Danny’s DNA Discoveries – Clitocybaceae of the PNW
Every clitocyboid mushroom, those of at least a certain size (caps >2.5 cm across or so) with decurrent gills and white spores (occasionally with an indented cap to boot) and no partial veil, was once placed in Clitocybe. The most similar stature is omphalinoid, usually reserved for smaller, similar mushrooms. The clitocyboid form has evolved independently many times, and many have been moved to new genera. Those that are left in Clitocybe usually grow on the ground and have inamyloid spores and clamp connections. Here is a summary of some of the segregate genera:
- are very similar, but often have an umbo that Clitocybe usually
Left inside the Clitocybaceae family are:
Clitocybe - the group that is left after all of the above were segregated out. The gills aren't always strongly decurrent and it's often a puzzle whether or not to look for an adnate gilled mushroom here or somewhere else.
Lepista - was split from Clitocybe for having spores that were warty (as well as always having a slightly coloured spore print which only happens sometimes in Clitocybe). Warty spores evolved three independent times inside the genus Clitocybe and therefore Lepista can't be thought of as a proper genus. They are all inside Clitocybe.
Collybia - tiny mushrooms growing on other decaying mushrooms, often Russula, and often with a sclerotium at the base of each stem. The gills are not decurrent so it is not obvious that they belong here. Other genera on rotting mushrooms are larger, with Asterophora being the closest lookalike, being fairly small but usually powdery, never with sclerotia, and with stockier stems a couple mm wide instead of 1 mm wide like Collybia). They are all inside Clitocybe too.
Singerocybe - these are just Clitocybe, at least the species reported from the PNW. I suppose it's possible the type species, Singerocybe viscida, once sequenced could turn out to deserve its own genus, but I doubt it.
Leucocalocybe - should never have been created as a genus. Its one species is a Clitocybe.
Dendrocollybia - a rare unique LBM with branches growing out of the stem. This is a distinct genus.
Collybia, Lepista, Leucocalocybe, Singerocybe, and Clitocybe all form one monophyletic group, with Clitocybe and Collybia being equally old names from 1821 (as tribes) and 1857 (as genera). By far more species are currently in Clitocybe, so that name will probably win out. This means that we should properly call all Lepista by the name Clitocybe, and we should even call all Collybia by that name too. These changes haven't been made yet. Dendrocollybia does in fact appear to be a distinct genus.
The authority on clitocyboids is Bigelow. His monograph will be needed to continue to demystify the many unknown species of Clitocybe.
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.
Clitocybe - click to expand
Smooth spores that are very close to white in print.
1. Skunk odor, large and grey
Clitocybe nebularis EU (=Clitocybe robusta NY) - the type species of the genus. It is a large grey mushroom that smells like skunk. It can grow in fairy rings. Clitocybe robusta, said to be a related white species, has four east coast ITS sequences that are identical to C. nebularis, so the colour does not seem to indicate a different genetic species. White forms are reported rarely from the PNW.
Clitocybe nebularis (sequenced and unsequenced) © Bruce Newhouse and Steve Trudell
2. Anise (black licorice) odor
Clitocybe odora EU - a beautiful large, pale blue mushroom that smells and tastes of black licorice. It's definietly a favourite.
Hygrophorus caeruleus ID - a waxy version of Clitocybe odora that may lack the licorice odor. You can read about it here. Something seems to infect Clitocybe odora and make weird fruiting bodies every once in a while. It has basically identical DNA to Clitocybe odora.
Clitocybe deceptiva ID - a small, watery grey-brown mushroom with a black licorice odor. Clitocybe fragrans (=Clitocybe obsoleta) is a similar sweet smelling mushroom rumoured from the PNW, but that DNA has not been found here yet, over a dozen sequences have all been C. deceptiva. Bigelow, the most trusted Clitocybe identifier, never reported C. fragrans north of CA. This species has large spores 6.5-9 x 4-4.5u.
Clitocybe 'idahoensis PNW03' - a couple OR collections and a CA collection with a similar sweet odor are a lookalike of C. deceptiva, but with smaller spores (4.5-6.5 x 2.3u).This is probably Clitocybe idahoensis, assuming it has a chalky white canescence when young, but we don't have the type sequence or sequences with white canescence to verify this.
Clitocybe odora © Andrew Parker and as Hygrophorus caeruleus © Richard Morrison, C. deceptiva © iNaturalist user hugebooter, C. 'idahoensis PNW03' © Jonathan Frank
3. Snowbank Clitocybe
Clitocybe glacialis WY - found in spring at elevation near snow, chalky white when fresh, revealing grey flesh below when rubbed or in age, it can undergo quite a transformation. This abundant species is one of the most recent sequences obtained on this page. It is related to C. 'amarescens PNW11, CA01 and C. brunneoceracea, which it resembles (strongly hygrophanous and grey-brown gills).
unsequenced Clitocybe glacialis © Danny Miller
4. Other chalky white species when fresh, but not truly hygrophanous nor farinaceous. Small spores.
Clitocybe 'rivulosa PNW07' - in grass, <6 cm across, chalky white when fresh, meaning it has a white bloom that can rub off revealing pinkish-orange tones. Small spores means it falls into the group with spores 4-5.5u long, not the group 6-8u long. It is thought to be quite poisonous. Our sequences are 1% different than EU sequences that Pablo thinks represent this species. This used to be called Clitocybe dealbata EU, but that poorly understood name has been abandoned in favour of C. rivulosa. Bigelow said that our NA species was not the same thing as found in Europe, and he declared that the proper name was probably Clitocybe sudorifica NY. We need to figure out what that is to test that theory.
Clitocybe 'rivulosa PNW02' - a lookalike grass species, so much so that Pablo used to think this species was C. rivulosa. It doesn't live in the same part of the tree, so either one of these lookalikes is poisonous and one isn't, or there might be a lot more poisonous species around too. It wouldn't make sense to me that both are poisonous but nothing else. One photo below shows it white and fresh, but the other photo shows is aged and waterlogged, where it has become tan coloured and difficult to place in this section. Fresh colleciton are needed for an accurate ID.
Clitocybe phyllophila EU (=C. cerussata EU, =C. dilatata EU) - found in forests and not grass, and larger (>6 cm across when fully grown). ITS DNA of both species is within a couple of bp of each other, so the latest paper agrees with those that considers them synonyms. It is called C. phyllophila when it has a spicy odor and off white spore print, and C. cerussata when it has a mild odor and white spore print, although intermediate collections are found. The only sequence verified photo is not fresh, so it is not white, but old and waterlogged and tan coloured. Fresh collections are needed for accurate ID. Note that Clitocybe dilatata EU is also a newer synonym of this. Locally, we've been using that name incorrectly when talking about Leucocybe connata. If you've heard the name C. dilatata, it was likely referring not to C. phyllophila but to Leucocybe connata.
There are lookalikes in the Lyophyllaceae family, Leucocybe (small white forest mushrooms <6 cm across, or clustered white mushrooms at the edges of trails and roads) and Atractosporocybe (spores more than twice as long as wide).
Clitocybe 'rivulosa PNW07' © Leah Bendlin (2 images), C. 'rivulosa PNW02' © Buck McAdoo (2 images)
Clitocybe phyllophila (atypical) © Buck McAdoo
5. White to buff with very pale gills - in forests, small spores, hygrophanous but not farinaceous
Clitocybe PNW01 - not farinaceous, small white spores (<6u). One collection was noted to be hygrophanous. One matching sequence was labeled Clitocybe diatreta, which has similar sized spores, but a tinted spore print.
Clitocybe PNW04 - our one PNW sequence (unphotographed) is a collection from the 70s by Joanne Lennox which was labeled Calocybe hebelomoides. It also matches one EU sequence labeled Clitocybe diatreta. Both of those are not farinaceous and small spored, so maybe this is too.
Clitocybe PNW05 - also not farinaceous smelling, perhaps slightly sweet. It was hygrophanous in at least one photo (the last one). Some collections have an indented cap and almost appear omphalinoid, making this less likely than the other two to actually be C. diatreta. Some collections we've examined show small spores, and others large spores, so I think there has been some kind of a mix-up that needs to be sorted out.
Clitocybe PNW01 © Richard Morrison, C. PNW05 © Daniel Winkler, and Yi-Min Wang (2 images)
6. Similar, but large spores
Clitocybe 'coniferophila/fuscidisca PNW10' - the disc can be darker than the margin, but that is not an uncommon feature. At least some photos show it can be hygrophanous, and usually no odor is noted so it is not distinctly farinaceous (rarely it is reported as somewhat farinaceous only when crushed). It should be noted that although many collections were examined and have large spores, three collections were reported to have small spores, so that needs to be sorted out. This keys out to Clitocybe coniferophila in Bigelow, which is noted to have a darker disc, but we have no prior sequences of it to test that theory. Joanne Lennox had a plan to name this Collybia fuscidisca back in the 70s, due to the darker disc. In the unlikely event that such an abundant species doesn't already have a name, I would call it Clitocybe fuscidisca. We're well on our way to having dozens of sequences of this.
'Clitocybe' salmonilamella CA - very similar but seems more likely to have an indented cap in age and may have a pink tinge to the gills and dries completely white (unlike most real Clitocybe), needs to move to Leucocybe. For more information, see that page.
Clitocybe 'coniferophila/fuscidisca PNW10' © Buck McAdoo, Jonathan Frank, and iNaturalist user solstice, 'Clitocybe' salmonilamella © Daniel Winkler
7. Watery grey-brown species, strongly hygrophanous, with grey-brown gills - many mushrooms matching this description are also found in the Lyophyllaceae. As a matter of fact, Bigelow called this group section Pseudolyophyllum. All of these match sequences identified as species that Bigelow keys out with grey-brown gills, so this is how I am going to treat them, but in practice you'll see the gills might look fairly white, complicating their easy identification. A key difference between species in Bigelow is how they smell, and whether the spores are large or small, something that has not yet been noted for all of our species and needs to be. Then we shall see what species Bigelow would have called them (see below for the full list).
Clitocybe PNW06 - it has been likened to C. vibecina and C. pseudodicolor (which have different odors, but both have large spores and a white spore print). One local collection had a mild odor like C. pseudodicolor. Large spores, >6u.
Clitocybe CA01 - with no farinaceous odor, photos showing it is umbilicate, and with spores mostly >6u (5.5-7x4-4.5) this keys out to Clitocybe concava EU in Bigelow. Not every photo shows the dark grey-brown colours. The odor has been reported as "faint pleasant" or "wood". A similar species is Tephroderma PNW01.
Clitocybe PNW13 - this species, found once in WA, is closely related to Clitocybe amarescens and Clitocybe CA01. This suggests it has a mild odor (confirmed in this collection) and spores mostly >6u long. It has an umbilicate cap, meaning that like CA01, it is a candidate for being Clitocybe concava. I do not know how to tell it apart from CA01.
Clitocybe 'amarescens PNW11' - a couple of WA sequences match Pablo's concept for Clitocybe amarescens, so perhaps this is that. It is described with a mild odor, bitter taste, and large spores with a pale yellow spore print. Locally, it was noted with a peculiar odor and mild taste. The large off-white spores, measured as 6-8x3.5-4u and 6-8x4-5u, do match. Perhaps this has been mistaken for the similar C. pseudodicolor EU (white spore print) or C. madefacta ID (vinaceous grey spore print).
Clitocybe PNW06 © Buck McAdoo, C. CA01 © Richard Morrison (2 images), C. PNW13 © Matthew Koons
Clitocybe 'amarescens PNW11' © Richard Morrison
Clitocybe brunneoceracea Australia - sometimes clustered. A few WA collections match an Australian type area sequence and likely represent this species, not previously known from the PNW. It may have another name too if Bigelow found it here. It does have an odor, reported here as sweet, melon or cucumber, but described in Australia as "slight phosphorus like". The spores are described as 5.5-6x2.2u, in other words, in between large and small. However, the photo on the right had spores that measured 5-6x3-4u, and a subsequent WA collection had spores 6-7 x 3.5-4.5u.
Clitocybe brunneoceracea © Jacob Kalichman and Richard Morrison
Little known species
Most of these reports come from Bigelow, so any collections matching the description of these mushrooms in his monograph should be sequenced, although experience has shown that Clitocybes are difficult to identify, and the same species will not always key out to the same result. It is highly probable that at least some of these will end in one of the many segregate genera and need to be moved once they are understood.
Clitocybes that need to be moved elsewhere (if I say maybe, my guess is based on the section Bigelow placed them in and where its neighbours ended up moving, because we have no DNA yet. We need collections).
Clitocybe adustiterricola OR - an omphalinoid on burned ground, in a unique section
of Bigelow with C. payettensis and C. albimontana (which is a
Lichenomphalia even though this can't be)
found on wood - so that's suspicious. I don't think they're actually in Clitocybe.
Clitocybe americana MI
Large, stocky species (all the rest are small to medium slender species)
Clitocybe crassa ID - large, grey brown with a short, bulbous stem, this
could possibly belong in
Clitocybe arvalis WA
Greyish-brown gills (this is how Bigelow keys them out but I can't vouch for how reliable a character this is).
Clitocybe murina WA - anise odor, with even bigger spores than C.
White gills and cap, hygrophanous or chalky - small spores 4-5.5u long
Clitocybe albidula NY
White gills and cap, hygrophanous or chalky - large spores 6-8u long
Clitocybe caperata ID
Usually large, tricholomatoid mushrooms, these have slightly coloured, warty spores. There are 3 monophyletic groups, each paraphyletic inside Clitocybe. Therefore, species with warty spores should probably all be called Clitocybe.
Lepista #1 - often large, white mushrooms, the core group of Lepista. They are more often found in grass or along roadsides and trailsides than in the deep forest.
Clitocybe luscina EU (=Lepista panaeola (panaeolus) EU, =Clitocybe subconnexa NY, =Lepista caespitosa EU) - a large, almost white species usually growing in grass, whose various names are due to various odors that different collections have had. This could be confused with Clitocybe brunneocephala, below, an even larger white mushroom with a short, thick stem, white spore print, and smaller warts on the spores that may be harder to detect. ITS DNA is the same for Lepista panaeola (sometimes spelled L. panaeolus), Clitocybe subconnexa, and Lepista caespitosa (which is not the same as Clitocybe caespitosa). The oldest name is Lepista panaeola. We don't have DNA of Clitocybe luscina yet, but the description is practically identical to the others, and it is now believed that it is an even older synonym. I would like EU DNA of L. luscina to prove it.
Clitocybe praemagna SK/CO - very similar, with somewhat larger spores. We don't have DNA yet, but as the spores are somewhat larger, this might not be yet another synonym of the above. We need collections to find out if it is distinct, or the same as Clitocybe luscina which has smaller spores, or the same as Clitocybe irina which has larger spores (described under Lepista #2), or the same as Clitocybe brunneocephala which has similar sized spores (described under Lepista #3).
Clitocybe densifolia EU - usually a smaller white mushroom (but not always) growing in the woods instead of grass, with smaller spores than L. luscina. When large it can be differentiated from C. luscina by its higher tendency to be in a forest instead of grass and smaller spores. When small, it can be differentiated from the many other small white Clitocybes by warty, coloured spores. Pablo Alvorado thinks he has figured out what sequences represent this species, and I am following him. We need local sequences to see if what has been reported is really this species.
Clitocybe luscina (sequenced and unsequenced) © Andrew Parker
Lepista #2 - another large, white mushroom in a second clade.
Clitocybe irina EU - a large, white clitocyboid much like C. luscina, but usually growing in forests not grass. It has larger spores. Clitocybe irini var. luteospora is described as having yellowish spores, but I don't know if it is genetically any different. Pablo thinks he knows what sequences of this look like, and I am following him. Many supposed collections of this turned out to be C. luscina, because the spore size wasn't checked. We need local collections to find out if our reports are true or if we have been mistaking L. luscina for this as well. Bigelow reported it from Idaho, and he always checked the spores under a scope for his collections, so I am inclined to believe this could be here.
Lepista #3 - mushrooms that often have purple shades, in a third clade. They are usually found in urban places, not the deep woods.
Clitocybe nuda EU - the blewitt (for blue hat), this large mushroom is purple in the cap, gills, stem and flesh when fresh, and smells slightly citrusy (some say like Tang powdered orange juice). It may be more common with hardwoods.
Clitocybe 'nuda PNW08' - this lookalike sequences quite distinctly and is probably stockier with thicker cap flesh. Reportedly, it will be more common with conifers and have a more unpleasant odor. We have sequences from CA and OR. In the PNW, the real C. nuda is more common, but down in CA, both might occur equally.
Clitocybe glaucocana EU - a uniformly pale pink to lilac large blewit-like mushroom. We have a BC sequence that matches EU sequences, but no local, sequenced photo.
Clitocybe personata EU (=Clitocybe saeva EU) - only the stem has purple tones when fresh. We have EU sequences, but no local sequences to prove it's really here as reported.
Clitocybe subalpina WA - nothing really is known about this large vinaceous grey-brown roadside mushroom, or how to tell it apart from the above. I don't even know that it belongs in Lepista #3 but I'm treating it here because of the vinaceous tones and warty, coloured spores. We'll need a type sequence and modern collections.
Clitocybe brunneocephala CA (=Leucocalocybe mongolica Asia) - usually in grass, this large white to tan capped mushroom with a short, thick stem is said to have the gestalt of a blewitt with no purple at all, even when fresh. It has the same medium sized spores (between C. luscina and C. irina) as Clitocybe praemagna in Lepista #1 and looks very similar to all three. We have CA and matching WA DNA. It is probably most easily confused with Clitocybe luscina, above (known by many other names), but that mushroom is not quite as big, has a longer stem, and has a slightly coloured spore print and more prominent warts on the spores. Asian sequences of Leucocalocybe mongolica are basically the same, so the genus Leucocalocybe should be abandoned, and although Tricholoma mongolicum, the former name of Leucocalocybe mongolica, is an older name than Clitocybe brunneocephala, since this species needs a name in Clitocybe, Clitocybe brunneocephala is the proper name for it.
Clitocybe sordida EU (=Clitocybe tarda MA) - a much smaller collybioid purplish-brown (but mostly in the gills) mushroom. West coast and east coast sequences of C. tarda match many EU sequences of C. sordida, which is probably the older synonym. The holotype sequence of Clitocybe tarda var. alcalina matches all of the above too, so it may not deserve distinct varietal status, at least as shown in ITS. At the time of this writing, Pablo Alvarado has a different concept of C. sordida. If that turns out to be correct, then our species should go by the name C. tarda.
Clitocybe nuda (cap faded) © Bruce Newhouse, C. 'nuda PNW08' © Alan Rockefeller, unsequenced C. glaucocana © Michael Beug
Clitocybe sordida © Ryan Downey, C. brunneocephala © David Arora and Edward Barge
Collybia and Dendrocollybia - click to expand
Of these small, decaying mushroom parasites, at least Collybia lives inside Clitocybe and technically need to be moved to that genus (or they to this).
Collybia tuberosa EU - the sclerotia are orange brown, and elongated seed shaped. Local and east coast sequences match EU sequences.
Collybia cookei EU - the sclerotia are more round and wrinkled in shape. We need local collections to compare with Europe. Back east their sequences are almost 1% different than EU sequences.
Collybia cirrhata EU - lack sclerotia. Local and east coast sequences match EU sequences very well.
Collybia bakerensis WA - is the last Collybia still needing to be moved somewhere else from the great exodus that happened after Gymnopus was erected. It is rare, and not local to those doing the moving, and got overlooked. It belongs in Collybiopsis.
Collybia tuberosa © NAMA and the Field Museum of Natural History, unsequenced C. cookei © John Plischke, C. cirrhata © NAMA and the Field Museum of Natural History
Dendrocollybia racemosa EU - long branches on the stem and round sclerotia. OR and BC sequences match most EU sequences and probably represent this species.
Dendrocollybia pycnoramella n.p. - not yet described, it has shorter, denser side branches and lobed, elliptical sclerotia. It is only known from one collection in CA and one in OR. The OR sequence is 15% different in ITS from D. racemosa.
Dendrocollybia racemosa © Jenny Lippert, D. pycnoramella n.p. © Noah Siegel
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