Lycoperdon echinatum
Encyclopedia
Lycoperdon echinatum, commonly known as the spiny puffball or the spring puffball, is a type of 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...

 mushroom in the genus Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...

. The saprobic species has been found in Africa, Europe, Central America and North America, where it grows on soil in deciduous woods
Temperate deciduous forest
A temperate deciduous forest, more precisely termed temperate broadleaf forest or temperate broadleaved forest, is a biome found in North America, southern South America, Europe, and Asia. A temperate deciduous forest consists of trees that lose their leaves every year...

, glades, and pastures. It has been proposed that North American specimens be considered a separate species, Lycoperdon americanum, but this suggestion has not been followed by most authors. Molecular analysis indicates that Lycoperdon echinatum is closely related to the puffball genus Handkea
Handkea
Handkea is a genus of puffball mushrooms in the Lycoperdaceae family. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia with distinct microscopic features, including a unique type of capillitium , with curvy slits instead of the usual pores...

.

The fruit bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

 of L. echinatum are 2 – wide by 2 – tall, supported by a small base, and densely covered with spines that are up to 0.6 cm (0.236220472440945 in) long. The spines can fall off in maturity, leaving a net-like pattern of scars on the underlying surface. Initially white in color, the puffballs turn a dark brown as they mature, at the same time changing from nearly round to somewhat flattened. Young specimens of L. echinatum resemble another edible spiny puffball, Lycoperdon pulcherrimum
Lycoperdon pulcherrimum
Lycoperdon pulcherrimum is a type of puffball mushroom in the genus Lycoperdon. It was first described scientifically in 1873 by Miles Joseph Berkeley and Moses Ashley Curtis....

, but this latter species does not turn brown as it ages. The fruit bodies are edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

 when young, when the interior is white and firm and before it has turned into a powdery brown mass of 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. Laboratory tests have shown that extracts of the fruit bodies can inhibit the growth of several bacteria that are pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

ic to humans.

Taxonomy and phylogeny

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

 in 1797. It was later reduced to a variety of Lycoperdon gemmatum (as L. gemmatum var. echinatum; L. gemmatum is now known as Lycoperdon perlatum
Lycoperdon perlatum
Lycoperdon perlatum, commonly known as the common puffball, warted puffball, gem-studded puffball or devil's snuff-box, is a species of puffball mushroom in the Agaricaceae family. A common species with a cosmopolitan distribution, it is a moderate-sized puffball with a round fruit body tapering to...

) by Elias Magnus Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

, but American mycologist 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...

, who extensively studied the North American distribution of the genus, raised it again to species level in 1879. He thought it worthy of status as a species distinct from L. gemmatum because of the different character of its warts, its much spinier appearance, and the smoother surface 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:...

 underneath the spines. Miles Joseph Berkeley
Miles Joseph Berkeley
Miles Joseph Berkeley was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology....

 and Christopher Edmund Broome
Christopher Edmund Broome
Christopher Edmund Broome was a British mycologist.-Background and education:C.E. Broome was born in Berkhamsted, the son of a solicitor. He was privately schooled in Kensington and in 1832 was sent to read for Holy Orders with the curate of Swaffham Prior in Cambridgeshire...

 wrote of the fungus in 1871, but believed their specimen, collected from Reading, Berkshire
Reading, Berkshire
Reading is a large town and unitary authority area in England. It is located in the Thames Valley at the confluence of the River Thames and River Kennet, and on both the Great Western Main Line railway and the M4 motorway, some west of London....

, by Hoyle, represented a new species, which they called Lycoperdon Hoylei. They wrote that their specimen agreed "exactly with an authentic specimen of Persoon's L. echinatum externally, who could, however, scarcely have overlooked the lilac spores." Despite the apparent difference in spore color, L. Hoylei is currently 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 L. echinatum. Utraria echinata, named by Lucien Quélet
Lucien Quélet
thumb|Lucien QuéletLucien Quélet was a French mycologist and naturalist who discovered several species and was the founder of the Société mycologique de France, a society devoted to mycological studies....

 in 1873, is another synonym for L. echinatum.

In 1972, Vincent Demoulin described the species Lycoperdon americanum on the basis of a specimen found in North Carolina
North Carolina
North Carolina is a state located in the southeastern United States. The state borders South Carolina and Georgia to the south, Tennessee to the west and Virginia to the north. North Carolina contains 100 counties. Its capital is Raleigh, and its largest city is Charlotte...

. Although he believed it to be a unique species, several authors consider it synonymous with L. echinatum. Phylogenetic analysis of the sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 and secondary structure
Secondary structure
In biochemistry and structural biology, secondary structure is the general three-dimensional form of local segments of biopolymers such as proteins and nucleic acids...

 of the ribosomal RNA
Ribosomal RNA
Ribosomal ribonucleic acid is the RNA component of the ribosome, the enzyme that is the site of protein synthesis in all living cells. Ribosomal RNA provides a mechanism for decoding mRNA into amino acids and interacts with tRNAs during translation by providing peptidyl transferase activity...

 (rRNA) genes coding for the internal transcribed spacer
Internal transcribed spacer
ITS refers to a piece of non-functional RNA situated between structural ribosomal RNAs on a common precursor transcript. Read from 5' to 3', this polycistronic rRNA precursor transcript contains the 5' external transcribed sequence , 18S rRNA, ITS1, 5.8S rRNA, ITS2, 28S rRNA and finally the 3'ETS...

 units suggests that Lycoperdon echinatum forms a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 with the puffball genus Handkea
Handkea
Handkea is a genus of puffball mushrooms in the Lycoperdaceae family. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia with distinct microscopic features, including a unique type of capillitium , with curvy slits instead of the usual pores...

, separate from 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...

 of Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...

, Lycoperdon perlatum. In previous analyses that used only the rRNA sequences for phylogenetic comparison, L. echinatum formed a clade with L. mammiforme
Lycoperdon mammiforme
Lycoperdon mammiforme is a rare, inedible type of puffball mushroom in the genus Lycoperdon, found in beech woodlands on chalk.The fruiting body is spherical to pear shaped, at first pure white with slightly grainy inner skin and an outer skin which disintegrates in flakes that are soon shed,...

, L. foetidum, and Bovistella radicata (now known as Lycoperdon radicatum
Lycoperdon radicatum
Lycoperdon radicatum is a type of puffball mushroom in the genus Lycoperdon....

), but separate from L. pyriforme.

The species is commonly
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...

 known as the "spiny puffball" or the "spring puffball"; Peck referred to the species as the "echinate puff-ball". The specific epithet echinatum is derived from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 word echinos (εχινος) meaning "hedgehog" or "sea-urchin".

Description

The fruit bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

 of L. echinatum are 2 – wide by 2 – tall, and are roughly spherical, or pear-shaped. The exterior surface is crowded with spines that may be up to 0.6 cm (0.236220472440945 in) long. According to 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...

, American specimens have more slender spines than European ones. Initially white and becoming dark brown in maturity, the spines are often joined at the tips in groups of three or four. In this form the puffballs resemble acorn caps of burr oak
Burr Oak
Burr Oak is a variant spelling of Bur oak . The term can refer to some places in the United States:*Burr Oak, Indiana*Burr Oak, Iowa*Burr Oak, Kansas*Burr Oak, Michigan*Burr Oak Township, Michigan*Burr Oak, Missouri...

, with which they may readily be confused. The spines slough off in age, revealing a somewhat net-like or reticulated surface. The fruit body has a small base that is an off-white or purple-gray color, and it may be attached to the growing surface
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...

 by thin white cords (rhizomorphs). The internal contents of the puffball contain 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...

, a mass of 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 and associated spore-producing cells. In young specimens the gleba is white and firm, but as the puffball ages, it turns yellowish and then brown to purple-brown and powdery. Mature specimens develop a pore at the top of the fruit body through which spores may be released.

The spores of L. echinatum, roughly spherical with warts on the surface, have diameters between 4 and 6 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...

. The capillitia (coarse thick-walled 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 in the gleba) are elastic, brown in color, contain small pores, and are 5–8 µm thick. The basidia (the spore-bearing cells) may be attached to two to four spores, and the sterigmata (tapered spine-like projections from the basidia that attach the spores) are up to 5 µm long.

Like most other puffball species, L. echinatum is edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

 when still young and while the gleba is still white and firm. Consumption of older specimens with a non-white gleba, or where the gleba has turned into a powdery spore mass, may cause stomach upsets. This species has a mild taste, and no distinguishable odor, although one source describes the smell of dried fruit bodies as similar to "old ham". One source notes that it is "well flavoured and tender when cooked", while another describes the texture (of edible puffballs in general) as "somewhat like French toast". Antonio Carluccio
Antonio Carluccio
Antonio Carluccio, OBE OMRI is an Italian chef, restaurateur and food expert, based in London.-Biography:Antonio Carluccio was born in Vietri sul Mare, Salerno, Italy. His father was a stationmaster, and he moved with his father's job when he was young and grew up in Piedmont...

 recommends sautéeing puffballs with other mushrooms. It is recommended to slice young puffballs intended for consumption with a longitudinal cut to ensure that the flesh is devoid of any internal structures, to avoid the possibility of confusion with potentially deadly Amanita
Amanita
The genus Amanita contains about 600 species of agarics including some of the most toxic known mushrooms found worldwide. This genus is responsible for approximately 95% of the fatalities resulting from mushroom poisoning, with the death cap accounting for about 50% on its own...

species.

Similar species

Lycoperdon pulcherrimum closely resembles L. echinatum, but its spines are stouter, do not turn brown in age, and the surface of the fruit body underneath the spines is smooth, not pitted. Alexander H. Smith
Alexander H. Smith
Alexander Hanchett Smith was an American mycologist known for his extensive contributions to the taxonomy and phylogeny of the higher fungi, especially the agarics.-Early life:...

 noted that in youth, they are "difficult if not impossible to distinguish from each other, but this will cause no inconvenience to those collecting for the table, since both are edible." In some areas the two species appear to intergrade, as specimens may be found whose spines turn brown but do not fall off. Young specimens of L. pedicellatum may also be difficult to distinguish from L. echinatum, but the former has a smooth outer surface when mature, and has spores attached to a pedicel (a narrow extension of the basidium on which the sterigmata and spores are formed) that is about 4–5 times as long as the spore. Lycoperdon compactum, found only in New Zealand, also resembles L. echinatum in appearance, but differs in having smaller spores, capillitia that are hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...

 (translucent) and septate (with partitions that divide the capillitia into compartments).

Habitat, distribution, and ecology

Lycoperdon echinatum can be found either solitary or in small groups. It typically grows on the ground in deciduous forests
Temperate deciduous forest
A temperate deciduous forest, more precisely termed temperate broadleaf forest or temperate broadleaved forest, is a biome found in North America, southern South America, Europe, and Asia. A temperate deciduous forest consists of trees that lose their leaves every year...

 and grassy areas, glades and pastures, on moss
Moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...

, humus, or woody debris. The fungus has been noted to have a preference for beech
Beech
Beech is a genus of ten species of deciduous trees in the family Fagaceae, native to temperate Europe, Asia and North America.-Habit:...

 woods. Fruit bodies may make their appearance anytime from the late spring to autumn. Older specimens are more likely to be overlooked, as their brown color blends into the surrounding environment of dead leaves and dead wood. The mushroom is used by various species of scuttle flies (family Phoridae) as larval
Larvae
In Roman mythology, lemures were shades or spirits of the restless or malignant dead, and are probably cognate with an extended sense of larvae as disturbing or frightening...

 food.

This species has been collected from eastern central Africa, China, Costa Rica, Iran, Japan, and Europe (including Britain, Bulgaria, the Czech Republic, Finland, Germany, Italy, Slovakia, Spain, Sweden, and Switzerland). In North America, it is "locally frequent" east of the Rocky Mountains
Rocky Mountains
The Rocky Mountains are a major mountain range in western North America. The Rocky Mountains stretch more than from the northernmost part of British Columbia, in western Canada, to New Mexico, in the southwestern United States...

.

It is considered a threatened species
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,...

 in the Aland Islands
Åland Islands
The Åland Islands form an archipelago in the Baltic Sea. They are situated at the entrance to the Gulf of Bothnia and form an autonomous, demilitarised, monolingually Swedish-speaking region of Finland...

 of Finland. A study of the species' distribution in Sweden reported that in the 1940s and 50s, it grew in beech
Beech
Beech is a genus of ten species of deciduous trees in the family Fagaceae, native to temperate Europe, Asia and North America.-Habit:...

 woods with broad-leaved grasses and herbs in top soils with soil pH
PH
In chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...

 levels between 5.0 and 6.6, but the populations have since decreased owing to the acidification during the last several decades. Fruit bodies collected near arsenic
Arsenic
Arsenic is a chemical element with the symbol As, atomic number 33 and relative atomic mass 74.92. Arsenic occurs in many minerals, usually in conjunction with sulfur and metals, and also as a pure elemental crystal. It was first documented by Albertus Magnus in 1250.Arsenic is a metalloid...

-contaminated sites have been shown to bioaccumulate
Bioaccumulation
Bioaccumulation refers to the accumulation of substances, such as pesticides, or other organic chemicals in an organism. Bioaccumulation occurs when an organism absorbs a toxic substance at a rate greater than that at which the substance is lost...

 arsenic, largely in the form of arsenobetaine
Arsenobetaine
Arsenobetaine is an organoarsenic compound that is the main source of arsenic found in fish. It is the arsenic analog of trimethylglycine, commonly known as betaine...

.

Antimicrobial activity

Using a standard laboratory method
Kirby-Bauer antibiotic testing
Kirby-Bauer antibiotic testing is a test which uses antibiotic-impregnated wafers to test whether particular bacteria are susceptible to specific antibiotics. A known quantity of bacteria are grown on agar plates in the presence of thin wafers containing relevant antibiotics...

 to determine antimicrobial
Antimicrobial
An anti-microbial is a substance that kills or inhibits the growth of microorganisms such as bacteria, fungi, or protozoans. Antimicrobial drugs either kill microbes or prevent the growth of microbes...

 susceptibility, methanol
Methanol
Methanol, also known as methyl alcohol, wood alcohol, wood naphtha or wood spirits, is a chemical with the formula CH3OH . It is the simplest alcohol, and is a light, volatile, colorless, flammable liquid with a distinctive odor very similar to, but slightly sweeter than, ethanol...

-based extract
Extract
An extract is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol or water. Extracts may be sold as tinctures or in powder form....

s of Lycoperdon umbrinum fruit bodies were shown in a 2005 study to have "significant" antibacterial activity against various human pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

ic bacteria, including Bacillus subtilis
Bacillus subtilis
Bacillus subtilis, known also as the hay bacillus or grass bacillus, is a Gram-positive, catalase-positive bacterium commonly found in soil. A member of the genus Bacillus, B. subtilis is rod-shaped, and has the ability to form a tough, protective endospore, allowing the organism to tolerate...

, Escherichia coli
Escherichia coli
Escherichia coli is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms . Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls...

, Salmonella typhimurium, Staphylococcus aureus
Staphylococcus aureus
Staphylococcus aureus is a facultative anaerobic Gram-positive coccal bacterium. It is frequently found as part of the normal skin flora on the skin and nasal passages. It is estimated that 20% of the human population are long-term carriers of S. aureus. S. aureus is the most common species of...

, Streptococcus pyogenes
Streptococcus pyogenes
Streptococcus pyogenes is a spherical, Gram-positive bacterium that is the cause of group A streptococcal infections. S. pyogenes displays streptococcal group A antigen on its cell wall. S...

, and Mycobacterium smegmatis
Mycobacterium smegmatis
Mycobacterium smegmatis is 3.0 to 5.0 µm long with a bacillus shape, an acid-fast bacterial species in the phylum Actinobacteria. It can be stained by Ziehl-Neelsen method and the auramine-rhodamine fluorescent method. It was first reported in November 1884 by Lustgarten, who found a bacillus...

. An earlier study (2000) had identified weak antibacterial activity against Enterococcus faecium
Enterococcus faecium
Enterococcus faecium is a Gram-positive, alpha hemolytic or nonhemolytic bacterium in the genus Enterococcus. It can be commensal in the human intestine, but it may also be pathogenic, causing diseases such as neonatal meningitis.Vancomycin-resistant E. faecium is often referred to as VRE.Some...

and Staphylococcus aureus. Although the specific compounds responsible for the antimicrobial activity have not been identified, chemical analysis confirms the presence of terpenoid
Terpenoid
The terpenoids , sometimes called isoprenoids, are a large and diverse class of naturally occurring organic chemicals similar to terpenes, derived from five-carbon isoprene units assembled and modified in thousands of ways. Most are multicyclic structures that differ from one another not only in...

s, a class of widely occurring organic chemicals that are being investigated for their potential use as antimicrobial drugs.

External links

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