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, elaphinus, primus, orestes, parilis n.p., pouzarianus, atromarginatus, laricinus, salicinus, petasatus, magnus, leucoborealis, brunneidiscus, washingtonensis, heterocystis, pellitus, nothopellitus, hongoi
Pluteus cervinus group - unclamped
Our most common species group, but only confidently identified microscopically by the absence of clamp connections.
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, but perhaps fewer fibrils on the stem. 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 suggested the opposite, also with a small sample size. So that teaches us the dangers of studies with small sample sizes. Note that pure white versions of Pluteus exilis have been found in WA twice, but I have no confirmed Pluteus cervinus collections lacking pigment.
Pluteus elaphinus MA - one WA collection was the first known west coast collection of this lookalike that might be told apart by a darker cap and more ornamentation on the cap and stem than the other clampless species.
Pluteus cervinus © Danny Miller, P. exilis (normal and albino) © Kieran Yeung and Yi-Min Wang, P. elaphinus © Danny Miller
Pluteus pouzarianus group - clamped
These three species in the have clamps, and are collected less frequently than the unclamped species. P. pouzarianus itself is an EU species that does not appear to occur here. Although some have tried to distinguish the species by cap colour, as you'll see from the photos, that doesn't appear to be reliable and they may have to be distinguished microscopically.
Pluteus primus EU - P. primus is a conifer species most often with a very dark cap but a microscope is required for proper confirmation. Several WA sequences match official EU and CA sequences quoted in the paper. One old collection had marginate gills. Some agarics seem to be capable of developing marginate gills in age and this may be one of them, as this has never been recorded for this species before.
Pluteus orestes CA - this clamped species has a medium to pale brown cap and is 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 supposedly a medium to pale brown capped conifer species from the coast and the cascades, but best distinguished microscopically. One BC sequence with a very dark cap is 4 bp and 1 indel different in ITS, but for now I'm assuming it represents P. parilis.
Pluteus primus © Sharon Squazzo and Yi-Min Wang
Pluteus orestes © Ron Pastorino and Sava Krstic
Pluteus parilis n.p. (pale version) © Julie Jones, P. aff parilis (dark version) © Christopher Ng
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.
probable Pluteus petasatus © Andrew Parker, P. petasatus © Bruce Newhouse
Pluteus leucoborealis Siberia - another large 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 type. Since then we found a better WA collection. It has a large pale or whitish cap with some brown, radial fibrils (although my specimen was probably old and 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 and Sharon Squazzo
Pluteus laricinus MI (P. atromarginatus EU misapplied?) - 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. Note that as seen with P. primus above, this is not the only species capable of having marginate gills due to weathering.
Pluteus laricinus © Michael Beug
Pluteus 'salicinus PNW11' - 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 Hispidoderma below has a dark capped bluing species that does not smell of radish nor have the "jester cap" cystidia. We need photographed, sequenced collections.
unsequenced Pluteus 'salicinus PNW11' © Erin Page Blanchard
Pluteus brunneidiscus CT (= P. washingtonensis WA, =P. heterocystis WA) - 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 fulvibadius, fulvobadius, romellii, chrysophlebius, rugosidiscus, aurantiorugosus, phaeocyanopus, thomsonii, tomentosulus, nanus, hispidulus, podospileus, inflatus
Pluteus fulvibadius (fulvobadius) OR (P. 'romellii' misapplied) - brown, yellow-brown (or occasionally just yellow) cap, often wrinkled and sometimes felty. It has a somewhat yellow stem and young gills. Similar Pluteus leoninus group mushrooms are hard to tell apart without a scope, but are more likely to have a felty/velvety cap and won't be wrinkled (as they are not in this section). Authors that studied PNW collections noted that P. fulvibadius 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. fulvibadius, but a recent paper has confirmed that this is the proper name of our local species. As to the spelling, Murrill originally spelled it "fulvibadius". Subsequent Latin scholars corrected it to "fulvobadius" for reasons that are unclear, so other Latin scholars have now argued to change it back. Some collections, at least in CA, have green tones in the young caps, but they do not have unique sequences and are considered the same species.
Pluteus vellingae CA - from CA, back east, and across northern Europe and Asia, but not yet known from the PNW. It is "challenging to differentiate" but a little smaller and its young gills are white or only slightly yellow. Perhaps we should keep an eye out for it.
Pluteus fulvibadius © Jonathan Frank and Andrew Parker (2 images)
Pluteus chrysophlebius SC - we have a similar but usually entirely bright yellow to orange-yellow mushroom with a bright yellow stem. We have somewhat reliable sequences from back east matching an OR sequence, but note that there are no green tones. See the next species as well.
Pluteus rugosidiscus ENA - a sister species said to have a green tinge to the cap, also from back east, differing by 5% in ITS. Many people thought this was a synonym of P. chrysophlebius. Collections with a green tinge to the cap have been reported from the PNW so we need greenish collections to sequence to see if we have this species here too.
Pluteus cf aurantiorugosus EU - a lovely deep orange capped species, but with a mostly white stem. 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. Our one WA collection matches EU sequences.
Pluteus chrysophlebius © Zane Walker (2 images), Pluteus aurantiorugosus © Peter Sheff
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 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 © Andrew Parker
Pluteus 'thomsonii PNW04' - this possibly nondescript brown capped species is supposedly especially susceptible to wrinkling and reticulation. It's possible the reticulation could be subtle in which case it will start to resemble P. brunneidiscus in section Pluteus or P. inflatus, described next. One local sequence and several CA sequences are 3% different from UK/EU sequences. There is a paper in the works to describe this sister species. An unsequenced collection with a strongly reticulated cap, thought to be this species, is shown below along with a sequenced collection that was very nondescript. We need collections with a strongly reticulated cap.
Unsequenced possible Pluteus 'thomsonii PNW04' © Danny Miller, P. 'thomsonii PNW04' © Yi-Min Wang
Pluteus 'nanus PNW01' - sequences of P. nanus 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 from all of them. Unfortunately, it is mostly a drab grey-brown mushroom best recognized microscopically, except for slightly bluing grey stem and roundish spores (many but not all look round from some angles, others are somewhat elliptical).
Pluteus 'nanus PNW02' - one WA collection also sequences about 4% different in ITS from all of the above. The photo shows perhaps a slight blue-grey tinged stem, but not nearly as much as the closely related P. phaeocyanopus above. It too has roundish spores.
Pluteus inflatus EU - also drab but without the blue tinge to the stem, also with roundish spores about 5.3–6.2u in diameter. It lacks pleurocystidia and was found once in WA in an urban setting. Members of the Pluteus podospileus complex vary by as much as 15% in ITS from each other, and this is the only member of that complex known from the PNW so far.
Pluteus 'nanus PNW01' (2 images) © Damon Tighe, Pluteus 'nanus PNW02' (2 images) and P. inflatus (2 images) © Yi-Min Wang
Pluteus 'hispidulus PNW12' - 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 'hispidulus PNW12' © 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 (= P. flavofuligineus NY) - yellow-brown to brown cap often with a yellow rim, pinkish stem becoming yellow. The similar Pluteus fulvibadius ('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. Most of our PNW sequences fall in the P. flavofuligineus clade, but one Idaho sequence falls within the P. leoninus clade, so if they are determined to be distinct species, both of them might occur here. We have no sequenced photographs yet.
Pluteus 'leoninus PNW10' - 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). One photo has an especially dark brown cap that lacks a yellow rim and looks especially like P. fulvibadius, 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 'leoninus PNW10' © NAMA and the Field Museum of Natural History, albino version © Andrew Parker
Pluteus umbrosus EU - Plutues granularis NY and Pluteus umbrosus EU are two very similar and closely related species with a dark brown scaly cap with the dark scales looking like radiating veins, and fibrils or scales on much of the stem too. Stems can be >5 mm thick. There are several species going by both these names, but Justo has provided sequences that may represent each species to help sort them out. One WA collection sequenced 5 bp different than what Justo calls P. umbrosus in the EU, and another WA collection sequenced a different 5bp different from it. But a CA sequence has ambiguous locations in those places, uniting the sequences. Either there are multiple species that can hybridize, or it's all one species. I'm going to assume for now that it's all one species. It can be variable though, as each collection looked different. The cap may be dark or pale brown, and the scales may be broken up all over the cap instead of radiating (see photos).
Pluteus granularis EU - Alaska has Justo's concept of P. granularis, but it has not yet been found in the PNW.
Pluteus 'plautus PNW06' - this relative of the EU P. plautus is stockier and darker than most other P. plautus relatives, covered next, making it hard to differentiate from P. umbrosus, although it may not be as likely to have scales like radiating veins. The second photo show an smaller, paler collection that looks much like the other P. plautus relatives and not P. umbrosus. It has been sequenced from BC, ID and OR.
Pluteus umbrosus © Danny Miller and Harte Singer, Pluteus 'plautus PNW06' ©Andrew Parker and Mary Berbee
Finally, rather nondescript, slender Hispidodermas from grey to tan to dark brown, with or without dark scales on the stem. Stems usually <5 mm thick. Caps often hygrophanous.
Pluteus 'plautus PNW07' - OR and CA. Our one photo has a tan, somewhat hairy, hygrophanous cap, and no scales on the stem except for some white pruinosity.
Pluteus 'plautus PNW08' - OR and CA. Our one photo has a somewhat pale grey cap with a dark brown disc, and a stem marked with white streaks and pruinosity.
Pluteus 'plautus PNW09' - one WA collection had a fairly pale cap and lacked scales on the stem. A BC collection had a dark cap and dark fibres on the stem. Clearly, more examples are needed of each to figure out how to recognize our local species in this complex.
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 in the P. plautus group.
Pluteus 'plautus PNW07' © Sava Krstic, Pluteus 'plautus PNW08' © Jonathan Frank, P. 'plautus PNW09' © Buck McAdoo and Zeenath Aleaf
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 surrecta UK - notable for growing parasitically on Clitocybe nebularis and perhaps other species. We have one local EWA sequence that matches three EU sequences, so we probably do have that species here.
unsequenced Volvariella surrecta © Noah Siegel
Volvariella terrea EU - a recent surprise from OR, matching the EU type sequence very well. This species may be parasitic on Agaricus xanthodermus, which has been found under the same oak tree as this collection.
Volvariella terrea © Bitty Roy
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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