© Fred Rhoades
Danny’s DNA Discoveries – Mycenaceae of the PNW
by Danny Miller
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Danny’s DNA Discoveries – Mycenaceae of the PNW
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Introduction
The mycenoid stature is named after the genus Mycena - the quintessential fragile, hollow stemmed little white spored mushrooms often with bell-shaped or conical caps. Marasmioids typically have flatter caps, distant gills (although some Mycena have few gills) and tougher stems. Collybioids can still be pretty small, and also flatter capped with tougher stems, but have gills more closely spaced. Resinomycena, Roridomyces, and Panellus are currently considered separate genera in the same family, but they actually live inside Mycena. They will probably have to become Mycenas, or Mycena will need to be split significantly. Favolaschia, while looking most distinct must likely also be considered part of Mycena unless Mycena is to be significantly split. This "family" has amyloid spores. Note that Hydropus, Mycopan and Leucoinocybe are difficult to distinguish from Mycena (usually with a different microscopic cap cuticle), and that Hemimycena s.l., Atheniella and Phloeomana (and 'Mycena' acicula and 'Mycena' oregonensis) have been separated from Mycena into genera in other familes on the basis of inamyloid spores. The two authorities on Mycena are Alex Smith for North America and Maas Geesteranus for Europe. Their monographs are indispensible. 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. |
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Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
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 Mycena - click to expand
The quintessential mycenoid small, fragile, bell-shaped to cone-headed white spored mushrooms, with amyloid spores. Hydropus and Mycopan have at least weakly amyloid spores as well and are separated microscopically by their different cap cuticle.
Club-shaped form of Mycena subvitrea OR - easily mistaken for Pterulicium/Macrotyphula, but growing from wood. Is this an asexual fruiting? I don't know.
Mycena subvitrea (club form) © Matthew Koons
Marginate gill edges (the cap itself may or may not colourful) Mycena strobilinoidea WA (=M. strobilinoides WA) - orange with orange marginate gills. It is a more uniform orange than the non-marginate gilled Atheniella aurantiidisca, and grows at higher elevations. Mycena aurantiomarginata EU -orange tinged brown cap with orange marginate gills. We have collections matching many EU type area sequences. Mycena sp. 'aurantiomarginata-PNW08' - some local sequences are almost 3% different in ITS. Mycena sp. 'CA17' - similar to M. aurantiomarginata, perhaps more towards brown than orange. Mycena citrinomarginata EU (=M. olivaceomarginata EU?, =M. olivaceobrunnea WA?, =M. chloranthoides OR?) - yellow-brown cap with yellow marginate gill edges. Worldwide sequences are within 4-6 bp of each other so it appears there may be some variation in ITS. Some EU sequences of the newer M. olivaceomarginata EU have the same ITS as M. citrinomarginata (and others are close but distinct), so they are likely either synonyms, or different but not differentiable in ITS. Some consider them separate. M. olivaceomarginata has tan gill edges, is found in grass, and has a red stem base. M. citrinomarginata has yellowish gill edges, is found in forests, and a pale stem base. Also, the local species M. olivaceobrunnea WA appears to have a similar ITS and may be a synonym of one or both. Some suspect M. chloranthoides OR is also a synonym of one or both. Those last two need their types sequenced. Finally, one collection with matching ITS had purple tinged gills but no gill margination.
Mycena strobilinoidea © Michael Beug, M. sp. 'aurantiomarginata-PNW08' © Steve Ness, M. citrinomarginata © Yi-Min Wang and Bitty Roy Mycena rubromarginata EU - reddish-grey cap, with dark red gill edges. More than a half dozen local sequences match a couple dozen EU sequences. It may look almost exactly like paler collections of M. purpureofusca, below. Mycena bulliformis CA - hard to distinguish from M. rubromarginata without a microscope, perhaps with a brighter cap when fresh. Described in 2016. Once it was found without marginate gill edges. M. elegantula CA - Smith called collections that sound a lot like M. bulliformis (red tones in the cap and pale pinkish-purple gill edges) by the name M. elegantula. Since then people have doubted or at least wondered what he really found. Could it have been M. bulliformis, which wasn't described at the time? We need sequences to find out what M. elegantula is. Mycena capillaripes NY - grey or brown cap with pink marginate gill edges (paler than in M. rubromarginata) with a bleach odor. We don't have NY type area sequences, but one EU, one NZ, and one CA sequence constitute one genetic species and this matches those pretty well. There's a good chance it will match with NY too. Be warned - not all observations with matching ITS show the pink marginate gill edges nor the bleach odor. When they do, this species is easy to identify. When they don't, it is not. Mycena sp. 'capillaripes-PNW01' - two WA sequences constitute a second sister species, 1.5% different. These are not closely related to the other bleach species. Our collections did not appear to have the bleach odor nor the pink marginate gills. Therefore, I don't know how to recognize it. The bleeding Mycenas may have red marginate gill edges, also see that section below.
M. rubromarginata © iNaturalist user pdawson9229 and Danny Miller, M. bulliformis @ iNaturalist user ellenkr & NAMA and the Field Museum of Natural History
Mycena capillaripes @ Buck McAdoo, M. sp. 'capillaripes-PNW01' @ Lauren Ré Mycena purpureofusca NY - greyish purple cap and stem fading to pale purplish grey, with purple marginate edges. Sequences of this species seem to match worldwide. Mycena viridimarginata, an olive-brown species with green marginate edges, appears to have the same ITS sequence. It should be looked for in the PNW. When not as darkly coloured, it may look almost exactly like M. rubromarginata, above. Mycena sp. 'rosella-NY01' - bright pink (but changing to orange-ish in age) with pink marginate gill edges. One collection seemed to stain reddish brown. We have 9 EU type area sequences showing us what this species' sequence likely is, and 1 EU sequence that is fewer than 6 bp different. NA sequences (including PNW ones) are in the second clade. Just in case it isn't all one species, NA collections along with the minority of EU collections, are being referred to as NY01. Mycena pterigena EU - very small and pink on fern fronds, usually with pink marginate gill edges. We have 3 EU sequences that probably show us what this is and two matching OR sequences.
Mycena purpureofusca © Vail Paterson and Michael Beug, unsequenced M. sp. 'rosella-NY01' © Noah Siegel, M. pterigena © Noah Siegel
Bleeding Mycenas - stems exude red, white, or clear milk. Most make up one genetic clade. Mycena haematopus EU - red cap with a beautiful scalloped edge, stem exudes red milk which can stain all parts red. It is usually on wood. The gill edges may be red marginate, keep reading. Mycena sp. 'haematopus-PNW11' - one OR sequence and one BC sequence of a similar species sequences quite distinct, and it does not even have support for being in the same clade as the others in this group. Smith described var. marginata with red gill edges as a distinct variety, so it should be investigated if one of our species has the red marginate edges and the other does not, or if there is some other way to tell the two apart. Mycena subsanguinolenta WA - a similar red species exuding red milk from the stem, but found on the ground, without the scalloped cap margin, and always with strongly red marginate gill edges. The type sequence is 5% different in ITS than EU sequences of M. sanguinolenta, so they are likely not synonyms as commonly believed. The east coast also appears to have their own unique species. We may have a species complex here, as our two modern collections are 4-5 bp and 3 indels different in ITS than the 1939 type sequence, but that sequence came from very old material. Also, a recent WA sequence had some ambiguous locations, but that didn't account for all the differences. For now, I'm assuming it's all one species. Mycena sanguinolenta EU - Smith also found this, with larger spores and pleurocystidia, and one WA collection matches many EU sequences. Most turn out to be M. subsanguinolenta.
Mycena haematopus © iNaturalist user hugebooter, M. subsanguinolenta © Steve Ness Mycena galopus EU (formerly 'NC01') - this plain grey/brown Mycena won't be recognized if you don't notice the white milk in the fresh stem. We have a BC sequence that matches dozens of worldwide sequences, and an OR sequence that is 4 bp different. Mycena sp. 'atroalboides-AK01' - this plain looking Mycena may exude water from the stem, but it isn't clear if this is typical or not. An AK sequence is a sister species to a PQ sequence (which may be the real thing), and there are WA, CA, and AK sequences of species AK01, another sister species, which likely represents reports of this species in the PNW.
unsequenced Mycena galopus © Noah Siegel (2 images), M. sp. 'atroalboides-AK01' © Noah Siegel
Mycena epipterygia group - viscid yellow stems, yellow or grey caps Mycena epiptyerygia var. epipterygia EU - described as a slender species with a yellow cap found on the ground. We have sequences from the EU and locally, and photos show that the caps might be grey as in M. epipterygioides. It should also be investigated if this variety can be found on wood like var. lignicola. Not all EU sequences match these sequences, so it is possible that once a type is chosen, this will not be the real thing and need a new name. Mycena epiptyerygia var. lignicola WA (including Mycena sp. 'epipterygia-BC01') - described with a yellow cap and growing on wood instead of the ground. Although strong farinaceous or iodoform odors are sometimes described in this species group, one collection smelled strongly fishy to me. The type collection does have distinct DNA, but sometimes collections with matching DNA are found on the ground and/or with a grey cap, so growth on wood may not be the correct way to differentiate this species. Species in this clade have a few differences in ITS from each other, and some go by the codename M. sp. 'epipterygia-BC01', but they all intermingle and for now I'm treating them all as this variety. Mycena epipterygia var. nov - one BC sequence of a collection with a pale grey cap doesn't quite match any other sequences, we should see if we find this sequence again. Mycena epipterygioides UK - the name Smith gave to collections without yellow in their cap but with a dark grey cap that didn't fade to white. Geesteranus doubts Smith's report that this is in America, and we have seen that yellow and grey capped collections have the same ITS sequence. We don't have any DNA of this from anywhere, but with the wide colour variation we are seeing in M. epipterygia, I doubt that reports of it represent a distinct species. Mycena viscosa EU - it's not clear what Smith collected and called by this name, as his description of them didn't quite match the actual description of M. viscosa. We need collections matching Smith's description to see if they do represent a unique species - more strongly unpleasantly farinaceous than the other species in this section and flesh that stains red. Mycena viscosa var. iodiolens OR - with an unpleasant odor that becomes like iodine after picking, described with a grey cap. We have the type sequence and another 1941 Smith collection sequenced, as well as one modern OR collection, and it probably needs promoting to species. Mycena nivicola CA - a much beefier mushroom, with a brown cap and a stem that turns from yellow to red-brown. You might not recognize that the mushroom is in this group when the stems aren't yellow anymore. The authors of this species strongly suspect that reports of the east coast Mycena griseoviridis represent this instead, because they are a microscopic match. However, Smith said that M. griseoviridis looked a lot like M. epipterygia but M. nivicola is quite different looking and easily distinguished. Mycena sp. 'CA15' - a brown cap and a yellowish stem that stains red-brown, much like M. nivicola, but more slender. Known from CA and WA. Mycena griseoviridis var. cascadensis WA - similar to M. nivicola, but the cap is yellow or grey, and the stem does not turn red-brown. We don't have any sequences yet, but the authors of M. nivicola suspect that this is a truly distinct species. We need collections.
Mycena epipterygia var. lignicola © Vail Paterson, unsequenced Mycena epipterygia (grey and yellow caps) © Sadie Hickey
Mycena viscosa var. iodiolens and M. nivicola © Jonathan Frank, M. sp. 'CA15' (from CA) © Sadie Hickey
Other Colourful Mycenas (not the usual white, grey, or brown - gills are not marginate). Mycena monticola OR - coral pink, with a darker pink stem than the similar pink Atheniella 'adonis PNW01'. We have the type sequence and some OR sequences without photos that match fairly well, although there's something odd about the sequences. Some differ by a couple %, but as many appear to come from the same source and are somewhat suspicious, I'm guessing they all represent the same species. We have one WA photographed sequenced collection. Mycena sp. 'PNW19' - one WA collection growing from a decayed cedar fence post had a pinkish grey cap and stem and a mild odor. A second collection had a bleach odor and a dark grey-brown cap. It sequences close to M. capillaripes, which is a grey mushroom with pink marginate gill edges and a bleach odor. We need more collections and a good description. Mycena sp. 'amicta-PNW13' - blue on the cap and stem when fresh. When the blue has faded, you can still recognize it by a peelable cap cuticle and hairs/pruinosity on the stem. There are multiple species going by this name in Europe. This species is 10% different than the most common EU species, but exactly matches one EU concept. Note that PNW13 was accidentally used twice in Mycena. Mycena sp. 'amicta-PNW14' - this species seems to have the blue in the cap fade much more quickly than in PNW13. It aslso has a peelable cap cuticle and pruinose stem. It is 2% different from the most common EU candidate. Even in the EU, DNA varies by 2% or so, so it's still possible this is the real thing, unless this complex is split into several species. Smith did note that this species was quite variable, so it's not surprising that there may actually be 2 species here. Mycena tubarioides EU - very small and pinkish, with a peelable cap cuticle and gill edge. Mycena pterigena, above, usually has marginate gill edges and no part is gelatinous.
Mycena monticola © Connor Dooley, Mycena sp. 'PNW19' © Kitty Lundeen-Ness
Mycena sp. 'amicta-PNW13' © Steve Ness, M. sp. 'amicta-PNW14' © Alan Rockefeller (2 images)
Mycena pura complex (genus Prunulus to some) - purple colours, rather large for a Mycena, with an odor of radish and usually pink or purple when fresh, although those colours can fade and many other colours are sometimes noted, like white, yellow, blue, or green. Although many varieties have been described for the different colours, the only colour that was found to have any taxonomic value was yellow (Harder, 2013). Mycena subaquosa OR is a local name for a white collection that might be the proper name for one of our species, either PNW15 or CA01. Most EU sequences are in a complex of 8 clades, with no known way to differentiate them yet (except for yellow clade V). The real Mycena pura is probably one of these clades. It's not clear how they will be split into multiple species, as the total variation is only 3% in ITS, although different genes do show the same separation. Mycena sp. 'pura-PNW15'/IN07' Harder clade VII/VIII - It's possible this is Mycena pura itself. Interestingly, clade VIII was only known in the EU in his study, not in NA, but the very close clade VII was known from the US. When you put all of our PNW15/IN07 sequences into a tree, there is no clear differentiation between clade VII and clade VIII, so those two clades may only represent one species. If they are distinct species, PNW15 and IN07 have not been applied properly to differentiate them. Either these codes will have to be combined or reapplied. Mycena sp. 'pura-CA01' (formerly PNW16) Harder clade X - Harder also found 3 more clades (IX, X and XI) that were outside of the main cluster of species, so the real species is less likely to be one of these, and CA01 almost certainly needs its own name. Clade X is mostly found in NA, not the EU. Mycena sp. 'pura-CA03' Harder clade II - one WA collection as well as many CA collections. Mycena sp. 'pura-PNW27' Harder clade VI - one darkly coloured WA collection is in this clade of US sequences, known from elsewhere in the US in Harder's study but not from the PNW until now. Not being present in the EU, and not being very close in ITS to any EU sequences, it definitely needs its own name. Mycena sp. 'pura-PNW25' - this species was not in Harder's study. It was known only from the east coast until found a few times in Spokane, WA. It seems to have very subtle lilac grey colours and is quite hygrophanous. Mycena sp. 'PNW39' - also not in Harder. One WA collection is a unique sequence so far. Mycena sp. 'IN03' - also not in Harder. It is known from back east, CA, CO, AK, and one OR collection.
Mycena sp. 'pura-PNW15' (purple and white forms) © Kitty Lundeen-Ness, and NAMA & the Field Museum of Natural History, M. sp. 'pura-CA01' © Steve Ness (2 images), M. sp. 'pura-PNW27' © Derek Hevel
Mycena sp. 'pura-PNW25' © Garrett Taylor and Sophie Deibel Mycena sp. 'pearsoniana-PNW23' - with arcuate/decurrent gills and a longer stem and a perhaps a somewhat flatter cap than most members of the Mycena pura group. We have one BC sequence and one CA sequence. The DNA is also found in Europe. Either this or PNW24 is the real thing from Mexico. Mycena sp. 'pearsoniana-PNW24' - A sister species is found in the EU, WNA, and ENA. I don't know which is the real Mexican species. In the east it is not umbonate like our WA photo. The type sequence of Mycena shengshanensis China is close to this sequence. Prunulus applanatus China (formerly Mycena sp. 'pearsoniana-PNW40') - another species in this complex. Some consider the Mycena pura group in a genus of its own, Prunulus. We'll have to wait to see hat happens to the rest of Mycena before adopting that.
Mycena sp. 'pearsoniana-PNW23' from CA © Dean Lyons, M. sp. 'pearsoniana-PNW24' © Yi-Min Wang Mycena pelianthina EU - an even stockier radish smelling Mycena that is collybioid, with purple marginate gill edges. We have a number of EU sequences but haven't found matching DNA in the PNW yet. We need collections. Mycena sp. 'IN06' (M. rutilantiformis misapplied) - similar, also with purple marginate gill edges. Local sequences match east coast type area sequences, so this could be M. rutilantiformis MO, but there are a couple competing possibilities for what M. rutilantiformis is. We need a type sequence. They say the stem apex can be yellowish in M. rutilantiformis but not M. pelianthina, and some of our collections do show a yellow stem. Mycena sp. 'pura-AB01' - BC, WA, and OR collections are a novel species near M. pelianthina, but it is more slender like M. pura and lacks marginate gill edges.
unsequenced M. pelianthina group member and M. sp. 'IN06' © Buck McAdoo, M. sp. 'pura-AB01' © Heather Dawson (2 images)
BLEACH odors Mycena alcalina EU - this has been a very popular name to use, but it was so poorly understood that Maas Geesteranus gave up on using that name and described 2 species to take its place - Mycena stipata and Mycena sylvae-nigrae. Both are EU species. So nobody should be using the name M. alcalina anymore. Mycena stipata EU - UNITE has 3 genetic species at 0.5% threshold. The third one is locked, so we can’t see it. Of the other two, if you go to 1% threshold, there’s only 1 species. So all 5 sequences are pretty close to each other, so we kind of know what Mycena stipata is, within 1%. Even though it is commonly reported from here, no PNW sequences match this so far, so there is no evidence that this is one of the species found in the PNW. Mycena silvae-nigrae EU (=Mycena sp. 'silvae-nigrae-IN02') - grey or brown. UNITE has 5 species concepts at 0.5%. If you go to 1.5%, they are all 1 species except for one outlier. So we kind of know (within 1.5%) what Mycena sylvae-nigrae is. It is about 4% different in ITS from M. stipata. M. silvae-nigrae was never reported from the PNW, but many local sequences match M. sylvae-nigrae within 2%. Some local sequences match some EU sequences very closely, so I am assuming we have the real M. sylvae-nigrae here, which has some variation in ITS. It is possible that it is a species complex, but I don't see the sequences falling into clades. This means that when M. alcalina was split into M. stipata and M. silvae-nigrae, we chose poorly when guessing that the one that occurred here was M. stipata. Mycena leptocephala EU - differs microscopically from M. silvae-nigrae, and usually less clustered than it. If you search UNITE for Mycena leptocephala, with 0.5% tolerance for a species, you get 7 different genetic species. The first one has been sequenced 541 times, surely a record (you usually see single digits or occasionally a couple dozen). This is clearly a very, very common species. Click on it and the map shows it was found 539 times in the EU, once in BC and once in AK. It was only identified as Mycena leptocephala four times. About a half dozen times it was only identified to genus. The other 530 times it was just a soil sample DNA blast (no mushrooms). So that is not overwhelming evidence that this is the real M. leptocephala. If we look at the other 6 genetic species, only one of them is found in the EU, so it’s the only other possibility of being the real species. This second possibility was found, identified and sequenced twice in Estonia. I’m going to assume the first species is the real thing, since it’s so common, even though I could be wrong. We don’t have a type sequence, and the first species was only identified as M. leptocephala 4 times vs. the second species being identified twice, so that is not quite as overwhelming as 541 sequences versus 2 sequences sounds at first, but still, let’s assume we now know what M. leptocephala is. Our 7 local sequences of M. leptocephala match very well with my assumed real EU sequences from the above paragraph, so I conclude that we likely have that species here. Our sequences are mostly within 1% and sometimes within 0.5% of EU sequences, although a pre-ITS region has a bunch of differences in one sequence, but I’m not going to make too big a deal out of that for now. Mycena sp. 'leptocephala-PNW20' - four collections stand out with ITS 2-3% different. This also has an alkaline odor. Mycena sp. 'leptocephala-PNW21' - one WA and one CA collection also stand out a few % different. The odor was not noted. This sequence matches a syntype sequence of Mycena tinctura, however other syntypes have different sequences, so Smith's type collection was a mixed collection and we probably shouldn't use that name for anything. Mycena abramsii CA - I don’t have any reliable sequences of this. There are some Korean sequences purporting to be this, but no California sequences. The reason people think this might be our common bleach smelling species instead of M. stipata is probably only because it is a local west coast species and M. stipata is from the EU, so odds usually favour using a local name over an EU name. But in this case, we do seem to have EU species here, so we are not in search of a new, local name for our collections. Also, M. abramsii is described from under oak on the ground, not on conifer wood, so the habitat is all wrong. Also, M. abramsii is described with a faint radish odor and only sometimes with a bleach odor. In other words, although it is believed to be common here, there is no proof that it is here at all. Geesteranus thought Mycena alnicola WA might be the same thing based on a radish odor among other things. That should be investigated, but we don't have any DNA yet. If you think you find this in the PNW, save it. Mycena capillaripes NY/sp. 'capillaripes-PNW01' - grey or brown, sometimes with pink marginate gill edges (and illustrated above in that section) and sometimes a bleach odor. We don't have NY type area sequences, but one EU, one CA, and one NZ sequence consistute one genetic species, and this WA sequence constitutes a second sister species, 1.5% different. I don't yet know if ours is the real one or not. It appears to be closely related to the above bleach species. Mycena sp. 'PNW19' - described above may be colourful, but apparently may also may be plain brown. If it is neither coloured nor with a bleach odor, it will be difficult to identify. Mycena sp. 'macrocystidiata-PNW02' - never reported from the PNW until one WA collection with a bleach odor had giant cystidia and seemed to be this species. It has unique ITS DNA, but we have no other sequences of this EU species to compare to to know if that's what it really is or not. It is not in the same clade with the other bleach species. Mycena subcana CA (was PNW10) - described below, often has a bleach odor even though it wasn't described with one.
Mycena silvae-nigrae © Steve Ness, M. leptocephala © Sadie Hickey, M. sp. 'leptocephala-PNW20' © Fred Rhoades
Mycena sp. 'leptocephala-PNW21' © Fred Rhoades, M. macrocystidiata © Buck McAdoo
Other odors Mycena sp. 'cinerella-PNW06' (M. brownii WA?) - strongly farinaceous, contrasting striations in the disc and arcuate gills. Local sequences are 1.5% different than a couple of EU sequences purporting to be that species. We do have a local name, M. brownii from WA, to use if it is determined that our species is not M. cinerella. Mycena tenax WA - viscid stem, peelable cap cuticle, strong rancid-farinaceus odor and taste. Gills not strongly arcuate. We have a half dozen OR sequences showing us what this species' sequence likely is, but no confirmed photographs yet. Mycena inclinata EU - another farinaceous species (some say odor of wet paint) with a slightly scalloped cap margin and a stem base that turns rusty brown. We have no local sequences yet to show if reports of this being here are true or not. Mycena odorifera NY - said to have a fruity or Matsutake odor. Viscid cap and stem, but not always obviously so. Arcuate gills. We have no DNA from anywhere yet. Mycena filopes EU - odor of iodine. A dozen worldwide sequences agree on what this species' ITS DNA is, including some from the EU type area, and we have local sequences that match them. Mycena sp. 'filopes-PNW12' - Two WA sequences and a couple CA sequences are 5% different and probably represent a sister species. The east coast has a local name, M. graveolens MI, that is thought to be a synonym of M. filopes, and I believe that to be correct, since M. filopes is found all over the east coast and PNW12 so far has not. So that is probably not PNW12. Mycena metata EU group - similar iodine odor, perhaps best differentiated microscopically. 70 EU sequences show us what this species is. Some sequences match very well, but others are about 1% different, although I treat them as the same species. Mycena sp. 'metata-PNW13' - one WA sequence is 3% different so I am treating it for now as a potential separate species. Note that PNW13 was accidentally used twice in Mycena. Mycena sp. 'PNW41' - sweet to rancid depending on age, and the gills stain reddish. Known from one BC collection.
Mycena sp. 'cinerella-PNW06' and M. filopes © Yi-Min Wang, M. sp. 'filopes-PNW12' © Alan Rockefeller, M. metata and sp. 'metata-PNW13' © Fred Rhoades
Viscid stems - growing on the ground unless otherwise specified. Not all species are close relatives of each other so they do not form a natural section like Smith hypothesized. Roridomyces roridus EU - see below, this species has a very slimy stem, dry cap, and arcuate gills and likely belongs in Mycena, not in a separate genus. Mycena sp. 'clavicularis-PNW04' - a grey/brown mushroom with a somewhat viscid stem that isn't as slimy as in Roridomyces. It also has a dry cap and arcuate gills and fruits in late summer. A number of local sequences are about 1% different in ITS than EU type area sequences. It should be investigated if these collections represent M. insignis, described below. Mycena clavicularis EU - one OR collection has a sequence close enough to some EU type area sequences that I'm tentatively calling it this, although we're not sure what that species is. Our collection had the expected dry cap. Mycena insignis WA - with a somewhat viscid cap, viscid stem, arcuate gills, and a pale, almost white colouration, that takes the pigment out of the conifer needles it grows out of, turning them white. No DNA yet, unless this species is M. 'clavicularis PNW04'. Mycena tenax WA - (also discussed in the "other odors" section). A strong rancid-farinaceus odor and taste. Also, a viscid stem and peelable cap cuticle. Gills not strongly arcuate. We have a half dozen OR sequences showing us what this species' sequence likely is, but no confirmed photographs yet. Mycena odorifera NY - (also discussed in the "other odors" section). It is said to have a fruity or Matsutake odor. Viscid cap and stem, but not always obviously so. Arcuate gills. We have no DNA from anywhere yet. Mycena vulgaris EU (=Mycena sp. 'vulgaris-CA01') - viscid cap (more so than M. insignis) and stem, peelable gill edge, arcuate/decurrent gills fruiting in the fall. Over 100 EU sequences show us what this is, and our sequences are a pretty close match, although some use a codename for our species even though the difference between some CA sequences is less than the differences between them and many EU sequences. Mycena melleodisca ME is thought to be a synonym. 'Mycena' (Mycopan) quiniaultensis WA - viscid cap and stem, somewhat dark grey when fresh, but gills not decurrent. It is illustrated under Mycopan, where it needs to be moved. Mycena sp. 'laevigata-PNW09' - a mostly white, entirely viscid Mycena growing on logs. It is larger than Hemimycena. It has practically the same ITS and LSU sequence as our two Mycena overholtsii species in the large species section below. Those species are larger, high elevation species with hairs around the stem base, one of which is a spring snowmelt species. Interestingly, the real EU species M. laevigata is 5% different in ITS and is distinguishable from M. overholtsii, but our local sister species isn't. Perhaps that means M. overholtsii and this undescribed species evolved here and the EU species was derived from it. Geesteranus suspected that our species might not be the same as the EU species, and it looks like he was right.
Mycena sp. 'clavicularis-PNW04' © Richard Morrison, Mycena clavicularis © Dan Morton
Mycena vulgaris © Lauren Ré, Mycena sp. 'laevigata-PNW09' © Richard Morrison and Matthew Koons
Small, white species with basal discs, bulbs or sugary/hairy caps or stems - the gills may be attached to a collarium allowing the stem to easily separate from the cap. Similar Hemimycena have inamyloid spores and usually lack the basal disc and hairs. Smith may have been right that these species make up a natural section in the tree. Resinomycena, below, is covered with a sticky, resiny sugary coating. Mycena stylobates EU - flat basal disc and some hairy thorns may project from the cap (look with a hand lens before you touch). This is the largest of these species -the caps may reach 1cm across and the stem is fairly long. CA and WA sequences match a couple of EU sequences. Mycena bulbosa EU - also fairly large (for this group) with a flat basal disc, but no hairs/thorns. The cap cuticle and gill edges are peelable. Smith did not find it here, but others since think they have. We have one EU sequence that might be this, showing up in the right place in the tree, but no local sequences. Mycena gaultheri WA - smaller cap (a few mm across) and shorter stem, with a yellowish cap and yellowish flat basal disc, no thorns, and growing on salal leaves. We don't have any DNA yet. Mycena aciculata CA (M. longiseta EU misapplied) - also a small cap with a small, rounded basal bulb, with hairs/thorns on both the cap and stem when fresh and before handling. This CA species is sometimes mistaken for the EU species M. longiseta, and in fact our local sequence is very similar to an EU sequence purporting to be M. longiseta, so I think this represents M. aciculata, although it would be nice to get more sequences or a type sequence to prove it. Mycena tenerrima UK (=M. adscendens EU) - also fairly small (usually only a few mm across) with a subtle basal bulb, and overall sugary appearance (with granules, not thorns). One BC sequence matches one EU sequence identified as this UK species, and so does a dirty WA sequence once it is cleaned up by hand. It would be nice to get macro photographs of the sugary appearance and a type sequence or additional UK/EU sequences to confirm this identification. One study may be showing that our species is more properly called Mycena nucicola instead. Mycena alphitophora Bermuda - differs microscopically from M. tenerrima and almost lacking any basal bulb. We have a NY sequence of a collection that matched microscopically and is the right place in the tree, but no local sequences yet. Mycena sp. 'clavularis-PNW22' - quite small, growing on moss covered bark of deciduous trees, with a large basal bulb, few gills and a collarium. It is 3% different in ITS than an EU type area sequence purporting to be M. clavularis. This species was never reported from the PNW before a WA collection was sequenced. In Europe, the species is reported with a viscid cap with separable cap cuticle and a possible nitrous odor.
Mycena stylobates © Yi-Min Wang (2 images), M. aciculata © Fred Rhoades, M. tenerrima © Yi-Min Wang
Mycena sp. 'clavularis-PNW22' unsequenced and sequenced © Matthew Koons
Large grey/brown species Mycena sp. 'maculata-IN01' (formerly PNW03) - a large species growing in clusters on wood, but the mushroom stains red when handled or in age. Our ITS DNA is 2% different than EU sequences. Mycena galericulata EU (=BC01) - a similar rather large (for a Mycena) species growing clustered on wood, but not staining red. Many local sequences match dozens of EU sequences somewhat closely, although this species has seems to have about 1% variability in ITS worldwide. Mycena sp. 'PNW42' - one candidate for Mycena megaspora MI, matching sequences from QC under that name. Mycena sp. 'PNW43' - a similar species Mycena sp. 'PNW44' - one OR sequence and some east coast sequences are 2% different than M. galericulata. No known differences yet. Mycena sp. 'PNW26' - a similar species with large spores and a long root-like stem growing in bogs.
Mycena sp. 'maculata-IN01' © Bruce Newhouse, M. galericulata © Bitty Roy, M. sp. 'PNW42' © Sandra Ruffner, M. sp. 'PNW44' © Jonathan Frank, M. sp. 'PNW26' © Connor Dooley Mycena hudsoniana WA - one of our larger spring Mycenas. We don't have the type sequence, but enough collections have been analyzed to know that we have accurately identified some local sequenced collections. Mycena sp. 'PNW28' - closely related to M. hudsoniana, but with only one known collection, from OR, I don't know how robust it typically is. Mycena cf robusta OR (=M. sp. 'IN22') - a common winter Mycena best recognized microscopically by distinctive cystidia, a few collections seem to match M. robusta, but we don't have a type sequence to prove it. One EU sequence purporting to be the older EU species M. aetites matches these sequences too. It should be investigated if that could be an older name for our species, but it is described differently, with a radish odor. Mycena overholtsii WY - a larger collybioid mushroom not always recognizable as a Mycena growing next to snow at high elevations in the spring, with hair around the stem base. The type sequence has practically the same ITS and matching collections also have the same LSU sequence as Mycena 'laevigata PNW09' (see the viscid stemmed section) and the below species and cannot be distinguished by those genes. Mycena sp. 'overholtsii-PNW30' - the winter low-mid elevation version of the species has the same ITS and LSU but looks different enough (white to dark grey), and the ecology is different enough, that I am treating it separately. It should probably be consdiered a distinct variety.
Mycena hudsoniana © Fred Rhoades, M. sp. 'PNW28' © iNaturalist user Tripp Mikich, M. cf. robusta (sp. 'IN22') © Bruce Newhouse
Mycena overholtsii © Daniel Winkler, M. sp. 'overholtsii-PNW30' © Heather Dawson
Tiny species on bark on tree trunks (occasionally on branches), probably not a natural group. Mycena meliigena USA complex (M. corticola EU misapplied) - a tiny species on bark with round spores. One WA sequence is within 1% of an ENA type area sequence and a number of EU sequences of M. meliigena. The oldest name for this species may be M. corticola, but as that is a poorly understood species, the name is being abandoned. M. meliigena is supposed to have a vinaceous brown cap, and some photos of our collection appear quite yellow, but others match closely enough to EU photos I have seen that I think we have the same species. Mycena madronicola OR - on bark of madrone, with elliptic spores. The gills are adnate to decurrent. We have an OR sequence from madrone bark that could be this species, but read on. Mycena pavsikensis EU - with the same ITS as the madrone collection above except for ambiguities, this is a moss species. We've found a BC collection matching the holotype. Mycena corticalis OR - a small species with more reliably arcuate gills than the others, growing on mossless western red cedar bark. We have the type sequence, but no photos. Spores almost round.
M. meliigena © Yi-Min Wang and Michael Beug, M. madronicola © Connor Dooley (2 images)
Plain grey or brown Mycenas - most are best differentiated microscopically in Smith's or Geesteranus' monographs. Mycena picta EU - this has a unique umbrella shape and gill arrangement that is easy to spot. Two local sequences match an EU sequence and probably represent this species. Mycena vitilis EU - one WA sequence matches over a dozen GenBank sequences. This species starts out grey/brown but can fade to almost white. They say the stem is flexible and can be swung without breaking. Mycena sp. 'PNW18' (=Mycena culmigena OR?) - small (cap ~3mm across), few gills, white cap with pale vinaceous tones, and growth on sedges and rushes. One OR collection is a micro match to Mycena culmigena OR, but we don't have any other sequences of it to compare to and prove it. It is also similar to Mycena riparia EU, but that hasn't been sequenced either. We also have a BC sequenced collection. Mycena sp. 'CA22' (=Mycena culmigena OR?) - another species that looks like M. culmigena, but it is not closely related to PNW18. Mycena sp. 'PNW05' - two collections (from OR and WA) have a unique sequence. This species appears to be pale grey , found once on wood and once on the ground. The OR collection had stem base tomentum. For the WA collection, the caps were 5-8 mm broad, and there were typical chielocystidia but no pleurocystidia. Spores 8.1 x 4.6 µm. Fred Rhoades keyed his collection out to Mycena pusilla CA, and Bitty Roy keyed out some collections to the probably related Mycena constans NY as the likely identity of this species, although unlike M. pusilla, it had never been reported from the PNW. Mycena sp. 'PNW07' - one OR sequence, an almost white Mycena growing on sticks on the coast. This seems to match Mycena piceicola OR. Mycena subcana CA (was PNW10) - may grow on conifer branches, not trunks, but usually only when still attached to the tree, so you won't find it on the forest floor. Elliptical spores, with differently shaped cheilocystidia than many other species. Many, many sequences match sequences of some of Smith's original collections (but we don't have a type sequence). The only problem with this is that it often smells bleachy, but it was described with no odor. Also, collections with similar ITS but in different habitats may be different species, more study is needed. One collection has ambiguous locations raising even more questions. Possible older synonyms to be investigated include Mycena fragillima CA, Mycena murina NY, and M. stannea EU. Mycena sp. 'PNW36' (=IN26?) - one WA collection with somewhat similar ITS to M. subcana seems ecologically and morphologically distinct. Some collections from back east called IN26 may be this. Mycena subvitrea OR - we have the OR type sequence and matching collections. As noted above, this has an odd club-shaped form. Mycena rebaudengoi EU - not known from the PNW until discovered by DNA in BC in grass near birch. Mycena sp. 'PNW31' - a unique sequence from ID, but the sequence seems clean. Mycena mirata NY - known from one BC collection Mycena sp. 'PNW32' - known from 2 WA collections. Mycena sp. 'HRL07' - two collections from WA with decurrent gills. Mycena sp. 'PNW33' - known from 1 WA collection Mycena sp. 'PNW35' - known from 1 OR collection Mycena sp. 'IN19' - one OR collection locally, but found across the US. Mycena sp. 'WA04' - known from WA and QC. Mycena sp. 'BC02' - one BC collection. Mycena sp. 'PNW45' - one BC collection.
Mycena picta © Sadie Hickey, M. vitilis © Yi-Min Wang, M. sp. 'culmigena-PNW18' © Connor Dooley
Mycena sp. 'PNW05' © Yona Celeste Riel and Fred Rhoades, M. sp. 'PNW07' © Ann Goddard
Mycena subcana © Fred Rhoades and Yi-Min Wang (2 images)
Mycena rebaudengoi © iNaturalist user angelique_k, M. sp. 'PNW31' © Krista Willmorth, M. mirata © Anne Polyakov
Mycena sp. 'PNW32' © Yi-Min Wang, M. sp. 'HRL07' © Yi-Min Wang (2 images)
Mycena sp. 'PNW33' © Yi-Min Wang (2 images)
Not actually in Mycena (either moved or not moved yet) Mycena rainierensis WA - believed to be a Fayodia, this may just be a newer synonym of the Fayodia that I believe to be present in the PNW, Fayodia gracilipes, but it's possible it could be a unique species as Smith said it had longer cheilocystidia and a paler colour. It should be sequenced to find out, and if it is unique, it needs to be moved to Fayodia. Mycena cineraria WA - this is the local name for one of our species of Gamundia. We need to figure out if it has an older name (Gamundia leucophylla is a possibility) and if it doesn't, then this species needs to be moved to Gamundia. Mycena acicula EU - received its own genus, Pruinomycea, but as the erection of that genus requires splitting Hemimycena even further, it might have been better to consider it as belonging to an expanded version of Hemimycena. Mycena oregonensis OR - needs to be moved to its own genus, or perhaps an expanded version of Phloeomana. Mycena quiniaultensis WA - needs to be moved to Mycopan.
Little known species (white or grey to brownish) For some of these, Smith reported them but Geesteranus disagreed, so these reports are mostly unverified. We may never know what some of these reports represent unless we sequence Smith's original collections. I haven't found reliable sequences of any of these, although some sequences are available that might represent them. More research needs to be done. I would appreciate type sequences and any collections that key out in Smith (found here) or Geesteranus to any of the following. Very small species (caps <1 cm across) with few gills
Mycena alcaliniformis NY (=M. subsupina CA) - Decurrent gills
Mycena fuliginella WA - decurrent gills Completely non-descript - good luck, you can begin on page 225 of Smith's monograph
Mycena algeriensis Algeria - some extralimital sequences match our sequence
of M. macrocystidiata |
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Favolaschia, Panellus, Resinomycena, and Roridomyces - click to expand
Multi-genes show these all probably belong in Mycena, even if Mycena is somewhat split. Favolaschia - easily recognized as tiny tan oysters with "pores" for gills. This genus is sister to all the others, but Mycena may still not be able to remain monophyletic if Favolaschia is kept distinct, as it likely will be. Panellus - pleurotoids with tiny lateral stems on wood. Difficult to separate from 'Panellus' mitis and Scytinotus, where some Panellus have been moved. Resinomycena - related miniscule white agarics a few mm across on woody debris, covered with sticky granules, but with a small central stem so not considered pleurotoid. Other granular Mycena are not sticky. Roridomyces - a plain grey Mycena with an extremely glutinous stem, dry cap and usually decurrent gills. Other viscid Mycena either have a sticky cap as well or their stem is not quite this ridiculously glutinous or the gills are not decurrent. Amyloid spores. Species mentioned: Favolaschia sp. Resinomycena saccharifera, kalalochensis, montana. Panellus stipticus, mitis, ringens, longinquus. Roridomyces roridus.
Favolaschia sp. 'PNW01' - our species does not match any of the species I know of in a 5-gene study of the genus. It was not known from the PNW until recently, and we finally got a sequence of a WA collection.
Favolaschia sp. 'PNW01' © Graham Steinruck
Panellus stipticus EU - the only local species left in Panellus, recognized by its tough orange fuzzy caps, slightly bitter taste and somewhat forked and interveined gills that glow greenish in the dark. The similar Phyllotopsis nidulans has no stem, is stinky, and doesn't taste bitter. One WA and CA sequence match EU sequences fairly well, better than east coast sequences which are 6 bp different, where they could potentially have a different species. Susan Libonati-Barnes attempted to describe our collections as var. occidentalis, so it should be investigated if ours deserves varietal status. Panellus longinquus and P. ringens are now in Scytinotus. P. mitis needs a new genus.
unsequenced Panellus stipticus © Noah Siegel
Resinomycena (Mycena) saccharifera EU - on hardwood debris, with few gills, a short stem and large spores. We have EU sequences, but no matching local sequences yet to prove the reports of it being here. Resinomycena sp. 'CA01' (=R. kalalochensis WA?) - one OR collection. This has smaller spores, matching the spore size of local species R. kalalochensis, but before this CA01 was discovered R. kalalochensis was assumed to be a synonym of R. saccharifera. We need to confirm what R. kalalochensis is. Resinomycena montana BC - on conifer debris with a moderate number of gills, a longer stem and even smaller spores. We have official collections from those whose described it sequenced from OR and CA.
Resinomycena sp. 'CA01' © Daniel Morton, R. montana © Connor Dooley
Roridomyces roridus EU - A WA and CA sequence match a half dozen EU sequences quite well.
Roridomyces roridus © Andrew Parker |
