Danny’s DNA Discoveries – Inocybaceae of the PNW
Recognized by a usually tattered appearance, sometimes called "fibre
heads". I mention unusual groups of mushrooms (like
Psathyrella) that have a cellular cap
cuticle, composed of spherical cells that can break up in any direction. In
contrast, Inocybe is the poster child for the typical kind of mushroom
cap. Made up of long fibrous material, the caps will usually only split
radially from the centre out to the edge of the cap. The smooth capped
species are hard to recognize, but the typical Inocybe cap is dry, fibrillose or scaly,
not hygrophanous and often umbonate. There
is often a small bulb at the base of the stem, and the stem apex may
be pruinose - these are important characters to note when trying to identify
one. They are mycorrhizal, and can also be recognized by their odor,
usually an unpleasant spermatic smell. They also usually have a
cortinate veil like Cortinarius, and
are actually very easily confused with some Telamonias, which usually
lack scales on the cap, lack the spermatic odor and never have a bulb. Most
Inocybes are suspected to be poisonous, containing muscarine, and due
to their being ubiquitous near almost every forested area, they are responsible
for many of the accidental mushroom poisonings by children and pets. Inocybe
spores may take a while to mature, leaving the gills white for a long time, so
it is hard to tell that they are brown spored until they are old. This is a good
reason to always consider other possible spore colours unless you have proven
what the spore colour is by taking a print. Very difficult to tell apart,
Inocybe is one of those genera that most identifiers cannot usually get to
species. Only a few species are colourful, and the rest are usually just shades
Inocybe metuloids © Danny Miller and nodulose spores © A and O Ceska
Ditte Bandini is one of the top experts in Europe, providing us many type sequences and other reliable sequences of European species. Our own Brandon Matheny (former PSMS educator who got his PhD at UW and is now at U Tennessee) is the undisputed North American expert on Inocybe, taking the reins from Daniel Stuntz, former UW mycologist before Joe Ammirati.
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.
Click here to download the FASTA data of all my DNA sequences
Summary of Interesting Results
Inosperma, Mallocybe and Pseudosperma - click to expand
The species without metuloids (and never with nodulose spores) probably represent the oldest lineages, before either of those traits evolved in the family. Although as a group these three genera are easy to separate from Inocybe with a microscope, they are more difficult to recognize individually. I think I would have preferred considering them all to be Inocybe, but then other mushrooms around the world that don't seem quite so Inocybe-like would have to be called Inocybe as well, and those people might have objected. For a more in-depth key to the individual genera and much more information, see Brandon's paper.
Species mentioned: Inocybe calamistrata, hirsuta var. maxima, mucidiolens, atrovirescens n.p., maculata, lanatodisca, fibrillosa, subdecurrens, dulcamara, terrigena, delecta, leucoblema, malenconii, breviterincarnata, flavella, holoxantha, niveivelata, obsoleta, occidentalis, bulbosissima, rimosa, sororia
Inosperma calamistratum group
These are recognized by being distinctly scaly all over, brown but with blue-green spots on the stem base and sometimes turning red where handled. I think a study showed that this genus contains neither Psilocybin (the bluing is unrelated) nor the poison muscarine unlike most in Inocbye s.s. The odor is raw fish, pine resin or green corn, not the usual spermatic odor. Here are the species we have in this group:
Inosperma 'calamistratum PNW01'
Inosperma maximum (=Inocybe hirsuta var. maxima) WA - usually a bit more robust, more red staining and fewer blue areas.
Inosperma mucidiolens/atrovirescens n.p. - when I. calamistratum var. mucidiolens was described from Nova Scotia (now raised to species status), it was said to be present in WA too, but WA sequences are 3 bp and 2 indels or so different from the Nova Scotia type sequence, and ours may be larger with a blackening stem so it is currently being studied to see if ours needs a new name. If so, it will be called Inocybe atrovirescens. Other genes may show a higher divergence than ITS does, supporting the idea that it needs a new name. It supposedly has a more moldy or green corn odor, whereas the others supposedly have more raw fish and pine resin odors, but that has not been confirmed for all the species mentioned above in the I. calamastratum group. I don't want to place too much imporance on one photo of each, but PNW01 was orange-brown, PNW02 was dark brown and PNW03 was olive brown.
Inosperma 'calamistratum PNW01' © Erica Son, I. 'calamistratum PNW02' © iNaturalist user fungi_fanatic, I. 'calamistratum PNW03' © Luca Hickey,
Inosperma maximum © Yi-Min Wang, I. mucidiolens/atrovirescens n.p. © Betty Hsu
Inosperma maculatum group
These nondescript brown fibrillose to scaly capped mushrooms are separated from Inocybe by their lack of metuloids, and from Mallocybe and Pseudosperma by having an almost naked stem (less fibrillose-scaly than most in the family). The basidia are all clear (not brown) under the scope. These species may have an umbo covered with bits of white veil material and a little stem bulb. They have a complex odor. We so far have not shown that the European I. maculatum itself is present in the PNW, but here are the two species we do have that have been mistaken for it:
Inosperma cf maculatum - our least uncommon species in this group appears to be quite distant genetically from the EU species an in need of its own name.
Inosperma lanatodiscum MI - similar, but a different shade of brown and with a different complex odor. We have a MA sequence that Kropp says is close enough to the type area of MI to represent the real thing. One dirty WA sequence is a pretty good match once cleaned up, but we could use more sequences.
Inosperma 'lanatodiscum PNW01' - 7 bp and 1 long indel different from I. lanatodiscum, we also appear to have a potential sister species in the PNW. It is yellowish, making it a lookalike of Pseudosperma, but perhaps not as straw coloured nor as tattered.
Inosperma maculatum group member © Danny Miller, I. 'lanatodiscum PNW01' © Yi-Min Wang
Mallocybe have more fibrillose stems than the Inosperma maculatum group and are not usually straw coloured and strongly conical like Pseudosperma is. Also unlike those two, these have basidia that are brown under the scope. Brandon will soon have a paper out giving names to the unnamed species and clarifying their differences. Stay tuned.
Mallocybe dulcamara EU and Mallocybe terrigena EU have been reported from the PNW, but they do not appear to be present here yet. Those collections were probably one of the lookalikes below.
Mallocybe fibrillosa NY (=M. subdecurrens NY?, =Mallocybe delecta EU?) - a wooly ochre species (yellow- to orange-brown). Brandon's east coast sequences of M. fibrillosa and M. subdecurrens are practically the same, different by a couple bp and a few ambiguous locations, so they may be synonymous, with M. fibrillosa being slightly older. However, he hypothesizes that the Mallocybe delecta EU may be the same, which is even older, but we have no sequences of that yet to prove it. This is one of the species we have been calling Mallocybe dulcamara EU in the PNW. We don't have reliable sequences of that species either, but it is not likely to be this.
Mallocybe leucoblema EU - our one WA collection sequences 4bp from EU sequences. It seems to be an interior Rocky Mountain species.
Mallocybe leucoloma EU - a slender yellow-brown species covered in copious white universal veil material when young. This European species alpine willow species is commonly found in the Rockies, but a single WA collection has been found for the PNW. ITS wasn't sequenced, but other genes match.
Mallocybe malenconii EU - a pale brown species with crowded gills. There are 2 species going by this name, but if the more common one turns out to be the real thing, that's the one found once in south eastern BC. It is normally found in the EU, but has occasionally been found back east too.
Mallocybe squamosodisca NY - this east coast species has been sequenced once in WA.
Mallocybe PNW02 - sequenced once on Salt Spring Island BC and in Oregon, and nowhere else yet. No other data so far.
Mallocybe PNW03 - known only from one WA collection, PBM1615, no ITS sequence.
Mallocybe PNW04 - known only from one WA collection, PBM2290, no ITS sequence.
Mallocybe PNW05 - three WA sequences (Swauk and Table Mountain)
Mallocybe PNW06 - one eastern WA sequence near the ID border.
Mallocybe PNW08 - perhaps with a scaly stem and a ring zone, as the 5 BC collections were mislabeled as Mallocybe terrigena.
Mallocybe PNW09 - Pemberton BC and Alaska have sequences of an additional species.
Mallocybe leucoblema © Andrew Parker, M. fibrillosa © Buck McAdoo, M. PNW05 © Shannon Adams, M. PNW06 © Andrew Parker
These are straw coloured, large, umbonate and quite tattered and may smell spermatic or like green corn. The stems are more fibrillose than in the Inosperma maculatum group and unlike Mallocybe, the basidia are clear, not brown under the scope. Kropp did a big study in Utah of this subgenus of Inocybe (now genus Pseudosperma) and found that instead of the 1 species in this group we thought we had, we actually have at least 9 lookalike species. Oh boy. Kropps' paper provides a key to many of the species in this genus.
Pseudosperma sororium MI - we have WA and ID sequences, but we need east coast sequences to prove we have the real thing.
Pseudosperma rimosum EU - reported often from the PNW, but no local DNA matches Kropp's EU DNA yet, so it may not be here. This was the only commonly known species in the genus for a long time, so all our species have long gone incorrectly by this name.
Pseudosperma breviterincarnatum UT - pink young gills. Kropp provided 6 sequences from WA that match his UT type sequence.
Pseudosperma cf flavellum EU - perhaps a brighter yellow cap. Kropp reported it From Idaho but did not provide the sequence, so we still need sequence proof that it is here. He did provide an official EU sequence to compare to, which disagrees with the sequence of most genbank collections labeled this. This should be sorted out.
Pseudosperma aff flavellum - we do have a Victoria BC sequence that clades close to the various concepts of P. flavellum, but doesn't match any of them. This could be what Kropp found in Idaho, or, if he found the real thing, this could be an additional species. See also PNW01.
Pseudosperma holoxanthum WA - also rather bright like P. flavellum. we have a NS paratype sequence but it would be nice to get local sequences of this local species.
Pseudosperma niveivelatum UT - white cap. we have the UT type sequence and 2 matching WA sequences.
Pseudosperma aff. obsoletum EU - pinkish- or brownish-grey cap (most others are yellow-brown). UT and WA sequences are a few % different than Kropp's EU sequences, so we appear to have an unnamed sister species.
Pseudosperma occidentale UT - the one lacking a distinct umbo. A WA sequence matches the UT type sequence.
Pseudosperma bulbosissimum EU - one old WA sequence matches Brandon's EU sequences.
Pseudosperma PNW01 WA - two Washington collections have a unique sequence and likely represent an undescribed species close to P. flavellum. 2 of the 3 collections looked like P. sororium, but one was bright yellow like P. flavellum.
Pseudosperma PNW02 OR - one Oregon collection has a unique sequence and likely represents an undescribed species. I cannot tell it apart from P. sororium.
Pseudosperma PNW03 WA - with bright yellow gills from the Olympic peninsula at 4,000', this WA collection is almost 10% different in ITS DNA than any known species.
Pseudosperma PNW04 WA - this one seems very stocky and grows with willow.
Pseudosperma PNW05 WA - this one (but we've only seen one so far) was strikingly papillate with a dark umbo and rooting stem.
Pseudosperma sororium, PNW01 and PNW02 © NAMA and the Field Museum of Natural History, P. PNW01 (2nd yellow image) © Connor Dooley, P. PNW03 © Steve Ness
Pseudosperma PNW04 © Shannon Adams, P. PNW05 © Yi-Min Wang
Inocybe - nodulose spores - click to expand
Inocybe itself probably evolved both metuloids and nodulose spores from the ancestral genera above.
Species mentioned: Inocybe albodisca, albodiscoides, alpigenes, assimilata, californica, castanea, ceskae, chelanensis, cicatricata, curvipes, decemgibbosa, fallax, grammata, heterochrominea, pseudoteraturgus, maritimoides, bufonia, rangiferi, intricata var. pallidistipitata, jacobi, lanuginosa, leptophylla, mixtilis, napipes, petiginosa, phaeocystidiosa, praetervisa, prominens, radiata, ranierensis, soluta, sphagnophila, stellatospora, suaveolens, subcarpta, umbratica, variabillima, xanthomelas
The macroscopic characters that separate species are cap colour (which shade of brown), how scaly the cap is, whether or not there is a stem bulb and whether or not the stem is almost entirely pruinose (if the species does not have a cortina when young), only at the apex (usually if the species has a cortina) or not at all. This will help get you to a smaller selection of species to choose from, but then you'll probably need the scope again. A pruinose stem has white dots and is usually white (left photo) while a fibrillose stem has longer, stringy white fibers and it is often brown (see right photo).
I. occulta pruinose stipe and I. 'assimilata PNW39' fibrillose stipe © Jacob Kalichman
1. Small species (caps <2 cm across)
Inocybe rufoalba EU (=Inocybe jacobi EU) - small, gills more distant than I. petiginosa. The stem is entirely pruinose. There is no agreement on what sequences of this are.
Inocybe petiginosa EU - very similar, but the gills are not as distant as in I. rufoalba. EU sequences seem to agree what this is, but no matching local sequences yet.
Inocybe PNW33/PNW34/PNW35 - we have no evidence of either of the previous two species in the PNW, but we do have a tight cluster of three small species, only about 1% different in ITS from each other. For now I am considering them separately, as there is more than one matching sequence in most cases. There are no good photographs of any of them yet, we need collections.
2. White umbonate species with stem bulb (and possible fragrant odor) - (smooth spored white species don't have a stem bulb and are much more common)
Inocybe umbratica EU (=Inocybe suaveolens WA) - whitish, umbonate, stem bulb, no veil and entierly pruinose stem. Sometimes it has been noted with a fragrant odor of sweet pea or lily of the valley, in which case it gets the name Inocybe suaveolens. When it has a normal Inocybe odor, it is called Inocybe umbratica, the older name. When a sequence of an original Stuntz collection of I. suaveolens was compared to the available EU sequences of I. umbratica, the sequences were the same, and Brandon confirms that they are the same species.
Inocybe umbratica © Ann Goddard
3. White disc, brown rim (not umbonate, stem bulb, mostly pruinose stem, relatively few nodules on the spores)
Inocybe albodiscoides (I. grammata and I. albodisca misapplied) - the white disc is not always this pronounced when old or weathered, but it is usually more pronounced than the several species described below that have pale universal veil material on their discs. Our sequences are 3% different than EU sequences of Inocybe grammata EU, which is what we used to call our species before it was described as distinct. Inocybe albodisca NY is a synonym of I. grammata and is also not found here. Our species has slightly larger mushrooms with slightly smaller spores than I. grammata, and is found with Doug fir and hemlock.
Inocybe albodiscoides © Sharon Squazzo
4. Dark brown, truly scaly cap and stem even when young, fleeting cortina (the I. lanuginosa group, similar to the smooth spored I. lacera from poor soils)
Inocybe lanuginosa EU - often found on rotten wood, not the ground. BC and WA sequences match many EU sequences. Many varieties have been described, but so far there is no evidence that they are genetically distinct.
Inocybe leptophylla USA - also less likely to be on wood with larger spores. We have 3 US sequences from TN, NY and WY that match and probably represent this species. No local DNA matches yet, even though it's commonly reported.
Inocybe PNW55 - known from 2 BC sequences in this group. No photos yet.
Inocybe stellatospora NY - this scaly capped mushroom is actually near the soluta/subcarpta clade, not in the lanuginosa clade. It is less likely to be on wood with differently shaped metuloids. One old WA Stuntz collection does match a couple of east coast sequences from Brandon.
Inocybe lanuginosa © Yi-Min Wang
5. Yellow-brown smoothish cap, stem bulb, entirely pruinose stem (no cortina). Similar smooth spored species are less common.
Inocybe ceskae EU (Inocybe mixtilis EU misapplied) -the caps can be almost entirely smooth when fresh, not looking like a typical Inocybe. We have 2 WA sequences matching the EU type sequence.
Inocybe occulta EU (Inocybe mixtilis EU misapplied) - difficult to distinguish from I. ceskae, previously both species went by the name I. mixtilis. The shape and size of the cystidia are somewhat different. We have a BC and WA sequence matching the EU type sequence.
Inocybe 'mixtilis PNW61' - overlooked in the study that described the two above species in the I. mixtilis complex, usually a spring/summer grass species (found once in spring, once in summer and once in fall) that was first discovered because it fatally poisoned a small dog in Vancouver, BC. We have one BC and two WA sequences. It is probably very difficult to differentiate from the other 2 species microscopically, but the time of year and/or grass habitat should do it.
Inocybe ceskae © Richard Morrison, I. occulta © Jacob Kalichman, I. 'mixtilis PNW61' © Yi-Min Wang
Inocybe praetervisa EU - perhaps a little larger, with the cap usually somewhat fibrillose. The stem may not be pruinose all the way down. However, the best way to differentiate it from the above 2 species might be by larger spores. The EU type sequence is dirty, but we have cleaner EU sequences, and 8 BC sequences and 1 OR sequence match it.
Inocybe phaeocystidiosa EU - with the larger spores of I. praetervisa but with the stem perhaps entirely pruinose (but hard to differentiate in practice). There are differences between the EU type sequence and our sequences, mostly in ITS2. It should be investigated if this is a species complex.
Inocybe 'xanthomelas PNW38' - very similar to I. phaeocystidiosa but perhaps with a stem that darkens on drying. Our many local BC, WA and OR sequences are a species >15% different in ITS from several EU sequences that likely represent this species.
Inocybe phaeocystidiosa © Yi-Min Wang, I. 'xanthomelas PNW38' © Ann Goddard
6. Plain brown, stem bulb, stem not pruinose or only at apex, cortina when young and may have pale universal veil material on cap. (I. assimilata/napipes clade)
Inocybe 'assimilata PNW39' - These species have fibrillose stems, with white stringy fibers, subtly different than the white pruinose dots of other species. I. assimilata is reported with pale universal veil material on the cap, and the closely related I. napipes isn't, but instead has spores that are more strongly nodulose. We haven't found either species in the PNW, despite both being reported as very common, but we have found this undescribed species differing from I. assimilata by about 10% in ITS. Perhaps reports of I. assimilata in the PNW represent this instead.
Inocybe PNW40 (I. napipes EU misapplied?) - this second local sister species, also different by at least 10% in ITS from I. assimilata (and even more different than I. napipes), represents two of the three supposed I. napipes collections that were sequenced from BC. Perhaps reports of I. napipes in the PNW represent this instead, but we'll need more collections to see if the situation is that simple or not, and how the actual species correlate with the presence of universal veil material on the cap and the spore shape. As you can see from the middle photos below, the two species can look the same and the spore photographs suggest that PNW40 has more nodulose spores, but this is all so far inconclusive.
Inocybe 'sphagnophila PNW31' - 2 OR and 1 WA sequence, 6% different in ITS than I. sphagnophila.
Inocybe 'alpigenes PNW63' - one BC sequence, the closest species is I. alpigenes, 5% different in ITS.
Inocybe 'assimilata PNW39' mushroom and spores © Yi-Min Wang, I. 'assimilata PNW39' © Jacob Kalichman, I. PNW40 mushroom and spores © Yi-Min Wang
Inocybe 'sphagnophila PNW31' © Ann Goddard (2 images)
7. Plain brown, no stem bulb, stem not pruinose or only at apex, cortina when young
Inocybe chelanensis WA - a high elevation spring species with pale universal veil material sometimes left on the cap, looking much like the above group but without the stem bulb. This species has the longest nodulose spores of any local species, up to 19x8u. We have a number of BC and OR sequences of this local species, but no DNA confirmed photograph.
Inocybe curvipes EU (=Inocybe radiata MA, =Inocybe variabillima Brazil) - quite variable in appearance, there are many more synonyms than just the ones given. It is found in urban landscaped areas (including grass or with non-native hardwoods) with the stem often darkening from the base up. Often it has a dark umbo, but it may be slender or stocky. But under the scope you can recognize it by pointy metuloids and oblong spores with only a few nodules that look more like the angles of Entoloma spores or tiny rocket ships. (Yes, some Inocybes are actually referred to as "rocket spore Inocybes"). A few WA sequences match many EU sequences fairly well. Some sequences in both areas differ by as much as 4 bp from each other, but so far it seems to all be one variable species.
Inocybe curvipes © Yi-Min Wang, Ben Woo, iNaturalist user marrstree and spores by Brandon Matheny
Inocybe soluta/subcarpta EU group - stem also darkening from the base up like I. curvipes, but usually with spores that are more strongly nodulose. PNW41 through PNW48 represent eight potential species in this group. None of our species seem to match sequences of either species found in the EU, so our species likely are in need of new names. I probably shouldn't refer to them all as members of the I. soluta/subcarpta group, as only PNW44 and PNW45 have strong support for being closely related. Mostly, I'm placing them all here because they appear to be morphologically similar.
PNW41 - one spring OR collection
Inocybe heterochrominea NS - is closely related to some of the above species and probably looks similar. We have the type sequence, and the authors reported it from WA too, but we have no local collections.
Inocybe PNW41 © Jonathan Frank, I. 'pseudoteraturgus PNW43' © NAMA and the Field Museum of Natural History, I. maritimoides © Shannon Adams, I. PNW44 © Noah Siegel
Reported locally, but no local sequence or confirmed photographs yet - any distinctive features known will be noted
Inocybe californica CA - stem bulb. Kauffman noted it from OR
Inocybe castanea NY - chestnut brown cap. We have BC and WA reports.
Inocybe cicatricata NA - usually small (<2.5 cm across). We have an epitype sequence to compare to. Stuntz thought he found it once in WA.
Inocybe decemgibbosa EU - stem bulb, entirely pruinose stem. We have a dozen matching EU sequence, but no local DNA. Smith reported it once from WA.
Inocybe fallax NY - perhaps an entirely pruinose stem. There are old reports by Kauffman
Inocybe intricata MA var. pallidistipitata WA - small size (<1.5 cm across), stem bulb, entirely pruinose stem. We have a half dozen ENA sequences of the type variety, but no sequences from here of our variety.
Inocybe prominens NY - stem bulb. There are old reports by Kauffman
Inocybe rainierensis WA - stem bulb. Locally described
There are more than 1,000 local sequences of Inocybe in GenBank, and I was not able to look at every single one of them, so more species exist.
Inocybe - smooth spores - click to expand
At least one giant clade of Inocybe lost their nodulose spores (they are smooth spored) but kept the metuloids. We need a multi-gene study to find out how many different clades lost their nodulose spores, but it may be that most of them are in one giant clade. Chances are you're going to need a scope to identify any Inocybe, so I don't feel too bad about making you use a scope to know which section to expand (something I usually try to avoid).
Species mentioned: Inocybe geophylla, whitei, pudica, armeniaca, insinuata, lilacina, pallidicremea, ionocephala, agglutinata, fuscodisca, virgata, fuscicothumata, obscuroides, cincinnata var. major, pyrotricha, griseolilacina, pusio, catalaunica, leiocephala, picrosma, kauffmanii, brunneolipes, semifulva, chalcodoxantha, submuricellata var. stenospermina, nitidiuscula, volvata, vaccina, laetior, cinnamomea, subdestricta, flocculosa, plurabellae, sindonia, melleiconica, oetziana, fraudens, olympiana, microlepidota n.p., griseoscabrosa, hotsoniana, glabripes, fuscidula, chondroderma, lanatopurpurea, lacera, monticola, pseudodestricta,nematoloma, eutheloides, hemileuca, pallidipes, praecox, siskiyouensis, splendens
The macroscopic characters that separate species are cap colour (which shade of brown), how scaly the cap is, whether or not there is a stem bulb and whether or not the stem is almost entirely pruinose (if the species does not have a cortina when young), only at the apex (usually if the species has a cortina) or not at all. This will help get you to a smaller selection of species to choose from, but then you'll probably need the scope again.
1. Inocybe geophylla group - smooth, often umbonate white or lilac caps, but some have brown discs and are hard to place in this group, and other whitish species like I. sindonia (see section 2) are not in this group. These present species all form a natural clade. They all have a cortina when young and lack a stem bulb. The stem apex may be pruinose. Our one known white nodulose spored species has a stem bulb.
Inocybe 'geophylla PNW05' -
abundant as a group, these five are usually slender, white,
often umbonate species. I. geophylla itself has never been recorded from the west coast.
Inocybe PNW62 - neither slender nor umbonate (stature closest to PNW11?) with distant gills.
Inocybe 'geophylla PNW06' © Vail Paterson (2 images)
Inocybe 'geophylla PNW11' © Ryan Downey, I. 'geophylla PNW12' © Sharon Squazzo, I. 'geophylla PNW13' © iNaturalist user hugebooter, I. PNW62 © Yi-Min Wang
Inocybe 'whitei PNW07' - a somewhat stockier white species, not usually prominently umbonate, that stains red where handled or in age. Our ITS sequences are 5% different than EU type area sequences provided by Brandon. Inocybe pudica is a synonym of I. whitei.
Inocybe 'armeniaca PNW08' - more slender and umbonate like the I. geophylla group of species, but reddening somewhat like PNW07. About 10% different in ITS from EU type area sequences provided by Brandon.
Inocybe 'insinuata PNW04' - a stocky chalky white species that is somewhat umbonate and somewhat bulbous. It is 4% different in ITS than Brandon's CA type area sequences.
Inocybe 'whitei PNW07' © Bitty Roy
Inocybe pallidicremea NS - lilac colours when fresh (otherwise much like the I. geophylla group above), slender and umbonate.
Inocybe 'pallidicremea PNW03' - one clade differs by 3 bp and 3 indels from the others in ITS, which isn't a whole lot, but they differ by 6 bp in LSU, which is a lot for that gene, indicating that it most likely is a distinct species.
Inocybe 'lilacina PNW02' - it is unknown how these two species differ from I. pallidicremea.
Inocybe 'lilacina PNW10' -
Inocybe ionocephala CA - a stockier lilac species that can develop a yellowish stem base. Fleeting cortina, stem apex pruinose.
I. pallidicremea © Sharon Squazzo, I. 'pallidicremea PNW03' © Ann Goddoard, I. 'lilacina PNW10' © iNaturalist user hugebooter, I. ionocephala © Bitty Roy
Brownish discs - not truly white, but usually paler on the margin than on the disc.
Inocybe agglutinata NY - brown disc, paler margin. Perhaps it can also be recognized by agglutinated fibrils on the cap (stuck together) even though the cap is dry and not viscid. This species will be the hardest to place in this group. We have many west coast sequences, but I would like an east coast sequence to verify that we have the same species.
Inocybe fuscodisca NY (=Inocybe virgata NA, =Inocybe fuscicothumata NS) - very distinct dark eye on the disc. Brown fibrils on a white ground colour on the cap and stem. We have local verified sequences, but no verified photographs. The I. lilacina group paper found that the type sequence of Inocybe fuscicothumata matched reliable west coast sequences of I. fuscodisca, and said that Inocybe virgata may also be an newer synonym.
Inocybe PNW09 - known from 6 BC sequences, sister to I. agglutinata, it's probably a brown species.
Inocybe PNW52 - known from 4 BC sequences, it is probably brownish (it was mistaken for I. geophylla, I. posterula and I. auricoma).
Inocybe agglutinata © Jacob Kalichman, possible I. fuscodisca © Kit Scates Barnhart
2. Pale, whitish caps, stem not bulbous
Inocybe 'sindonia PNW23' - pale capped, at least on the rim. The real I. sindonia has a cortina and the stem is only pruinose at the top, but PNW23 and PNW24 haven't been examined closely enough to know for sure. They are ~10% different in ITS from Bandini's official EU sequences of I. sindonia.
Inocybe 'sindonia PNW24' - a second species in the complex, equally distinct in ITS. It has been examined microscopically.
Inocybe melleiconica NS - this closely related species is described with an entirely pruinose stem without a cortina. We have a bunch of BC and WA sequences matching an NS type sequence.
Inocybe fraudens EU - a stocky whitish species with a cortina and non-pruinose stem with reddening flesh and a fruity or matsutake odor that does not appear to be in the same section as the other three. We have the type sequence, but no local sequences to see if the couple of old reports of this species being here are true.
Inocybe 'sindonia PNW23' © Buck McAdoo, I. 'sindonia PNW24' © Fred Rhoades, I. melleiconica (2 images) © Yi-Min Wang
3. Possible lilac stem apex when fresh (sometimes hard to detect, and sometimes not present, making this group difficult to recognize). Most seem capable of having true recurved scales on the cap and possibly the stem too. Almost all have a cortina when young, no stem bulb and are pruinose at most at the stem apex, if at all. These form a natural clade with the possible exception of I catalaunica, I. griseolilacina, I. pusio, and I. laetior. The colourful Inocybe cinnamomea is also in this clade, but you wouldn't know it as there are no purple tones on the stem.
Inocybe obscuroides EU (=Inocybe cincinnata var. major EU) - with dark brown fibrils on the stem. Bandini showed that I. cincinnata var. major, reported from the PNW, is a newer synonym of I. obscuroides. This DNA is not known from the PNW yet. See the next entry.
Incybe cincinnata EU - known from 4 BC sequences, I think what local people have been calling Inocybe cincinnata var. major is just the type variety of Inocybe cincinnata. Stuntz called it Inocybe obscura, but that species really is obscure and nobody knows what it is. ITS can vary by almost 2% in this species worldwide, and our DNA does not quite match EU DNA, but for now it is all considered one species.
Inocybe pyrotricha WA - with reddish stem scales. We have the type sequence and a recent collection with a matching sequence, but the photo was lost.
Inocybe 'griseolilacina PNW28' - this group often has a grey-brown cap (others have a more plain brown cap) and lacks dark brown fibrils on the stem. Some BC and WA sequences match one of the two sister species in Europe going by this name. It's possible this represents I. griseolilacina. The one spore measurement we have is on the high end for that species, averaging 10.3 × 5.5 µm.
Inocybe 'griseolilacina PNW29' - some BC and WA sequences are match the second EU species, 2% different in ITS than the first. It is possible that no geographical, morphological nor ecological differences will be found and these could be the same thing.
Inocybe pusio EU - we have a few EU concepts purporting to be this, but no matching local DNA to any of them yet. Perhaps one of the unnamed species is being mistaken for this.
Inocybe catalaunica (=Inocybe leiocephala WA?) - we have the EU type sequence of I. catalaunica, but that DNA is not found in the PNW yet. I. catalaunica is not in the same clade as the rest of the "lilac stem apex" species in this group, so I am doubting the synonymy. Perhaps one of the unnamed species below is I. leiocephala instead. Being a local species, I. leiocephala is here and we need to figure out its ITS sequence.
Inocybe laetior WA - unrelated, entirely pink stem (no bulb and entirely pruinose). Red-brown smoothish cap. We have the type sequence, but no recent sequences nor photographs. Its description seems distinct and unique and I wasn't sure where to key it out so I put it here in this group.
Inocybe 'griseolilacina PNW28' © Richard Morrison, I. 'griseolilacina PNW29' © Jacob Kalichman
Inocybe PNW17 - (Brandon's cincinnata sp. 1) from oak and pine with an appressed scaly conical cap. Two WA sequences and over a dozen matching BC sequences are in this clade, but I have not yet observed lilac colours in the upper stem.
Inocybe PNW18 - (Brandon's cincinnata sp. 7) a similar spring species from conifers that might also be pointy capped, and might be recognized microscopically by exceptionally skinny metuloids and spores 9.5-11.5 x 5-6u. Two WA sequence and 10 BC sequences.
Inocybe PNW50 - (Brandon's cincinnata sp. 4) from oak and pine. Known from 3 BC sequences.
Inocybe PNW60 - (Brandon's cincinnata sp. 2) from Doug fir. Known from 2 BC sequences. A sister species' DNA has only been found in soil samples in the PNW so far, not fruiting. If it is ever found, it will be Brandon's cincinnata sp. 3.
Inocybe PNW17 © Jacob Kalichman, I. PNW18 © Luca Hickey and Yi-Min Wang (2 images)
4. Truly scaly cap (but no lilac stem apex), no stem bulb, stem pruinose at most at apex (veil present). Many species are capable of scaly discs when old, but these are generally scaly at all ages.
Inocybe lacera EU - dark brown scaly cap, stem nowhere pruinose (but strongly fibrillose) like the nodulose spored I. lanuginosa group. It has unique very long, cylindrical shaped spores (15x5u). It is often found in very poor soil (versus on wood for the I. lanuginosa group). Bandini provided sequences and we have dozens of matching sequences from BC and one from WA.
Inocybe flocculosa EU - medium brown scaly cap, stem pruinose at apex only. We have dozens of EU sequences that match each other, but no local sequences yet to see what local reports of this are.
Inocybe microlepidota n.p. - a small (<2cm across) grey scaly cap. It has not been described yet, we have a WA photo and sequence.
Inocybe griseoscabrosa NY - this species was reported once from the PNW long ago, but a modern find matching the description turned out to be the undescribed sister species that will be called I. microlepidota described above. It may be that the old report of I. griseoscabrosa was also I. microlepidota, so we need modern collections. We have east coast type area sequences of I. griseoscabrosa to compare to.
Inocybe lacera © Yi-Min Wang, Inocybe microlepidota n.p. © Jacob Kalichman
5. Yellow-brown smoothish caps, stem bulb, much of the stem pruinose (similar to the nodulose spored I. mixtilis group, which are more common)
Inocybe picrosma WA - yellow-brown, stem bulb, at least the stem apex pruinose. A unique resinous odor. We have the type sequence and matching WA collection
Inocybe kauffmanii USA - yellow-brown cap, stem bulb and stem entirely pruinose. Brandon has provided sequences from the west coast; I don't know which coast the type is from. We should get east coast sequences to make sure they're the same, but so far this seems to be a species with variable ITS, as some west coast sequences are as much as 6bp different from each other.
Inocybe chalcodoxantha NS - also a yellow-brown cap, stem bulb and entirely pruinose stem but with a pale umbo. We have the NS type sequence, but no local sequences yet to confirm. It was described from both NS and WA, so I expect it is here.
Inocybe semifulva NS - two toned smoothish cap, tawny on the disc and yellow at the margin. Sometimes a stem bulb, stem pruinose at the top. We have the NS type sequence, but no local sequences yet to confirm. It was described from both NS and WA, so I expect it is here.
Inocybe volvata WA - supposedly red-brown capped but our photographed collection is yellow brown. Small rounded stem bulb and pruinose at the stem apex. The cortina is capable of leaving a volva on the stem base. Brandon provided a reliable sequence, we have sequenced it from WA and BC.
Inocybe picrosma © Richard Morrison, I. kauffmanii © Yi-Min Wang, I. volvata © Jacob Kalichman (2 images)
6. Yellow- to cinnamon-orange to red-brown fibrillose caps (not just plain brown)
Inocybe submuricellata NY var. stenospermina NS - yellowish cap, no stem bulb, pruinose at stem apex. We have the NS type sequence, but no local sequences of this mushroom described from NS and WA. We don't have the type variety sequence to know if this is a valid variety or needs to be elevated to species.
Inocybe cinnamomea CA - with cinnamon coloured gills, but quite a variable cap - from pale to dark cinnamon, and a fibrillose (not pruinose) stem without a bulb, it is usually a little more brightly capped than the typical Inocybe, almost like Cortinarius, but those are not this tattered. We have the CA type sequence, many BC sequences and three matching WA sequences that were photographed. However, some collections seem to have spores much larger than expected, 9-12 x 5-7u, while other collections with identical sequences have the expected spore size of 7-9 x 4-5.5u.
Inocybe PNW14 - one sequence from OR, the mushroom resembles Inocybe cinnamomea, but is not closely related.
Inocybe praecox WA - a low elevation spring species, stem entirely pruinose often with a bulb. We have reliable DNA. It is described as yellow-brown, but some reliable photos are more colourfully orange-brown. We need photographed, sequenced collections.
Inocybe cinnamomea (2 images) © Yi-Min Wang, I. PNW14 © Jenny Lippert, unsequenced I. praecox © Richard Morrison
Inocybe olympiana WA - yellow-brown to cinnamon coloured cap, farinaceous odor, fibrillose (not pruinose) stem with a bulb, seemingly preferring old growth forests. We have the type sequence and a recent BC sequence, but no sequenced verified photographs yet.
Inocybe chondroderma WA - this orange-brown capped species with a paler margin has no stem bulb and is pruinose at the apex only. It is the only species that turns turquoise from the chemical PDAB. We have the type sequence and many matching BC sequences, but no photo yet.
Inocybe vaccina EU - orange-brown cap, no stem bulb, entirely white pruinose stem. Bandini provided a sequence and an ENA sequence is 2% different. We have no local sequenced collections yet to prove that this is what our bright orange collections are.
unsequenced Inocybe olympiana © Steve Trudell, unsequenced I. chondroderma © A and O Ceska, possible I. vaccina © Steve Trudell
Inocybe monticola UT - red brown cap sometimes with pale universal veil material on the disc, no stem bulb, entirely pruinose stem, found in the spring at high elevations. We have the UT type sequence and many matching BC sequences.
unsequenced Inocybe monticola © Shannon Adams
7. Brightly coloured gills
Inocybe 'flocculosa bright yellow gills' - what is this? It resembles I. flocculosa var. crocifolia. No sequences yet.
Inocybe 'flocculosa bright yellow gills' © Danny Miller
8. Brown smoothish cap, entirely pruinose stem, no stem bulb
Inocybe brunneolipes NS - we have the NS type sequence, but no local sequences yet to confirm. It was described from both NS and WA, so I expect it is here.
9. Brown fibrillose cap, stem bulb, entirely pruinose stem?
Inocybe 'splendens PNW01' - the several species that resemble I. splendens do not clade together, including this one, not closely related to I. splendens. The presence of the stem bulb has not been verified.
Inocybe 'splendens PNW01' © Yi-Min Wang
Inocybe PNW19 - perhaps with an umbo, stem bulb and pruinose stem. One BC and one WA sequence.
Inocybe PNW20 - also appears to have a stem bulb and pruinose stem. Spores 10-11.5 x 5-6µ. One OR and one WA sequence.
Inocybe PNW22 - it has a stem bulb and an apparently pruinose stem. A handful of BC and WA sequences, some of these were mistakenly identified as Inocybe nitidiuscula.
Inocybe PNW19 © Jacob Kalichman (2 images), Inocybe PNW20 © NAMA and the Field Museum of Natural History, Inocybe PNW22 (2 images) © Yi-Min Wang
10. Brown fibrous cap, no stem bulb, pruinose at most at apex only - this is the largest and most indistinct group of species.
Inocybe 'nitidiuscula PNW15' - this BC sequence is too dirty to know if it that species or an undescribed sister species. We need more collections. I. nitidiuscula is reported from the PNW and Bandini has provided reliable EU sequences.
Inocybe subdestricta MI - We have one Stuntz sequence from NS that might be this, and one Brandon sequence from TN that might be it instead. Others think it is a synonym of Inocybe nitidiuscula. We need local collections to see what local reports of this really are.
Inocybe pseudodestricta EU - 5 BC sequences match many EU sequences fairly well, but there is a bit of variation in ITS.
Inocybe glabripes EU (=Inocybe fuscidula EU) - Bandini synonymized them. We have the type sequence of I. glabripes, but no local sequences to prove the many reports of I. fuscidula, which may have been something else.
Inocybe plurabellae EU - actual shade of brown said to be extremely variable. It may be a spruce species. One local Victoria BC sequence is 1.5% different than the EU type sequence, but the paper could not find any ecological or morphological differences, so it is being considered the same species.
Inocbye hotsoniana WA - stem not pruinose. We have the type sequence and matching BC and OR sequences, but no photos.
Inocybe oetziana EU - we have some BC and WA sequences that match fairly well to the EU type sequence (except for one long chunk of indels) and other sequences that vary by up to 6bp, mostly in ITS2. The various sequences don't seem to separate into groups of species, so for now I am assuming that this is all one species with variable ITS, especially ITS2.
Inocybe oetziana © Jacob Kalichman and Yi-Min Wang
Inocybe 'lanatopurpurea PNW26' - while I. lanatopurpurea itself is known for wooly universal cap material and purple tones in the cap, stem and flesh, our related species may not exhibit the same features. Our collections seem to have relatively smooth brown caps. Perhaps the best clue to recognizing it is that both WA collections had sparse crystals on the metuloids. The spores measure 9.5-11.5 x 5-6µ. Sequences of PNW26 are almost 3% different in ITS than the EU type sequence of I. lanatopurpurea.
Inocybe 'lanatopurpurea PNW30' - one BC and one OR sequence are 1.5% different in ITS from PNW26.
Inocybe 'lanatopurpurea PNW26' © Yi-Min Wang and Jacob Kalichman, Inocybe 'lanatopurpurea PNW30' © Jonathan Frank
Inocybe PNW16 - two WA collections sequence near a group of species around I. nitidiuscula, I. alberichiana and I. laurina, but is probably itself undescribed.
Inocybe PNW16 © Jacob Kalichman and Richard Morrison
Inocybe PNW27 - one OR sequence and 18 BC sequences
Inocybe PNW27 © Ann Goddard
Unknown appearance (no photos or description available)
Inocybe PNW21 - one Victoria BC sequence, no photos. Their position in the tree tells me these species are likely smooth spored.
Inocybe PNW51 - known from 21 BC sequences, no photos.
Inocybe PNW56 - known from 5 BC sequences, no photos.
Inocybe PNW57 - known from 5 BC sequences, no photos.
Inocybe PNW58 - known from 3 BC sequences, no photos.
Inocybe PNW59 - known from 12 BC sequences, no photos.
no DNA available yet, but described with smooth spores and reported from the PNW
Inocybe eutheloides NY -
Unknown spore shape
Inocybe PNW37 - orange brown cap, no stem bulb. From our one known photo, the stem appears pruinose at the apex but probably not for most of the length, but I can't be sure. It appears in the tree with nodulose spored species, but it was mistaken for a smooth spored species although I don't know if they checked the spores or not. It was sequenced once from BC and once from OR and mistakenly shown in Bandini's tree as I. lampetiana, a smooth spored species, but it is nothing like that type sequence.
Inocybe nematoloma EU - an OR sequence matches an EU sequence, but it's not one that Bandini provided so the ID isn't proven.
Inocybe PNW53/PNW54 - 2 other species in the I. nematoloma complex, known from a few BC sequences each
Inocybe PNW37 © iNaturlist user winterwren22
There are more than 1,000 local sequences of Inocybe in GenBank, and I was not able to look at every single one of them, so more species exist.
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