab. III: 4a & b) be selected as the lectotype
Lectotype
In botanical nomenclature and zoological nomenclature, a lectotype is a kind of name-bearing type. When a species was originally described on the basis of a name-bearing type consisting of multiple specimens, one of those may be designated as the lectotype...

.

The specific epithet is derived from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 words colum, "strainer", and forma, "shape". Berkeley's vernacular
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 name "Cullenden puff-ball" refers to a colander; Gray
Samuel Frederick Gray
Samuel Frederick Gray was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray.-Background:...

 called it the "sievelike pill-box". The generic name is from the Greek words μνριος "countless" and στομα "mouth". It is commonly known as the "salt-and-pepper shaker earthstar" or the "pepperpot".

Description

The fruit bodies start their development underground or buried in leaf debris, and have a strand of mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 at the base. As they mature, the exoperidium (the outer tissue layer of the peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...

) curves backwards and splits open into rays. This movement pushes the fruit body above the surface of the substrate
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...

. There are 8–14 rays that are unequal in size, with tips that are often rolled inward. Fully opened individuals can reach dimensions of 2 – across from ray tip to tip. The rays are made of three distinct layers of tissue. The thick pseudoparenchymatous layer is fleshy and thick when fresh, pale beige but becoming yellow to brown as it matures, and often cracking and peeling off in the process. The exterior mycelial
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 tissue layer, often matted with fine leaf debris or dirt, usually cracks to reveal the middle fibrous layer, which is made of densely packed hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

e that are 1–2.5 μm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...

 wide. The base is concave to vaulted, often showing a large umbilical scar where the mycelial strand was attached. The spore sac, or endoperidium, have very short, inconspicuous pedicels, shaped like flattened spheres, 1 – in diameter. It is gray-brown in color, and minutely roughened with small subreticulate warts. There are usually one to six evenly dispersed mouths (ostioles, or stomata), mainly on upper half of the spore sac. They are roughly circular and have fringed (fimbriate) margins. The fruit bodies are inedible.

Like many earthstars, the fungus uses the force of falling raindrops to help disperse the spores; the spores are ejected in little bursts as falling raindrops strike the outer wall of the spore sac. The gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

 is brown grayish-brown. It has a "cottony" or "hairy" texture that when compressed, allows the endoperidium to flex quickly, creating a puff of air that is forced out through the ostiole. This generates a cloud of spores that are lifted in the air, facilitating dispersal. There are several to many columellae (sterile tissue usually in the base of the gleba, and extending up into the gleba), 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 openings at the top of the spore case, but rather, terminate at some distance from them in the gleba. The capillitium (coarse thick-walled cells found in the gleba) are long, slender, free, tapering, unbranched, and 2–5 μm thick, with thick walls. The spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s are spherical, nonamyloid
Amyloid (mycology)
In mycology the term amyloid refers to a crude chemical test using iodine in either Melzer's reagent or Lugol's solution, to produce a black to blue-black positive reaction. It is called amyloid because starch gives a similar reaction, and that reaction for starch is also called an amyloid reaction...

, and are ornamented with warts up to 2 μm high. They measure 4–5 μm diameter (without ornamentation), and 5.4–7.0 μm including the warts.

Similar species

Myriostoma coliforme is the largest earthstar fungus, and the only one that produces numerous openings on the top of its spore sac, and numerous stalks that support the spore sac. Historically, it was thought that the openings might have been a result of insect action. This was addressed in an early report of the fungus by Woodward (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 ciliae) 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.

Ecology, habitat and distribution

The fungus grows in groups in well-drained or sandy soil, often in the partial shade of trees. It is saprobic, deriving nutrients by decomposing organic matter. It can be found in both deciduous and mixed forests, gardens, along hedges and grassy road banks as well as grazed grasslands. In the Northern Hemisphere, it tends to grow on well-drained south-facing slopes, while it prefers similar habitat on south facing slopes in Australia. In Europe, its major habitat is Riparian
Riparian zone
A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the fifteen terrestrial biomes of the earth. Plant habitats and communities along the river margins and banks are called riparian vegetation, characterized by...

 mixed forests along the great rivers, dominated by Salix alba and Populis alba galleris. In Hawaii, it has been collected at elevations above 2000 m (6,561.7 ft) where it appears to favor the mamame (Sophora chrysophylla) forest. The fungus is widespread in Europe, having been found in Czechoslovakia, Germany, Italy, The Netherlands, Poland, Portugal, and Sweden—its most northerly location; it is generally rare in northern Europe. It has also been found in the Canary Islands
Canary Islands
The Canary Islands , also known as the Canaries , is a Spanish archipelago located just off the northwest coast of mainland Africa, 100 km west of the border between Morocco and the Western Sahara. The Canaries are a Spanish autonomous community and an outermost region of the European Union...

, Morocco (in the Mamora forest), and in South Africa. In South America, the fungus has been collected in the Galapagos Islands
Galápagos Islands
The Galápagos Islands are an archipelago of volcanic islands distributed around the equator in the Pacific Ocean, west of continental Ecuador, of which they are a part.The Galápagos Islands and its surrounding waters form an Ecuadorian province, a national park, and a...

, Argentina, Brazil, and Chile. The North American distribution ranges from Canada to Mexico (Ajos-Bavispe National Forest Reserve). In the Middle East
Middle East
The Middle East is a region that encompasses Western Asia and Northern Africa. It is often used as a synonym for Near East, in opposition to Far East...

, it has also been found in Afghanistan and Iran. It is also in Australia (where it may be an introduced species
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

) China, and India.

Myriostoma coliforme is listed as one of the 33 threatened
Threatened species
Threatened species are any speciesg animals, plants, fungi, etc.) which are vulnerable to endangerment in the near future.The World Conservation Union is the foremost authority on threatened species, and treats threatened species not as a single category, but as a group of three categories,...

 fungi in Europe, and is listed on 12 Regional Red List
Regional Red List
A Regional Red List is a report of the threatened status of species within a certain country or region. It is based on the IUCN Red List of Threatened Species, an inventory of the conservation status of species on a global scale...

s. In the United Kingdom, it was considered extinct in mainland Britain due to lack of collections since 1880, but it was found in Suffolk
Suffolk
Suffolk is a non-metropolitan county of historic origin in East Anglia, England. It has borders with Norfolk to the north, Cambridgeshire to the west and Essex to the south. The North Sea lies to the east...

 in 2006 near Ipswich
Ipswich
Ipswich is a large town and a non-metropolitan district. It is the county town of Suffolk, England. Ipswich is located on the estuary of the River Orwell...

, one of its original localities. It is also found in the Channel Islands
Channel Islands
The Channel Islands are an archipelago of British Crown Dependencies in the English Channel, off the French coast of Normandy. They include two separate bailiwicks: the Bailiwick of Guernsey and the Bailiwick of Jersey...

.

External links

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK