Danny’s DNA Discoveries – Entoloma s.l. of the PNW
The Europeans place every species on this page inside Entoloma, but in North America, some acknowledge separate genera, as there is often (but not always) a fairly clear way to separate them and otherwise the number of species becomes unwieldy. Even if I treated them all as Entoloma, I would be separating the page into sections for each subgenus, so I might as well call each section a genus. David Largent's 1994 comprehensive treatment of the family on the west coast is the basis for much of what we know of our local species.
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:
Entoloma (Rhodopolia) - click to expand
Entoloma in the sense I use it are medium to large (tricholomatoid), smooth, lubricious capped species with grey or brown tones. There are clamps in the cap cuticle. They are very difficult to differentiate from each other without a scope. Nolanea are smaller and more slender (collybioid or mycenoid) with similar cap texture and colour, without clamps in the cap cuticle. The other genera on this page are more colourful or not lubricous. Rarely, an Entoloma might be a tiny omphalinoid species, in which case it will be confused with Paraeccilia.
Species mentioned: Entoloma alnobetulae, alpicola, brunnescipes, clavaformipes, grande, griseoavellaneum, griseum, heracleodora, lividoalbum, majaloides, saussetiense, lividum, lupinum, lyophylloidium, myrmecophilum, nidorosum, politum, pseudocostatum, pseudolividum, rhodopolium, sericatum, sinuatum, speculum, subpolitum, subsaundersii, subsinuatum. Nolanea abbreviatipes.
Although I am treating this section as the genus Entoloma, the issue is complicated. By some accounts, the type species of Entoloma is Entoloma prunuloides, so my section Prunuloides would need to be called Entoloma, and this section would need to be called Rhodopolia. To keep more names stable, an exception could be made to conserve this section as Entoloma, but I cannot yet predict how this will eventually shake out.
It's possible it will later be discovered that some of these need to move to other genera, but preliminary analysis suggests that all of these are inside Entoloma s.s. Although we don't have any sequences of some of them, the sections they've been placed in, or the species they've been compared to, suggest they do belong here.
Entoloma politum EU - an omphalinoid Entoloma, with a mostly non-striate stem and mild to nitrous odor. This was the type species of the genus Eccilia, but since it is inside Entoloma, that genus name is not needed. We have a number of EU sequences and no matching local collections yet, but keep reading.
Entoloma subpolitum WA - a very similar omphalinoid but with a striate stem and farinaceous odor. No sequences yet. Keep reading.
Entoloma 'politum PNW01' - Our only local sequence from this group is 1.5% different in ITS. This could be E. subpolitum, but for now I am assuming it isn't as Dave Largent, who described E. subpolitum, sees significant differences in this collection. Some think that Entoloma maculatum NC and Entoloma triviale NA are synonyms of E. politum, but they may not be. We need sequences of both of those and E. subpolitum to investigate further.
Entoloma 'politum PNW01' © Richard Morrison
Entoloma lividum group - possible yellow gills when young
Entoloma grande NY - Largent thought that this was the white capped Entoloma lividum group member to be found in the PNW. There is conflicting information about whether or not the gills will be yellow when young. We don't have any DNA to compare to, so we need reliable east coast sequences and local collections.
Entoloma subsinuatum ME - although never reported from the PNW, this turned out to be the sequence of a (probably) white capped BC collection in the Entoloma lividum group. The gills of this species were originally reported as white, but a modern study showed the gills are probably yellow when young. We need more collections to see if E. grande occurs here too, as the spore sizes overlap.
Entoloma pseudolividum WA - this was described as our darker capped member of the Entoloma lividum group with yellow young gills but we have no DNA of it yet. It has been described as being much like E. flavifolium NY, which we do have DNA of. We need local collections.
Entoloma rhodopolium group, etc. - some common species
Entoloma rhodopolium EU - usually farinaceous with roundish spores. Largent commonly reported six forms from the west coast, but we have not yet found DNA of this species here. We need local collections to see what they are.
Entoloma 'griseum PNW05' - this species, also farinaceous with roundish spores like E. rhodopolium, is the DNA that turns up most often in local Entoloma collections. That makes me wonder if collections of this are being mistaken for E. rhodopolium. We are not sure what the DNA of E. griseum looks like. Our DNA matches one of 2 possibilities so far, and may not be the right one. Rough spore measurements for our species are 8-9x7.5-8.5u and 8-9x6.5-9u.
Entoloma nidorosum EU - a closely related species but nitrous smelling . We have found one WA collection with ITS DNA only a couple of base pairs different from EU sequences.
Entoloma sericatum EU - supposedly with a slight nitrous odor only when fresh, then farinaceous. We have dozens of EU sequences and BC and WA sequences that match quite well.
Entoloma abbreviatipes WA (=Nolanea abbreviatipes) - with a short stem. It was originally described as a Nolanea, as it is smallish and without clamps, but the type sequences inside Entoloma instead, which I cannot yet explain. It can be a rather small species, but the photo of our one photographed collection (2 bp different than the type sequence, but not all of ITS was obtained) shows it as somewhat stocky like an Entoloma. The newer EU species Entoloma griseorugulosum has a similar type sequence, 3 bp and 2 indels different, one of the indels a long chunk.
Entoloma cf griseum © Daniel Winkler and Sharon Squazzo, E. nidorosum © Midy Liou
Entoloma sericatum (2 photos) © Richard Morrison, E. abbreviatipes © Yi-Min Wang
Especially large, stocky species.
Entoloma lividoalbum EU (=E. saussetiense EU) - a quite stocky grey-brown capped species. There are so many similar species almost every collection sequences to a different species, but in 2015 Kokkonen thought he got a sequence of the real thing, and Brandrud agreed in a 2018 paper, so let's go with that. It's the same as the newer species E. saussetiense. We haven't found that DNA locally yet, but keep reading.
Entoloma lividoalbum forma inodoratum CA - Largent described a local distinct form, forma inodoratum CA for collections lacking the farinaceous odor. We need collections of that to see if they have the same ITS sequence as farinaceous collections. Keep reading.
Entoloma 'lividoalbum PNW02' - a probably somewhat farinaceous BC sequence is >1% different than E. lividoalbum. One WA sequence with no odor detected had too short a sequence to tell if it matched the BC sequence or EU sequences. It should be resequenced.
Entoloma 'lividoalbum PNW03' - another very short WA sequence is different enough from any other known sequence that it may represent another species in this group. It too should be resequenced. Further studies are need to tell how many species are in this group here and what their proper names should be.
Entoloma 'majaloides PNW06' - this stocky species, found once in summer in WA, has a sequence almost 2% different than sequences that match the short type sequence.
Entoloma subsaundersii CA - a large, stocky species that may remain partially buried. We don't have ITS DNA yet nor any sequenced local collections.
Entoloma 'lupinum PNW04' - this related species may or may not be stocky, stay tuned. It is 1.5% different in ITS from the type sequence of E. lupinum.
Entoloma 'lividoalbum PNW02' © Paul Kroeger, E. 'lividoalbum PNW03' © Jan Agosti, E. 'majaloides PNW06' © Yi-Min Wang (2 images)
Locally described so definitely present, but still little understood
Entoloma griseoavellaneum WA - known only from the type collection, it has an unusually fibrillose cap and a fragrant odor. We need collections and sequences.
Entoloma heracleodora WA - a smoky green capped Entoloma with a farinaceous and slightly celery odor. We need collections and sequences.
Entoloma lyophylloidium WA - a nearly black species under fir at high elevations.
Entoloma pseudocostatum WA - with interveined gills.
Poorly understood non-local species that have been reported here
Entoloma alnobetulae EU - we have a very short, dirty type sequence, but hopefully that's enough to recognize it by. It's very similar to another short type sequence of Entoloma bisporigerum. We need local collections to confirm what the PNW reports of this are.
Entoloma alpicola EU - formerly a variety of E. clypeatum. Different EU sequences are clearly different species, so I don't know what the DNA of this looks like. We need reliable EU sequences as well as local sequences to see what reports of this species really are.
Entoloma brunnescipes CA - the stem discolours brownish-grey in age. There is one questionable report of this CA mushroom from Idaho. We need local collections to confirm its presence here, and also CA collections because we don't have its DNA yet.
Entoloma clavaformipes FL - with a club shaped stem base. This FL mushroom was reported from BC but we need both FL and local collections to figure out what its sequence is and to verify that it does occur here.
Entoloma mymecophilum EU - with a blackish, matted-felty cap. There is one questionable rumour of it maybe being in WA, so be on the lookout for it. We have EU type area DNA to compare to, but it hasn't been found here yet.
Entoloma speculum EU - supposedly with a very pale cap. every EU sequence of this is a different species so I don't know what the DNA looks like. Neither do we have any sequences of local collections.
Nolanea - click to expand
Nolanea are small and slender (colybioid or mycenoid), smooth, lubricious capped species with grey or brown tones without clamps in the cap cuticle. They are very difficult to differentiate from each other without a scope. Entoloma are larger (tricholomatoid) and have clamps in the cap cuticle. One species may be omphalinoid, easily confused with Paraeccilia and the omphalinoid Entoloma politum group. The other genera on this page are more colourful or not smooth and lubricious.
Species mentioned: Nolanea abbreviatipes, bicoloripes, californica, cetrata, clandestina, cuspidifera, edulis, farinogusta, fructifragrans, fusciceps, fusco-ortonii, hebes, hirtipes, holoconiota, incanosquamulosa, latifolia, minutostriata, obscurata, occidentalis, seattlense, washingtonense, papillatoides, pseudopapillata, proxima, proxima forma inodorata, pseudostrictior, pseudostrictia, pusillipapillata, sericea, staurospora f. discoloripes, conferenda, staurospora var. incrustata, staurospora var. farinacea, strictior, subcapitata, solstitialis, subsolstitialis, substrictior, subviolaceoverna, terrea, undatomarginata, undulata, verna var. isodiametrica. Entoloma leptotus, propinquum, rhodocyclix, psammophilohebes.
'Nolanea' abbreviatipes WA - with a short stem, can be small like a Nolanea, and lacks clamps, but the type sequenced inside Entoloma, which I cannot yet explain. See that genus.
1. Omphalinoid - pale grey with few gills
'Entoloma' rhodocyclix EU - an omphalinoid Nolanea resembling the Entoloma politum group and Paraeccilia. One BC soil sample sequence is 2 bp different from three UK sequences provided by Vila in a 2014 paper, but a fruiting body had never been found. Perhaps until now. This OR collection resembles it and is being sequenced now. I think it would need to be moved to Nolanea, although Vila's paper couldn't place it definitively inside Nolanea (but his sampling was poor). My ITS tree shows it inside Nolanea.
'Entoloma' cf. rhodocylix © Connor Dooley
2. Spores angled so strongly they resemble a 4-pronged jumping jack. Farinacceous.
Nolanea conferenda EU (=N. staurospora EU) - a matching sequence has been found in BC. Largent described (using the name N. staurospora) forma discoloripes from CA, var. incrustata from WA, and var. farinacea from WA on the basis that all some collections have a stem that bruises brown in places instead of becoming entirely dark brown and tiny microscopic differences. The latter two varieties are said to grow in smaller groups (of 1-4 instead of 10 more more) and are now thought to be synonymous with each other. A BC sequence is only 1 bp different than EU sequences of N. conferenda, so I don't know that the different varieties are genetically distinct. We should sequence more collections matching the various varieties and the types.
Nolanea conferenda © Anna Mladzyanovskaya
3. Pruinose stem over 1/3 of the length, not just at the apex. (Spring, odorless).
Nolanea holoconiota ID ('PNW03') - spring, pale conical cap, odorless with a pruinose stem. We have many local sequences of a species that appears to match N. holoconiota, our most abundant spring species. Even though we have no type sequence to prove it, since this is the only species said to be pruinose over an extended portion of the stem, we have confidence that this is it.
Nolanea cf holoconiota PNW03 © Buck McAdoo
4. Other non-farinaceous, at most faintly so.
Nolanea bicoloripes WA - no DNA yet.
Nolanea 'bicoloripes PNW05' - we have one OR sequence of a collection with this name, but I don't know what it really is. There is a second collection with the same name, but it is inside Cyanula and can't be what it is called.
Nolanea clandestina var. clandestina EU - we have dozens of EU sequences, but no local collections sequenced yet, which we need to see if our darker collections truly are the EU type variety.
Nolanea clandestina var. oculobrunnea CA - with paler colours than the type variety, is 4% different in ITS and truly is distinct. We have CA and WA sequences.
Nolanea clandestina var. oculobrunnea © Yi-Min Wang
Nolanea edulis var. edulis CA - awaiting DNA, we need collections.
Nolanea edulis var. concentrica OR - with the cap olive grey and concentrically rivulose. Many collections do show a concentric indentation around the umbo, but they aren't always olive-grey. I wonder if, when we get DNA of the type variety, this variety will truly be distinct. Entoloma vindobonense EU was recently described and only differs by 2-3 bp plus an ambiguous location or two. I am assuming they are distinct, but it doesn't matter as our name is older.
Nolanea edulis var. concentrica © Buck McAdoo, Bitty Roy, Jonathan Frank and Yi-Min Wang
Nolanea latifolia OR (=Entoloma fuligineocinerum) - we have the type sequence, and a few recent matching sequences. The lumpers had to give it a different name in Entoloma because Entoloma latifolia was taken.
Nolanea latifolia © Gabriela d'Elia and Yi-Min Wang (3 images)
Nolanea strictior NY - we have more than a dozen east coast sequences, but no local collections yet to prove it is here.
Nolanea subcapitata WA - we have a the type sequence and a number of matching WA sequences.
Nolanea subviolaceoverna WA - with the type sequence only 2 bp different from more than a dozen EU sequences of N. verna EU, this may not be a distinct species. N. verna is said to have longer spores, so this relationship should be investigated.
Nolanea verna EU var. isodiametrica OR - if one local sequence is to be believed, this local variety is 10% different than the many EU sequences we have of N. verna, so it needs to be promoted to species. We need a type sequence or more reliable sequences to verify this. If the OR sequence is not this, what is it?
5. Distinctly farinaceous
Nolanea cetrata EU - Largent examined collections with 3 ranges of spore sizes and number of spores per basidia, and described forma mediospora from WA and forma minimospora from CA, but EU sequences of the type form, Russian and a half dozen PNW sequences of unidentified forms, and the paratype sequence of forma minimospora from WA all match within a single base pair, so I do not think the forms have any genetic significance.
Nolanea cetrata © Yi-Min Wang
Nolanea hebes EU (=Entoloma leptotus EU) - E. leptopus is thought to be the same species. E. hebes and N. hebes are the older names and take precedence. We have a dozen EU sequences all within 4 bp of each other, but we need local collections.
Nolanea hirtipes EU - a sharply conical cap, very long and slender stem that can stain brown, and strong rancid-farinaceous odor make this is an attractive, distinctive Nolanea. Some EU sequences match ours almost exactly, even though in the EU it is said to typically have a darker cap and not stain in the stem.
Nolanea 'hirtipes PNW02' - >1% different in ITS, but quite small in comparison. It still has the conical cap and very long stem, but has a strongly striate cap and fewer gills.
Nolanea hirtipes and 'hirtipes PNW02' (2 images) © Yi-Min Wang
Nolanea incanosquamulosa WA (=Entoloma brunneosericeum EU?) - we have the type sequence and a recent matching sequence. The more recently described Entoloma brunneosericeum from the EU has an type sequence that only differs by 1 bp and may be a newer synonym.
Nolanea incanosquamulosa © Luca Hickey
Nolanea obscurata WA - we have the type sequence, but no photos yet. It is 3% different from Nolanea cetrata in ITS.
Nolanea sericea EU - in grass, a dark robust species, umbonate, farinaceous. One WA sequence matches the EU epitype sequence, but I'd expect we'd have many sequences of this by now given how abundant it supposedly is. We should investigate how common it really is.
Nolanea fusciceps WA - an ITS1 sequence of a 1925 collection that is the type (or might as well be) is 4 bp different than N. sericea, but distinct. One recent CA collection was photographed and sequenced, providing ITS2.
Nolanea sericea (2 images) © Eric Chandler, N. fusciceps from CA © Julien Pometta
6. unknown odor so far
Nolanea cuspidifera EU - not previously known in the PNW, but one OR collection in sphagnum moss is a good match to that species and matches one EU sequence labeled by that name.
Nolanea cuspidifera © Connor Dooley
Nolanea solstitialis EU - one WA sequence is 1 bp different than a trusted EU sequence. But see below
Nolanea subsolstitialis WA - Largent described this new species from WA as much like N. solstitialis, but did not report N. solstitialis from the PNW. We need the type sequence or other collections of N. subsolstitialis to verify that it is indeed different than N. solstitialis, given that we have found a local collection with DNA so close to N. solstitialis.
Nolanea solstitialis © Yi-Min Wang
Nolanea 'terrea PNW06' - one OR collection has a sequence 5-7 bp and a couple of indels different than almost a dozen EU and ENA sequences of N. terrea. It was farinaceous.
Nolanea solstitialis © Danny Miller
Nolanea 'psammophilohebes PNW08' - three OR sequences are between 3 and 6 bp different from the type sequence of Entoloma psammophilohebes. It should be investigated if ours is the same species, and if so, would need to be renamed to Nolanea.
Nolanea 'psammophilohebes PNW08' © Jonathan Frank
Nolanea PNW01 - two recent sequences (OR + WA) are a unique species. Perhaps it is one of the below poorly understood species.
Nolanea PNW01 © Tom Bruns
Nolanea PNW07 - a unique WA sequence. No notes yet, perhaps it is one of the below poorly understood species.
Nolanea PNW07 © Yi-Min Wang
Little known species without any DNA yet. The ones described from the west are definitely here.
Nolanea fructifragrans WA - unique sweet/floral fruity odor instead of the usual farinaceous or mild odor.
Nolanea californica CA -
Nolanea fusco-ortonii CA -
Nolanea minutostriata CA -
Nolanea occidentalis WA (=Entoloma seattlense WA, =E. washingtonense WA) -
Nolanea papillatoides CA (=N. pseudopapillata Largent CA) -N. pseudopapillata was already taken (can you believe that?) so it had to be renamed.
Nolanea proxima forma inodorata CA -
Nolanea pseudostrictior CA (Nolanea pseudostrictia?) - when Nolanea strictia was corrected to N. strictior some say this should be corrected too.
Nolanea undatomarginata CA (=N. undulata Largent) - the first name for it, N. undulata, was already taken so it was renamed.
Nolanea farinogusta EU -
Nolanea proxima forma proxima (=Entoloma propinquum CA) - The name Entoloma proximum was already taken, so the lumpers had to come up with a unique name in Entoloma.
Nolanea pusillipapillata CA -
Nolanea substrictior CA -
Leptonia - click to expand
These beautiful scaly capped blue to purple-black mostly slender mushrooms are hard to distinguish from 'Cyanula', the group of Leptonias that have to be split into a new genus. The true Leptonia are more likely to be found attached to wood, which does not as typically happen with Cyanula, which are more likely to be reported in grass. However, both genera are commonly reported from the forest floor. Also, Leptonia caps are usually the same kind of scaly throughout the cap, although the scales can be more concentrated on the disc. Cyanula caps can be scaly on the disc, but more fibrillose or even smooth towards the edge. Finally, unlike Cyanula, Leptonia usually have clamps in the cap cuticle. Thanks to Ben McCormick's 2021 bachelor's thesis that incorporated DNA, Leptonia are some of the best understood Entolomas in the PNW.
Leptonia may have only one single local species that is not blue at least somewhere on the fruiting body.
Species mentioned: Leptonia cyanea var. occidentalis, tjallingiorum var. laricinum, insueta, lampropus, subeuchroa, convexa, zanthophylla, occidentalis, occidentalis var. metallica, occidentalis var. fibrillosipes, coelestina, subcoelestina, violacea, dichroa, subgracilis, violaceonigra. Entoloma kauffmanii.
Leptonia 'not blue PNW01' - this olive-yellow-brown species really resembles something in the 'Cyanula' formosa group and I do not yet know how to differentiate it. My ITS tree shows a much better than even chance that this non-blue species is inside Leptonia, which is surprising, as I believe it would be the only non-blue Leptonia. A multi-gene study should be done to verify its position.
Leptonia 'PNW01 non-blue' © Danny Miller
Non-slender species growing directly on rotten wood.
Leptonia tjallingiorum var. laricinum Russia (=L. cyanea var. occidentalis CA) - growing on rotten conifer wood, capable of being our largest Leptonia. The stem is fibrillose and the cap usually has brown tones as well as blue. DNA showed that we do not appear to have the type variety of L. tjallingiorum in the PNW, but instead the very similar var. laricinum, although it does not necessarily grow with larch. L. cyanea var. occidentalis is a newer synonym, sharing the same ITS DNA and had been tentatively separated out as a unique species based on slight differences in colour and odor.
Leptonia violaceonigra WA - said to be a medium sized, pure blue species with bluish young gills growing on rotten wood. We do not have any sequences and need local collections.
Leptonia tjallingiorum var. laricinum © Andrew Parker
These are exceptionally small species that are pure blue, growing on the ground.
Leptonia coelestina EU - a very small species (caps 1cm across and stems 1mm thick) with white young gills. It grows on the ground sometimes in moss or grass, not on wood. Rare clamps in the cap cuticle. We have Russian DNA of this EU species, but it has not been sequenced locally. We need collections.
Leptonia subcoelestina WA - very similar to L. coelestina, but with a blue tinge to the young gills. No clamps in the cap cuticle (but clamps are found elsewhere). We still do not have DNA of this species yet and need collections to show that it indeed is distinct from L. coelestina.
Leptonia violacea EU (=Entoloma kauffmanii EU) - also very similar to L. coelestina, with white young gills and more abundant clamps in the cap cuticle. We do not have any sequences yet from anywhere to know what this is or if it really occurs in the PNW as reported. We need collections.
Somewhat slender species that are usually pure blue, growing on the ground.
Leptonia dichroa EU - Ben's study found this species for the first time in the PNW. It is a slender species that has a granular or fibrillose bluish cap (not truly scaly) that is often a different shade of blue from the stem. It grows on the ground, not on wood.
Leptonia occidentalis var. occidentalis OR - a somewhat slender species with a striking dark steel blue-black cap and stem without brown tones. The gills are usually bluish when young as well. It grows on the ground, not on wood. This variety has a bald stem and lilac young gills. We believe we now have ITS DNA of this very rare local species.
Leptonia occidentalis var. metallica WA (=Leptonia occidentalis var. fibrillosipes WA) - this variety has a fibrillose stem and bluish-grey young gills. These two additional varieties were described based on slight differences in spore size and the fact that the former variety had a more iridescent stem, but a collection matching each had the same ITS DNA so the two varieties are being synonymized.
Leptonia tjallingiorum var. laricinum © Jessica Javor, L. occidentalis var. occidentalis © Scott Loring and Drew Henderson, L. occidentalis var. metallica © Ben McCormick
These species have at least a hint of brown or grey in the caps, which are not pure blue. Growing on wood or the ground.
Leptonia lampropus EU var. insueta WA - a slender species with a mostly brown cap perhaps with orange tones, and a dark blue stem with occasional black scales. It grows on duff or woody debris. L. insueta was provisionally reduced to a variety of L. lampropus, as their ITS DNA is within 1% of each other and their features are practically identical except for geographic range.
Leptonia subeuchroa OR (=Leptonia convexa CA, =Leptonia zanthophylla WA) - a slender species with some red-brown tones on the disc only in age (the rim stays blue). The young gills are whitish and it may grow on woody debris or on the ground. This seems to be our least uncommon species.
Leptonia subgracilis WA - greyish-brown cap with no blue on the rim that is concentrically grooved and the usual blue stem. It grows on humus not on wood. We don't have the type sequence but we think we have one ITS1 sequence of what is probably this species, except for the fact that our collection didn't have the tomentose stem apex. It would be nice to find and sequence a collection with that character to double check. Leptonia approximatum from northern CA differs only by more sharply angled spores and is to be look for here too.
Leptonia lampropus var. insueta © Ben McCormick, L. subeuchroa © Yi-Min Wang, L. subgracilis © A & O Ceska
Cyanula - click to expand
Cyanula needs to be split from Leptonia, but the genus hasn't formally been erected yet. These colourful, scaly capped mostly slender mushrooms are hard to distinguish from Leptonia when they are blue. Other colours besides blue are typically not found in Leptonia. It should be noted that even brown Cyanula have interesting pigments that makes them recognizable and stand out as from among all the "ordinary" brown mushrooms of the world. Cyanula are typically found on the ground, sometimes in grass, whereas Leptonia are more often found attached to wood and aren't as often reported from grass. However, both genera are commonly reported from the forest floor. Cyanula caps can be scaly on the disc, but more fibrillose or even smooth towards the edge, whereas Leptonia caps are usually the same kind of scaly throughout the cap. Finally, unlike Leptonia, Cyanula don't have clamps in the cap cuticle.
Species mentioned: Leptonia acutoumbonata, albida, albinella, anatina, asprella, atrifucata, badissima var. badissima, badissima var. longisporum, caesiocincta, chalybea, coacta, decolorans, decolorans forma cystidiosa, earlei, exalbida, exilis, foliocontusa, formosa, formosa var. microspora, fuligineomarginata, gracilipes, grisea, incana, lividocyanula, lutulenta, microspora, nigrosquamosa var. californica, ovatospora, parva, pseudobulbipes, rectangula, rosea var. marginata, rostrata, serrulata, sodalis, striatula forma farinacea, strictipes, subnigra, subrubinea, subviduense, subviduense var. marginata, trichomata, turci, umbilicata, undulatella. Entoloma hesleri, subfurfuraceum, squamatum, nigroviolaceum var. striatulum, unbiliciforme.
Sequenced local collections - unnamed species could be one of the reported species in the unsequenced section below.
Blue cap and stem
Cyanula 'serrulata PNW07' - blue/black marginate gill edges like the EU species Cyanula serrulata, but we have DNA of that and this is not it. Does C. serrulata occur here too, as reported? It hasn't been verified so far. C. caesiocincta below has a mostly brown cap.
Cyanula PNW04 - lacks the strongly coloured gill edges (but a slight gill margination should be investigated as a couple photos appear to show that and it is closely related to C. serrulata). Unlike other Cyanula, has been found growing on rotten wood in 2 of the four collections.
Cyanula PNW05 - one WA and one OR sequence are unique, and were incorrectly labeled Leptonia convexa, which may mean this species has a blue cap that fades to brown in places, while still retaining some blue colours. No known photos.
Cyanula PNW08 cf atrifucata WA/cf foliocontusa CA/cf striatula forma farinacea WA (=Entoloma nigroviolaceum UK var. striatulum TN forma farinacea WA) - four OR collections, with at least one of each of these three names, had the same sequence. It is not known which species they actually are (or if it's none of the above). All three have been locally reported and we need reliable collections of each of them.
PNW06 cf rectangula CA - one OR sequence with this name has a unique
sequence. We need to figure out what it is, and if it's not C. rectangula,
what that species' sequence actually looks like as it has been reported locally.
It has an almost black cap and stem.
Cyanula 'serrulata PNW07' (on rotten wood) © NAMA and the Field Museum of Natural History, C. PNW04 © Steve Ness
Brown cap, blue stem
Cyanula PNW03 - a single WA collection
Cyanula PNW09 - not all photos looked like the same species, one was striate and entirely orange, and that should be double checked.
Cyanula PNW03 © Sharon Squazzo, C. PNW09 © Danny Miller
Orange/brown cap and stem
Cyanula formosa EU - larger and not umbilicate. EU and ENA and WNA sequences all seem to agree what this species is
Cyanula 'formosa PNW02' - one sequence from WA is 3% different and may represent a distinct sister species. The spores seemed roughly the same size.
Cyanula formosa var. microspora CA - with smaller spores, still needs to be sequenced to see if it is genetically distinct from the type variety. It is even possible that this could be PNW02, but at first glance, those spores did not appear smaller.
Cyanula PNW01 - smaller than C. formosa and umbilicate. Not all photos of this species looked the same, one had a brown cap and blue stem and should be double checked.
Cyanula formosa © Yi-Min Wang, C. 'formosa PNW02' © Yi-Min Wang, C. PNW01 © Buck McAdoo and Yi-Min Wang
Species reported from the PNW without local DNA yet (unnamed species above could be one of these. All species are currently named Leptonia and/or Entoloma)
Cyanula incana EU - we have EU and ENA sequences that match, but no local DNA yet.
Pink cap and stem
Cyanula rosea ENA? var. marginata WA -
Cyanula subrubinea CA -
Blue cap and stem
Cyanula chalybea var. chalybea EU - I have official EU sequences to compare to and we haven't found this DNA here yet.
Cyanula decolorans forma cystidiosa TN -
Cyanula decolorans forma decolorans NZ - no ITS, but other genes might be available
Cyanula nigrosquamosa TN var. californica CA - no ITS, but other genes might be available
Cyanula parva NY -
Cyanula subviduensis var. marginata CA -
Cyanula subviduensis var. subviduensis CA -
Almost black cap and stem
Cyanula subnigra FL -
Cyanula trichomata WA - no ITS
Brown cap, blue stem
Cyanula caesiocincta EU - black marginate gill edges, cap mostly brown with some blue on the rim. One Kokkonen sequence may represent this.
Cyanula acutoumbonata WA -
Cyanula anatina EU - we have one short purported sequence that might be from the UK to compare to
Cyanula asprella EU - we have 2 competing sequences, maybe from the UK, as to what this might be
Cyanula coacta WA -
Cyanula gracilipes MA -
Cyanula lividocyanula UK -
Cyanula microspora WA -
Cyanula sodalis EU -
White species (easily confused with Alboleptonia, but those have clamps in the cap cuticle which is composed of entangled hyphae instead of erect hyphal hairs)
Cyanula albida NC (=Entoloma hesleri?) - I should note that there is confusion over two mushrooms named Entoloma albidum, one of which needed changing to Entoloma hesleri. I'm currently following IF in thinking that ours is E. hesleri, especially since a sequence we have labeled E. albidum is in section Rhodopolia and not Cyanula.
Cyanula albinella NY - we have a TN sequence purporting to be this to compare to
Orange/brown cap and stem
Cyanula fuligineomarginata NY - reddish brown, the others are mostly brown, but an "interesting" brown.
Cyanula badissima var. badissima and var. longisporum WA -
Cyanula exalbida CA -
Cyanula exilis EU - no ITS, but other genes might be available
Cyanula earlei Cuba (=Entoloma subfurfuraceum Cuba) -
Cyanula grisea NY (=Entoloma squamatum NY) -
Cyanula lutulenta CO -
Cyanula ovatospora WA -
Cyanula pseudobulbipes WA -
Cyanula rostrata WA -
Cyanula strictipes MA -
Cyanula turci EU - EU + Asia + NA sequences seem to match, no local sequences yet
Cyanula umbilicata NY (=Entoloma umbiliciforme NY) -
Cyanula undulatella NY -
Alboleptonia - click to expand
Slender, all white mushrooms. Rarely, a Cyanula will be all white, but those lack clamps in the cap cuticle and don't have a cap cuticle of entangled hyphae, but instead of erect hyphal ends)
Species mentioned: Alboleptonia earlei, ochracea, sericella var. lutescens, adnatifolia
Alboleptonia 'sericella PNW01' (A. sericella var. lutescens EU misapplied?, =A. adnatifolia CA?) - said to discolour yellow, our least uncommon species is said to be A. sericella var. lutesens EU, but a sequence of what is probably our most common species does not match a couple of EU sequences purporting to be the type variety of A. sericella. This probably means that our species needs a new name. Alboleptonia adnatifolia CA is a name that is currently assumed to be a synonym of A. sericella, but perhaps it is the correct name for this or another of our Alboleptonias.
Alboleptonia ochracea WA - said to discolour orange-brown. A collection matching this description from CA was sequenced, so we might have a sequence of this now. We need local collections.
Alboleptonia earlei Cuba - said to remain white and not discolour. We have no sequences at all yet from anywhere, nor any photos.
Alboleptonia 'sericella PNW01' © NAMA and the Field Museum of Natural History, A. cf. ochracea© Joann Olson
Prunuloides - click to expand
A large, viscid, blue, farinaceous Entoloma. The cap and stem are usually blue, but the colour may fade in age to brown, making it hard to identify. Rarely, it seems to fruit lacking any blue pigmentation even from the start.
Species mentioned: Entoloma bloxamii, medianox, madidum, caesiolamellatum
Entocybe seems to be near this clade in multi-gene studies, even though it is not so in my ITS only tree. This would mean one could consider Prunuloides and Entocybe as a single genus if one wanted to. Furthermore, by some accounts, the type species of Entoloma is Entoloma prunuloides, so this section would need to be called Entoloma, and my Entoloma section would need to be called Rhodopolia. To keep more names stable, an exception could be made to conserve that section as Entoloma, but I cannot yet predict how this will eventually shake out.
'Prunuloides' medianox CA - still officially in the genus Entoloma. Previously this has incorrectly been called Entoloma bloxamii and Entoloma madidum, which are similar EU species that have not been found here. California has a second species that is also blue in the gills, a sister species of the EU species 'Entoloma' caesiolamellatum, which should be looked for in the PNW.
unsequenced 'Entoloma' medianox © Noah Siegel, 'Entoloma' medianox © NAMA and the Field Museum of Natural History
Entocybe - click to expand
Entocybe nitida has a beautiful blue opaque cap and stem. It is medium sized with a dry cap and little odor, smaller than the large 'Prunuloides' medianox (with a viscid cap and stronger farinaceous odor) but larger than the many blue Leptonia and 'Cyanula'.
Entocybe trachyospora complex members are smaller with a pointy transulcent cap, and may have a hint of blue in the cap and a bent, silver, striate stem. When all the blue is faded it is difficult to separate from Nolanea, and when it hasn't, it is difficult to separate from Leptonia and 'Cyanula'.
Species mentioned: Entocybe nitida, trachyospora
Entocybe seems to be near Prunuloides in multi-gene studies, even though it isn't so in my ITS only trees. This would mean one could consider Prunuloides and Entocybe as a single genus if one wanted to.
Entocybe nitida EU - local collections from BC and WA match EU DNA.
unsequenced Entocybe nitida © Steve Trudell
Entocybe trachyospora WA complex - although four varieties are described with varying amounts of blue, all ITS DNA so far is basically identical. If you find a collection of a variety that is not represented here, save it so we can see if any of the varieties have unique DNA. They have not all been systematically tested yet.
Entocybe trachyospora © iNaturalist user mdziak, Danny Miller and Yi-Min Wang
Claudopus and Paraeccilia - click to expand
Claudopus contains the pleurotoid (oyster) Entolomas, but DNA is showing that the concept of Claudopus may need to be expanded to include at least some omphalinoid Entolomas (the type species of Paraeccilia may be inside Claudopus). The partially secotioid Entoloma abortivum is also inside Claudopus.
Species mentioned: Claudopus byssisedus, byssisedus var. microspora, parasiticus. Paraeccilia minutissima, nucisapora, perundata, rusticoides, sericeonitida var. sericeonitida, sericeonitida var. ligniphila. Entoloma abortivum.
Claudopus byssisedus EU - a farinaceous, fibrillose little brown oyster with tufts of rhizomorphs at the base of the eccentric stem. Different EU sequences are clearly different species, so I don't know which is the real species. We have no local sequences yet to know how our collections fit in. We need some.
Claudopus parasiticus EU - supposedly the fibrillose little grey oyster species that grows on chanterelles from a white mycelial mat, but we have no idea yet what its DNA looks like. We need local collections with large spores (see below).
Claudopus 'byssisedus var. microsporus' EU - has smaller spores than the above two species. Two local sequences supposedly of Claudopus parasiticus from OR match EU sequences of C. byssisedus var. microsporus exactly (and are 4 bp from Chinese sequences). Perhaps this is the proper identity of our local collections of C. parasiticus and our local spore size has not been properly checked, or perhaps there is more than one species here or perhaps they are the same species (although that would surprise me). One was growing on a chanterelle, as expected, but the other was just a fuzzy white mycelial mat on a Suillellus (no fruiting bodies). This sequence is not near any of the candidates for Claudopus byssisedus, so it needs promotion to species level.
Claudopus 'byssisedus var. microsporus' on a chanterelle and as fuzz on a Suillus © Jenny Lippert and Jonathan Frank
Paraeccilia sericeonitida EU - one sequence with this name suggests that it may be inside Claudopus. Other Paraeccilia sequences may be elsewhere. We need local and EU collections to make this determination.
Paraeccilia rusticoides EU - no sequences yet from anywhere to verify reports that this species occurs here.
Paraeccilia sericeonitida var. ligniphila ID - the following are locally described and therefore do occur here, but we need sequences of all of them to verify what genus they are in and that they are all distinct.
Paraeccilia minutissima WA -
Paraeccilia nucisapora WA -
Paraeccilia perundata CA - not exactly locally described, but it has been reported from ID.
'Entoloma' abortivum New England - recognized by some secotioid fruit bodies and some regular fruit bodies, the regular fruit bodies with decurrent gills. This species appears to interact with Armillaria. One OR sequence matches an east coast sequence so it seems we do have this species here, but it probably needs to be moved to Claudopus.
'Entoloma' abortivum regular and secotioid form © Jonathan Frank
Pouzarella - click to expand
Small, conical, fibrillose, usually unpleasant smelling mushrooms that resemble dark
Nolaneas. The stem base is usually strigose.
Microscopically, the cap hyphae have incrusted, thickened, brownish walls.
Species mentioned: Pouzarella fernandae, fulvostrigosa, versatilis.
Pouzarella 'versatilis PNW01' - while it had a sequence unlike other sequences with this name, this collection from BC was originally identified as P. versatilis. It's unknown if reports of that EU species are represented by this sequence, or if that species should still be looked for.
Pouzarella fernandae EU -
Pouzarella fulvostrigosa UK - two additional Pouzarellas which have been reported from the PNW, but there is no DNA data yet. We need collections.
Pouzarella 'versatilis PNW01' © Steve Joya
Rhombisporium s.l. and Inocephalus - click to expand
Fibrillose caps, with what is referred to as a scummy disc. Pouzarella and Trichopilus caps also have fibrillose/scaly caps. Formerly all placed in Inocephalus, it is now known that some species belong in a separate genus, which I am calling Rhombisporium. Rhombisporium is currently only a section of Entoloma and for those who do not lump every species on this page into Entoloma, it still needs to be promoted to genus.
Species mentioned: Inocephalus appressus, azureus, fabaceolus, furfuraceidiscus, minutopilus, perfuscus, rhombisporus. Entoloma cocles.
'Rhombisporium/Cocles' PNW01 - DNA shows that this species and Entoloma cocles are basal to Rhombisporium, meaning it's unclear if they should be part of Rhombisporium or a separate genus. They are very dark species with blackish-brown fibrillose caps and grey stems. Microscopically, these two species have heterodiametric spores, meaning they are not "roundish" like true Rhombisporium species, but "elliptical" with a length/width ratio of >1.25 (but the angles on the spores complicate things and make words like "round" and "ellipse" not really apply). Previously, no species was known closer than 18% different in ITS from E. cocles, until this species showed up, only differing by 10-12%.
'Rhombisporium/Cocles' PNW01 © Buck McAdoo
Rhombisporium PNW02 - WA and CA sequences are clearly inside Rhombisporium, as is the next sequence.
Rhombisporium PNW03 - only known from CA so far.
'Rhombisporium' PNW02 © Fred Rhoades, 'Rhombisporium' PNW03 © David Largent
The following species have been described from the PNW (with the exception of Inocephalus rhombisporus) and therefore definitely exist here, but without any DNA data on them yet, I don't yet know if they're properly placed in Inocephalus or 'Rhombisporium', or if they are all unique genetic species. It's possible one of the above unnamed species represent reports of one of the following.
Inocephalus appressus WA -
Inocephalus azureus WA -
Inocephalus fabaceolus WA -
Inocephalus furfuraceidiscus ID -
Inocephalus minutopilus WA -
Inocephalus perfuscus WA -
'Rhombisporium' (Inocephalus) rhombisporus EU - we have a neotype sequence, and it is the type species of section Rhombisporium.
Trichopilus - click to expand
Scaly/fibrillose caps, characters also found sometimes in Inocephalus/Rhombisporium and Pouzarella, but Trichopilus have distinctive cap cystidia not found in those other genera.
Species mentioned: Trichopilus jubatus, plebeioides.
Trichopilus cf jubatus EU -
Trichopilus cf plebeioides EU - both species have been reported from the PNW, but we need local collections to confirm them. One possible collection of T. plebeioides was sequenced from CA, matching an unconfirmed sequence from Lebanon.
Trichopilus cf. plebeioides © Harte Singer
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