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Psathyrella

Psathyrella is a large genus of saprotrophic fungi in the family Psathyrellaceae (order Agaricales), encompassing approximately 400 species of small to medium-sized mushrooms worldwide in its current circumscription, commonly known as brittlestems due to their fragile stipes and often drab, hygrophanous caps that fade with age.^[1] These mushrooms typically feature dark brown to purplish-black spore prints, smooth-walled basidiospores with an apical pore, and the presence of pleurocystidia and cheilocystidia on the gills.^[2] They are primarily decomposers of organic matter, such as decaying wood, leaf litter, humus, or occasionally dung, and are found in diverse habitats including forests, grasslands, and urban areas across temperate and tropical regions.^[1] The genus was established by Elias Magnus Fries in 1838, with the type species P. gracilis, though taxonomic revisions have significantly altered its circumscription over time.^[3] Early monographs, such as those by Alexander H. Smith (1972) for North America and Jan Kits van Waveren (1985) for Europe, recognized hundreds of species based on morphology, but molecular phylogenetic studies have revealed that the traditional broad sense of Psathyrella was polyphyletic, with species distributed across at least 11 clades in 2008 analyses.^[1] Subsequent revisions, such as Örstadius et al. (2015), narrowed the genus to a monophyletic core (Psathyrella s.s.) within five major clades of Psathyrellaceae, leading to the transfer of many taxa to genera like Coprinellus, Candolleomyces, and Parasola; further reclassifications (e.g., Wächter & Melzer 2020) have continued this process.^[1]^[4] More recent work (e.g., Voto et al., 2019) has refined subsections like Spadiceogriseae using combined ITS and Tef-1α data.^[2] Despite their ecological role in nutrient cycling, Psathyrella species are generally overlooked by foragers due to their inconspicuous appearance and inedibility, with no significant medicinal or economic value reported.^[1] Identification often requires microscopic examination of spore features and cystidia, as macroscopic traits overlap with related genera such as Coprinellus and Panaeolus. Ongoing taxonomic research continues to describe new species and resolve synonymies, with over 900 names documented in databases like MycoBank, though many remain unverified or reclassified.^[3]

Taxonomy

History and classification

The genus Psathyrella was originally described by Elias Magnus Fries in 1838 as a subdivision (stirps XXVIII) under the genus Agaricus in his Epicrisis Systematis Mycologici, based on species with fragile fruiting bodies, dark brown spores, and a tendency toward deliquescence similar to those in Coprinus. Fries initially grouped many of these taxa within Coprinus due to shared features like rusty-brown to black spore prints and partial gill deliquescence in some species, but Psathyrella species were distinguished by their general lack of full auto-digestion (deliquescence) of the lamellae, a key trait of true Coprinus. In 1872, Lucien Quélet elevated the group to generic rank as Psathyrella (with the type species P. gracilis), emphasizing the fragile, brittle nature of the stipe and cap derived from the Greek psathyros (friable).^[5] Throughout the 20th century, significant taxonomic revisions occurred as species were transferred from the broad Coprinus sensu lato to Psathyrella based on morphological differences, particularly spore ornamentation and the absence of true deliquescence; notable contributions include Alexander H. Smith's 1972 monograph The North American Species of Psathyrella, which described over 400 species and formalized many transfers.^[6] Historically placed in the family Coprinaceae alongside Coprinus, the genus was reclassified into the newly erected family Psathyrellaceae in 2001, reflecting distinctions in spore morphology (e.g., germ pores and hilar appendages) and early molecular evidence separating non-deliquescent lineages.^[7] Phylogenetic analyses using nuclear ribosomal DNA (nrDNA) sequences in the 2000s confirmed the monophyly of Psathyrellaceae and positioned Psathyrella in close relation to Coprinellus and Coprinopsis, though the genus itself proved polyphyletic with multiple clades warranting further generic segregations. These studies, such as those by Padamsee et al. (2008), highlighted convergent evolution of traits like spore color and ecology across the family, leading to refined boundaries. Subsequent research has transferred numerous species to segregate genera such as Candolleomyces (now comprising about 60 species as of 2025) and established new genera like Heteropsathyrella (2021), further narrowing Psathyrella's circumscription.^[1]^[8]^[9] As of 2025, approximately 400 species are recognized in Psathyrella, though estimates vary due to ongoing molecular revisions and synonymy resolutions.

Etymology

The genus name Psathyrella derives from the Greek psathuros (ψαθυρος), meaning "friable" or "brittle", alluding to the fragile stipe and gills typical of species in this group.^[10] In 1872, French mycologist Lucien Quélet emended the original name Psathyra (proposed by Elias Magnus Fries) to Psathyrella by appending the diminutive suffix "-ella", a convention frequently employed in mycology to denote smaller or related forms.^[3] Common English names like "brittlestems" or "brittle caps" stem directly from this trait of structural delicacy, emphasizing the ease with which the fruiting bodies disintegrate.^[11]^[10] Since Quélet's emendation, the etymological understanding of Psathyrella has remained unchanged, with consistent references to its Greek roots and mycological nomenclature in subsequent literature.^[3]

Description

Macroscopic characteristics

The fruit bodies of Psathyrella species are typically small and fragile, with a brittle consistency that distinguishes the genus. The cap (pileus) measures 1–5 cm in diameter, starting conical to campanulate and expanding to convex or nearly flat, often with a small central umbo; it is thin-fleshed, hygrophanous (absorbing water and changing color), and features striate (grooved) margins due to translucent tissue. Colors range from pale tan or clay-brown when moist to darker gray-brown or reddish-brown when dry, frequently appearing two-toned with a darker center.^[12]^[6] The gills (lamellae) are adnate or adnexed, occasionally with a short decurrent tooth, close to crowded, and narrow, initially pale whitish before darkening to brown or blackish-brown as spores mature; they are delicate and prone to breaking easily when handled.^[12]^[6] The stem (stipe) is slender and central, 3–10 cm long and 0.1–0.5 cm thick, hollow or stuffed, fragile enough to snap cleanly, with a surface that is smooth to fibrillose or pruinose (powdery) especially at the apex; it lacks a volva at the base and typically has no annulus, though evanescent fibrillose or membranous remnants from a partial veil may cling to the cap margin or upper stem in young specimens.^[12] Odor and taste are generally mild or indistinct, rarely strong or disagreeable across the genus. Fruit bodies appear solitary to gregarious, often in small groups or clusters on substrates, but do not form fairy rings.^[13]^[14]

Microscopic characteristics

The basidiospores of Psathyrella are typically ellipsoid to ovoid or subcylindrical, measuring 6–12 µm in length by 3.5–6.5 µm in width, with a quotient (Q) often ranging from 1.3–2. They appear dark brown to blackish in 5% KOH mounts, feature smooth walls, and possess a distinct hilar appendix along with a broad apical germ pore measuring 0.5–1.5 µm in diameter.^[15]^[6]^[16] Spore prints are characteristically dark purplish-brown to black, and the spores exhibit no amyloid reaction in Melzer's reagent.^[15]^[17] Basidia are clavate, predominantly 4-spored (occasionally 2-spored), and measure approximately 15–30 µm in length by 6–10 µm in width, typically bearing basal clamp connections.^[15]^[18] Cheilocystidia are abundant on gill edges, varying from cylindrical and clavate to utriform or lageniform, with dimensions of 20–80 µm long by 6–24 µm wide; they are thin-walled and may occur in two forms, including smaller clavate elements.^[15]^[17]^[18] Pleurocystidia, when present, are similar in shape and size but scattered on gill faces, and their absence or presence serves as a species-level diagnostic trait.^[15]^[16] The pileipellis is structured as a hymeniderm composed of erect, cylindrical to globose hyphae, often transitioning to a paraderm or, less commonly, a cutis with diverticulate elements; elements are typically 10–40 µm wide and hyaline to pigmented.^[15]^[19] A key diagnostic feature distinguishing Psathyrella from related genera like some Coprinopsis species is the lack of metachromatic granules in the spores.^[6]^[16]

Habitat and ecology

Distribution

Psathyrella species exhibit a cosmopolitan distribution, occurring on all continents except Antarctica. The genus is most diverse in the temperate regions of the Northern Hemisphere, particularly Europe and North America, where historically over 400 species were described for North America (Smith 1972), though many have been reclassified following molecular studies.^[6]^[1] They are also common in Asia, with more than 50 species reported from China and at least 40 from India; the genus is present in Australia across multiple states and in South America, though records are sparser in arid and polar regions.^[20]^[6]^[21]^[22]^[19] In temperate zones, Psathyrella species typically appear seasonally from spring to fall, while in tropical areas they fruit year-round. Some species have been introduced to new regions via human activities, including in urban parks and grassy areas. As of 2025, approximately 400–600 species are described worldwide, with ongoing discoveries in biodiversity hotspots such as Asia.^[23]^[19]^[1]

Ecological roles

Psathyrella species are primarily saprotrophic fungi that play a crucial role in ecosystems by decomposing lignocellulose in substrates such as wood, leaf litter, soil, and dung, thereby facilitating nutrient recycling in forests and grasslands.^[1] As decomposers, they break down complex organic compounds, releasing essential nutrients like carbon, nitrogen, and phosphorus back into the soil for uptake by plants and other organisms.^[24] This process is enhanced by their production of enzymes, including laccases and lignin-degrading peroxidases, which target cellulose and lignin, accelerating decomposition rates.^[24] Within the genus, ecological niches vary: some species are lignicolous, colonizing dead wood; others are terricolous, growing on soil or humus; and several are coprophilous, specializing in dung decomposition.^[1]^[25] Additionally, certain species, such as Psathyrella carbonaria, act as post-fire specialists in phoenicoid niches, fruiting exclusively in burned areas where they contribute to soil stabilization and early succession by aggregating soil particles through their mycelial networks.^[26] No mycorrhizal associations with higher plants are known for Psathyrella, distinguishing them from many other basidiomycetes; however, potential roles as minor pathogens on weakened plants remain unconfirmed.^[1] Fruiting in Psathyrella is typically triggered by environmental cues like increased moisture from rainfall and moderate temperatures, leading to the rapid development of basidiocarps that last only 1-3 days before deliquescing. These short-lived structures release billions of dark brown to black spores, enabling effective dispersal primarily via wind, though insects may assist in some species, particularly in aquatic habitats.^[27] This high-volume spore production ensures colonization of new substrates, supporting the genus's role in nutrient cycling across diverse ecosystems.^[28]

Species

Diversity

The genus Psathyrella is estimated to comprise approximately 400–500 species worldwide, with particularly high diversity observed in humid forest ecosystems where saprotrophic niches are abundant. Many additional species remain undescribed, primarily due to cryptic speciation that is only revealed through molecular analyses, leading to an underestimation of true richness.^[1] Ongoing taxonomic revisions since 2020 have further split the genus, transferring species to new genera like Olotia and refining its circumscription. Endemism in Psathyrella is pronounced in Europe, the type region of the genus, and North America, where extensive taxonomic surveys have documented hundreds of species, including regional endemics like P. cystidiosa in Minnesota.^[29] In contrast, endemism appears lower in tropical regions historically, but recent molecular studies using markers such as ITS have uncovered novel lineages, suggesting increasing diversity as research expands beyond temperate zones.^[30] Most Psathyrella species are not considered globally threatened, though some wood-associated taxa are declining due to habitat loss from deforestation and urbanization; as of 2025, only a few, such as P. atlantica (Data Deficient) and P. cystidiosa (locally endangered), have been assessed using IUCN criteria by the Global Fungal Red List Initiative, with the majority unlisted.^[31]^[32] Diversity in Psathyrella is driven by high morphological plasticity, which has resulted in overlooked synonyms and convergent traits across lineages, complicating traditional classifications.^[28] Recent molecular phylogenies, particularly those employing ITS sequencing, have resolved species complexes by splitting morphologically similar taxa, revealing polyphyletic groups and new clades.^[33] Significant research gaps persist, including incomplete monographs for much of the genus, with comprehensive treatments limited to regions like North America and Europe; the Southern Hemisphere, including South America and Australia, lacks detailed surveys, hindering global cataloging efforts.^[34]

Selected species

Psathyrella aquatica is a unique saprobic species known for being the first reported gilled mushroom to fruit underwater, discovered in Oregon, USA, in 2012. It occurs in small streams and rivers, growing submerged on decaying wood or plant debris in cool, flowing water.^[35] The cap is 0.5–2 cm in diameter, convex to flat, translucent and pale tan to whitish when submerged, with a smooth, hygrophanous surface. The stem is slender, 1–4 cm long and 0.1–0.2 cm thick, fragile, and whitish, while the gills are distant and pale, maturing to brown. Microscopically, it features smooth basidiospores 8–11 × 4–5.5 µm with a germ pore. Due to its aquatic habitat, it is rarely encountered and not considered edible.^[36] Psathyrella piluliformis, the common stump brittlegill, is a small agaric distributed worldwide in temperate regions, including North America, Europe, and parts of Asia, typically fruiting in dense clusters on decaying hardwood stumps and woody debris.^[13] The cap is 1–2 cm across, initially spherical and button-like when young, expanding to hemispherical or convex with a reddish-brown to chestnut hue that fades when dry, often covered by a thin white veil in immature stages.^[37] The stem is brittle, 2–6 cm tall and under 0.3 cm thick, whitish with a fibrillose base, and the gills start pale tan before turning dark brown from maturing spores.^[38] As a wood-decomposing saprobe, it serves as an indicator of advanced decay in urban and forested settings, though it is not coprophilous as sometimes misreported.^[39] Psathyrella corrugis (formerly known as P. gracilis), the type species of the genus, is a widespread lignicolous fungus found on decaying wood in forests across North America and Europe. It fruits singly or in groups on hardwood logs and stumps.^[40] The cap measures 1–5 cm in diameter, starting conical and expanding to convex with a wrinkled or corrugated surface when dry, colored reddish-brown to dark brown when moist and fading to pale tan. The stem is 4–10 cm long and 0.3–0.6 cm thick, fragile, whitish to brownish, often with rhizomorphs at the base. The gills are close, adnate, and turn dark brown with age. Spores are 7–9 × 4–5 µm, smooth with an apical pore. It is inedible and serves as a model for studying the genus's morphology.^[41] Psathyrella prona is a grassland-inhabiting species noted for its small size and occurrence in open areas such as woodland glades, fields, and lawns, with records spanning Europe and recent reports from Asia, including Northeast China.^[42] The cap is 1–3 cm wide, convex to flat with a persistent central dark spot or umbo, colored pale red-brown to gray-brown and striate at the margin when moist.^[43] The stem is 3–7 cm tall and slender, whitish and fragile, while the gills are distant and brownish.^[42] Its ellipsoid spores measure 7–9 × 4–5 µm, smooth with a germ pore, distinguishing it from related taxa in grassy habitats.^[44] While most Psathyrella species are inedible due to their extreme fragility and lack of flavor, a few contain trace amounts of psilocybin, potentially inducing weak hallucinogenic effects if consumed in quantity, though such instances are rare and not characteristic of the genus.^[45] These mushrooms are frequently employed in mycology education to demonstrate key diagnostic features, such as dark-spored prints and diverse cystidial structures, making them accessible models for studying saprobic agarics.^[6]

References

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Psathyrella multipedata, Clustered Brittlestem mushroom - First Nature
Psathyrella, the genus name, is the diminutive form of Psathyra, which comes from the Greek word psathuros meaning friable; it is a reference to the crumbly ...
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Brittlestems (Genus Psathyrella) - iNaturalist
Psathyrella is a large genus of about 400 species, and is similar to the genera Coprinellus, Coprinopsis, Coprinus and Panaeolus, usually with a thin cap ...
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Jun 29, 2015 · A number of. Psathyrella species are coprophilous and grow on dung of various animals (Singer 1986, Pegler. 1986, Amandeep et al. 2013). Kirk et ...
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The Genus Psathyrella (MushroomExpert.Com)
Under the microscope, species of Psathyrella have spores that are brown or dark gray when mounted in KOH and feature smooth walls, as well as an apical pore.
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Psathyrella piluliformis (MushroomExpert.Com)
Psathyrella piluliformis grows in clusters on hardwoods, has a white veil on young caps, a brown to reddish-brown cap, and a fragile, whitish stem.
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[PDF] Psathyrella (Fr.) Quél. LEIF ÖRSTADIUS & HENNING KNUDSEN
Pileipellis a hymeniderm, sometimes with a transition to a paraderm, rarely a cutis. Veil most often present, either as hyphae with cells differing in size, ...
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PSATHYRELLA in the Pacific Northwest
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Twenty seven species of Psathyrella have been found in Northeast China. Amongst them, P. conica, P. jilinensis, P. mycenoides and P. subsingeri are described as ...
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Apr 27, 2021 · Psathyrella were made, of which four species viz. Psathyrella ... As many as 40 species of this genus are reported from India (Natarajan et al.
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Psathyrella ammophila, Dune Brittlestem mushroom - First Nature
The appearance of these salt-tolerant fungi is very variable, dependent on ... Season. June to November in Britain and Ireland. Culinary Notes. Even if ...
[22]
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Aug 6, 2025 · ... Psathyrella atroumbonata ) cultivated on lignocellulose waste was investigated. Mn2+ and Ca2+ ions stimulated growth of both fungi and ...
[23]
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Three new species are described: Psathyrella fimiseda, P. merdicola, and P. scatophila. P. stercoraria is described as a new species in order to validate the ...
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Psathyrellas are associated with the cycling of organic matter, such as wood, leaf litter or dung, and may occur in or outside woods. Psathyrella cystidiosa was ...<|control11|><|separator|>
[29]
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Psathyrella atlantica
Without further data the species could be anything from Least Concern to Critically Endangered and so it is assessed as Data Deficient. Geographic range.
[31]
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Apr 23, 2016 · Psathyrella cystidiosa was listed as an endangered species in Minnesota in 1996, previously thought to be endemic to Minnesota. Psathyrella ...
[32]
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[33]
Psathyrella candolleana (MushroomExpert.Com)
Psathyrella candolleana is a saprobe, usually found in the vicinity of recently dead hardwood trees--often in urban areas, but also in the woods.
[34]
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[35]
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Habitat. Scattered to gregarious along trails, in gardens or in grassy areas; fruiting from late fall to mid-winter. Edibility. Described as edible, but ...Missing: distribution | Show results with:distribution
[36]
Psathyrella piluliformis, Common Stump Brittlestem mushroom
Psathyrella piluliformis is a very common wood-rotting fungus in broadleaf woodlands, where it is found on and around the stumps of dead deciduous trees.Missing: emended Quélet 1888
[37]
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[38]
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It is distinguished by a clustered fruiting habit, usually at the base of hardwood stumps, a hygrophanous brown cap with appendiculate veil fragments that in ...
[39]
Parasola conopilea (MushroomExpert.Com)
Like species in Psathyrella (as traditionally defined), Parasola conopilea features a fragile stem, a dark spore print, and a hygrophanous brown cap. But ...
[40]
Parasola (Psathyrella) conopilus, Conical Brittlestem mushroom
The Conical Brittlestem has a shiny red-brown, conical cap (2-3.5cm) when moist, a long white stem (7-14cm), and grows on leaf litter under deciduous trees.Missing: characteristics | Show results with:characteristics
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California Fungi: Parasola conopilea - MykoWeb
Parasola conopilea has a conical, reddish (when young) hygrophanous cap and yellowish stipe, found in woodchips, rich soil, and waste places. Its edibility is ...
[42]
Psathyrella prona | NatureSpot
It occurs on soil, often among grass in woodland glades, fields and gardens. Late summer and autumn.Missing: spores | Show results with:spores
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Psathyrella prona - mushrooms of Russia
In St. Petersburg, Leningrad Oblast, and north-western Russia, Psathyrella prona has been found growing in a variety of habitats, including deciduous and mixed ...Missing: spores | Show results with:spores
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