Pholiota - click to expand
Everything else.
Species
mentioned: Pholiota highlandensis, carbonaria, fulvozonata, molesta,
subsaponacea, brunnescens, luteobadia, subangularis, flammans, kauffmaniana,
aurivella, adiposa, limonella, abietis, connata, filamentosa, hiemalis,
terrestris, gummosa, melliodora, squarrosoides, squarrosa, kodiakensis,
flavopallida, lurida, lenta, scamba, spumosa, baptistii, baptistiae,
velaglutinosa, rubronigra, humii, decorata, lubrica, lubrica var. luteifolia,
rufodisca, avellaneifolia, fulviconica, ferrugineolutescens, alniphila, occidentalis, mixta,
agglutinata, fulvodisca, nubigena, abruptibulba, acutoconica, aurantioflava,
bakerensis, brunnea, canescens, crassipedes, curcuma, ferruginea,
galerinoides, gruberi, iterata, lactea, luteola, macrocystis, milleri,
multifolia, olympiana, pallida, paludosella, mutans, polychroa, populicola,
proximans, pseudopulchella, pulchella var. pulchella, pulchella var.
brevipes, rivulosa, scabella, scamboides, sienna, sipei, subangularis,
subechinata, subflavida, subfulva, subgelatinosa, sublubrica, subminor,
subochracea, subpapillata, tuberculosa, P. curvipes, umbilicata, velata,
verna, vialis, vinaceobrunnea
(Burned ground/wood species)
Pholiota highlandensis NY (P. carbonaria
EU/CA, P. carbonicola CA, P. fulvozonata ID) -
usually slender, stem <4 mm thick (but some are thicker). No recent local
sequences yet, but we have all three type sequences and an old, original WA
sequence, and they are officially synonymized.
Pholiota molesta ID (P.
subsaponacea ID) - usually squat, stem >4 mm thick (but some
are thinner). We have the type sequences and 4 WA sequences used in the study
that synonymized them.
Pholiota brunnescens OR (P.
luteobadia MI) - the only one where the stem stains brown. All
4 recent PNW burn sequences are this species, so maybe it's our most common
species locally. There are a handful of ambiguous locations in some sequences,
including the type sequence, as well as some run length differences (indels) but
besides that there's very little variation among the sequences. An ecological
and morphological study showed them all to be the same species.
They are common as a group
but only recently did we learn to reliably tell them apart so some are
undoubtedly less common than others.
Crassisporium funariophilum EU ('Pholiota' subangularis ID) - is
in a different family. It
has a dry cap and reddish, thick walled spores. It is usually
restricted to the spring whereas the Pholiotas can be found spring or fall.

Pholiota highlandensis © Brandon Matheny, P.
molesta © Brandon Matheny, P. brunnescens © Steve
Trudell
Here's a (hopefully) interesting story about how the correct name of Pholiota
highlandensis/carbonaria/carbonicola/fulvozonata got chosen among the four competing names that turned out to
all refer to the same thing. In 1818 Elias Fries, by some accounts only the
world's second mycologist, found it, presumably somewhere near Sweden, and named
it Agaricus carbonarius, because all gilled mushrooms were called
Agaricus at the time and because the epithet means "burn associated". Then
in 1872, Charles Peck, America's most prominent early mycologist, found it in
Highland Falls, NY, and named it Agaricus highlandensis. Then in 1944,
Alexander Smith, one of America's most prominent mycologists at the time, found
it in California and called it, well... he accidentally referred to it as
Pholiota carbonaria half the time and Pholiota carbonicola the other
half of the time in the paragraph he wrote about it. Then in 1951 Singer got
around to changing the name of the European Agaricus carbonarius to
Pholiota when he realized that nobody had moved that one yet to its proper
genus (moving the thousands of Agaricus species to their new proper genus
is something that had slowly been going on for many decades already). But he
didn't know that Smith had already used the name Pholiota carbonaria, so
most people think he erred in reusing the name Pholiota carbonaria and
that he should have picked a new name. In 1952, Singer got around to changing
Agaricus highlandensis to Pholiota highlandensis, but he didn't
provide enough information about what he was doing, so the name change didn't
stick until 1990 when somebody made it official. Then in 1962, Singer saw the
confusion about the name that Smith had chosen and selected Pholiota
carbonicola as the official name of Smith's mushroom to try and avoid the
problem that would make his 1951 renaming invalid, since that name was not in
conflict with anything else, like Pholiota carbonaria was. Finally, in
1968 Smith found and named Pholiota fulvozonata in Idaho, which turned
out to also be the same thing, but that name is clearly not the oldest name
among them so it can be discounted.
Normally, as Agaricus carbonaria was found and described way sooner
than any of the others (1818), you would expect the official name chosen from
among all the synonyms would be Pholiota carbonaria, which might be a valid choice if we accept Singer's view that we should consider Smith to
have named his 1944 mushroom Pholiota carbonicola and not Pholiota
carbonaria. But in "Pyrophilus taxa of Pholiota" in 2018, the authors, who tested the DNA of all of the above as
well as microscopy and ecology, and concluded they were all the same, seemed to
take the view that the bold type faced name in larger font at the top of the
page, Pholiota carbonaria, was the name Smith really meant to use, and
that Pholiota carbonicola was a typo, and you can't just pretend that
Smith meant to use Pholiota carbonicola instead of Pholiota carbonaria
because that would conveniently avoid a duplicate name. So therefore,
Pholiota carbonaria only dates back to 1944. Pholiota highlandensis
dates back to 1872 through its synonym Agaricus highlandensis. True,
Agaricus carbonarius dates back to 1818, but it's not an Agaricus, is
it? It's a Pholiota. They declared the official name Pholiota
highlandensis.
But the story doesn't stop there. Since then, it has been argued that P.
carbonaria and P. carbonicola are not two competing names for the
same mushroom, but instead simply orthographic variants of each other (minor
spelling differences of the same thing). If you accept this view, then Singer
was allowed to choose either one of them as the official spelling, it would not
matter which name Smith intended: the clarifying author, Singer, gets to decide,
and you have to follow his lead. The other name disappears and is no longer
around to conflict with anything. So now it is accepted by most that there was
no Pholiota carbonaria Smith in existence when Agaricus carbonaria
Fries was recombined into Pholiota carbonaria. Now the official name has
gone back to being Pholiota carbonaria, and even the authors of the 2018
paper have conceded.
I am bucking the trend and have listed Pholiota highlandensis as the
official name on this page for now. Now you have a bit of insight into what it's like to be a nomenclaturist (somebody
who studies the proper names of things) as opposed to just a taxonomist (someone
who studies how organisms are actually related to each other but doesn't
necessarily worry too much about what they're called). If you actually did
really enjoy this discussion, and you find that you have a strong opinion one
way or the other right now, perhaps being a nomenclaturist is for you.
(brightly coloured orange-yellow species, scales not prominently erect and
not tightly clustered)
Pholiota flammans EU (=Pholiota
kauffmaniana WA?) - a striking bright yellow to orange species
covered in bright yellow scales. EU sequences differ by 2 bp and 6
somewhat suspicious indels from each other. A WA and OR sequence match each
other but are between 2 and 5 bp and a half dozen to a dozen suspicious indels
from the EU sequences. I'm not quite sure what to make of that, but for now I'm
assuming it's one species worldwide with some variability in ITS, although it
could be investigated if the colour forms show any genetic distinction.

Pholiota flammans © Steve Trudell and Ben Woo
Pholiota adiposa EU (=Pholiota aurivella EU,
=P.
limonella NY?) - a similar bright yellow mushroom with brown scales
that remind me of onion skin. One old pre-genetics study suggested that what
we have locally is usually the east coast P. limonella (90% of the time) with
most of the rest being P. aurivella and rarely, P. adiposa. However, worldwide
sequences vary in a bunch of ambiguous locations as well as other places, with
no rhyme or reason to which of the 3 names are attached to the sequence, so they
may all be the same thing. The three BC, WA and OR sequences we have differ from
each other in 4-5 ambiguous locations and 3-4 other places that don't seem
ambiguous yet, definitely within the noise of the worldwide sequences, so my
guess is ours (and Europe's) most common species is just one species,
Pholiota adiposa. It is now commonly accepted that P. aurivella is
the same thing. We need more collections from back east and here to determine if
P. limonella is the same thing too, and if not, if we have both here.
Pholiota abietis WA/P. connata MI/P. filamentosa EU/P. hiemalis ID -
other supposed group members that have been reported from the PNW without any
genetic information known. We need more collections and more studies to figure
out the exact identity of the species found here, but it's reasonable to assume
we have more than one species here since the spore sizes of some are reported to
be quite different.

Pholiota cf aurivella © Michael Beug
(Unusual for Pholiota, mostly found on the ground)
Pholiota terrestris OR - found in clusters on the ground,
often prominent erect fibrillose scales on a brown cap and stem. Hard to recognize as a Pholiota
because the wood it is associated with is always buried. We have a number of PNW
sequences of this native which all match, but as it is a species with
variable colours and scaliness, this could be investigated to make sure it's
all the same species.
Pholiota gummosa EU (P. melliodora OR) - this is a sister species of
Pholiota terrestris and also often found in clusters on the ground,
but it is pale yellow-brown and the scales might tend to wash off more
easily. The type
sequence of P. melliodora differs from reliable EU sequences mostly in
ambiguous locations, and is probably a synonym. We have 2 recent WA collections
as well that were pale yellow-brown, one with a greenish tint. Smith placed P.
gummosa, P. melliodora and P. terrestris in three different
subgenera.

Pholiota terrestris © Leah Bendlin, Pholiota gummosa ©
Paul Dawson
(Prominently erect scales, tight clusters)
Pholiota squarrosoides NY -
clustered on wood, with prominent erect scales. We have a half dozen east coast sequences, and a few local
sequences that match, one of ours is 4 bp different. This species is recognized
by a viscid cap and sweet odor.
Pholiota cf squarrosa EU/kodiakensis
AK -
Very similar, yet not closely related, is P. squarrosa with a dry cap,
garlic odor, and yellow young gills that turn greenish in age. We have over a hundred EU sequences, so we know
what this is, but that sequence has never been recorded in North America, so what
are our local sightings of this? One possibility is Pholiota kodiakensis,
a similar sister species from Alaska with a short type sequence and a matching
recent sequence. This appears to be the same as the newer EU species Pholiota
lundbergii. We need local collections to find out which of those two we
have.

Pholiota cf squarrosa and cf squarrosoides (unsequenced) © Danny Miller
(pale, almost white cap colours, perhaps small)
Pholiota lenta EU - an
all white Pholiota with some scales
on the cap and especially the stem. We have reliable EU sequences and one
short but matching sequence of an original Smith collection from 1946 from OR.
This is an especially pale member of the large Pholiota lubrica
clade, detailed below, and easier to recognize than most of them.
Pholiota scamba EU -
one of our smallest Pholiotas, pale capped and scaly
mostly on the stem.
Confusable with other LBMs like Galerina. One WA sequences matches a bunch of
reliable EU sequences. This species is on a branch of its own.

probable P. lenta © Andrew Parker, P. scamba © Daniel Winkler
(bright yellowish-brown 'spumosa' cap colour - indistinctly scaly)
Pholiota spumosa EU (P. baptistii/baptistiae ID,
P. subflavida WA?, P. vialis WA?) -
yellowish-orange-brown cap usually with a darker plain brown disc,
usually indistinctly scaly. Similar
colours to Flammula, above, but a different stature. Sequences from all
over the PNW match many EU sequences quite well. The Tian/Matheny paper
sequenced the local type of P. baptistii (formerly incorrectly spelled P. baptistiae) and found it is a synonym.
The colour difference between this and members of the Pholiota lubrica
group, below, can be subtle, but this species has larger spores than any
of those. P. subflavida is thought by some to be the same thing. P.
vialis was even thought by Smith himself to perhaps be a duplicate of this
too. This should be confirmed.

probable P. spumosa © Steve Trudell
(cap colour not bright yellowish like P. spumosa - the large P. lubrica clade)
Pholiota lenta EU -
described above, is a scaly, almost white member of this clade.
Pholiota flavopallida ID (=P. lurida MI) -
pale whitish rimmed Pholiotas often with a yellow-brown
disc and
no scales anywhere. The rim seems to be more well defined than in
some of the below species and the colouring seems somewhat irregular (although a two-toned cap is quite common). We have the type sequence
of P. lurida, and a couple sequences that Tian identified as P.
flavopallida from China, but that is far away from Idaho so the
identifications might not have been correct. However, that does show that
sequences of collections matching the microscopic characters of P.
flavopallida (pleurocystidia sometimes have thickened walls) match sequences
of P. lurida (without thickened walls) and that is a very slight
microscopic difference, the kind which usually doesn't have any genetic
significance. Therefore, I think they probably are synonyms, with P.
flavopallida being the older name.

Pholiota flavopallida © Yi-Min Wang
Pholiota velaglutinosa OR (=P. rubronigra CA?) - the type sequences
match, and they are described as differing by one having a glutinous
partial veil, although that seems like a significant thing to be different or be
overlooked if they are the same species. Also, P. rubronigra was supposed
to be fragrant. Smith also found P. rubronigra in Idaho, so if they are
not the same, we might have both species here, but it's going to be hard to
recognize. It's more common in the south of our region and California.

probable Pholiota velaglutinosa © Dimitar Bojantchev
Pholiota humii ID - one of many species that has a
dark orange-brown disc
and a paler margin (but the disc may not as defined as in P. flavopallida
above and the colours seem more regular). It has no
scales anywhere and is is differentiated from similar species below by
supposedly subtly different
shades of brown, but may need microscopy for confirmation. Collections of this
species have been long going by the name P. decorata (described next)
because the guide books have been confused about whether or not that species has
scales (it does). The type sequence of P. humii is dirty in one spot, but we
have other sequences to fill in what the missing part is (that match exactly everywhere else).
Pholiota humii © Yi-Min Wang
Pholiota decorata WA -
has scales on both the cap and stem that may disappear in age, a less
fragrant odor, and pleurocystidia with thinner walls. We don't have the type sequence, but we do
an original Smith collection and other trustworthy sequences. Its sister species is the less common and
paler
Pholiota lenta, above.

P. decorata © NAMA and the Field Museum of Natural History
Pholiota lubrica EU (lubrica var. luteifolia ID? P. rufodisca ID?) - we have some reliable EU
sequences of P. lubrica and a paratype sequence of P. rufodisca
from NM is very close, differing in at least 3 locations (one of them a run)
from the closest EU sequences (and in a couple extra places from another EU
sequence). We don't have the ID holotype sequence to know that the paratype
matches it. If the species are the same, P. lubrica is by far the older
name. We have no data for whether or not P. lubrica var. luteifolia from
ID and WA is the same species or not. P. lubrica is said to have a
somewhat scaly stem and white
veil and gills when young. P. lubrica var. luteifolia and P.
rufodisca are said to have yellow veil and gills when young, but
differ in the fact that the latter has pleurocystidia with thickened walls (and
we've already seen that does not necessarily indicate a separate genetic
species).
Pholiota avellaneifolia ID - similar
somewhat scaly stem, with a yellow veil but
avellaneous (smoky) gills when young, this does seem to be a separate genetic
species. We have the type sequence and a good match with a recent WA collection,
differing in only 2 locations near the beginning. Until this collection this
species was little known and in these photos the veil is still intact so I
believe it is a young specimen showing the young smoky gills even before the
spores mature.

Pholiota avellaneifolia © Leah Bendlin
Pholiota fulviconica FL (=P. ferrugineolutescens CA?) - the holotype
sequence of P. ferrugineolutescens is the same as an isotype sequence
from Idaho of P. fulviconica, but we don't have the holotype sequence
from Florida, so it's possible they aren't the same species. But assuming they
are, the CA name is a newer synonym. However, they are not described as being
the same, with P. fulviconica being described as almost identical to
P. spumosa and P. ferrugineolutescens being described as very similar
to P. lubrica but with a stem that stains yellow. This is suspicious. I don't know that either have ever
been photographed in colour.
Pholiota alniphila OR (Pholiota occidentalis OR?) - these are
differentiated from P. decorata etc. and P. lubrica etc. above by
yet more subtly different shades of brown. We have a paratype
sequence from ID of P. occidentalis that I assume is correctly
identified, and we have sequences of modern collections from OR and BC. Some
authors synonymize it with the older Pholiota alniphila, if that's not
true, then we have 2 species here. Conceivably even three, since there is also a
Pholiota occidentalis var. luteifolia that we don't have sequences of to
know if it is genetically different.
Pholiota mixta EU (P. agglutinata ID, P. fulvodisca ID) - Another
species with a red-brown disc and pale rim, somewhat
indistinctly scaly.
The stem base stains brown in age. Both local
type sequences turned out to be the same as the EU species P. mixta, and they
have been officially synonymized. They seem to only have ever been separated by
subtly different shades of brown on the disc.
Pholiota aff mixta - one OR sequence differs by 3% from all the others
and could be a unique sister species, it seems (at least in our only photo) with
more pale colours showing than for P. mixta itself.

Pholiota aff mixta © Jonathan Frank
Pholiota nubigena CA - a
gastroid Pholiota from CA. We don't
have the type sequence, but we have a reliable CA sequence of this distinctive
mushroom also reported from the drier areas of the PNW. This appears to be in
the large Pholiota lubrica clade with the above species.

probable Pholiota nubigena © Steve Trudell
Little known species
In order to learn more about these, we'll need either a type sequence or
sequences of new collections that key out to one of these species in Smith's
Pholiota monograph so we can get an idea of what these species might be.
It's possible that some may not even belong in Pholiota.
Here is a copy of Smith's monograph you can use
to ID your unusual Pholiota collections that don't exactly match one of the
above (and I didn't give very good descriptions, so I admit it's going to be
difficult to feel confident of a perfect match).
Pholiota abruptibulba OR - with an abrupt stem bulb and other
microscopic differences, said to be related to P. occidentalis, P.
verna and P. fulvodisca.
Pholiota acutoconica OR - a very conical mushrooms said to be related
to P. fulviconica and P. lenta.
Pholiota aurantioflava ID - described as somewhat like P. gummosa.
Pholiota bakerensis WA - described as somewhat like P. lubrica.
Pholiota brunnea ID -
Pholiota canescens OR - this is similar to
Tubaria confragosa
and might be a Tubaria. The real T. confragosa probably does not
occur here, and our species needs a new name, but Tubaria canescens is
already taken so we cannot use this epithet for our local species.
Pholiota crassipedes WA -
Pholiota curcuma SC - Smith reported this from ID too, and it might be
related to Phaeomarasmius.
Pholiota ferruginea WA - said to be somewhat like P. lubrica.
Pholiota galerinoides WA - this could be related to Kuehneromyces.
Pholiota gruberi ID -
Pholiota iterata OR -
Pholiota lactea WA - possibly related to Phaeomarasmius.
Pholiota luteola ID -
Pholiota macrocystis ID -
Pholiota milleri ID -
Pholiota multifolia MO - reported by Smith from WA
Pholiota olympiana WA -
Pholiota pallida ID - probably a Kuehneromyces
Pholiota paludosella OH - reported by the Key Council from the PNW
Pholiota mutans MI - Smith wasn't clear if this was present in the PNW
or not. It might be a Meottomyces, perhaps the same as
Pholiota pattersoniae CA.
Pholiota polychroa OH - reported once from BC
Pholiota populicola WA - probably a Kuehneromyces
Pholiota proximans MI - reported by Smith from ID, found recently in
WA. Now officially Phaeomarasmius
proximans, see that page.
Pholiota pseudopulchella OR -
Pholiota pulchella var. pulchella WA and var. brevipes ID -
Pholiota rivulosa ID -
Pholiota scabella OR - Tian and Matheny sequenced the type and it is
now Stropharia scabella
Pholiota scamboides ID -
Pholiota sienna OR -
Pholiota sipei OR -
Pholiota subangularis ID - now officially
Crassisporium funariophyllum
Pholiota subechinata WA - possibly a Phaeomarasmius
Pholiota subfulva NY - reported by Smith from ID
Pholiota subgelatinosa OR -
Pholiota sublubrica ID - said to resemble P. lubrica (photo
in MM)
Pholiota subminor OR -
Pholiota subnigra WA - now known to be the local name for at least one
of our species in the Cyclocybe erebia group.
Pholiota subochracea WA - Tian and Mathaney sequenced an old OR Smith
collection (but not the type) and discovered it belongs in Hypholoma,
where it was originally placed before Smith changed his mind and moved it to
Pholiota.
Pholiota subpapillata ID - may belong in Keuhneromyces
Pholiota tuberculosa EU (P. curvipes EU) - not found by Smith, but
reported in the PNW since then, assumed to be the same species as P. curvipes.
Probably not a Pholiota, one possibility is Simocybe, another is
Phaeomarasmius.
Pholiota umbilicata ID -
Pholiota velata ID -
Pholiota verna WA - said to resemble P. decorata (photo in
MM)
Pholiota vinaceobrunnea ID - also said to be close to P. decorata.
Also, Stropharia scabella OR