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Psathyrellaceae

Psathyrellaceae is a family of basidiomycete fungi in the order Agaricales, consisting primarily of agaricoid species with gilled fruiting bodies, dark brown to black spore prints, and often fragile, deliquescent tissues that autodigest into an inky fluid in many taxa.^[1]^[2] The family, established as Psathyrellaceae Locq. ex Locq. (1972) and later emended by Vilgalys, Moncalvo & Redhead (2001),^[3] encompasses approximately 90% of all known deliquescing mushroom species and is distinguished by its polyphyletic genus Psathyrella, which serves as a repository for numerous small, brown-spored agarics.^[1] Key genera within Psathyrellaceae include Psathyrella (over 400 species, often psathyrelloid with non-deliquescent gills), Coprinopsis and Coprinellus (coprinoid with deliquescing lamellae), Parasola, and Lacrymaria, reflecting a diversity of morphologies from small, conical-capped mushrooms to larger, veil-bearing forms.^[2]^[4] Species exhibit convergent traits such as utriform cystidia, a hymeniderm pileipellis, and spores with a prominent germ pore, though autodigestion—mediated by enzymatic lysis—is a hallmark in coprinoid lineages but absent in many psathyrelloid ones.^[2]^[5] Ecologically, Psathyrellaceae are cosmopolitan saprotrophs, decomposing lignocellulosic substrates in terrestrial habitats worldwide, including decaying wood, leaf litter, soil, grassy lawns, bogs, and dung, with peak fruiting in temperate and tropical regions during moist seasons.^[6] Their rapid development and autodigestion facilitate efficient spore dispersal, contributing to nutrient cycling in forest floors and urban greenspaces, though some species face taxonomic challenges due to cryptic diversity revealed by molecular phylogenies.^[2]

Description

Macroscopic Features

Members of the Psathyrellaceae are typically small to medium-sized agarics characterized by fragile fruiting bodies with thin, brittle stipes and caps; many species, particularly in coprinoid genera such as Coprinopsis, deliquesce into an inky liquid upon maturation, while psathyrelloid taxa often lack this trait.^[7] This autodigestion primarily affects the gills and cap margins in deliquescent species, contributing to the family's ephemeral nature, while the stipe usually persists. The overall texture is notably brittle, leading to rapid decay shortly after spore dispersal, which underscores their saprotrophic lifestyle.^[8] The cap (pileus) is generally convex to umbonate, measuring 1-10 cm in diameter, and hygrophanous, changing color with moisture levels—often appearing darker when wet. Colors range from grayish-brown to dark brown, with striate margins visible under moist conditions due to translucent tissue. The surface is smooth to slightly fibrillose or scaly, emphasizing the delicate structure common to the family.^[9] Gills (lamellae) are adnate to free, close to crowded, and initially pale before turning dark gray to blackish from the abundant dark spores; in deliquescent species, they autodigest progressively from the margin inward. This darkening and liquefaction are key visible traits, though the spore pigmentation itself is detailed microscopically.^[7] The stem (stipe) is slender and fragile, typically 3-15 cm tall and 1-5 mm thick, hollow or stuffed, with a smooth to fibrillose surface that is usually pale. It often bears remnants from a partial veil, such as fibrillose patches or a fugacious annular zone.^[8] Odor is typically mealy or indistinct, and taste mild, aligning with the unassuming external profile.^[10] A partial veil is present in some species, forming evanescent patches on the cap margin or a fragile ring on the stem, but it is often fugacious and leaves minimal traces. These features collectively highlight the family's adaptation for quick spore release in decaying organic matter.^[7]

Microscopic Features

The microscopic features of Psathyrellaceae are critical for distinguishing members of this family from other dark-spored agarics, particularly through the examination of basidiospores, reproductive structures, and tissue arrangements under light microscopy. Basidiospores in Psathyrellaceae are typically dark brown to black in mass, with spore prints ranging from dark brown to purplish brown or blackish, measuring 5-15 µm in length, and exhibit smooth or slightly rough surfaces; they are often elliptical to subglobose in shape, featuring a prominent hilar appendix and a distinct germ pore that facilitates germination.^[11] These spores are non-amyloid, lacking a positive reaction with iodine reagents, and their pigmentation arises from melanin-like compounds deposited in the spore walls.^[12] The spore pigments display metachromatic properties, shifting color under certain stains, and notably bleach when treated with concentrated sulfuric acid (H2SO4), a reaction that helps differentiate Psathyrellaceae from families like Cortinariaceae, where pigments are more stable; in KOH, spores appear brown or dark gray.^[13] ^[14] This bleaching occurs due to the solubility of the pigment granules in acidic solutions.^[15] Basidia are predominantly 4-spored, clavate in form, and range from 15-30 µm in length, with thin-walled sterigmata that bear the spores; occasional 2- or 3-spored variants occur but are less common across the family.^[16] Cystidia are abundant and serve as key diagnostic elements: cheilocystidia and pleurocystidia are typically utriform to lageniform, measuring 30-60 µm tall, with necks often 5-10 µm wide and apexes that may bear crystalline appendages; these structures are hyaline, thin-walled, and project from the gill edges and faces, aiding in spore dispersal and species identification.^[11] The gill trama is irregular in organization, composed of loosely interwoven, hyaline hyphae without the radial elongation seen in families like Agaricaceae, contributing to the fragile nature of the lamellae.^[17] The pileipellis varies between a cutis of repent, cylindrical hyphae or a hymeniform structure with erect, vesiculose elements, sometimes up to 50 µm tall in species with cellular layers; this arrangement often includes clamped connections at hyphal septa throughout the family.^[16]

Taxonomy

History

The genus Psathyrella was established by Lucien Quélet in 1872 to accommodate small, fragile agarics that superficially resembled species of Coprinus but lacked the deliquescent (auto-dissolving) lamellae characteristic of that genus.^[6] Prior to this, such taxa were often included within the broadly conceived family Coprinaceae sensu lato, which encompassed a diverse array of dark-spored, often coprophilous or lignicolous mushrooms with varying degrees of gill auto-digestion.^[18] In the early 20th century, many species now recognized as belonging to Psathyrella were misclassified or lumped into other genera, such as Psilocybe and Deconica, due to overlapping morphological features like dark spores and collybioid habits, leading to taxonomic confusion in pre-molecular classifications.^[18] A significant advancement came with Rolf Singer's comprehensive 1986 monograph The Agaricales in Modern Taxonomy, which provided a detailed infrageneric classification of Psathyrella based on morphological traits, including spore shape, veil remnants, and habitat preferences, while maintaining its placement within Coprinaceae.^[19] The family Psathyrellaceae was formally established in 2001 by Rytas Vilgalys, Jean-Marc Moncalvo, and Scott A. Redhead, drawing on preliminary molecular phylogenetic data (primarily ribosomal DNA sequences) that demonstrated the polyphyly of Coprinaceae sensu lato and separated psathyrelloid taxa into a distinct clade, with Psathyrella designated as the type genus.^[20] This reclassification narrowed the scope of Coprinaceae, which, following the transfer of Coprinus sensu stricto to Agaricaceae, became a synonym of that family in 2001, thereby elevating the status of Psathyrellaceae as an independent lineage within the Agaricales.^[21]

Current Classification

Psathyrellaceae is a family of fungi within the order Agaricales, classified under the phylum Basidiomycota, class Agaricomycetes.^[22] The family encompasses approximately 20 genera and over 2000 species, many of which exhibit fragile, deliquescing fruiting bodies with dark spores. The type genus is Psathyrella (Fr.) Quél., established in 1872.^[23] Significant taxonomic revisions have shaped the current framework, notably the 2015 multi-locus phylogenetic study by Örstadius et al., which demonstrated the polyphyly of Psathyrella sensu lato and segregated it into multiple genera, including the introduction of three new ones: Kauffmania, Homophron, and Typhrasa.^[24] This work refined the boundaries of Psathyrellaceae by incorporating segregates from the former Coprinaceae, emphasizing molecular markers such as ITS, LSU, SSU, and RPB2. Recent 2025 updates further expand the family, with Yang et al. describing the new genus Cantonopsathyra and two new species, alongside Zheng et al. introducing Candolleomyces yunnanensis based on morphological and molecular evidence.^[25]^[26] The family is formally recognized in authoritative databases as of 2025, including Index Fungorum and MycoBank, which list Psathyrellaceae at the family rank with Psathyrella as the type.^[27] Ongoing molecular studies indicate potential further subdivisions among psathyrelloid species, particularly in unresolved clades within genera like Psathyrella and Candolleomyces.^[28]

Phylogeny

Molecular Studies

Early molecular studies on Psathyrellaceae utilized nuclear ribosomal DNA sequences, including the internal transcribed spacer (ITS) region and nuclear large subunit (nLSU) rDNA, to delineate the family's boundaries from Coprinaceae. In 2001, analyses of these markers revealed a distinct clade comprising Psathyrella and related genera, separate from the Coprinus comatus group, justifying the erection of Psathyrellaceae as a new family within Agaricales. This work built on prior nLSU data showing strong phylogenetic support (≥90%) for the separation, highlighting the family's monophyly based on ribosomal gene divergence. A comprehensive multi-locus phylogenetic study in 2015 expanded this foundation by sequencing ITS, LSU, RPB1, RPB2, and EF1-α across more than 100 species, employing Bayesian inference and maximum likelihood methods. These analyses confirmed Psathyrellaceae as monophyletic within the Agaricales suborder Agaricineae and demonstrated that Psathyrella s.l. is paraphyletic, necessitating taxonomic revisions. The study introduced three new genera—Homophron, Kauffmania, and Typhrasa—along with 18 new species, refining the family's internal structure through robust tree topologies with high posterior probabilities and bootstrap support.^[24] Recent investigations in 2025 have further refined these phylogenies using multi-locus approaches in journals such as MycoKeys and Phytotaxa, confirming expansions in genera like Ephemerocybe and describing additional new species and one new genus within Psathyrellaceae. For instance, phylogenetic analyses incorporating ITS, LSU, and tef-1α sequences supported the delimitation of novel taxa and provided notes on Ephemerocybe's relationships, emphasizing ongoing resolution of generic boundaries. Although mitogenome data have been integrated in broader Agaricales studies to explore deeper evolutionary relationships, their application to Psathyrellaceae remains preliminary, with nuclear markers continuing to dominate family-level inferences. Key findings across these works uphold the dark-spored clade's integrity but reveal internal diversity in genera such as Coprinopsis, where subclades indicate potential for further taxonomic subdivision pending additional sampling.^[29]

Relationships to Other Families

Psathyrellaceae occupies a position within the order Agaricales, forming part of the core dark-spored clade alongside Agaricaceae, from which it diverged following the reclassification of certain Coprinus species, such as Coprinus comatus, into the latter family based on molecular evidence.^[24] Recent multi-locus phylogenetic studies employing nuclear ribosomal and protein-coding genes have reinforced this placement, showing Psathyrellaceae as a distinct but sister lineage to the expanded Agaricaceae, with shared evolutionary history in saprotrophic dark-spored agarics.^[29] These analyses, incorporating up to several dozen loci, highlight the family's monophyly within Agaricales, distinct from more distant groups like the Tricholomataceae.^[30] Members of Psathyrellaceae share key traits with Agaricaceae, including dark brown to black spore deposits and a predominantly saprotrophic lifestyle on decaying organic matter, reflecting their common ancestry in nutrient-recycling niches.^[24] However, Psathyrellaceae is distinguished by its characteristically fragile basidiomata that often deliquesce and basidiospores featuring a prominent germ pore, adaptations not uniformly present in Agaricaceae.^[4] In contrast to Galeropsidaceae, which includes coprophilous genera like Panaeolus lacking autodigestion (deliquescence) and typically exhibiting non-poroid, mottled spores, Psathyrellaceae emphasizes terrestrial saprotrophy with poroid spores and frequent tissue breakdown.^[31] Similarly, Bolbitiaceae differs through genera like Conocybe and Bolbitius, which often display bulbous stipe bases, viscid or wrinkled pilei, and spores with rugulose ornamentation rather than the smooth, poroid forms typical of Psathyrellaceae.^[32] Phylogenetic reconstructions indicate that Psathyrellaceae arose from a common ancestor approximately 50–60 million years ago, coinciding with the diversification of major deliquescent lineages such as Coprinopsis and Parasola during the Eocene, a period of rapid agaric evolution.^[33] Some species exhibit mycoparasitic tendencies, linking the family ecologically to Agaricaceae hosts; for instance, Psathyrella epimyces deforms fruiting bodies of Coprinus comatus through tissue invasion.^[34] Recent multi-locus studies from 2025 further affirm the family's monophyly, supporting its stable placement amid ongoing refinements in Agaricales taxonomy.^[29]

Ecology and Distribution

Habitat and Life Cycle

Psathyrellaceae species are predominantly saprotrophic fungi that colonize nitrogen-enriched substrates, including animal dung, decaying wood, grassy lawns, gardens, and disturbed soils, where they function as primary or secondary decomposers in terrestrial ecosystems.^[4] Many genera, such as Psathyrella and Coprinopsis, exhibit a lignicolous or terricolous habit, breaking down complex polymers like lignin and cellulose to facilitate nutrient cycling, while a subset are coprophilous, thriving on herbivore dung and contributing to the decomposition of organic waste in grasslands and pastures.^[4]^[35] Although most members are free-living saprotrophs, rare cases of mycoparasitism occur, as seen in Psathyrella epimyces, which parasitizes the fruiting bodies of other agarics like Coprinus comatus by deforming host tissues to extract nutrients.^[34] The life cycle of Psathyrellaceae follows the typical hymenogastralean pattern of basidiomycetes, beginning with mycelial colonization of suitable substrates through vegetative hyphae that often feature clamp connections for dikaryotic growth. Fruiting body formation, or basidiomata development, is typically triggered by environmental cues such as rainfall in cool, moist conditions, particularly during late summer to autumn in temperate regions, leading to rapid maturation within 1-3 days.^[36] Basidiospores, which possess a distinct germ pore for germination, are forcibly discharged from basidia on the gills and may deliquesce in many species, where the cap and gills autolyse into an inky fluid to prevent desiccation and enhance spore release in ephemeral habitats.^[33] Post-dispersal, germinated spores form monokaryotic hyphae that eventually fuse in mating to reestablish the dikaryotic mycelium, completing the cycle.^[37] These fungi exhibit short-lived basidiomata, lasting from hours to a few days, which aligns with their occupation of transient, nutrient-rich niches.^[33] Spore dispersal occasionally involves interactions with invertebrates, such as mycophagous insects that consume and excrete viable spores, particularly in coprophilous species where dung-inhabiting arthropods facilitate spread across landscapes.^[38] This mode supports their role in ecosystem dynamics, recycling nutrients in disturbed or enriched environments while adapting to seasonal wet periods for reproduction.^[36]

Global Distribution

The family Psathyrellaceae exhibits a cosmopolitan distribution, with species recorded across all continents except Antarctica.^[39] This widespread occurrence is attributed to the family's saprotrophic lifestyle, allowing adaptation to diverse substrates like decaying wood, dung, and soil. The greatest species diversity is concentrated in the temperate regions of the Northern Hemisphere, particularly in Europe, North America, and Asia, where approximately 500 species of Psathyrella alone have been documented globally, with over 100 recognized in Europe.^[39]^[11] In contrast, diversity is lower in tropical and subtropical zones, though the family is present in areas such as Australia, South America, and Southeast Asia, with new species discoveries expanding known ranges in these regions.^[40]^[41]^[42] Regional hotspots for Psathyrellaceae include urban and disturbed habitats worldwide, such as lawns, gardens, wood chips, and roadside verges, where species like Psathyrella candolleana thrive due to nutrient-rich, anthropogenic environments.^[43]^[44] In Europe, high species richness is evident in deciduous forests and grasslands, while in North America and Asia, similar temperate ecosystems support dense populations. Endemism is notable in certain areas, with many species restricted to regional locales; for instance, recent studies have identified endemics in Yunnan Province, China, including Candolleomyces yunnanensis, highlighting localized diversity in Asian montane forests.^[26] Human activities have facilitated introductions, enabling some species to establish in non-native urban settings globally.^[45] Psathyrellaceae species predominantly occupy mesic to humid climatic zones, favoring moist soils and organic-rich substrates in forests, grasslands, and wetlands.^[46] Their altitudinal range spans from sea level to elevations exceeding 2000 meters, as observed in highland collections from the Alps and Asian plateaus.^[47]^[48] Threats to natural populations include habitat loss from deforestation, urbanization, and soil disturbance, particularly in biodiversity hotspots like the Atlantic Forest and European grasslands; however, the family's resilience in anthropogenic sites often buffers overall declines.^[49]^[50]^[51]

Genera

Psathyrella

Psathyrella is the type genus of the family Psathyrellaceae, comprising approximately 400 species of small to medium-sized, collybioid to tricholomatoid agarics that are typically non- or partially deliquescent and grow as saprotrophs on wood or soil.^[4] These fungi are characterized by fragile basidiomata with a hygrophanous pileus that is usually campanulate to convex, measuring 1-5 cm in diameter, and often brown with striations when moist. The lamellae are adnexed to adnate, dark brown to blackish, and the stipe is slender, fragile, white to pale, and typically exannulate. Microscopically, the basidiospores are ovoid to ellipsoidal, smooth, thick-walled, 7-12 µm long, dark brown to blackish in deposit, and equipped with a prominent germ pore; cheilocystidia are always present and often prominent, while pleurocystidia may be absent or sparse, and clamp connections are common at basidia bases.^[52] Following phylogenetic revisions since 2015, the core Psathyrella (sensu stricto) has been delimited to around 200 species within the /Psathyrella clade, excluding segregate genera such as Coprinellus and Kauffmania, while retaining its cosmopolitan distribution across temperate and tropical regions as wood- or litter-decomposing saprotrophs.^[16] The genus exhibits high morphological diversity, with species varying in pileus texture from smooth to fibrillose and in habitat preferences from terrestrial litter to lignicolous clusters, though all share the non-ink-cap habit of persistent, non-deliquescent gills. No species are considered toxic, though they are generally regarded as inedible due to their fragile texture and insubstantial flesh.^[14] Representative species include Psathyrella candolleana, a common terrestrial form often found on dung or nutrient-rich soil in grasslands and woodlands, featuring a pale tan to whitish hygrophanous cap up to 4 cm broad and ellipsoid spores 8-10 × 4-5 µm.^[53] Another widespread example is Psathyrella piluliformis, which grows gregariously on hardwood stumps and decaying wood, distinguished by its chestnut-brown cap 2-5 cm across with veil remnants and subcylindrical spores 10-13 × 5-6 µm.^[54] Recent taxonomic work in 2025 has added variants and new descriptions, such as those related to Candolleomyces yunnanensis from subtropical Asian regions, highlighting ongoing refinements in species delimitation through molecular and morphological analyses.^[26]

Coprinopsis

Coprinopsis is a genus of mushrooms in the family Psathyrellaceae, comprising approximately 100 species recognized as true ink-caps characterized by complete deliquescence of the gills, where the lamellae autodigest into a black, inky liquid at maturity to facilitate spore dispersal.^[55] These species were transferred from the polyphyletic genus Coprinus in a major taxonomic revision in 2001, based on molecular phylogenetic analyses of nuclear ribosomal DNA sequences that placed them firmly within Psathyrellaceae, distinct from Coprinus sensu stricto in Agaricaceae.^[55] The genus was originally described by Petter Karsten in 1881 but revived and emended in this revision to encompass the clade featuring a hyphal pileipellis and specific veil structures.^[56] Morphologically, Coprinopsis species typically exhibit caps measuring 2–10 cm in diameter, initially cylindrical to bell-shaped, often gray-brown in color, and expanding with age before deliquescing from the margin inward. The spores are smooth, dark brown to black, measuring 8–15 µm in length, distinctly truncate at the end due to a broad germ pore, and produced on basidia with abundant fusiform cheilocystidia and usually pleurocystidia present on the gill edges and faces.^[56] This full autodigestion of the gills contrasts with the partial deliquescence seen in Coprinellus species, which have non-truncate spores lacking such a prominent germ pore.^[55] The diversity of Coprinopsis centers on saprotrophic species primarily associated with dung or decaying wood, with many coprophilous forms thriving in nutrient-rich, disturbed habitats like grasslands and pastures, while others are lignicolous on stumps or buried wood in forests and urban areas.^[57]^[58] Some species, such as Coprinopsis atramentaria, are considered edible when young but cause severe alcohol intolerance due to the mycotoxin coprine, which inhibits aldehyde dehydrogenase and leads to symptoms resembling disulfiram reactions if alcohol is consumed within 48–72 hours.^[59] Representative examples include Coprinopsis atramentaria, a common medium-sized species (cap up to 7 cm) forming dense clusters on wood or soil, and Coprinopsis lagopus, a larger ink-cap (up to 15 cm tall) with woolly stem scaling, often solitary on dung; smaller species like Coprinopsis variegata grow in clusters on wood debris, featuring scaly caps under 3 cm.^[58]^[60] Unlike the grassland-specialized, diminutive Parasola species, Coprinopsis taxa are generally larger and more tied to organic-rich substrates like dung and wood.^[55] Recent molecular studies in 2025 have confirmed distinct subclades within Coprinopsis, particularly in complexes like the Coprinopsis alnivora group, using ITS and LSU sequence data to delineate new species such as C. austrolilacina and C. persicina based on phylogenetic divergence and subtle morphological variations in veil and spore traits.^[41] These analyses reinforce the genus's monophyly and highlight ongoing taxonomic refinements driven by genomic evidence.^[61]

Coprinellus

Coprinellus is a genus of approximately 80 species of small, fragile agarics within the family Psathyrellaceae, distinguished by the presence of a granular or mealy veil that leaves mica-like remnants on the cap surface and partial deliquescence of the gills, where the lamellae liquefy but the cap does not fully autodigest.^[62] These mushrooms are saprotrophic decomposers, typically fruiting in dense clusters or troops on decaying wood, leaf litter, soil, or occasionally dung, contributing to nutrient recycling in forest and grassland ecosystems.^[63] The genus was established to accommodate species previously placed in Coprinus section Micacei, emphasizing their veil anatomy and lack of complete auto-digestion compared to related genera.^[62] Morphologically, Coprinellus species feature caps measuring 1–3 cm in diameter, often conical to bell-shaped when young, expanding to convex with age, and colored in shades of reddish-brown, tawny, or ochre, adorned with fine, glistening granular scales from the veil.^[64] Stipes are slender, 3–10 cm tall and up to 0.5 cm thick, white to pale, and fibrillose, sometimes with a basal mycelial mat. Gills are close, adnate to decurrent, initially pale but turning grayish-black as they deliquesce from the margin inward. Microscopically, basidiospores are dark brown to blackish, smooth, ovoid to mitriform, measuring 6–11 × 4–7 µm, with a small central germ pore; cystidia are abundant on the gill edges and cap surface, aiding in veil formation.^[63] These traits, particularly the veil remnants and spore morphology, differentiate Coprinellus from smoother-capped relatives.^[62] The genus exhibits significant diversity, with species adapted to temperate and subtropical habitats worldwide, often forming extensive, radiating clusters that enhance spore dispersal. Notable examples include Coprinellus micaceus, the common fairy inkcap, which is widespread on buried wood or grassy areas, produces small but edible fruitbodies (though of limited culinary value due to bitterness and size), and features prominent mica-like granules on its honey-brown cap.^[64] Another representative is Coprinellus radians, recognized for its dense, radiating troops on soil or wood, with elliptical spores around 7 × 4 µm and a smooth, brownish cap lacking heavy veiling.^[65] Coprinellus domesticus, or firerug inkcap, stands out for its taller, slender stature (stipe 4–10 cm), grayish-brown bell-shaped cap up to 4 cm, and striking orange mycelial mat on substrates like burned wood or litter.^[66] Phylogenetic analyses, based on multi-locus data including ITS, LSU, and β-tubulin sequences, divide Coprinellus into major clades such as Domestici/Micacei (with veiled species) and Setulosi (setulose or haired forms), though sections are not always monophyletic.^[62] Recent studies in Asia, particularly a 2024 investigation of the "fairy inkcap" complex in China, have highlighted substantial undescribed diversity, identifying nine lineages across 17 provinces, including four new species (C. aureodisseminatus, C. austrodisseminatus, C. parcus, C. velutipes) and a new subspecies (C. disseminatus subsp. orientalis), with high genetic divergence (Fst > 0.7) underscoring the region's subtropical richness.^[67] These findings, drawn from 74 collections spanning 1998–2023, emphasize Coprinellus's evolutionary adaptability in Asian ecosystems.^[67]

Parasola

Parasola is a genus of small, veil-less coprinoid mushrooms in the family Psathyrellaceae, comprising approximately 30–40 recognized species worldwide, though taxonomic revisions continue to refine this number. These minute agarics superficially resemble diminutive Coprinus species but are distinguished by partial deliquescence of the gills rather than full auto-digestion of the fruitbody. They are saprotrophic fungi that decompose organic matter and are commonly encountered in grasslands and lawns.^[68]^[69]^[70] Morphologically, Parasola species feature caps typically less than 2 cm in diameter, convex to subcylindrical when young, often pale brown to ochraceous, and translucent-striate due to their hygrophanous tissue, which reveals the underlying gills as radiating lines. The gills are free or nearly so, sometimes with auriculate attachment (a small ear-like connection to the stipe), crowded and ventricose, starting whitish and turning grayish-black as they mature and partially dissolve. Basidiospores measure 7–10 µm (varying by species, e.g., 7.0–10.8 × 5.8–9.8 µm in some), dark reddish-brown, often angular (ovoid to heart-shaped) with a central or eccentric germ pore and a characteristic plage—a darker spot near the hilum.^[68]^[69] These fungi thrive in urban and suburban environments, including lawns, paths, and disturbed grasslands, as well as coprophilous (dung-associated) or terricolous (soil-inhabiting) habitats, contributing to nutrient cycling in grassy areas. They are generally considered non-edible due to their small size and fragile nature. Representative species include Parasola auricoma, noted for its golden to orange-brown cap with a central umbo and long reddish hairs, often found in woodchip mulch or grassy edges; and Parasola conopilea, characterized by a conical, reddish-hygrophanous cap and yellowish stipe, commonly reported across Europe on lawns and meadows.^[68]^[71]^[69] Recent taxonomic studies have expanded the genus, with additions from 2023–2025 including Parasola papillatospora, a saprotroph from soil in temperate deciduous forests of Europe (e.g., under oaks and beeches), distinguished by its papillate-apex spores and variable shapes; and eight new species from China, highlighting ongoing diversity in Asia through morphological and molecular analyses. These updates underscore Parasola's global distribution and adaptability to varied microhabitats.^[69]^[70]

Other Genera

In addition to the core genera Psathyrella, Coprinopsis, Coprinellus, and Parasola, the Psathyrellaceae encompasses approximately 20 genera overall, many resulting from segregations of the historically polyphyletic Psathyrella, contributing to a family diversity of nearly 2000 species. These minor genera highlight the family's morphological and ecological variability, often featuring specialized traits that distinguish them from the typical agaricoid forms dominant in the major genera. Among these, Candolleomyces represents a key segregate, characterized by the absence of pleurocystidia and dark-spored basidiomata typically found in soil as saprobes; recent discoveries include two new species from Pakistan described in 2025, further emphasizing their role as terrestrial decomposers in Asian ecosystems.^[72] Lacrymaria stands out for its lacrimose gills, which exude tear-like droplets of dark liquid when moist, a feature linked to its hygroscopic spores and aiding in spore dispersal.^[73] Ephemerocybe exemplifies short-lived fruiting bodies, with 2025 phylogenetic notes clarifying its position and underscoring the ephemeral nature of its agaricoid structures in damp habitats.^[25] Other notable minor genera include Gasteroagaricoides, which exhibits secotioid, puffball-like basidiomata with enclosed gills and powdery spore masses, diverging from the open-hymenophore typical of agaricoid members. Olotia, a monotypic genus, features robust, pigmented cystidia and is adapted to specialized microhabitats, contributing to the family's underrepresented diversity in non-agaricoid forms. A 2025 study in Phytotaxa introduced the new genus Cantonopsathyra with two species from southern China, expanding Asian representation through phylogenetic analyses of psathyrelloid clades.^[25] The significance of these minor genera lies in their revelation of ongoing taxonomic challenges, as molecular data continue to uncover unresolved paraphyly within smaller groups, prompting further refinements to family boundaries and emphasizing the need for integrated morphological and genetic approaches.

References

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Taxonomy browser (Psathyrellaceae) - NCBI - NIH
Taxonomy ID: 184208 (for references in articles please use NCBI:txid184208). current name. Psathyrellaceae. NCBI BLAST name: basidiomycete fungi. Rank: family
[2]
[PDF] Molecular Phylogenetics and Evolution
These nodes represent ancestors of the major genera recognised in Psathyrellaceae, except for Node 5. (=Node 1 plus Node 4) and Node 6 (root of Psathyrellaceae) ...
[3]
The mushroom family Psathyrellaceae: Evidence for large-scale ...
The genus Psathyrella (Fries) Quélet consists of 400–600 species of mushrooms (Hawksworth et al., 1995, Kirk et al., 2001, Vašutová, 2006). Species of ...
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[PDF] A new genus and four new species in the /Psathyrella s.l. clade from ...
May 26, 2021 · The species in the family Psathyrellaceae can be roughly divided into two types by mac- romorphology: psathyrelloid and coprinoid.
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Iugisporipsathyra reticulopilea gen. et sp. nov. (Agaricales ...
Jun 22, 2022 · Species of Psathyrellaceae are cosmopolitan and often grow on decaying logs, woody debris, humus or soil, in woodlands, lawns or bogs and can ...
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[9]
PSATHYRELLA in the Pacific Northwest
Flesh is fragile. Gills are attached and not decurrent. The stem, typically whitish in color, is slender and fragile, often snapping evenly and cleanly in half.
[10]
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
May 16, 2019 · Macro-morphological characteristics of the Psathyrella basidiomes: scales of velar origin on pilei tops and margins, and beige-coloured gills ( ...
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[PDF] Two new coprophilous varieties of Panaeolus (Psathyrellaceae ...
Jun 28, 2013 · (Psathyrellaceae, Agaricales) from Punjab, India. Mycosphere 4(3) ... Taxonomic descriptions. Panaeolus africanus var. diversistipus ...<|separator|>
[12]
[PDF] PDF - Fungal Systematics and Evolution
Jan 25, 2019 · Cresyl blue and Cotton blue were utilised to highlight the ortho-/metachromatic reactions in the spores. Dried fragments were rehydrated in ...
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The Genus Psathyrella (MushroomExpert.Com)
... genus (see the reference list below). Species Pages. Psathyrella bipellis · Psathyrella candolleana · Parasola conopilea · Psathyrella cystidiosa · Psathyrella ...
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A new genus and four new species in the /Psathyrella s.l. clade from ...
May 26, 2021 · Discussion. The species in the family Psathyrellaceae can be roughly divided into two types by macromorphology: psathyrelloid and coprinoid.
[15]
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Apr 13, 2018 · It is characteristic of fragile basidiomata, hygrophanous pileus, brown to black brown spore print, always present cheilocystidia and ...
[16]
(PDF) Species Diversity of the Genus Psilocybe (Basidiomycotina ...
Aug 10, 2025 · Moreover, 53 species of Psathyrella, many times related with Psilocybe ... Morphologically, Psilocybe differs from Deconica by having blackish ...
[17]
The Agaricales in Modern Taxonomy - Rolf Singer - Google Books
Bibliographic information ; Title, The Agaricales in Modern Taxonomy ; Author, Rolf Singer ; Edition, 4, illustrated ; Publisher, Koeltz Scientific Books, 1986.
[18]
Coprinus Pers. and the disposition of Coprinus species sensu lato
Feb 1, 2001 · Redhead, S. A., Vilgalys, R., Moncalvo, J.-M., Johnson, J. & Hopple, J. S., Jr.: Coprinus Pers. and the disposition of Coprinus species ...
[19]
Understanding the Evolutionary Processes of Fungal Fruiting Bodies
Uncertainties in topology and support values have been attributed to the undersampling of Psathyrella taxa (Redhead et al. 2001, Vasutova et al. 2008). Our ...
[20]
Psathyrellaceae - GBIF
Classification ; kingdom; Fungi ; phylum; Basidiomycota ; class; Agaricomycetes ; order; Agaricales ; family; Psathyrellaceae ...<|control11|><|separator|>
[21]
Psathyrellaceae - Wikipedia
The Psathyrellaceae are a family of dark-spored agarics that generally have rather soft, fragile fruiting bodies, and are characterized by black, dark brown ...
[22]
Psathyrella - Mycobank
Fungi > Dikarya > Basidiomycota > Agaricomycotina > Agaricomycetes > Agaricomycetidae > Agaricales > Agaricineae > Psathyrellaceae > Psathyrella. Synonyms.
[23]
Introducing one new genus and two new species of Psathyrellaceae ...
May 22, 2025 · Introducing one new genus and two new species of Psathyrellaceae (Basidiomycota, Agaricales), with notes on Candolleomyces and Ephemerocybe.
[24]
Candolleomyces yunnanensis (Psathyrellaceae, Agaricales), a ...
Jul 25, 2025 · Candolleomyces yunnanensis is a novel species from Yunnan Province, China, belonging to the Psathyrellaceae family. It is a new species with ...<|control11|><|separator|>
[25]
Names Record - Index Fungorum
Record Details: Psathyrellaceae Vilgalys, Moncalvo & Redhead, in Redhead, Vilgalys, Moncalvo, Johnson & Hopple, Taxon 50(1): 226 (2001).
[26]
Phylogenetic studies on the genus Candolleomyces ... - MycoKeys
Jan 16, 2025 · Wächter and Melzer (2020) established the genus Candolleomyces D. Wächt. & A. Melzer (family Psathyrellaceae Locq.) to accommodate 25 species ...
[27]
Molecular phylogenetics and taxonomy in Psathyrellaceae ...
Apr 11, 2015 · Psathyrella is a large genus in Agaricales with 1,004 records in Index Fungorum (www.indexfungorum.org) including synonyms, varieties, forms, ...
[28]
Morphology and multi-locus phylogeny reveal four new ... - MycoKeys
Aug 28, 2025 · The family Psathyrellaceae Vilgalys, Moncalvo & Redhead was established by Redhead et al. ... Species within the family have a cosmopolitan ...
[29]
Morphological characterization of clades of the Psathyrellaceae ...
Aug 7, 2025 · ... Its young fruiting bodies have white to pink free lamellae that turn dark brown at maturity, a stipe with an annulus, and creates brown ...
[30]
(PDF) Studies in the Panaeolus papilionaceus complex (Agaricales ...
Apr 9, 2024 · Studies in the Panaeolus papilionaceus complex (Agaricales, Galeropsidaceae): two new species discovered in the Dominican Republic and Australia ...
[31]
Molecular Phylogeny and Morphology Reveal Four New Species of ...
This study investigates Conocybe (Bolbitiaceae, Agaricales) on the Plateau, based on specimens collected over the past decade, using morphological and molecular ...Missing: differences | Show results with:differences
[32]
Transcriptomics Reveals the Putative Mycoparasitic Strategy of ... - NIH
Psathyrella epimyces causes the deformation of sporocarp tissue of ... More broadly, other species within the Entolomataceae are suspected mycoparasites ...
[33]
[PDF] Psathyrella (Psathyrellaceae, Agaricales) species collected on dung ...
Jun 29, 2015 · The genus Psathyrella (Fr.) Quél., belonging to the family Psathyrellaceae Readhead,. Vilgalys & Hopple, is characterized by grayish, pale ...
[34]
A Thirty-Year Survey Reveals That Ecosystem Function of Fungi ...
Nov 27, 2012 · Mushroom fruiting, the reproduction of fungi, has broad implications for forest health, terrestrial biomass turnover, and global carbon ...<|control11|><|separator|>
[35]
Life History and Developmental Processes in the Basidiomycete ...
Two main types of mycelia are distinguished in its life cycle, the infertile monokaryon and the fertile dikaryon (71, 176).<|separator|>
[36]
[PDF] Fungus Consumption (Mycophagy) by Small Animals
Such fungi depend on mammal and invertebrate mycophagy for spore dispersal. These and other fungi, in turn, are important in the diet of many animals and in ...
[37]
Phylogenetic studies in Psathyrella focusing on sections Pennatae ...
The basidiomycete genus Psathyrella (Psathyrellaceae, Agaricales) ... (Redhead et al. 2001). These three genera are included in the Psathyrellaceae ...
[38]
(PDF) A New Tropical Species of Psathyrella (Psathyrellaceae ...
PDF | Basidiocarps of a hitherto undescribed Psathyrella species were collected during a sampling excursion in North Vietnam. They grew on deciduous.
[39]
(PDF) Two new species of the Coprinopsis alnivora ...
Aug 5, 2025 · Combined morphological and phylogenetic studies of collections of the family Psathyrellaceae from Australia have revealed two novelties in ...Missing: hybridization | Show results with:hybridization
[40]
A New Species of Psathyrella (Agaricales, Coprinaceae) from ... - jstor
dark brown or reddish-brown in Melzer's reagent, wall 0.3-0.4 Am thick, with a hyaline subcylindric pore and a well-developed hilar appendix. Basidia 20-25 ...
[41]
Psathyrellaceae - Burke Herbarium Image Collection
Psathyrellaceae. 8 genera 102 species. Show only taxa with photos. Order by: Scientific name. Common name. Display as: Index to genera: Coprinellus, Coprinopsis ...Missing: list | Show results with:list
[42]
Psathyrella corrugis - California Fungi - MykoWeb
Psathyrella corrugis is recognized by a brown hygrophanous cap (striate when fresh) that soon fades to pale buff-brown, the disc usually dingy yellowish-brown.Missing: macroscopic features texture
[43]
Genera and species within the Psathyrellaceae - iNaturalist
Aug 4, 2016 · The same data led to the recognition of a number of related genera within the psathyrellaceae: Coprinellus, Coprinopsis and Parasola. These are ...
[44]
Psathyrella cystidiosa - Rare Species Guide - Minnesota DNR
Both were found in deciduous forests. The mushroom occurs in shade and may have a preference for thoroughly moistened areas prior to fruiting. Biology / Life ...
[45]
Altitudinal upwards shifts in fungal fruiting in the Alps - Journals
Dec 24, 2019 · Here, we take advantage of two national datasets of fungal fruit body records from Switzerland and Austria, to explore altitudinal distributions ...
[46]
Psathyrella atlantica
Habitat and Ecology Psathyrella atlantica is a saprotrophic species, growing in litter deposited on soil. This species forms caespitose basidiomata composed of ...
[47]
Updated Distribution of Endangered Psathyrella ammophila in Poland
Mar 1, 2021 · The greatest losses of such habitats have been observed in Southern Europe, which is connected with the development of tourism and strong urban ...
[48]
Psathyrella cystidiosa - The Global Fungal Red List Initiative
Apr 23, 2016 · The species is found in moist soil or attached to dead branches of deciduous trees. A search for this mushroom in the same location conducted ...Missing: cycle | Show results with:cycle
[49]
(PDF) Diversity of genus Psathyrella (Psathyrellaceae, Agaricales ...
Jan 4, 2021 · In this paper,. 17 collections belonging to 14 species of genus Psathyrella has been worked out for their external,. internal details and key ...
[50]
Psathyrella candolleana, Pale Brittlestem mushroom - First Nature
This mushroom occurs in all kinds of woodlands and woodland clearings as well as on timber buried in damp grassland.
[51]
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.
[52]
Coprinus Pers. and the Disposition of Coprinus Species sensu lato
Aug 9, 2025 · The family Psathyrellaceae Vilgalys, Moncalvo & Redhead was established by Redhead et al. (2001) , with Psathyrella (Fr.) Quél. as the ...
[53]
Two new psathyrelloid species of Coprinopsis (Agaricales ...
Sep 8, 2021 · The amplification primers of LSU nrDNA ( LSU ) were LROR and LR5 ( Vilgalys and Hester 1990 ), the ITS nrDNA ( ITS ) regions were ITS1 and ITS4 ...Materials And Methods · Results · Coprinopsis Pusilla G. Rao...
[54]
[PDF] Taxonomic study on coprophilous species of Coprinopsis ...
Jan 18, 2014 · Redhead, Vilgalys and Moncalvo in Taxon 50(1): 229, 2001. Figs. 5 – 6. Basionym – Agaricus lagopus Fries in Systema Mycologicum 1: 312, 1821 ...
[55]
Coprinopsis atramentaria (Common Ink Cap) - MycoAnatomical
Apr 19, 2024 · HABITAT: Growing terrestrially or directly on substrate, typically stumps, in either urban or wooded environments as scattered to dense clumps ...
[56]
Alcohol intolerance with Coprine-like syndrome after consumption of ...
Alcohol intolerance after consumption of wild mushrooms is a recognized phenomenon. This is best understood with Coprinus atramentarius.Missing: Coprinopsis atramentaria source
[57]
Full article: Phylogenetic analyses of Coprinopsis sections Lanatuli ...
Abstract: Sections Lanatuli and Atramentarii of the genus Coprinopsis contain some of the best known and most important agaric species, including C. cinerea and ...<|control11|><|separator|>
[58]
[PDF] Third report of coprinoid fungi (Psathyrellaceae, Agaricales) in the ...
Feb 22, 2025 · All the species in that clade are characterized by three-dimensional spores or, when the widths in the two views barely differ, by the distinct ...
[59]
Phylogeny and species delimitation in the genus Coprinellus with ...
... Redhead et al. 2001). These include several taxa of the cortinati group ... Psathyrellaceae, which has been referred to as coprinoidization (CitationNagy et al.Taxonomy · Discussion · C. Bisporus Clade<|control11|><|separator|>
[60]
The genus Coprinellus (Basidiomycota; Agaricales) in Pakistan with ...
Sep 11, 2018 · The genus Coprinellus is one of the important lineages of coprinoid mushroom in the family Psathyrellaceae. Species-level taxonomy in ...
[61]
Coprinellus micaceus (MushroomExpert.Com)
Microscopic Features: Spores 7-11 x 4-7 µ; subelliptical to mitriform; smooth; with a central pore. Basidia 4-spored; surrounded by 3-6 brachybasidia.
[62]
[PDF] Coprinellus radians (Desm.)Vilg. Hopple & Johnson
Mar 7, 2018 · Spores: basidiospores. 7 x 4 µm,, elliptical, smooth, dark brown. Diagnostic macroscopic feature: this is one of several inky caps that ...
[63]
Coprinellus domesticus: The Retro Inky (MushroomExpert.Com)
Stem: 4-10 cm long; up to 1 cm thick; equal, with a slightly swollen base; smooth; white; hollow; sometimes with a volva-like rim at the base; usually arising ...
[64]
Species clarification of fairy inkcap (“Coprinellus disseminatus”) in ...
in 1994 and 1999, Redhead et al. (2001) confirmed the taxonomic status of the genus Coprinellus, which includes C. disseminatus and C. micaceus (Bulliard) ...
[65]
https://www.herbarium.iastate.edu/files/fungi/Coprinellus%2520radians%2520%28Desm.%29Vilg.%2520Hopple%2520&%2520Johnson.pdf
[66]
https://www.mushroomexpert.com/coprinellus_domesticus.html
[67]
Emendation of morphology and infrageneric standards of Parasola ...
May 30, 2025 · Context extremely thin, brown in center and almost unseen at margin, odor and taste not distinctive. Lamellae crowded, free and remote from ...
[68]
California Fungi: Parasola conopilea - MykoWeb
Parasola conopilea can be distinguished by its conical, reddish (when young) hygrophanous cap and yellowish stipe.
[69]
Two novel species of Candolleomyces (Psathyrellaceae ... - Phytotaxa
Mar 7, 2025 · Detailed description of the two new species along with field photographs, illustrations and molecular phylogenetic analyses are provided.Missing: Ephemerocybe | Show results with:Ephemerocybe
[70]
Lacrymaria lacrymabunda, Weeping Widow mushroom - First Nature
Lacrymaria lacrymabunda (Bull.) Pat. - Weeping Widow. Phylum: Basidiomycota - Class: Agaricomycetes - Order: Agaricales - Family: Psathyrellaceae.