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Panellus, Bio-luminescent (Panellus stipticus)

Panellus stipticus Species Guide

Panellus stipticus

Panellus stipticus is a small, fan-shaped shelf fungus native to dead hardwood logs across the northern temperate world, bioluminescent in its eastern North American form. Its bioluminescence is produced by a self-recycling chemical system that has operated continuously for 160 million years, since the Jurassic period. The glow comes from the gills, the mycelium, and the spores themselves, and requires only oxygen to sustain.

Panellus stipticus (Bull.) P. Karst., 1879 — Family: Mycenaceae — Order: Agaricales

Species Panellus stipticus
Family / Order Mycenaceae / Agaricales
Type Saprotrophic white-rot fungus
Bioluminescence Eastern North America only
Range Northern temperate zone
Season Sept–Nov; year-round in SE US

Panellus stipticus, known as the bitter oyster, luminescent panellus, and astringent panus depending on where you are in the world, is among the most scientifically remarkable fungi in the northern temperate zone. Despite its small size and total inedibility, it is the subject of over a century of bioluminescence research, two sequenced genomes, and a 2025 study that definitively resolved why some populations glow while others, morphologically identical, do not. Panellus stipticus is cultivable, persistent, and genuinely bizarre — an ideal species for anyone interested in observing living bioluminescence firsthand.

What Is Panellus stipticus?

Panellus stipticus is a saprotrophic white-rot fungus — meaning it decomposes dead wood by enzymatically dismantling both lignin (the structural polymer that gives wood its rigidity) and cellulose. White-rot fungi are the only organisms known to achieve complete lignin mineralization; without them, the forest floor would accumulate undecomposed wood indefinitely. P. stipticus is particularly associated with recently fallen hardwood logs and stumps, where it functions as an early-succession decomposer, establishing before longer-lived species outcompete it.

The species carries four recognized English common names, with usage split by geography: "bitter oyster" and "luminescent panellus" are most common in North America; "bitter oysterling" is the preferred British name; "astringent panus" appears in older mycological literature. None of these names dominates globally, which is why the scientific name Panellus stipticus is the universal identifier. The specific epithet comes from the Greek styptikós meaning astringent — a reference to its traditional use in China as a hemostatic (blood-stopping) agent and to the harsh, mouth-drying bitterness of fresh specimens.

The most interesting fact about Panellus stipticus: Morphologically identical individuals of the same species glow green in the dark in eastern North America — but are completely dark in Europe, Asia, and western North America. A 2025 genomic study confirmed that this reflects the presence or complete absence of a 10-kilobase luciferase gene cluster. Luminescent and non-luminescent strains are the same biological species and remain interfertile. Why luminescence was retained in some populations and lost in others, and whether there is any selective pressure involved, remains an open scientific question.

The fruit bodies of Panellus stipticus are small, fan-shaped, and tough. They emerge from hardwood logs in overlapping clusters that persist through wet and dry cycles — drying to near-paper hardness in drought and reviving fully when rain returns. This desiccation tolerance is unusual among gilled mushrooms and makes the species detectable year-round. The gills are crowded and connected by numerous cross-veins, the cap surface is velvety and buffish-tan, and the taste is acutely bitter and astringent, with some geographic populations causing notable throat constriction and nausea. The species is inedible everywhere it grows.

Interested in this species? Out-Grow carries a liquid culture.

Panellus stipticus Liquid Culture

How Is Panellus stipticus Classified?

Panellus stipticus was first described by French botanist Jean Baptiste Bulliard in 1783 as Agaricus stypticus. The current combination was established by Finnish mycologist Petter Karsten in 1879. The species is the type species of the genus Panellus — the species against which all other Panellus species are compared taxonomically. One recurring spelling variant, Panellus stypticus (with a y), appears in older literature; the accepted spelling per Index Fungorum is stipticus with an i.

Family placement has been revised significantly. P. stipticus was placed in Tricholomataceae for most of the twentieth century — a large wastebasket family for white-spored gilled fungi. Multi-gene phylogenetic analysis subsequently showed a close relationship between Panellus, Dictyopanus, and the core Mycena clade. The current consensus, reflected in Index Fungorum, GBIF, and multiple phylogenetic publications, is placement in Mycenaceae. The California Fungi database (MykoWeb) notes that recent multi-gene phylogenies suggest P. stipticus may eventually be transferred to genus Mycena itself — a discussion ongoing at the time of writing.

Rank Name
Kingdom Fungi
Phylum Basidiomycota
Class Agaricomycetes
Subclass Agaricomycetidae
Order Agaricales
Suborder Marasmiineae
Family Mycenaceae
Genus Panellus P. Karst., 1879
Species Panellus stipticus (Bull.) P. Karst., 1879

The extensive synonym list for Panellus stipticus reflects its unusual pleurotoid morphology — a laterally-attached, shelf-like growth form that successive authors assigned to different genera before the current placement was stabilized. Early synonyms include Agaricus stypticus Bull. (1783), Crepidopus stipticus Gray (1821), Pleurotus stipticus P. Kumm., Panus stipticus (Bull.) Fr., and Lentinus stipticus (Bull.) J. Schröt. (1885). The Index Fungorum LSID is urn:lsid:indexfungorum.org:names:355858.

Common online error: Several websites classify Panellus stipticus in Panellaceae — this is incorrect and reflects outdated or unsupported taxonomy. The current consensus classification is Mycenaceae. Some sites also incorrectly describe the species as causing brown rot — it causes white rot, a critical distinction for understanding what substrates it can colonize.

How Do You Identify Panellus stipticus?

In eastern North America, the bioluminescence of Panellus stipticus is its most distinctive field characteristic — though it requires at least five minutes of dark adaptation to observe, and camera long-exposure settings to photograph reliably. In regions where the species does not glow (Europe, Asia, Pacific coast), identification relies on a combination of morphological features.

Macroscopic Features

Cap 5–32 mm; fan- or kidney-shaped; dry, velvety; buff to ochre-tan; margin inrolled when young
Gills Adnate to decurrent; crowded; narrow; cross-veined; pale golden-tan; 40–70 complete gills
Stem Up to 5–20 mm long; lateral or eccentric; velvety; may be absent; pale tan to rusty brown
Flesh Thin, tough, leathery; whitish to pale brown; no distinctive odor; bitter/astringent taste
Spore Print White
Substrate Dead hardwood logs and stumps; occasionally conifer; in overlapping clusters

A critical identification feature is the distinctive reticulate (net-like) pattern on the gill surface caused by numerous cross-veins connecting adjacent gills. This cross-venation, combined with the white spore print, lateral attachment, and velvety cap surface, separates P. stipticus from most lookalikes. The bitter taste is reliable and immediate. The cap surface can develop cracked or areolate (broken into irregular sections) texture with age, and may show faint concentric zoning reflecting seasonal growth pulses. Dried specimens bleach to pale tan but revive and regain color and texture when moistened — a behavior unusual for gilled mushrooms.

Microscopic Features

Spores of Panellus stipticus measure 2.5–4 × 1.5–2 µm (per MushroomExpert), are ellipsoid to slightly sausage-shaped, smooth-walled, hyaline in KOH solution, and amyloid — they react positively to Melzer's reagent, a dye that turns blue-black in the presence of starch-like polysaccharides in cell walls. This amyloid reaction is described as weak and should be confirmed carefully if used as an identification criterion. Basidia measure 15–25 × 2–3.5 µm, are clavate (club-shaped), four-sterigmate, and have clamp connections at their base. Cheilocystidia (sterile cells on the gill edge) are cylindric to filiform and often diverticulate — meaning they bear small finger-like projections at the apex. Clamp connections are present throughout all tissues.

Lookalike Species

Crepidotus mollis

The most frequent confusion species. Differentiated by brown spore print (versus white in P. stipticus), lacks bioluminescence, and gills are not connected by cross-veins. Cap surface is often slightly gelatinous when fresh.

Schizophyllum commune

White to gray, densely hairy cap that superficially resembles P. stipticus in pale dried specimens. Easily separated: S. commune has split gill-folds rather than true gills, and its gills appear to split lengthwise. No bioluminescence.

Panus rudis

Distinctly larger; cap reddish-brown fading to pinkish-tan with lilac tinges when young. No bioluminescence. White spore print and lateral attachment are shared, but the larger size and lilac coloration distinguish it.

Panellus mitus and P. pusilis

Closely related Panellus species with similar morphology. P. mitus is a rare West Coast species; P. pusilis is distributed in Florida. Neither is bioluminescent. Definitive differentiation may require microscopy.

Where Does Panellus stipticus Grow?

Panellus stipticus is a common species across the northern temperate zone, found on the dead wood of a wide range of hardwood trees. It grows on logs, stumps, fallen branches, and occasionally on wounds in living trees. As an early-succession decomposer, it is most commonly associated with recently dead wood and is not typically recorded from timber stands more than 20 years old.

Region Status Preferred Hosts
Eastern North America Common; bioluminescent Oak, birch, maple, hickory, ash, American hornbeam
Pacific Coast North America Present; non-bioluminescent Hardwoods; occasionally conifers (pine)
Europe Common; non-bioluminescent Oak, birch, alder, beech, hazel, chestnut, ash
Asia (Japan, Korea, China) Present; non-bioluminescent Various hardwoods
Australasia Present; non-bioluminescent Various hardwoods

Primary fruiting season in Europe and eastern North America is September through November. Secondary fruiting can occur in spring. Because the fruit bodies are exceptionally desiccation-tolerant — drying hard and reviving after rain — P. stipticus can be found year-round, with fresh fruiting in the southeastern US continuing through winter. Slug predation on fruit bodies is common, and slugs may serve as spore dispersal agents. The species holds no conservation concern anywhere in its range.

Can You Cultivate Panellus stipticus?

Panellus stipticus is fully cultivable on hardwood substrates. It will colonize wood-based materials, produce visible bioluminescent mycelium, and develop recognizable fruiting bodies under appropriate conditions. It is not cultivated for food — the mushrooms are inedible — but is actively grown as a novelty organism for observing its bioluminescence. For mycology hobbyists, a colonized block or log in a dark room provides one of the most visually striking displays in the fungal world.

Important distinction: Liquid culture mycelium in solution does not luminesce. Bioluminescence in Panellus stipticus requires growth on a solid substratum. Luminescence ceases within approximately one hour of anaerobic (oxygen-deprived) conditions. Once transferred from liquid culture to agar or hardwood substrate, the mycelium will begin to luminesce as it colonizes the surface — provided the culture is from a bioluminescent eastern North American strain.

Substrate

Hardwood Sawdust Oak, beech, or maple; preferred for fruiting body production
Hardwood Logs Plug spawn method; most natural fruiting; 6–12+ month colonization timeline
Wood Chips Suitable; less consistent than sawdust blocks
Grain (rye, wheat) Mycelium colonizes grain but fruiting bodies abort or develop abnormally; not recommended for fruiting
Agar Media Breadcrumb agar (BCA) outperforms MEA for bioluminescence intensity; lower malt concentration better for long-term luminescence
Avoid Activated charcoal in media — reduces luminescence by 22–42% and suppresses colony size

Panellus stipticus is a wood decomposer by biology — its enzyme toolkit is optimized for lignin and cellulose in hardwood. Grain alone cannot support normal development because the species lacks the substrate cues and nutrient balance it needs to form fruiting bodies. Log cultivation under natural outdoor conditions has the advantage of lower contamination risk because the outdoor microbial community provides natural competitive exclusion.

Cultivation Parameters

1

Substrate Preparation

Sterilize hardwood sawdust blocks or prepare freshly cut hardwood logs (4–6 inch diameter preferred). Sawdust blocks require autoclave sterilization; logs can be inoculated directly with plug spawn after cutting.

2

Inoculation

Use liquid culture to inoculate grain spawn, which is then transferred to hardwood substrate. Alternatively, inoculate agar plates directly for bioluminescence observation without fruiting.

3

Colonization

Incubate at 65–80°F (18–27°C) in complete darkness. Dark incubation produces denser growth and maximizes luminescence. Colonization is slow: expect 3–5 weeks for grain spawn, months for logs.

4

Fruiting Trigger

Lower temperature to 60–75°F (15–24°C). Increase fresh air exchange. Maintain high humidity (85–95% RH). On logs, autumn outdoor conditions naturally trigger fruiting. No cold shock protocol is established.

5

Observing the Glow

Allow 5+ minutes of complete dark adaptation before viewing. Camera long-exposure settings are needed to photograph the glow. Bioluminescence is maximized at 22°C and pH 3–3.5 — conditions naturally occurring in weathered hardwood.

6

Culture Maintenance

Store colonized agar plates at 35–43°F (2–6°C), sealed to prevent desiccation. Transfer to fresh media every 1–2 months. Cultures maintained too long on PDA show reduced growth vigor; BCA is preferred for maintaining luminescence intensity.

Contamination Risks

Panellus stipticus is significantly slower to colonize substrate than most cultivated fungi, creating a high contamination window. Trichoderma spp. (a family of aggressive green mold contaminants) and bacterial wet rot are the primary risks during the extended colonization period. Strict sterile technique is essential for grain spawn production. Log cultivation outdoors carries lower risk because competitive fungal communities naturally exclude many lab contaminants.

What the Out-Grow Liquid Culture Contains & How to Use It

Out-Grow's Panellus stipticus liquid culture is a bioluminescent eastern North American strain, selected for its luminescence capability. The 10cc syringe contains actively growing mycelium suspended in nutrient solution. Use it to inoculate sterilized grain jars for spawn production, which can then be transferred to hardwood sawdust blocks or log inoculation plugs for bioluminescent display. You can also inoculate agar plates directly — breadcrumb agar is recommended over standard MEA for maximizing luminescence intensity. Store in a cool, dark location between 35–43°F (2–6°C). Mycelium in liquid culture will not glow; luminescence begins once the mycelium establishes on solid substrate and is exposed to oxygen.

What Compounds Does Panellus stipticus Contain?

The chemistry of Panellus stipticus is dominated by its bioluminescence system, which is the best-characterized aspect of the species. Its broader bioactive chemistry — antioxidants, antimicrobials, cytotoxic compounds — has not been studied; this is an explicitly open research gap.

The Bioluminescence Pathway

The biochemical mechanism underlying the glow in Panellus stipticus was fully decoded by Oliveira, Yampolsky et al. in 2018 (PNAS 115:1415). It is a four-enzyme recycling cycle based on the phenylpropanoid metabolite caffeic acid — a compound widespread in plants and fungi. The four enzymes are encoded by genes designated luz (luciferase), hisps (hispidin synthase), h3h (hispidin-3-hydroxylase), and cph (caffeoylpyruvate hydrolase):

1. Caffeic acid → hispidin (via hispidin synthase, HispS)
2. Hispidin → 3-hydroxyhispidin (the fungal luciferin) via H3H
3. 3-hydroxyhispidin + O&sub2 → oxidized by luciferase (Luz) → caffeylpyruvic acid + green light at ~525 nm
4. Caffeylpyruvic acid → recycled to caffeic acid via CPH

The system is self-sustaining as long as oxygen is available — caffeic acid is regenerated after each light-emission event. This makes fungal bioluminescence energetically distinct from firefly bioluminescence, which consumes luciferin irreversibly.

Panal (sesquiterpene)

Isolated from P. stipticus fruiting bodies in 1988 by Shimomura et al. A cadinene keto-aldehyde sesquiterpene. Originally investigated as a potential luciferin precursor. Its precise relationship to the hispidin pathway identified in 2018 has not been fully reconciled in published literature — an open research question.

Species-specific

PS-A & PS-B

1-O-decanoylpanal and 1-O-dodecanoylpanal; acyl derivatives of panal characterized in 1993. Initially treated as luciferin precursors; the relationship between these compounds and the 2018-identified hispidin pathway is unresolved.

Species-specific

Luciferase gene cluster

~10 kb cluster encoding luz, hisps, and h3h genes. Present in bioluminescent eastern NA strains; completely absent from non-bioluminescent Korean strain genome (KUC8834). Phylogenetically conserved across mycenoid lineage. Emission maximum at 525 nm.

Genomic

Laccase

Copper-containing phenoloxidase; primary enzyme for lignin ring cleavage in white-rot fungi. Confirmed in P. stipticus. Catalyzes hydroxylation of phenolic compounds. Relevant to bioremediation applications.

Enzymatic assay

Phenolic / astringent compounds

Responsible for the bitter, mouth-drying taste. Specific compounds have not been identified by GC-MS or other analytical methods. This is an explicitly open research gap — no published study has characterized the volatile or taste compounds of P. stipticus.

Uncharacterized

Superoxide dismutase (SOD)

SOD activity in the fruit body modulates luminescence by quenching superoxide anion (O&sub2−), a molecule involved in the bioluminescent reaction. Higher SOD activity correlates with reduced luminescence intensity.

Biochemical

Bioremediation Activity

The laccase and related white-rot enzyme system of Panellus stipticus has been investigated in two bioremediation contexts. Liquid culture mycelium reduced phenolic concentration in olive mill wastewater by 42% after 31 days of incubation. In a separate screening study of approximately 1,500 basidiomycete strains for dioxin degradation capacity, P. stipticus strain 99-334 was selected as an effective 2,7-dichlorodibenzo-p-dioxin (2,7-DCDD) degrading strain. Both findings are from single laboratory-scale studies; neither has been developed beyond the preliminary stage. These represent the only published bioactivity data for this species.

Evidence note: Panellus stipticus has no published in vitro cytotoxicity, antimicrobial, antioxidant, or anti-inflammatory studies. No human clinical trials exist. There are no marketed supplements derived from this species. Any claims about medicinal properties beyond the traditional hemostatic use would be extrapolation from unrelated species and should be treated accordingly.

Is Panellus stipticus Safe to Handle?

Panellus stipticus is consistently described as inedible rather than toxic in standard mycological references. No confirmed alkaloids, amatoxins, or recognized mycotoxins have been reported. The species deters consumption effectively through its intense bitterness and leathery texture rather than through systemic toxicity. No contact dermatitis, skin irritation, or respiratory hazard has been reported from handling.

An honest safety caveat: at least one mycology resource notes reports of nausea and vomiting following consumption of specimens from certain geographic areas, attributing these symptoms to the harsh astringent compounds rather than to confirmed toxins. This is consistent with the documented geographic variation in taste intensity — populations outside North America (particularly in Japan, New Zealand, and Russia) produce more pronounced throat constriction effects. The species has never been part of any culinary tradition; the absence of documented poisoning cases primarily reflects the absence of consumption rather than an established safety profile. The species should be treated as inedible.

What Makes Panellus stipticus Remarkable?

Several features of Panellus stipticus biology are genuinely unusual, some with no parallel in any other organism.

A Bioluminescence Switch Within a Single Species

The geographic bioluminescence pattern in P. stipticus — glowing in eastern North America, dark everywhere else — is unique among known bioluminescent organisms. The 2025 genomic study by Rabara and Xie provided definitive confirmation: the bioluminescent strain genome contains a ~10 kb luciferase gene cluster; the non-bioluminescent Korean strain genome shows complete absence of all four bioluminescence genes (luz, hisps, h3h, cph). Dot-plot analysis of the two genomes confirms high overall conservation — the strains are clearly the same species. The mechanism and timing of luminescence loss in non-luminescent populations, and whether any selective pressure is involved, remain open questions.

160 Million Years of Continuous Bioluminescence

The luciferase gene cluster in P. stipticus traces back to the last common ancestor of the mycenoid and marasmioid clade of Agaricales approximately 160 million years ago — the Jurassic period, when the earliest flowering plants were appearing. The fungal bioluminescence chemistry (based on 3-hydroxyhispidin derived from caffeic acid) is fundamentally different from all other known bioluminescent systems. Firefly luciferin, bacterial luciferin, and jellyfish photoprotein are chemically unrelated to fungal luciferin. Fungi arrived at the same functional outcome — biological light emission — through a completely independent evolutionary pathway.

A Self-Recycling Light System

Unlike firefly bioluminescence, in which luciferin is consumed and must be continuously resynthesized, the fungal bioluminescence pathway is a closed cycle. Caffeic acid is regenerated after each light-emission event by caffeoylpyruvate hydrolase. The system can theoretically sustain indefinitely as long as oxygen is available and the caffeic acid pool is maintained. This is energetically more elegant than any other known animal bioluminescence system and distinguishes fungi as the only organisms known to achieve recycling bioluminescence.

A Diurnal Glow Clock

Cultured mycelia of P. stipticus show a pronounced diurnal periodicity in bioluminescence, with maximum luminescence consistently between 6 and 9 pm regardless of whether cultures are maintained in continuous light, continuous darkness, or a normal day/night cycle. This implies an endogenous circadian rhythm for bioluminescence that persists independent of external light cues. The molecular basis of this rhythm has not been characterized — making it one of the more fascinating unsolved problems in fungal biology.

Bioluminescence as an Environmental Sensor

The sensitivity of P. stipticus bioluminescence to chemical contamination has been validated as a biosensor system. Metal salts (copper sulfate, mercury chloride, zinc sulfate) reduce bioluminescence by 68–72% within 30 minutes of exposure. Nitrate salts reduce it by 15% within 90 minutes. Herbicide (atrazine) and pesticide (permethrin) exposure reduces it by 12–13%. A concentration-dependent luminescence reduction is detectable for lead oxide. A greater than 10% reduction in bioluminescence can be visually detected within 130 minutes for all tested contaminant concentrations. This makes living cultures of P. stipticus a potentially practical, low-cost tool for environmental toxicity screening.

White Rot and the Carbon Cycle

White-rot fungi including P. stipticus are the only organisms known to achieve complete lignin mineralization — the full breakdown of the aromatic polymer that constitutes roughly 30% of the dry weight of all woody plants. Without white-rot fungi, the planet's carbon cycle would be fundamentally different: vast quantities of recalcitrant woody carbon would accumulate rather than returning to CO&sub2 and soil nutrients. P. stipticus plays a specific and replaceable role in early-succession wood decomposition as a pioneer species on fresh deadwood.

Also available as a culture plate from Out-Grow.

Panellus stipticus Culture Plate

Frequently Asked Questions About Panellus stipticus

Does all Panellus stipticus glow in the dark?

No. Bioluminescence in Panellus stipticus is geographically restricted to eastern North American strains. European, Asian, Pacific coast North American, and Australasian strains are morphologically identical but completely dark. A 2025 genomic study confirmed that this reflects the presence or complete absence of a ~10 kb luciferase gene cluster. If you want to observe bioluminescence, make sure you source a culture confirmed to be from a bioluminescent eastern NA strain — such as Out-Grow's offering.

Why doesn't my liquid culture of Panellus stipticus glow?

Bioluminescence in Panellus stipticus requires growth on a solid substratum and adequate oxygen exposure. Mycelium suspended in liquid culture does not luminesce regardless of strain. Once transferred to agar, sawdust, or hardwood substrate, the mycelium will develop bioluminescence as it colonizes the surface. Allow at least five minutes of dark adaptation before checking — the glow is faint and requires your eyes to adjust.

What is foxfire, and is Panellus stipticus the cause?

Panellus stipticus is one of the best-known causes of the "foxfire" or "glow wood" phenomenon — the eerie green glow observed emanating from rotting logs at night in eastern North American forests. The same phenomenon has been documented in folklore across temperate regions for centuries. However, P. stipticus is not the only bioluminescent wood-rot fungus; approximately 132 fungal species are known to produce bioluminescence, several of which can colonize the same type of dead wood. A glowing log in the field could involve multiple bioluminescent species.

What is the best substrate for cultivating Panellus stipticus?

For fruiting body production and the most vigorous bioluminescence display, hardwood sawdust (oak, beech, or maple) in sterilized blocks is recommended. Hardwood logs inoculated with plug spawn produce the most natural fruiting but require 6–12+ months of colonization outdoors. Grain spawn can be used to build up mycelium for substrate transfer, but fruiting bodies on grain alone typically abort. For agar culture, 10% breadcrumb agar (BCA) produces better growth and higher bioluminescence than standard malt extract agar (MEA), based on peer-reviewed data from a 2025 study.

Is Panellus stipticus edible or medicinal?

Panellus stipticus is inedible. The mushrooms are too small, too tough, and too bitter to eat, and the species has no culinary tradition anywhere in its range. Traditional Chinese medicine documented an external use as a hemostatic (blood-stopping) agent, which is plausible given the astringent chemistry, but no modern scientific validation of this use exists. No human clinical trials have been published for any application. No antioxidant, antimicrobial, or cytotoxic studies have been conducted on this species. It has no marketed supplement form.

What family does Panellus stipticus belong to?

Panellus stipticus belongs to family Mycenaceae within order Agaricales. Many online sources still list it in Tricholomataceae (a now-fragmented wastebasket family) or, incorrectly, Panellaceae. The current consensus, based on multi-gene phylogenetic analysis and reflected in Index Fungorum and GBIF, is Mycenaceae. There is ongoing scientific discussion about whether P. stipticus should eventually be transferred to genus Mycena itself, though this reclassification has not yet been formalized.