Lycoperdon echinatum | |
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Mature specimen with brown spines | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Class: | Agaricomycetes |
Order: | Agaricales |
Family: | Agaricaceae |
Genus: | Lycoperdon |
Species: | L. echinatum |
Binomial name | |
Lycoperdon echinatum Pers. (1797) | |
Synonyms | |
Lycoperdon echinatum | |
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Glebal hymenium | |
Lacks a stipe | |
Spore print is brown | |
Ecology is saprotrophic | |
Edibility is edible or inedible |
Lycoperdon echinatum, commonly known as the spiny puffball or the spring puffball, is a type of puffball mushroom in the family Agaricaceae. The saprobic species has been found in Africa, Europe, Central America, and North America, where it grows on soil in deciduous woods, 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 L. echinatum is closely related to the puffball genus Handkea .
The fruit bodies of L. echinatum are 2–4 cm (0.8–1.6 in) wide by 2–3.5 cm (0.8–1.4 in) tall, supported by a small base, and densely covered with spines that are up to 0.6 cm (0.2 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 , but the latter species does not turn brown as it ages. The fruit bodies are edible when young, when the interior is white and firm and before it has turned into a powdery brown mass of spores. Laboratory tests have shown that extracts of the fruit bodies can inhibit the growth of several bacteria that are pathogenic to humans.
The species was first described by Christian Hendrik Persoon in 1797. [1] It was later reduced to a variety of Lycoperdon gemmatum (as L. gemmatum var. echinatum; L. gemmatum is now known as Lycoperdon perlatum [2] ) by Elias Magnus Fries, [3] but American mycologist Charles Horton Peck, 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 underneath the spines. [4] Miles Joseph Berkeley and Christopher Edmund Broome wrote of the fungus in 1871, but believed their specimen, collected from Reading, Berkshire, 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." [5] Despite the apparent difference in spore color, L. Hoylei is currently considered synonymous with L. echinatum. [6] Utraria echinata, named by Lucien Quélet in 1873, [7] is another synonym for L. echinatum. [8]
In 1972, Vincent Demoulin described the species Lycoperdon americanum on the basis of a specimen found in North Carolina. [9] Although he believed it to be a unique species, [10] several authors consider it synonymous with L. echinatum. [11] [12] [13] [14] Phylogenetic analysis of the sequence and secondary structure of the ribosomal RNA (rRNA) genes coding for the internal transcribed spacer units suggests that Lycoperdon echinatum forms a clade with the puffball genus Handkea , separate from the type species of Lycoperdon , Lycoperdon perlatum. [15] In previous analyses that used only the rRNA sequences for phylogenetic comparison, L. echinatum formed a clade with L. mammiforme , L. foetidum , and Bovistella radicata (now known as Lycoperdon radicatum [16] ), but separate from L. pyriforme . [17] [18]
The species is commonly known as the "spiny puffball" [13] or the "spring puffball"; [19] Peck referred to the species as the "echinate puff-ball". [4] The specific epithet echinatum is derived from the Greek word echinos (εχινος) meaning "hedgehog" or "sea-urchin". [20]
The fruit bodies of L. echinatum are 2–4 cm (0.8–1.6 in) wide by 2–3.5 cm (0.8–1.4 in) tall, and are roughly spherical, or pear-shaped. [21] The exterior surface is crowded with spines that may be up to 0.6 cm (0.2 in) long. According to Curtis Gates Lloyd, American specimens have more slender spines than European ones. [22] Initially white and becoming dark brown in maturity, the spines are often joined at the tips in groups of three or four. [23] In this form the puffballs resemble acorn caps of burr oak, with which they may readily be confused. [11] 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, [21] and it may be attached to the growing surface by thin white cords (rhizomorphs). [24] The internal contents of the puffball contain the gleba, a mass of spores 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. [13] Mature specimens develop a pore at the top of the fruit body through which spores are released when hit by falling raindrops. [24]
The spores of L. echinatum, roughly spherical with warts on the surface, have diameters between 4 and 6 μm. [13] The capillitia (coarse thick-walled hyphae in the gleba) are elastic, brown in color, contain small pores, and are 5–8 μm thick. [23] 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. [25]
Like most other puffball species, L. echinatum is edible when still young and while the gleba is still white and firm. [13] Consumption of older specimens with a non-white gleba, or where the gleba has turned into a powdery spore mass, may cause stomach upsets. [26] This species has a mild taste, and no distinguishable odor, [21] although one source describes the smell of dried fruit bodies as similar to "old ham". [27] One source notes that it is "well flavoured and tender when cooked", [19] while another describes the texture (of edible puffballs in general) as "somewhat like French toast". [26] Antonio Carluccio recommends sautéeing puffballs with other mushrooms. [28] To avoid possible confusion with potentially deadly Amanita species, it is recommended to slice young puffballs with a longitudinal cut to ensure that the flesh is devoid of any internal structures. [29]
Lycoperdon pulcherrimum closely resembles L. echinatum, but its spines are stouter, [21] do not turn brown in age, and the surface of the fruit body underneath the spines is smooth, not pitted. [13] Alexander H. Smith 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." [30] In some areas the two species appear to intergrade, as specimens may be found whose spines turn brown but do not fall off. [11] 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. [31] Lycoperdon compactum , found only in New Zealand, also resembles L. echinatum in appearance, but differs in having smaller spores, capillitia that are hyaline (translucent) and septate (with partitions that divide the capillitia into compartments). [32]
Lycoperdon echinatum can be found either solitary or in small groups. It typically grows on the ground in deciduous forests and grassy areas, glades and pastures, on moss, humus, or woody debris. The fungus has been noted to have a preference for beech woods. [19] [23] Fruit bodies may make their appearance anytime from the late spring to autumn. [13] [21] Older specimens are more likely to be overlooked, as their brown color blends into the surrounding environment of dead leaves and dead wood. [30] The puffball is used by various species of scuttle flies (family Phoridae) as larval food. [33]
This species has been collected from eastern central Africa, [34] China, [35] Costa Rica, [36] Iran, [37] Japan, [38] and Europe (including Britain, [39] Bulgaria, [40] the Czech Republic, [41] Finland, [42] Germany, [43] Italy, [44] Slovakia, [45] Spain, [46] Sweden, [47] [48] and Switzerland [49] ). In North America, it is "locally frequent" east of the Rocky Mountains. [19]
It is considered a threatened species in Åland (Finland). [42] A study of the species' distribution in Sweden reported that in the 1940s and 50s, it grew in beech woods with broad-leaved grasses and herbs in topsoils with soil pH levels between 5.0 and 6.6, but the populations have since decreased owing to soil acidification during the last several decades. [48] Fruit bodies collected near arsenic-contaminated sites have been shown to bioaccumulate arsenic, largely in the form of arsenobetaine. [50]
Using a standard laboratory method to determine antimicrobial susceptibility, methanol-based extracts of Lycoperdon umbrinum fruit bodies were shown in a 2005 study to have "significant" antibacterial activity against various human pathogenic bacteria, including Bacillus subtilis , Escherichia coli , Salmonella typhimurium , Staphylococcus aureus , Streptococcus pyogenes , and Mycobacterium smegmatis . [51] An earlier study (2000) had identified weak antibacterial activity against Enterococcus faecium and Staphylococcus aureus. [52] Although the specific compounds responsible for the antimicrobial activity have not been identified, chemical analysis confirms the presence of terpenoids, [51] a class of widely occurring organic chemicals that are being investigated for their potential use as antimicrobial drugs. [53]
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.
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. It was formerly classified within the now-obsolete order Lycoperdales, as the type genus which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split. Lycoperdon is now placed in the family Agaricaceae of the order Agaricales.
Lycoperdon perlatum, popularly known as the common puffball, warted puffball, gem-studded puffball or devil's snuff-box, is a species of puffball fungus in the family Agaricaceae. A widespread species with a cosmopolitan distribution, it is a medium-sized puffball with a round fruit body tapering to a wide stalk, and dimensions of 1.5 to 6 cm wide by 3 to 10 cm tall. It is off-white with a top covered in short spiny bumps or "jewels", which are easily rubbed off to leave a netlike pattern on the surface. When mature it becomes brown, and a hole in the top opens to release spores in a burst when the body is compressed by touch or falling raindrops.
Lycoperdon umbrinum, commonly known as the umber-brown puffball, is a type of Puffball mushroom in the genus Lycoperdon. It is found in China, Europe, Africa, and North America.
Calvatia craniiformis, commonly known as the brain puffball or the skull-shaped puffball, is a species of puffball fungus in the family Agaricaceae. It is found in Asia, Australia, and North America, where it grows on the ground in open woods. Its name, derived from the same Latin root as cranium, alludes to its resemblance to an animal's brain. The skull-shaped fruit body is 8–20 cm (3–8 in) broad by 6–20 cm (2–8 in) tall and white to tan. Initially smooth, the skin (peridium) develops wrinkles and folds as it matures, cracking and flaking with age. The peridium eventually sloughs away, exposing a powdery yellow-brown to greenish-yellow spore mass. The puffball is edible when the gleba is still white and firm, before it matures to become yellow-brown and powdery. Mature specimens have been used in the traditional or folk medicines of China, Japan, and the Ojibwe as a hemostatic or wound dressing agent. Several bioactive compounds have been isolated and identified from the brain puffball.
Apioperdon pyriforme commonly known as the pear-shaped puffball or stump puffball, is a saprobic fungus present throughout much of the world. Emerging in autumn, this puffball is common and abundant on decaying logs of both deciduous and coniferous wood. It is considered a choice edible when still immature and the inner flesh is white. It is often called Lycoperdon pyriforme, but was transferred to Apioperdon in 2017 based on phylogenetic and morphological differences. It is the only species in the genus.
Bovista is a genus of fungi commonly known as the true puffballs. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the species of Bovista are now placed in the family Agaricaceae of the order Agaricales. Bovista species have a collectively widespread distribution, and are found largely in temperate regions of the world. Various species have historically been used in homeopathic preparations.
Bovista nigrescens, commonly referred to as the brown puffball or black bovist, is an edible cream white or brown puffball. Phylogenetic relationships between Bovista nigrescens and species of Lycoperdaceae were established based on ITS and LSU sequence data from north European taxa.
Handkea utriformis, synonymous with Lycoperdon utriforme, Lycoperdon caelatum or Calvatia utriformis, is a species of the puffball family Lycoperdaceae. A rather large mushroom, it may reach dimensions of up to 25 cm (10 in) broad by 20 cm (8 in) tall. It is commonly known as the mosaic puffball, a reference to the polygonal-shaped segments the outer surface of the fruiting body develops as it matures. Widespread in northern temperate zones, it is found frequently on pastures and sandy heaths, and is edible when young. H. utriformis has antibiotic activity against a number of bacteria, and can bioaccumulate the trace metals copper and zinc to relatively high concentrations.
Bovista aestivalis is a species of small puffball in the family Agaricaceae. It is generally found in the coastal regions of California, but was reported from Korea in 2015. This fungus is often confused with Bovista dermoxantha, because of its similar peridium, and Bovista plumbea. The surest way to tell the species apart is to examine the spores and exoperidium, respectively, with a microscope.
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.
Calvatia sculpta, commonly known as the sculpted puffball, the sculptured puffball, the pyramid puffball, or Sierran puffball, is a species of puffball fungus in the family Agaricaceae. Attaining dimensions of up to 8 to 15 cm tall by 8 to 10 cm wide, the pear- or egg-shaped puffball is readily recognizable because of the large pyramidal or polygonal warts covering its surface. It is edible when young, before the spores inside the fruit body disintegrate into a brownish powder. The spores are roughly spherical, and have wart-like projections on their surfaces.
Arachnion is a genus of gasteroid fungi in the family Agaricaceae.
Mycenastrum is a fungal genus in the family Agaricaceae. The genus is monotypic, containing one widely distributed species, Mycenastrum corium, known by various common names: the giant pasture puffball, leathery puffball, or tough puffball. The roughly spherical to turnip-shaped puffball-like fruit bodies grow to a diameter of 6–24 cm (2–9 in). Initially covered by a thick, felted, whitish layer, the puffballs develop a characteristic checkered skin (peridium) in age. When the internal spore mass, the gleba, is firm and white, the puffball is edible, although some individuals may suffer mild gastrointestinal symptoms after eating it. As the spores mature, the gleba turns first yellowish then purplish brown. Spores are released when the peridium eventually splits open into irregularly shaped sections. Microscopically, the gleba consists of spherical, dark brown spores with rounded bumps on their surfaces, and a capillitium—intricately branched fibers that form long thorn-like spines. The puffball grows on or in the ground in prairie or desert habitats. Although widely distributed, it is not commonly encountered. Mycenastrum corium is a threatened species in Europe.
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.
Handkea excipuliformis, commonly known as the pestle puffball or long-stemmed puffball, is a species of the family Agaricaceae. A rather large puffball, it may reach dimensions of up to 15 cm (5.9 in) broad by 25 cm (9.8 in) tall. Widespread in northern temperate zones, it is found frequently on pastures and sandy heaths.
Lycoperdon marginatum, commonly known as the peeling puffball, is a type of puffball mushroom in the genus Lycoperdon. A common species, it is found in Europe and North America, where it grows on the ground. It is characterized by the way that the spiny outer layer peels off in sheets.
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.
Bovista pila, commonly known as the tumbling puffball, is a species of puffball fungus in the family Agaricaceae. A temperate species, it is widely distributed in North America, where it grows on the ground on road sides, in pastures, grassy areas, and open woods. There are few well-documented occurrences of B. pila outside North America. B. pila closely resembles the European B. nigrescens, from which it can be reliably distinguished only by microscopic characteristics.
Calvatia pachyderma, also known as the elephant-skin puffball or thick-skinned puffball, is a species of edible fungus. This mid-sized, spring-fruiting puffball is known from relatively dry, open places near human settlements. The appropriate binomial name, taxonomic placement, and geographic distribution "have been much debated and are the subject of controversy".