Danny’s DNA Discoveries – non-waxy Hygrophoraceae of the PNW (Marasmiineae)
There's not just one clade of this family that "looks waxy" or one that "doesn't", it's more complicated than that. A 4-gene study suggests that the waxy genera - Cuphophyllus, Hygrophorus and Hygrocybe s.l. (incl. Gliophorus, Humidicutis and Gloioxanthomyces in the PNW) either became waxy in 3 or more separate events, or all of the genera on this page lost their waxiness in several different events. Nothing much unites these genera, unfortunately, although some are still quite colourful like the waxies (but others are very drab). There's no good way without DNA to know you have a mushroom on this page unless you go through a rather complicated key.
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:
Ampulloclitocybe - click to expand
Clitocyboid. Grey or brown slightly umbonate (yet depressed in age) caps, strongly decurrent white gills. Resembles Cantharellula (below) but without the forking gills that stain red in age and without amyloid spores. (Pseudoclitocybe in the Tricholomatineae has no umbo, no forking, no staining and amyloid spores). Lacking amyloid spores, this genus is especially difficult to separate from Clitocybe. If the umbo is obvious, that will help.
Species mentioned: Ampulloclitocybe clavipes, avellaneialba, avellaneoalba. Clitocybe subclavipes.
Ampulloclitocybe 'clavipes PNW01' - grey or brown cap with a small umbo (yet depressed in age), strongly decurrent white gills, a club stem and often a grape bubble gum odor when fresh (but sometimes not). Bigelow reported Clitocybe clavipes EU (now Ampulloclitocybe clavipes) as very common here, and the odorless, slightly microscopically different Clitocybe subclavipes NY (nobody has bothered to move it to Ampulloclitocybe yet) as being here as well. He noted that out west there was less of a bulb on the stem because the stems were overall thicker to begin with. Well, that character is significant, as our species is likely undescribed, 2% different in ITS. The two named species may be distinct, although only 3 bp and 2 indels separate them, but neither of them have been proven to be here. Our species needs to be described with a thicker stem (making the bulb less pronounced) and that it may or may not have the odor at easily detectable levels. Ampulloclitocybe sub-sub-clavipes anyone?
Ampulloclitocybe avellaneialba WA - (formerly misspelled A. avellaneoalba), usually with a darker grey or brown cap that may have a slightly ribbed cap margin. Lacking the bulb and the odor. This species may occur on or near rotting wood. Don't mistake them for the thinner-fleshed Gerronema atrialbum. Not always darker than A. 'clavipes PNW01', as you can see, the darker of the two A. 'clavipes PNW01' photos is much like the paler of the two A. avellaneialba photos. Then you have to consider the amount of odor and stem bulb.
Ampulloclitocybe 'clavipes PNW01' © Yi-Min Wang, NAMA and the Field Museum of Natural History, A. avellaneialba © Yi-Min Wang and Yi-Min Wang
Arrhenia s.l. - click to expand
There are two very distinct looks for Arrhenia, and some species of the first type don't have strong support for belonging to this genus.
Species mentioned: Arrhenia acerosa, subglobisemen, lobata, auriscalpium, retiruga, chlorocyanea, epichysium, hohensis, latispora, obscurata, onisca, peltigerina, pubescentipes, rainierensis, sphagnicola.
My ITS only tree shows that Arrhenia, as well as Lichenomphalia below, fall into 2 distinct clades far enough from each other and it appears that a few of the species might need to be moved to their own genus. In this case, a few of the omphalinoid Arrhenias appear to be only distantly related to the others. I would like a multi-gene study to verify this, as mine is not the only study to show that all Arrhenias don't necessarily belong together. All oyster-like Arrhenias belong in Arrhenia proper.
1. Moss associates with eccentric stems and possibly poorly developed gills
These species don't appear to all group together (they may have gained or lost this shape in more than one event) so we probably can't give them their own genus. We just have to accept that there are two very distinct looks for Arrhenia. Similar looking mushrooms are found in Rimbachia (Tricholomatineae) and Muscinupta (Hymenochaetales), but those are pure white.
Arrhenia 'subglobisemen PNW06' - brown, short lateral stem, fairly well developed gills. It was reported that we have Arrhenia acerosa EU here, but three sequences turn out to be 2% different than A. subglobisemen instead and much further away from A. acerosa. If anybody thinks A. acerosa is really here, save a collection because we have EU DNA to compare to. It is a complex where ITS DNA can vary by 2%, and has spores that vary widely in shape, often being almost round like in A. subglobisemen, so that may have created the confusion, and if the spore shape corresponds with certain ITS sequences in the complex, it could also mean A. acerosa will be split up into more than one species.
Arrhenia cf. lobata EU - brown, stemless, poorly developed wrinkle-like "gills". Reported from the PNW, and we have EU DNA to compare to, but no collections to prove it's really here. We need some, especially since it's similar to the previous species.
Arrhenia 'auriscalpium PNW07' - long lateral stem, slightly veined underside. One North American sequence is 2% different than type area EU sequences. We don't have local sequences (we need them) but ours is probably a sister species and not the real thing.
Arrhenia 'retiruga PNW08' - paler tan or grey felty cap, stemless, almost smooth underneath. One WA sequence is 3% different than 2 competing concepts in the EU for what this species is. Ours likely needs a new name.
Arrhenia 'subglobisemen PNW06' © Kitty Lundeen-Ness, A. 'auriscalpium PNW07' © Ben Woo, A. 'retiruga PNW08' © NAMA and the Field Museum of Natural History
2. Brown (rarely green) omphalinoid mushrooms
'Clitocybe' atroviridis WA (=Arrhenia chlorocyanea EU?) - This unique, beautiful dark green omphalinoid, found on the ground often in moss, was described from WA as Clitocybe atroviridis but that is thought to be a synonym of the older EU species Arrhenia chlorocyanea. We don't have any EU sequences of that mushroom to compare to, and given that our local species of other Arrhenias has turned out to be different than the EU counterpart that was thought to be the same, we need EU sequences to find out for sure. If not, we'll need to rename ours to Arrhenia atroviridis.
unsequenced Arrhenia chlorocyanea © Ben Woo
'Arrhenia2' telmatiaea UK (='Omphalina' fusconigra UK) - recognizable by its dark brown smooth cap and growth in sphagnum. It is in the "other" clade. Two OR sequences match the type sequence of this newly described species. One theory is that Bigelow's reports of Arrhenia onisca, which could not have been that species as it is a synonym of Lichenomphalia ericetorum, represent A. telmatiaea.
'Arrhenia2' philonotis EU - a paler brown smooth cap and growth in sphagnum. Also in the "other" clade. Two OR sequences match the type sequence.
'Arrhenia2' telmatiaea and 'Arrhenia2' philonotus © Connor Dooley
'Arrhenia2' bigelowii MI (=Arrhenia gerardiana var. fusca NY) - recognized by having a scaly cap in sphagnum moss. One WA collection matches the type sequence. Formerly, it was thought that this species/variety always had a darker cap than Arrhenia gerardiana, but our local collection confirms this is not the case. Dark capped collections are A. bigelowii. Not-so-dark capped collections could be either A. bigelowii or A. gerardiana (indistuingishable so far except for by DNA). Therefore, local reports of A. gerardiana need confirmation as it is not yet confirmed by DNA from the PNW. This is in the "other" clade of Arrhenia.
'Arrhenia2' bigelowii © Luca Hickey (2 images)
Arrhenia 'latispora PNW09' - a moss species (not just sphagnum?) with wide spores (7.8-9.5 x 6.5u) and said to have a geranium odor at least in the EU, where it was described. 2 collections from OR match the EU epitype sequence, however the species is described as a pleurotoid oyster-like mushroom, very different than what ours looks like. Stay tuned.
Arrhenia 'obscurata PNW11' - Arrhenia obscurata UK (=Omphalina atrobrunnea UK) is found on sandy soil or moss, reported from ID and BC. We don't have any UK sequences, but Voitk provides some ENA sequences. Our one WA collection is at least 2% different in ITS. It's possible ours will match true UK sequences of A. obscurata, but until we prove that, I'm giving it a code.
Arrhenia 'latispora PNW09' © Connor Dooley, and NAMA and the Field Museum of Natural History, Arrhenia 'obscurata PNW11' © Matthew Koons
The following lookalike dark brown omphalinoid species, from the ground or moss, are reported from here, and below them I show four sequenced local collections that have not been matched up with anything specific yet.
'Arrhenia2' epichysium EU - it should be investigated if reports of this dark brown species on wood are really Arrhenia telmatiaea, above, or something else. We need collections from wood apart from sphagnum. We have some sequences of A. epichysium from the EU to compare to and it is a complex of at least 2 species. It's in the "other" clade.
Arrhenia cf peltigera NY - on Peltigera lichen. Bigelow claims to have found this in Idaho, but we need local collections to sequence to verify if it's really here. We don't even know what its DNA looks like, the three east coast sequence that purport to be this are all different sister species.
'Arrhenia onisca' EU - Bigelow reported this from the PNW on moss or burned moss, but it's now known that this name is a junior synonym of Lichenomphalia ericetorum, so nobody knows what Bigelow really found. My leading theory is Arrhenia telmatiaea, described above.
'Arrhenia sphagnicola' EU - This name too has officially been declared a synonym of Lichenomphalia ericetorum, so it is no longer being used. This name has been used for reports of A. bigelowii and A. telmatiaea back when A. sphagnicola was incorrectly thought to be a synonym of Omphalina gerardiana and O. fusconigra, but this name should not be used anymore.
Arrhenia rainierensis WA - on soil or moss with interesting shaped spores. Described from here but we don't have any reliable sequences. I don't even know which clade it's in.
Arrhenia pubescentipes ID - on soil or moss. No local sequences, so no idea which clade it's in.
'Arrhenia2' hohensis WA - on soil or moss. Described from here but we have no sequences. One mycologist suggested it might be similar to or the same as the EU species Arrhenia obatra, if so, that species appears to be one of those possibly needing to be moved to the new genus, so maybe A. hohensis does too, but maybe not.
The first two local species below are in the true Arrhenia clade. The last two are in the "other" clade. They have slightly different 5.8s sequences than other Arrhenias, which is unusual, but as I've seen it in more than one sequence, I think the sequences are good.
Arrhenia PNW01 - 1 collection from OR
Arrhenia PNW10 - 2 collections from WA, in Polytrichum moss, with a somewhat greasy cap with a wavy margin.
'Arrhenia2' PNW03 - a very dark, terrestrial species not with moss. Found 3 times in WA so far. Bigelow found it in Idaho and mistook it for A. peltigera.
'Arrhenia2' 'epichysium PNW12' - very dark, terrestrial, on non-sphagnum moss. Found once in WA. It is close to one possible concept of the EU species A. epichysium, which is found on wood.
'Arrhenia2' PNW04 - 1 collection from WA
'Arrhenia2' PNW05 - on soil, with highly contrasting colour striations on the cap. 1 collection from OR.
Arrhenia PNW01 OR © Bitty Roy, Arrhenia PNW10 © iNaturalist user fish_narc (2 images)
'Arrhenia2' PNW03 © Yi-Min Wang, 'Arrhenia2' 'epichysium PNW12' © Larry Martin, 'Arrhenia2' PNW04 © Yi-Min Wang, 'Arrhenia2' PNW05 © Connor Dooley
Cantharellula - click to expand
Clitocyboid. Grey or brown slightly umbonate (yet depressed in age) caps, strongly decurrent white gills. So far that describes Ampulloclitocybe, but these have forking gills that stain red in age and amyloid spores. (Pseudoclitocybe in the Tricholomatineae has no umbo, no forking, no staining and amyloid spores).
Species mentioned: Cantharellula umbonata, oregonensis.
Cantharellula umbonata EU - one CA sequence is a couple indels different and has a few ambiguous locations, but it seems to be a dirty sequence at the beginning so I'm not sure we're seeing all the differences. We probably have this EU species here, but some local collections will be needed to be sure. East coast sequences match the EU very well except for a few ambiguous locations.
'Cantharellula' oregonensis OR - In 1942 Singer thought that Murrill's 1913 Clitocybe oregonensis should be called Cantharellula oregonensis, but it doesn't have forking gills and doesn't stain red, so I don't think he was right. Then he himself changed his mind in 1961 and called it Pseudoclitocybe oregonensis. I think that might be right, see the Pseudoclitocybe page for more information.
Cantharellula umbonata © Steve Trudell
Cantharocybe - click to expand
A large and stocky, bright yellow, strongly decurrent mushroom often with an off centre stem.
Species mentioned: Cantharocybe gruberi.
Chrysomphalina - click to expand
Colourful (almost waxy), small mushrooms growing on wood with dry caps. They also have decurrent gills an may be hygrophanous.
Species mentioned: Chrysomphalina aurantiaca, grossula, chrysophylla
Chrysomphalina aurantiaca OR - small, completely orange little waxy, with many local sequences.
Chrysomphalina grossula EU - yellow-green little waxy. One or two consecutive and equal chunks, four nucleotides long (GCAT) may be missing from our DNA compared to European DNA, but as a single event could drop all those characters, that probably doesn't represent enough of a change to constitute ours being a separate species. Otherwise, there is only 1 or 2bp difference.Chrysomphalina chrysophylla EU - brown capped, golden yellow little waxy. Washington and California DNA matches European DNA, so our local species is the real thing.
Chrysomphalina aurantiaca and grossula © Kit Scates Barnhart, Chrysomphalina chrysophylla © A and O Ceska
Lichenomphalia and Protolichenomphalia - click to expand
One of two known PNW basidiolichen genera (algae usually pair with ascos to form a lichen). The other is Multiclavula, a club. Yellowish tan to brown omphalinoids with grainy or leafy lichen material at the stem base (unlike the very similar Arrhenia, Omphalina, etc.). Our most abundant species may need a new genus.
'Lichenomphalia' ericetorum EU (=Lichenomphalia umbellifera) - dull colours, granular green lichen material at the base. Local sequences match EU sequences within a couple of bp plus a few ambiguous locations. However, this species is not closely related to the type species of Lichenomphalia, described next, and has been placed in its own subgenus, Protolichenomphalia. My ITS-only studies show that it may belong in its own genus, and some other studies have shown that as well "depending on the methods, genes and taxa used" as quoted from another study. A multi-gene study will have to resolve the issue, in the meantime, there is a chance it will have to be renamed to Protolichenomphalia ericetorum. The other interesting thing about this species is that Lichenomphalia umbellifera turned out to refer to 'Marasmius' epiphyllus (and there's a whole story about that on that page), so that name can no longer be used for this species. The proper name is now L. ericetorum, what used to be just a junior synonym.
Lichenomphalia cf hudsoniana Nunavut, Canada - more brightly coloured, with leafy green lichen material at the base. It has a somewhat coloured spore print (orange-yellow) and a pubescent stem, growing on tundra in the far north. It's unclear how far down BC it can be found. We need local collections to prove it's here since we only have ENA and Scandinavian DNA. This is the type species of the genus, and gets to stay in Lichenomphalia.
Lichenomphalia cf oreades NH (=Clitocybe albimontana NH) -brown to dark brown cap and stem (that is pubescent at first) with paler gills growing from moss and sandy soil. The brown colours make it look exactly like several Arrhenia and one Omphalina, except this species will have granular green lichen material at the stem base. Bigelow reported this from WA, but we need a modern collection to prove it. It is a true Lichenomphalia, as we have the type sequence of L. oreades. One EU sequence of the older Lichenomphalia velutina matches ours (but other sequences with that name from elsewhere do not). The type of L. velutina is a drawing, not a real mushroom, so nobody knows for sure what it is and until that is sorted out, I am not ready to state that we should use that older name. The folks who got the type sequence of L. oreades also got the type sequence of L. grisella, thought to be a newer synonym of L. velutina, and it turns out that L. grisella is not the same species as L. oreades. We need local collections to see if Bigelow really found Lichenomphalia oreades when he discussed Clitocybe albimontana. For now, I'm assuming so.
Lichenomphalia ericetorum © Steve Trudell
Pseudoarmillariella - click to expand
Medium sized thin-fleshed funnel shaped Clitocyboids on conifer logs. Yellowish-tan radially streaky cap, occasional forked gills that are sometimes quite yellow. Amyloid spores. No other fair-sized white spored mushroom quite like it grows out of logs, even though if you found it growing from the ground you'd be stumped.
Spodocybe - click to expand
Small somewhat felty grey Clitocyboids.
Species mentioned: Spodocybe senilis, trulliformis.
Spodocybe 'trulliformis PNW01' - probably 6-7% different than the EU Spodocybe trulliformis and in need of a new name. This species has distinct felty scales on a grey cap.
Spodocybe 'senilis PNW02' - This doesn't seem to be as dark grey nor as felty/scaly as PNW01. It may have been wet, but this species seems to have a streaky, watery-translucent grey cap, easily differentiated from PNW01, and the cap is barely felty if at all. This kind of reminds me of Clitocybe senilis, reported from the PNW as like Spodocybe trulliformis but less scaly on the cap. It should be investigated if reports of that species are this present species. We don't have any sequences of the EU species Clitocybe senilis to compare to.
Spodocybe 'trulliformis PNW01' © Danny Miller and Richard Morrison, S. 'senilis PNW02' © Connor Dooley and Yi-Min Wang
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