Danny’s DNA Discoveries – Omphalotaceae of the PNW (Marasmiineae)
Click here for my Pictorial Key to the Collybioids, where most Omphalotaceae are found.
This is one of the largest families in the Marasmiineae sub-order, containing many species of collybioid stature (and sometimes marasmioid stature). Usually, a white spored mushroom with a tough, cartilaginous stem is somewhere in this sub-order (not necessarily here) but many species could not be properly placed until DNA came along. There is a vast "white spored multitude" of species, as they say in Mushrooms of the Redwood Coast, and it can be very difficult to identify a mushroom even to sub-order never mind family never mind genus. This is definitely a case of "learn the species to learn the genera" as often, nothing much unites them.
Given that Campanella from the Marasmiaceae appears to be closer to this family than the family it is currently placed in, and the fact that there is not much genetic distance between the two families, the Omphalotaceae may not deserve their own family and it may be best to think of the Omphalotaceae as part of the Marasmiaceae, the family that contains most other mushrooms of marasmioid stature.
These have not been well studied in the PNW, and we keep seeing species we don't recognize, so I knew we would find surprising new species when we starting sequencing them. They are called collybioid because all mushrooms of that stature used to be called Collybia. Then we found out that the group was very artificial and those mushrooms were split, and it turned out that the first and true Collybias were atypically small and actually part of Clitocybe in the Tricholomataceae, so many collybioid mushrooms were moved to Gymnopus. Then DNA discovered that some Gymnopus were inside Marasmiellus, and some former Marasmiellus didn't belong there. Finally, it was discovered that Marasmiellus was a word that shouldn't be used, since Collybiopsis is an older synonym for that genus. There is no good way to tell Gymnopus and Collybiopsis apart without DNA, so I am treating them in one section.
All three garlic mushroom clades are in this family. You can identify your garlic-like sub-order Marasmiineae mushrooms this way:
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
Here are some of the newest, most interesting results of the study:
Gymnopus and Collybiopsis (Marasmiellus) - click to expand
The main genera for many collybioid mushrooms (white spores, slender stature and never that large, possible tough stem, attached gills that aren't usually decurrent). Some marasmioid mushrooms are found here too (even smaller size, distant gills, dark stem bottoms). Some of the collybioids smell of garlic and will be confused with Mycetinis and Paragymnopus, below, which are smaller marasmioids. Uniting these two genera (and making them hard to tell apart) is the fact that although they have typical repent, filamentous hyphae in the cap cuticle (and not spherical cells) the hyphae are often knobby or branched with interesting outgrowths.
mentioned: Gymnopus dryophilus, aquosus, alpinus, junquilleus, subsulphureus, ocior, earleae, vernus, erythropus, spongiosus, semihirtipes, fuscopurpureus,
alkalivirens, brassicolens, dysodes, impudicus, polyphyllus,
putillus, fusipes, contrarius, quinaultii, striatipes. Marasmius androsaceus.
Collybiopsis peronata, confluens, confluens subsp. campanulatus, bakerensis, luxurians, subpruinosus, villosipes, eneficola, hasanskyensis, californica,
pluvius, vaillantii. Marasmiellus papillatus, filopes. Microomphale
1a. Gymnopus - collybioid (caps can be >2.5 cm across), odorless, not turning green in KOH (easily confused with section 3 - collybioid Collybiopsis)
Gymnopus dryophilus EU - a somewhat hygrophanous warm orange-brown cap and stem, named after oak but found ubiquitously throughout any forest. No bulb and white rhizomorphs.
Gymnopus aff. dryophilus EU - Some local sequences match very well to the type sequence, but most of them differ by 7 bp or so in ITS2, although they are still almost an exact match (only 1 bp different) in ITS1. It's a little unusual to find such asymmetry in areas that supposedly randomly mutate. A 2013 Antonin paper could find no morphological or ecological differences between them, and for now it is all considered one species, only distinguishable by ITS2, but I am noticing that these "alternate clade" collections often have yellow or orange-brown (the cap colour) on the stems, whereas the collections that better match the type sequence have white or orange-brown on the stems, without yellow. That usually makes the caps of 'aff. dryophilus' darker than the stem. Hmmm...
Gymnopus aquosus EU - a lookalike we never noticed was here (we undoubtedly thought they were G. dryophilus) until DNA showed us it was. It supposedly has a stem bulb and pink rhizomorphs. ITS DNA of G. aquosus varies a bit more than the usual 0.5% around the world, but it's all considered one species.
Gymnopus 'aquosus PNW04' (alpinus EU/junquilleus NC/subsulphureus KS complex) - some sequences of G. aquosus are about 1% different, practically exactly the same as the ITS DNA of a few sister species that all share the same ITS DNA. Those species have a very dark red-brown cap (G. alpinus) or are entirely bright yellow (G. junquilleus and G. subsulphureus). At least two of them have been shown to interbreed, so perhaps our local collections with this sequence do not represent a separate species but represent G. aquosus of the first group with "pure" ITS DNA hybridizing with one of the different looking species but still retaining its own character. This is an interesting puzzle. We need collections with this different DNA to compare features.
Gymnopus dryophilus and aff. dryophilus (yellow stem) © Bruce Newhouse and Yi-Min Wang, G. aquosus © Buck McAdoo, G. 'aquosus PNW04' © Buck McAdoo
Gymnopus ocior EU - a darker, colder wine-coloured cap, possibly with yellowish gills, growing in disturbed urban places like wood mulch. This species has serrated gill edges like Rhodocollybia, in fact it is believed that Rhodocollybia extuberans is a synonym of this species, but see below under Rhodocollybia for a discussion on that name. This was also not known from the PNW until we sequenced collections that were either mistaken for G. dryophilus or that were not recognized. Two WA sequences match EU sequences quite well.
Gymnopus 'earleae PNW05' GA (Gymnopus vernus EU misapplied?) - usually a darker chestnut brown colour than G. dryophilus, but may also be orange like the latter, found in spring as well as fall. (It is similar to the green KOH species G. aff. spongiosus grp below, but usually less striate. It is also similar to but less striate than Collybiopsis villosipes, further below, but that species has a hairy stem). This was also not previously known from the PNW, having also been mistaken for G. dryophilus or just not recognized. There are 2 competing concepts from eastern North America about what G. earleae is. Out west we find one of them, but I can't say for sure if it's the real one or the unnamed lookalike. We need more GA sequences, if not the type to find out. It's possible that reports of the similar G. vernus EU represent this (it's a very similar spring mushroom) but I don't know, so if you think you find the real G. vernus here, save it.
Gymnopus ocior © Buck McAdoo, G. 'earleae PNW05' © Buck McAdoo
'Gymnopus' striatipes NY - not a Gymnopus, but in the Rhodocollybia collection of genera described below.
1b. Gymnopus - collybioid, odorless, turning green in KOH
Gymnopus erythropus EU - notable for its strongly two-toned stem grading from dark reddish brown below to paler orange above, but some collections don't have much of a pale upper stem and are difficult to identify. The cap fades from darker brown to lighter brown. Many local sequences match many EU sequences. Occasionally collections smell like rotting cabbage, even though this species is not in the garlic clade covered next. Those collections have identical sequences to the ones that don't have an odor.
Gymnopus spongiosus PNW06/PNW07 ('fuscopurpureus'?) -dark chestnut brown and strongly striate. G. 'earleae PNW05' above doesn't turn green in KOH and is usually less striate. Collybiopsis villosipes below doesn't turn green in KOH and has a hairy stem. Two different groups sequence 5 bp and 1 indel apart from each other in ITS1 only (about 3% different) and may represent two different species. We need ITS2 of the second species and more collections of the first to sort this out. The closest named species are G. spongiosus (with a completely hairy stem) and G. semihirtipes (with a hairy stem bottom). PNW06 is smooth stemmed and did not turn green in KOH (we need more collections to check them). PNW07 has an entirely pubescent stem and did turn green in KOH. Gymnopus fuscopurpureus EU and Gymnopus alkalivirens VI have been reported from the PNW but we don't have reliable sequences of either of those, although it starting to seem unlikely that either species is found in the PNW.
Gymnopus 'fuscopurpureus PNW08' - starting out warm brown, but the cap, gills and stem seem to grey in age. Bald stem. KOH turns green. One WA sequence matches an AZ sequence, but there are many competing concepts for what a sequence of the EU species G. fuscopurpureus really is, and this seems unlikely to be it. So what is it?
Gymnopus erythropus © Steve Trudell, G. aff. spongiosus PNW06 and PNW07 © Buck McAdoo, G. 'fuscopurpureus PNW08' © Buck McAdoo
Gymnopus PNW01 - Distant ochre gills, orange brown cap, possible fuzzy stem, turns green in KOH. Spores (5.5-6.8) x (3.6-3.7)u.
Gymnopus PNW02 - Close gills. Orange brown cap with a vinaceous brown disc. Known from one WA sequence. A sister species to PNW01 that should turn green in KOH but our only collection did not appear to.
Gymnopus PNW01 © iNaturalist user joemat, green KOH reaction © Yi-Min Wang, G. PNW02 © Buck McAdoo
1c. Gymnopus - collybioid, garlic/cabbage odor (but note that G. erythropus above may smell of rotten cabbage and 'Micromphale' arbuticola is a small marasmioid)
Gymnopus impudicus EU (Gymnopus dysodes misapplied?) - dark red brown cap (when fresh), possibly paler stem, everything fading in age, distant gills and garlic/onion odor found in the wild. The very similar Gymnopus dysodes (with a very striate cap, much larger spores and found in woodchips) was reported from the PNW too but we need collections to see if it really is here or if those were cases of mistaken identity of G. impudicus (the caps weren't very striate so I have my doubts that G. dysodes is really here). One other thing to check when figuring out which (or both) species is here is whether or not KOH turns green with either species, as it is supposed to at least with G. dysodes, even though it doesn't clade with other KOH positive species. We have a couple of EU sequences of G. impudicus that could be the real thing that 2 WA collections match, but we don't have a type sequence to prove it.
Gymnopus 'polyphyllus PNW03' - a pale yellow-tan cap with very crowded gills and a strong garlic odor. There was one vague 30 year old report of it in the PNW, and we finally found something that looks like it, but it is an unnamed relative and not the real thing described from NY. We should keep collecting it to see if the real thing is also here. Our local collection smelled more like rotting cabbage than garlic.
Gymnopus cf. brassicolens EU - much like G. erythropus but always smells like rotting cabbage/garlic. Its two toned stem is almost blackish purple on the bottom. It is reported from CA and to be looked for in Oregon, but no local collections yet.
'Micromophale' arbuticola CA - a much smaller marasmioid garlic mushroom growing on the bark of madrone. It has yet to be moved to Gymnopus but should, in fact it is in the same clade with all these others, probably closest to Gymnopus foetidus, it just unfortunately evolved to be smaller than the others. We need ITS sequences of it; we don't have any yet. Our common 'Micromphale' perforans was moved to Paragymnopus when Micromphale was split between the two genera.
Gymnopus impudicus © Buck McAdoo, G. 'polyphyllus PNW03' © Colin Meyer
2. "Horsehair" Gymnopus - marasmioid (caps ~1cm across), especially wiry black "horsehair" stem
Gymnopus androsaceus EU - smaller than the other Gymnopi, with a mostly black (except possibly at the top) "horsehair" stem that is very thin, wiry and black. The odor is mild. A brownish cap fades in age, but is usually not as pale as in the similar Pseudomarasmius, below, whose stem may not be as wiry. Collybiopsis californica, also small, lacks a truly wiry stem. Paragymnopus species sometimes have a rotten cabbage odor, but if they don't, they will be very difficult to differentiate as their stems can be somewhat wiry too. Formerly in Marasmius. One Idaho sequence and a recent WA sequence are at least 6 bp different (1%) than numerous EU sequences, but no micro nor ecological differences can be found. East coast sequences are also about 1% different than both of those, so I suspect G. androsaceus has a little more genetic variation in ITS than usual, caused by geographical isolation.
Gymnopus androsaceus © Richard Morrison
3. Collybiopsis - collybioid (caps can be >2.5cm across) (easily confused with section 1a - collybioid Gymnopus without KOH reaction and without garlic odor)
Collybiopsis peronata UK - usually yellow tones, a hairy stem base and distant gills. Introduced to the PNW around the 90s but it really loves it here as it quickly became one of our most abundant, ubiquitous mushrooms. DNA worldwide matches DNA from Scotland, the type area.
Collybiopsis peronata © Fred Rhoades and Steve Trudell
Collybiopsis confluens EU -Collybiopsis confluens subsp. campanulatus NY - tan to orange-brown colours (brighter when fresh), somewhat of a collar around the stem where it attaches to the cap that can look like a ball-in-socket joint, and crowded gills. The North American subsp. campanulatus is more common here, but you can occasionally find the original EU species as well. There doesn't seem to be a good way to tell them apart without sequencing, they even appear to interbreed, but with the populations 3.25% apart in ITS, they are distinct genetically.
Collybiopsis confluens (unknown subspecies) © Steve Trudell
Collybiopsis PNW02 - a decorated stem that is black at the base. Two OR and two WA collection are probably an unnamed species. One collection didn't look like the others (the cap was very pleated and the entire stem black) but now we have enough collections to know it's typical look.
Collybiopsis PNW03 - this appears to have a two or three toned stem like Gymnopus erythropus but almost black as the bottom colour like Gymnopus cf brassicolens above, but this species doesn't have the garlic odor. Only found once so far in southern OR.
Collybiopsis PNW02 © Yi-Min Wang, Collybiopsis PNW03 © Jonathan Frank
Collybiopsis luxurians NC - our largest species, brown cap and pale brown stem (here shown with darker stems than normal), growing sometimes clustered on rotten wood and in urban places like wood chips, gardens and grass. Our sequences match NC sequences.
Collybiopsis luxurians © Eric Chandler
Collybiopsis cf. subpruinosus Jamaica - a tan coloured species whose cap can become quite pleated. There are one set of sequences from WA, CA and Chile that all match, but on the other hand, another sequence from Puerto Rico is 10% different, and that's a lot closer to Jamaica, the type area, so I'm not sure which is the real one.
Collybiopsis villosipes Australia - a colder darker brown, with similarly coloured gills. Somewhat striate. Stem covered in fine hairs. It does not have yellow tones like Collybiopsis peronata. KOH does not turn green. It could still be confused with one of the undescribed species on this page. They say it probably evolved here on the west coast and then travelled the world where it was first described in Australia. Not all sequences appear to be the same, but some sequences have 5 ambiguous locations accounting for most of the differences.
Collybiopsis cf subpruinosa © Buck McAdoo, Collybiopsis villosipes © Ann Goddard
Collybiopsis eneficola NFLD - pale tan (almost white) in all stages of freshness with subdistant gills and a hairy stem. Joanne Lennox found this in Idaho back in the 70s and called it Collybia tomentipes, but it finally got described for real in 2014 in Newfoundland. Her sequence is only of ITS1 but it matches the NFLD type sequence exactly. It has not been found since so we don't have a local photograph. We need recent local collections. It may be a typically small species, caps <2.5 cm across.
Collybiopsis eneficola Newfoundland © Andrus Voitk
'Collybia' bakerensis WA - a small, pale collybioid growing on large conifer logs, it is fairly non-descript with a little different stature than Mycena (Mycenaceae) and Clitocybula (Porotheleaceae) (found in other families of the Marasmiineae sub-order) that are also white with white spores and non-decurrent gills that grow on larger pieces of wood. Being a rather small Collybiopsis and growing directly on logs, it is not easily identified. This species still needs to be moved to the genus Collybiopsis. We don't have a type sequence, but we have a few local collections that all agree.
unsequenced 'Collybia' bakerensis © Andrew Parker
4. Collybiopsis - marasmioid (smaller with distant gills) (most easily confused with Pseudomarasmius below and the the usually darker capped Paragymnopus that lack a strong odor)
Collybiopsis californica CA 1987 - smaller and more delicate than other Collybiopsis, with a pale cap, distant gills and more brightly coloured red-brown stem that is somewhat two toned. Described from oak litter in California. Note that Marasmiellus californicus refers to something else, Paragymnopus sequoiae.
'Marasmiellus' pluvius BC 1982 - almost the same, described from conifer litter in British Columbia. This still needs to move to Collybiopsis.
Well, guess which sequence three WA collections on conifer litter matched? C. californica, not M. pluvius. The only M. pluvius sequence we have is from Tennessee, so I have a theory that actually, the two species are the same and can live on either kind of litter and that the TN species is undescribed. Stay tuned.
Collybiopsis californica © Connor Dooley
Collybiopsis 'hasanskyensis PNW01' - typical reddish brown cap fading to very pale, but the stem usually retaining some colour, occasionally almost two toned, which makes it much like Gymnopus erythropus above, but this species appears to have very distant gills. Two WA collections surprised us by being 7 bp and 3 indels different than official sequences of this Russian species. That may be enough to imply that ours needs its own name.
Collybiopsis 'hasanskyensis PNW01' © Yi-Min Wang (2 images) and Connor Dooley
'Collybiopsis' 'omphaloides PNW04' - a whitish cap and multi-toned stem ranging from black at the bottom to reddish brown or paler at the top. Probably odorless. It somewhat resembles Collybiopsis californica, for one. It was found once in OR, this species is 4% different than a NZ sequence that might be Marasmiellus omphaloides. However, these sequences are more than 25% different in ITS from any other known sequences. What happened? Did horizontal gene shift change ITS in these species only? We will have to sequence other genes to find out, but for now I'm going to assume this is a species of Collybiopsis.
'Collybiopsis' 'omphaloides PNW04' © Connor Dooley
Collybiopsis cf. vaillantii EU - perhaps a paler cap, more decurrent gills, and stem that only darkens at the bottom (compared to the previous two, which it is not that closely related to). Reported from the PNW, but not proven, perhaps reports of this represent one of the above two species. We need local collections, but we also need EU sequences to compare to, as we only have one purported ENA sequence to compare to right now, which might be misidentified.
'Marasmiellus' papillatus NY - this small nondescript species was reported from the PNW but not proven, we don't have any DNA yet so we don't even know if it really belongs in Collybiopsis. We need collections.
'Marasmiellus' filopes NY - this mildy garlicky small species was reported from the PNW but not proven, we don't have any DNA yet so we don't even know if it really belongs in Collybiopsis. We need collections.
5. Other Gymnopi to look for
'Gymnopus' quinaultii WA - is small. Nobody is sure what this local species is, and there is no DNA yet, so we desperately need collections to find out for sure what genus it is really in and if it really is a unique species. Most of the other small Gymnopus described in the same paper turned out to belong to Paragymnopus (below).
'Gymnopus' putillus - wine-red cap and gills contrast with a very pale entirely pubescent stem. Reports of this species being here have not been verified, nor do we even know what it really is. One ID collection from 100 years ago with this name turned out to sequence inside Rhodocollybia (see Rhodocollybia PNW07 below).
Gymnopus cf. fusipes - has a spindle shaped rooting stem, distant gills and spots red-brown all over. Nobody really believes the vague report that it might be in the PNW, but keep an eye out for it anyway.
'Gymnopus' contrarius NY - this probably needs a genus of its own (it is sister to Neonothopanus and Omphalotus), so it certainly is a contrary species, but thankfully it is not recorded from the PNW yet. Just in case it ever is, I'm mentioning it here.
Mycetinis - click to expand
Garlic odors - smaller marasmioid mushrooms than the garlic Gymnopi, and with two-toned black bottomed stems. Similar to Paragymnopus which have rotten cabbage odors, but Mycetinis has spherical cap cuticle cells.
Species mentioned: Mycetinis scorodonius, scorodonius forma diminutivus, allaceus, salalis, copelandii, applanatipes
Mycetinis scorodonius EU/scorodonius forma diminutivus WA - with a bald stem. It is typically found here with a cap <1 cm in the small forma diminutivus (of which we have the type sequence and a BC and WA sequence), but there are reports of the larger M. scorodonius (caps >1 cm) being found here as well. They share similar ITS DNA, differing by around a couple of bp.
Mycetinis salalis BC -caps >1 cm, tomentose stem. This is our most common species growing on salal debris, differing by host and microscopically (one of the "ridiculously long spored" species).
Mycetinis cf copelandii CA -the other "ridiculously long spored" species, although not as ridiculously long as M. salalis. All data so far indicates the two ridiculously long spored species have the same ITS DNA. This is by far the older species. It may be found in at least southern Oregon. We need collections to prove it.
Mycetinis applanatipes CA - from high elevations with red fir, probably also with a tomentose stem. It differs microscopically from all of the above. It was reported from OR (and BC?), but we need collections to prove it. We have an official CA sequence to compare to. It is officially a rare red listed species in CA.
Mycetinis scorodonius forma diminutivus © Yi-Min Wang, M. salalis © Steve Trudell
Paragymnopus - click to expand
May have a rotten cabbage odor, but recent studies are showing that they also may not, making their identification more difficult than we thought. They are small, brownish capped marasmioid mushrooms with tri-coloured stems that are black on the bottom. When strongly odiferous, they are most similar to Mycetinis, but those have odors more consistenly described as garlic instead of rotten cabbage, although the odors are similar. Mycetinis can easily be differentiated microscopically by a spherical cap cuticle. The small Collybiopsis californica, Gymnopus androsaceus (with a blackish wiry stem) and Pseudomarasmius pallidocephalus (with a pale brown cap) are easily confused with those Paragymnopus that lack the strong odor.
Species mentioned: Paragymnopus perforans, perforans subsp. transatlanticus, ponderosae, pinophilus, sublaccatus, sequoiae, Gymnopus bulliformis, fragillior, glabrosipes
Paragymnopus perforans EU/perforans subsp. transatlanticus PQ - the European and eastern North American species long assumed to be the species found here too, but as we now have at least 5 more species described from the west coast, the reports that this species is actually here are now doubted and need confirmation. Genetically, there seems to be a bit of a difference between EU and ENA sequences, although in ITS only the picture is not very clear. As this is the only species in the genus to reliably smell like rotten cabbage, we need strong smelling local collections to find out what they are.
Paragymnopus sequoiae CA (=Paragymnopus sublaccatus BC?) - BC, OR and CA ITS sequences of both species match very closely and seem to represent the same sister species of Paragymnopus perforans, although the authors did not synonymize them. P. sequoiae is the older name. It is reported as developing a latent rotten cabbage smell. The odor of P. sublaccatus was not noted. Both local collections (WA + OR) with matching ITS of this group appear to have completely lacked the rotten cabbage odor.
Paragymnopus ponderosae CA (=Paragymnopus pinophilus NC?) - We have an Idaho sequence officially recognized as P. ponderosae and many type locale sequences of P. pinophilus collected by the author of that species that certainly represent P. pinophilus. Their ITS is almost identical, but the mycologists who moved them both into Paragymnopus did not synonymize them yet. As they were both described in the same publication, I'm not sure which name has priority if/when they are synonymized. Until then I will use the local west coast name, P. ponderosae. P. ponderosae was described without the odor.
'Gymnopus' fragillior WA - no official DNA nor photo, this was described on the basis of a single dried collection that looked different from other collections of P. perforans. It probably needs to be moved to Paragymnopus. It did possess a garlic-type odor. One recent collection from WA without any odor but that matched 'Gymnopus' fragillior microscopically was sequenced. Its sequence did not match any of the above, so it could be this species. If it is not 'Gymnopus' fragillior, we have to figure out what it really is.
'Gymnopus' bulliformis WA - no DNA nor photo, also described on the basis that one dried collection looked different from other collections of P. perforans. It probably needs to be moved to Paragymnopus. Unfortunately, the odor was not recorded.
'Gymnopus' glabrosipes WA - no DNA nor photo, also described on the basis that one dried collection looked different from other collections of P. perforans. It probably needs to be moved to Paragymnopus. Unfortunately, the odor was not recorded.
Paragymnopus sequoiae © iNaturalist user winterwren22, 'Gymnopus' cf fragillior © Buck McAdoo
Rhodocollybia, Connopus, Pseudomarasmius, etc. - click to expand
Connopus aff. acervatus (bagleyensis n.p.) is a very tightly clustered burgundy collybioid.
Pseudomarasmius pallidocephalus is a small, pale capped marasmioid with a dark stem that is fairly thin but not as wiry as Gymnopus androsaceus. Easily mistaken for Collybiopsis californica and the somewhat darker capped Paragymnopus.
Rhodocollybia are larger and fleshier, with fleshy (not tough) stems that may be long and rooting. The spore print is somewhat coloured. The gill edges are often serrated. Rhodocollybia may need to be split, with at least 'Rhodocollybia' subsulcatipes possibly needing a new genus. 'Gymnopus' striatipes also needs a new genus. The only species guaranteed to stay in Rhodocollybia are the Rhodocollybia maculata group of species. The others clade with that type species with weak support at best.
Species mentioned: Rhodocollybia butyracea, butyracea forma asema, badiialba, olivaceogrisea, maculata, oregonensis, unakensis, extuberans, subsulcatipes, Gymnopus striatipes, Collybia cylindrospora
Connopus aff. acervatus (bagleyensis n.p.) - burgundy caps that fade, but the stems still remain bright, recognized by being very tightly clustered in large quantities. Joanne Lennox back in the 70's was going to call our species Collybia bagleyensis, but she never finished formally describing it. It turns out she might have been right that our species is not the same as 'Collybia' acervata EU, as other people have since noticed differences in our collections from EU and ENA collections, and our ITS DNA differs by more than 3%. In Joanne's honour, I would like to call our species Connopus bagleyensis.
Connopus bagleyensis n.p. © Steve Trudell
Pseudomarasmius pallidocephalus MI - a small marasmioid with a dark stem, easily confused with the small Collybiopsis species above. The stem is not quite as wiry or "horsehair" like as Gymnopus androsaceus, also above, which it is paler than when fresh.
probable Pseudomarasmius pallidocephalus © Andrew Parker
'Gymnopus' striatipes NY (=Collybia cylindrospora OR) - a clustered tan capped species with a grooved stem and gill edges that are not serrated. This may need its own genus. It's possible that a multi-gene study could show it belongs in Rhodocollybia, although it would be one of the only Rhodocollybia without serrated gill edges.
'Gymnopus' striatipes © Jairul Rahaman
Rhodocollybia butyracea EU - dark brown cap, stem not rooting like other species but the gill edges are serrated. Our local DNA matches EU DNA quite well.
Rhodocollybia asema EU - similar, with a paler cap. This was formerly a form or variety of R. butyracea, but the DNA is too different so it was recently elevated to species. There is geographical isolation in this species: EU, ENA and WNA sequences are <1% different from each other in ITS, but for now I'm treating them all as the same species, even though here in the west, ours supposedly does not dry with a very dark disk like EU collections.
Rhodocollybia badiialba WA - with a medium brown cap, but the colour is variable enough that it is easily mistaken for both R. butyracea and R. asema. It is best differentiated by its roundish spores and pink FeSO4 reaction of the gills.
Rhodocollybia butyracea © Buck McAdoo, R. asema © Buck McAdoo, R. badiialba © Buck McAdoo
Rhodocollybia 'olivaceogrisea PNW06' - olive grey cap, serrated gill edges, stems do not appear to root. Our one collection appears to be stocky with an umbo. It has a sequence 10 bp and 1 indel different from official collections of the Costa Rica species R. olivaceogrisea was recently found in WA. The Costa Rica species was found under oak and ours is clearly a conifer species based on the photo. It perhaps needs its own name.
Rhodocollybia 'olivaceogrisea PNW06' © Buck McAdoo
Rhodocollybia maculata group - all varieties have serrated gills and somewhat rooting stems, and will have one or both of the following two characters: odor sweet like almond or cherry and red staining flesh wherever handled. Six varieties have been reported from the PNW:
var. maculata EU - pale cap, no odor, staining red. The only
variety with roundish spores.
Rhodocollybia oregonensis OR - very similar with a two toned cap with a vinaceous brown disk and paler rim. Sweet odor and red staining. I don't know why this variety got its own name.
So far, we have found 5 genetic species in the PNW, and I'm trying to match them up with the above 7 varieties. They all probably deserve their own species name as the genetics are quite different for each. They do all form a clade together with Rhodocollybia oregonensis, so conceivably R. oregonensis could be reduced to a variety of R. maculata instead.
PNW01 - var. scorzonerea? This is easy to recognize when bright yellow, but the yellow may be very subtle, resembling var. occidentalis but with a more uniform cap colour. When not very yellow, it also resembles var. maculata, but that has round spores, and var. immutabilis, but that doesn't stain red. PNW01 has the sweet odor plus the red staining. I do not think this is the real var. scorzonerea as this DNA is not found in Europe yet. So this may need a new name, and the real var. scorzonerea will have to be demystified and its presence checked for in the PNW.
PNW02 - var. fulva? The odor and staining and state of the gill edges were not noted, so this is just a guess. We need collections with a dark vinaceous brown cap, no odor, and with red staining. If this isn't var. fulva, then the locally described var. fulva is still unsequenced.
PNW03 - var. occidentalis may have a white rim and slightly coloured disc, or an entirely pigmented cap but the colour will be not uniform, but somewhat splotchy. PNW01 and PNW05 have a more uniformly coloured cap. I feel fairly confident that this really is var. occidentalis.
PNW04 - Rhodocollybia oregonensis (two toned wine coloured disc with paler brown rim). Sweet odor at least when fresh. It stained red. I feel fairly confident that this really is R. oregonensis.
PNW05 - var. maculata? This has the round spores that only var. maculata has, plus it is a macroscopic match as well, with a pale cap and red staining but with no sweet odor. However, it doesn't match EU sequences so our collections probably need a new name.
No collections have been sequenced yet purporting to be var. immutabilis nor var. nigra. We still need collections of those.
Keep reading for one more lookalike.
Rhodocollybia maculata var. 'scorzonerea PNW01' © Danny Miller and Yi-Min Wang, PNW02 (cf. var. fulva) © NAMA and the Field Museum of Natural History, var. occidentalis PNW03 © Buck McAdoo
R. oregonensis PNW04 © Yi-Min Wang and Alan Rockefeller, PNW05 (cf. var. maculata) © Buck McAdoo
'Rhodocollybia' subsulcatipes WA - described with even gill edges (not serrated) and no staining reaction, but our photos show serrated gill edges and slight staining (near the top of the stem on the right hand fruiting body). Round spores. It was also described with a vinaceous tan cap, rooting stem, and sweet odor. It looks like it belongs to the Rhodocollybia maculata/oregonensis group of species (in the sense those either stain red or smell sweet). A modern collection, and our only photographs, show a stocky, Phaeocollybia-like mushroom with a pinkish-tan conical cap, long stem, and round spores that were coloured like Rhodocollybia. I was able to identify it because Joanne Lennox, who transferred the 1944 species Collybia subsulcatipes to Rhodocollybia in 1979, saved one of her contemporary collections which we sequenced and it matches this pictured modern collection. It shows up in a clade by itself in my ITS tree near Connopus, and probably needs its own genus.
'Rhodocollybia' subsulcatipes © Sienna McDonald
Rhodocollybia cf unakensis TN/Rhodocollybia cf extuberans EU - more species with a two-toned wine coloured disc with a paler brown rim, but these supposedly do not smell sweet nor stain red. Both of these species have been reported from the PNW, but some think we only have one species here that has gone by both names. I have a couple of east coast sequences that might be R. unakensis, but not with any confidence. I also have sequences and a collection of Gymnopus ocior (see above in the Gymnopus section), which is thought to be the correct name for Rhodocollybia extuberans. We need local collections of both to investigate this.
Rhodocollybia PNW07 - a 100 year old collection from ID sequenced inside Rhodocollybia. It was labeled Gymnopus putillus, a species with a wine-red cap and gills contrasting with a very pale entirely pubescent stem. That species is not reported to have serrated gill edges nor a rooting stem like Rhodocollybia should.
Omphalotus and Lentinula
Not typically found in the PNW, but here for completion.
Omphalotus olivascens CA - the glow in the dark poisonous jack-o-lantern, perhaps found in southern OR.
Lentinula aff. edodes - shiitake, often cultivated here and might escape. Worldwide sequences vary by up to 6%, so I don't know if it's a complex of species or not.
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