Danny’s DNA Discoveries – Pluteus, Volvopluteus and Volvariella of the PNW
This part of the Pluteaceae family is easy to recognize, having evolved free gills and pink spores. Pluteus grow on wood and do not have a volva at the bottom of the stem. Volvopluteus and Volvariella do have a sac volva (like some Amanita) and are found in a variety of habitats like wood, cultivated soil, grass, and parasitic on other mushrooms. Volvopluteus has a smooth, viscid cap. Volvariella has a fibrillose, dry cap. They are often tall, thin, statuesque mushrooms, whose caps may flatten out. This stature, the free gills and the volva that some have, are shared by their sister family, Amanitaceae, which did not evolve the pink spores. Melanoleuca is the other common genus in the Pluteaceae, but with attached gills and white spores, the only feature they have in common with their other family members is their tall statuesque, often flat capped nature, which is sometimes key to identifying a Melanoleuca.
abundant common uncommon rare - colour codes match my Pictorial Key and are my opinions and probably reflect my bias of living in W WA. Rare species may be locally common in certain places at certain times.
Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
Pluteus section Pluteus - click to expand
No sac volva. Growing on wood. This section smells and tastes like radish and has pleurocystidia on the gill faces that are fat and spindle shaped and look like they are wearing a jester cap.
Species mentioned: Pluteus cervinus, exilis, primus, orestes, parilis n.p., pouzarianus, atromarginatus, laricinus, salicinus, petasatus, magnus, leucoborealis, brunneidiscus, washingtonensis, heterocystis, pellitus, nothopellitus, hongoi
Pluteus cervinus group - unclamped
Pluteus cervinus EU - the so-called "deer" mushroom, because the cap reminded somebody of a deer pelt. Black fibrils are usually on the stem. No clamp connections under the microscope. Usually a medium brownish-grey, but the cap can be darker or lighter like other species found below with clamps, so you need to check for clamps to be certain you have one of these two species.
Pluteus exilis CA - very, very difficult to distinguish without DNA sequencing. A recent DNA paper declared that only P. exilis had been found so far in the PNW, but both species were in California, but they had a very small sample size (four collections). So I started sequencing all the deer mushrooms I could find in OR and WA and all 6 turned out to be P. cervinus and none were P. exilis, so my report is the opposite, also with a small sample size, saying that we only have P. cervinus in the PNW and not P. exilis. So that teaches us the dangers of studies with small sample sizes.
Pluteus cervinus, and probable Pluteus exilis (brown and white variants) © Danny Miller and Richard Morrison
Pluteus pouzarianus group - clamped
Pluteus primus EU - these three species in the Pluteus pouzarianus complex have clamps. That is an EU species that does not appear to occur here itself. P. primus is a conifer species often with a dark cap but might have a medium cap like P. cervinus and P. exilis, and might even have a pale cap like the next two species, so a microscope is required for proper confirmation. One WA sequence of mine matches official EU and CA sequences quoted in the paper, so this species does occur here, but it was rather pale capped and easily confused with the next two.
Pluteus orestes CA - this clamped species is a medium to pale capped and found in the cascades on conifers. Best distinguished microscopically. OR and WA sequences match CA type area sequences.
Pluteus parilis n.p. CA - this clamped species is not described yet, but probably will be from CA. I have WA and OR sequences that match the CA sequences that it will be described from. It is a medium to pale capped conifer species from the coast and the cascades. Best distinguished microscopically. One BC sequence is 4 bp and 1 indel different in ITS, but for now I'm assuming it represents P. parilis.
Pluteus primus (dark, microscopic match and paler genetic match) © Danny Miller and Kate Draper
Pluteus orestes © Ron Pastorino and Sava Krstic
Pluteus parilis n.p. (pale version) © Julie Jones
Pluteus petasatus EU - very similar to the above, but without black fibrils on the stem. Whitish cap is overlain with black fibrils. On urban area hardwoods. No clamps. CA and WA sequences match the EU exactly. Justo concluded that the type of Pluteus magnus was the same as this, although they could not obtain DNA to prove it.
Pluteus petasatus © Andrew Parker
Pluteus leucoborealis JP - another difficult to identify species that was not known from the PNW (but was reported in California) until I found it in my yard and sequenced it and it matched the Japanese type. It supposedly also has a pale or whitish cap with some brown, radial fibrils (although my specimen had a uniform tan cap), and a pale stem (much like P. petasatus) but is more often found in wild areas instead of urban areas.
Pluteus leucoborealis © Danny Miller
Pluteus laricinus ('atromarginatus') MI - black marginate gill edges set this species apart from our others. We have long called this by the EU name P. atromarginatus, but those sequence are at least 4 bp and one long chunk of 5 indels different. One BC sequence and 2 OR sequences are basically a perfect match to the type sequence of the more recent Michigan species P. laricinus. The next closest match (3 bp different from the type sequence) is the MI species P. atropungens. Justo considered them all potential synonyms but the clades do separate clearly, so I'm not so sure. If they are all the same, the oldest name is P. atromarginatus, but for now it's safest to call them P. laricinus.
Pluteus laricinus © Michael Beug
Pluteus aff. salicinus EU - medium brown cap with a white stem that turns blue when handled from psilocybin. Our one BC sequence is 4% different than the dozens of EU sequences we have of this, so it is likely an undescribed sister species. Section Hisidoderma below has a dark capped bluing species that does not smell of radish nor have the "jester cap" cystidia.
Pluteus aff salicinus © Erin Page Blanchard
Pluteus brunneidiscus (washingtonensis, heterocystis) CT - a sometimes smaller 5cm deer mushrooms, often with a dark cap and again without black fibrils on the stem. This photo shows an interesting white ground colour showing through that is atypical. With hardwoods. Has clamp connections. Justo examined the Washington type collections of P. washingtonensis and P. heterocystis and find them great microscopic matches to P. brunneidiscus, and probably not separate species, although DNA could not be obtained to prove it. But I have shown that sequences from BC, WA, OR and ID all match east coast sequences of P. brunneidiscus, so our collections matching this description throughout the PNW always match the DNA of P. brunneidiscus, so I think that closes the case.
Pluteus brunneidiscus © Jonathan Frank
There has been speculation that Pluteus pellitus, Pluteus nothopellitus or Pluteus hongoi could conceivably be present in the PNW, but no reports nor even rumours yet. But if you think you find one, let me know.
Pluteus section Celluloderma - click to expand
No sac volva. Growing on wood. This section does not smell of radish and does not have the jester capped pleurocystidia. The microscopic cap cuticle is composed of rounded elements, which makes the cap susceptible to wrinkling. (There may also be elongated elements like section Hispidoderma, but that section should not have any rounded elements). Usually they are colourful and have a smooth cap, but they might have a felty or fibrillose cap like section Hispidoderma, so if you don't have a microscope, also check that section.
Species mentioned: Pluteus fulvobadius, romellii, chrysophlebius, rugosidiscus, aurantiorugosus, phaeocyanopus, thomsonii, tomentosulus, nanus, hispidulus
Pluteus fulvobadius OR ('romellii') - brown, often wrinkled and sometimes felty cap and bright yellow stem and young gills. Similar Pluteus leoninus group mushrooms are more likely to have a felty/velvety cap but won't be wrinkled (as they are not in this section). Authors that studied PNW collections noted it had differently shaped cystidia than the very similar EU mushroom Pluteus romellii, and suggested we should probably be using our local species name instead of the EU name (as everybody has been using the EU name for our collections). I can confirm that DNA sequences from BC, OR and WA are distinct in ITS from EU sequences of P. romellii by about 1% or so, and our local sequences mostly differ from each other in ambiguous locations only. We don't have a type sequence of P. fulvobadius to prove it, but it seems that the studies were correct and we should use the name P. fulvobadius.
Some collections, at least in CA, have green tones in the young caps. We should study if these represent a genetically different species.
Pluteus fulvobadius © Jonathan Frank
Pluteus cf rugosidiscus/chrysophlebius ENA - we have a similar mostly yellow mushroom often with green tinges on the cap, that might have brown in the cap, making it somewhat confusable with P. fulvobadius, above. We have somewhat reliable sequences of both mushrooms from back east (they are sister species differing by about 5% in ITS), but no local collections to find out which of the two occur here, although the theory is that P. rugosidiscus might be the one found here as green tones are often present in our caps and one theory is that P. rugosidiscus might be more likely to have the green tones. We will need local sequences to find out, though.
Pluteus cf rugosidiscus © Andrew Parker
Pluteus cf aurantiorugosus EU - a lovely orange capped species, often with orange in the stem as well. East coast sequences differ from EU sequences by 4-6 bp or so, but a study was unable to find any ecological or morphological differences so it is thought they are all one species. We need local sequences to see where our collections fall on the spectrum.
Pluteus cf aurantiorugosus © Michael Beug
Pluteus phaeocyanopus CA - a dark brown cap with a paler stem that turns blue when handled. Interestingly, this species is in a different section than our paler capped bluing species, Pluteus salicinus, which means that Psilocybin has evolved in Pluteus multiple times. It is a CA oak species probably found in at least OR as well, but we need local collections to match up with CA sequences.
Pluteus phaeocyanopus in CA © Christian Schwarz
Pluteus cf thomsonii UK - this dark brown cap is especially susceptible to wrinkling and reticulation. It's possible the reticulation could be subtle in which case it will start to resemble P. brunneidiscus. We don't have any local sequences yet (nor any from North America) to compare to EU sequences to verify this is what our species is.
Pluteus cf thomsonii © Danny Miller
Pluteus tomentosulus NY - this cottony capped all white Pluteus could be mistaken for a Volvariella if the lack of volva isn't noted, and since many Pluteus seem to have albino forms, every all-white collection is not necessarily this. Sequences from WA and OR match sequences from ENA, where is was described. One microscopic analysis failed to find rounded elements in the cap cuticle, but the genetics clearly place this species in section Celluloderma.
Pluteus tomentosulus © Danny Miller
Pluteus aff nanus EU - sequences of this species in Europe differ by as much as 2%, so it may be one species with some genetic variation or it may be a species complex. Several forms and varieties have been described, so perhaps they correspond to the different sequences. Three sequences in California and one in Oregon are a sister species about 4% different, even more likely to be considered separate from any of the EU varieties, and not the real thing. I don't have a photo, but keep an eye out for this rare Pluteus. Unfortunately, it is mostly a drab grey-brown mushroom best recognized microscopically, except for possibly a bluish-grey stem.
Pluteus aff hispidulus EU - One lone OR collection sequences about 10% away from a lone EU sequence labeled Pluteus hispidulus, but Justo saw the sequence and concluded ours was most likely a species in the P. hispidulus group. It is only known from this one collection. It can be differentiated from other species in this section by a distinctly fibrillose dark brown cap. Similar looking species are found in section Hispidoderma.
Pluteus aff hispidulus © Leah Bendlin
Pluteus section Hispidoderma - click to expand
No sac volva. Growing on wood. This section does not smell of radish and does not have the jester capped pleurocystidia. The microscopic cap cuticle is composed of elongated elements. (Section Celluloderma may also have elongated elements, but it should have at least some rounded elements too). Usually they have a fibrillose or velvety cap, but not always, so if you don't have a microscope, also check section Celluloderma.
Species mentioned: Pluteus leoninus, flavofuligineus, umbrosus, granularis, plautus, latifolius
Pluteus leoninus EU group (flavofuligineus NY) - yellow-brown to brown cap often with a yellow rim, pinkish stem becoming yellow. The similar Pluteus fulvobadius ('romellii') is often going to have a wrinkled cap because of its placement in section Celluloderma (and usually, but not always, has a more uniformly dark brown cap and a brighter yellow stem). P. leoninus may have a felty cap texture, which was formerly attributed only to P. flavofuligineus but as P. leoninus has also been found with a felty cap, and since sequences of both are within 3-4 bp and a couple indels of each other (but with no ecological or other morphological differences), Justo and others consider them synonyms, with the EU name P. leoninus being older.
Pluteus leoninus - One Idaho sequence is within the clade, so the real Pluteus leoninus does occur in the PNW.
Pluteus aff. leoninus - Two OR and WA sequences are a half dozen bp and a half dozen indels different, far enough away that Justo considers this and a couple of other clades in ENA and Japan to possibly be an additional sister species, but morphological and ecological features will have to be checked (no known differences yet). The coloured photo has an especially dark brown cap that lacks a yellow rim and looks especially like P. fulvobadius, so a close look at the cap texture would probably be necessary for a correct ID. The albino photo shows that white Pluteus can be very hard to ID as so many of them seem to have white forms.
Pluteus aff leoninus © NAMA and the Field Museum of Natural History, albino version © Andrew Parker
Pluteus cf granularis NY/cf umbrosus EU - two very similar and closely related species with a warm brown scaly cap and fibrils or scales on much of the stem too. Stems can be >5 mm thick. The brown colours may be warm, but there is usually no actual yellow anywhere as in Pluteus aff. leoninus, above. Reports of both the NY species and the EU species have yet to be confirmed by DNA, so we need samples. Several sequences from each type area match each other, so it's possible they are the same species, but Justo said that the species found both on the east coast and in Europe may be P. granularis and provided a separate EU sequence that may be P. umbrosus. CA reports possibly more than one genetic species in this group. Let's get PNW collections and see what we have here. We do know that Alaska has Justo's concept of P. granularis.
Pluteus cf granularis (unsequenced) © Andrew Parker
Plutues plautus group EU - similar cold brown scaly cap, with or without dark scales on the stem. Stems usually <5 mm thick. Caps often hygrophanous.
Pluteus aff plautus #1 - BC and OR have sequences of sister species #1 (to the real EU species, which has not been found here yet) that from our only photo, seems to have a dark brown scaly cap and dark scales over the entire stem.
Pluteus aff plautus #2 - OR and CA have sequences of sister species #2. Our one photo has a tan, somewhat hairy, hygrophanous cap, and no scales on the stem except for some white pruinosity.
Pluteus aff plautus #3 - OR and CA have sequences of sister species #3. Our one photo has a somewhat pale grey cap with a dark brown disc, and a stem marked with white streaks and pruinosity. Clearly, more examples are needed of each to figure out how to recognize our local species in this complex.
Pluteus aff. plautus #1 © NAMA and the Field Museum of Natural History, #2 © Sava Krstic, #3 © Jonathan Frank
Pluteus latifolius WA - locally described somewhat vaguely with a cold brown scaly cap like Pluteus plautus, but with a stem covered in short tomentose hairs, it has never been collected (or looked for) since the 1911 Seattle type collection. We need to get a type sequence or recent collections that match microscopically to figure out what it is. It could be one of the undescribed species mentioned above.
Volvopluteus - click to expand
Sac volva. Growing on the ground, but often in rich soil or grass. The cap is viscid and smooth.
Species mentioned: Volvopluteus gloiocephalus, Volvariella speciosa
Volvopluteus gloiocephalus (Volvariella speciosa) EU - our least rare species was recently moved to a new genus when it was discovered that the viscid, smooth capped Volvariellas were not so closely related to the others. It is found worldwide and a BC sequence matches quite well. It used to be called Volvariella speciosa, but it turned out that V. gloiocephalus was the same thing and the most valid, oldest name.
Volvopluteus gloiocephalus © Noah Siegel
Volvariella - click to expand
Sac volva. Growing on wood, the ground, rich soil, grass or other mushrooms. The cap is dry and fibrillose.
Species mentioned: Volvariella volvacea, bombycina, surrecta, pusilla, hypopithys, smithii
Volvariella cf volvacea EU - the store bought "paddy straw" mushroom found in many Asian dishes may have escaped into the wild here as it has been reported. It has a fairly dark grey cap, whereas the others have mostly white to light grey caps. There are at least two genetic sister species in the EU going by this name, so more work is needed to figure out which one is the real one, and which one might occur here.
Volvariella cf bombycina EU - quite large with an equally impressive large volva, and growing directly on hardwood instead of the ground like most others. I have reliabe EU sequences but we still need local sequences to compare to to make sure this is what is found here.
Volvariella cf bombycina © Noah Siegel
Volvariella cf surrecta UK - notable for growing parasitically on Clitocybe nebularis and perhaps other species. We have one Spanish sequence that may correctly identified, but we don't have local sequences to compare to yet, and we don't know for sure that the Spanish sequence really represents this.
Volvariella cf surrecta © Noah Siegel
Volvariella cf hypopithys EU / Volvariella cf smithii WA / Volvariella cf pusilla EU
Small, white Volvariellas growing on the ground. They are supposedly separated by the first two having a pubescent stem (the third with a smooth stem), by V. smithii having an off white (pinkish or brownish) disc and volva and narrower spores, and by V. pusilla being more often in urban soils and grasses. These differences seem slight. We don't have any local sequences of any of them, but a bunch of sequences from the EU and Quebec match within 1% and are sometimes called V. pusilla and sometimes V. hypopithys, so this may be all one species with a bit of genetic variation. Since EU and eastern NA sequences match, perhaps ours will match too. If there's only one species, the oldest name by far is Volvariella pusilla. But with notably narrower spores, our endemic Volvariella smithii may be a distinct species. We'll need local collections to find out if we have 1, 2 or 3 species here.
Volvariella cf hypopithys © Danny Miller
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